Lincoln University

Biological invasions cause ecological and economic impacts across the globe. However, it is unclear whether there are strong patterns in terms of their major effects, how the vulnerability of different ecosystems varies and which ecosystem services are at greatest risk. We present a global meta-analysis of 199 articles reporting 1041 field studies that in total describe the impacts of 135 alien plant taxa on resident species, communities and ecosystems. Across studies, alien plants had a significant effect in 11 of 24 different types of impact assessed. The magnitude and direction of the impact varied both within and between different types of impact. On average, abundance and diversity of the resident species decreased in invaded sites, whereas primary production and several ecosystem processes were enhanced. While alien N-fixing species had greater impacts on N-cycling variables, they did not consistently affect other impact types. The magnitude of the impacts was not significantly different between island and mainland ecosystems. Overall, alien species impacts are heterogeneous and not unidirectional even within particular impact types. Our analysis also reveals that by the time changes in nutrient cycling are detected, major impacts on plant species and communities are likely to have already occurred.

Many agroecosystems are unfavorable environments for natural enemies due to high levels of disturbance. Habitat management, a form of conservation biological control, is an ecologically based approach aimed at favoring natural enemies and enhancing biological control in agricultural systems. The goal of habitat management is to create a suitable ecological infrastructure within the agricultural landscape to provide resources such as food for adult natural enemies, alternative prey or hosts, and shelter from adverse conditions. These resources must be integrated into the landscape in a way that is spatially and temporally favorable to natural enemies and practical for producers to implement. The rapidly expanding literature on habitat management is reviewed with attention to practices for favoring predators and parasitoids, implementation of habitat management, and the contributions of modeling and ecological theory to this developing area of conservation biological control. The potential to integrate the goals of habitat management for natural enemies and nature conservation is discussed.

Summary Humans have traded and transported alien species for millennia with two notable step‐changes: the end of the Middle Ages and beginning of the Industrial Revolution. However, in recent decades the world has entered a new phase in the magnitude and diversity of biological invasions: the Era of Globalization. This Special Profile reviews the links between the main drivers of globalization and biological invasions and examines state‐of‐the‐art approaches to pathway risk assessment to illustrate new opportunities for managing invasive species. Income growth is a primary driver of globalization and a clear association exists between Gross Domestic Product and the richness of alien floras and faunas for many regions of the world. In many cases, the exposure of these economies to trade is highlighted by the significant role of merchandise imports in biological invasions, especially for island ecosystems. Post‐1950, technical and logistic improvements have accelerated the ease with which commodities are transported across the globe and hindered the traceability of goods and the ease of intercepting pests. New sea, land and air links in international trade and human transport have established novel pathways for the spread of alien species. Increasingly, the science advances underpinning invasive species management must move at the speed of commerce. Increasing transport networks and demand for commodities have led to pathway risk assessments becoming the frontline in the prevention of biological invasions. The diverse routes of introduction arising from contaminant, stowaway, corridor and unaided pathways, in both aquatic and terrestrial biomes are complex. Nevertheless, common features enable comparable approaches to risk assessment. By bringing together spatial data on climate suitability, habitat availability and points of entry, as well a demographic models that include species dispersal (both natural and human‐mediated) and measures of propagule pressure, it is possible to generate risk maps highlighting potential invasion hotspots that can inform prevention strategies. Synthesis and applications . To date, most attempts to model pathways have focused on describing the likelihood of invader establishment. Few have modelled explicit management strategies such as optimal detection and inspection strategies and assessments of the effectiveness of different management measures. A future focus in these areas will ensure research informs response.

Although research on human-mediated exchanges of species has substantially intensified during the last centuries, we know surprisingly little about temporal dynamics of alien species accumulations across regions and taxa. Using a novel database of 45,813 first records of 16,926 established alien species, we show that the annual rate of first records worldwide has increased during the last 200 years, with 37% of all first records reported most recently (1970-2014). Inter-continental and inter-taxonomic variation can be largely attributed to the diaspora of European settlers in the nineteenth century and to the acceleration in trade in the twentieth century. For all taxonomic groups, the increase in numbers of alien species does not show any sign of saturation and most taxa even show increases in the rate of first records over time. This highlights that past efforts to mitigate invasions have not been effective enough to keep up with increasing globalization.

Biological invasions are a global consequence of an increasingly connected world and the rise in human population size. The numbers of invasive alien species - the subset of alien species that spread widely in areas where they are not native, affecting the environment or human livelihoods - are increasing. Synergies with other global changes are exacerbating current invasions and facilitating new ones, thereby escalating the extent and impacts of invaders. Invasions have complex and often immense long-term direct and indirect impacts. In many cases, such impacts become apparent or problematic only when invaders are well established and have large ranges. Invasive alien species break down biogeographic realms, affect native species richness and abundance, increase the risk of native species extinction, affect the genetic composition of native populations, change native animal behaviour, alter phylogenetic diversity across communities, and modify trophic networks. Many invasive alien species also change ecosystem functioning and the delivery of ecosystem services by altering nutrient and contaminant cycling, hydrology, habitat structure, and disturbance regimes. These biodiversity and ecosystem impacts are accelerating and will increase further in the future. Scientific evidence has identified policy strategies to reduce future invasions, but these strategies are often insufficiently implemented. For some nations, notably Australia and New Zealand, biosecurity has become a national priority. There have been long-term successes, such as eradication of rats and cats on increasingly large islands and biological control of weeds across continental areas. However, in many countries, invasions receive little attention. Improved international cooperation is crucial to reduce the impacts of invasive alien species on biodiversity, ecosystem services, and human livelihoods. Countries can strengthen their biosecurity regulations to implement and enforce more effective management strategies that should also address other global changes that interact with invasions.

Bemisia tabaci has long been considered a complex species. It rose to global prominence in the 1980s owing to the global invasion by the commonly named B biotype. Since then, the concomitant eruption of a group of plant viruses known as begomoviruses has created considerable management problems in many countries. However, an enduring set of questions remains: Is B. tabaci a complex species or a species complex, what are Bemisia biotypes, and how did all the genetic variability arise? This review considers these issues and concludes that there is now sufficient evidence to state that B. tabaci is not made up of biotypes and that the use of biotype in this context is erroneous and misleading. Instead, B. tabaci is a complex of 11 well-defined high-level groups containing at least 24 morphologically indistinguishable species.

ABSTRACT In the literature, the terms species richness and species diversity are sometimes used interchangeably. We suggest that at the very least, authors should define what they mean by either term. Of the many species diversity indices used in the literature, the Shannon Index is perhaps most commonly used. On some occasions it is called the Shannon–Wiener Index and on other occasions it is called the Shannon–Weaver Index. We suggest an explanation for this dual use of terms and in so doing we offer a tribute to the late Claude Shannon (who passed away on 24 February 2001).

Abstract With the growing body of literature assessing the impact of invasive alien plants on resident species and ecosystems, a comprehensive assessment of the relationship between invasive species traits and environmental settings of invasion on the characteristics of impacts is needed. Based on 287 publications with 1551 individual cases that addressed the impact of 167 invasive plant species belonging to 49 families, we present the first global overview of frequencies of significant and non‐significant ecological impacts and their directions on 15 outcomes related to the responses of resident populations, species, communities and ecosystems. Species and community outcomes tend to decline following invasions, especially those for plants, but the abundance and richness of the soil biota, as well as concentrations of soil nutrients and water, more often increase than decrease following invasion. Data mining tools revealed that invasive plants exert consistent significant impacts on some outcomes (survival of resident biota, activity of resident animals, resident community productivity, mineral and nutrient content in plant tissues, and fire frequency and intensity), whereas for outcomes at the community level, such as species richness, diversity and soil resources, the significance of impacts is determined by interactions between species traits and the biome invaded. The latter outcomes are most likely to be impacted by annual grasses, and by wind pollinated trees invading mediterranean or tropical biomes. One of the clearest signals in this analysis is that invasive plants are far more likely to cause significant impacts on resident plant and animal richness on islands rather than mainland. This study shows that there is no universal measure of impact and the pattern observed depends on the ecological measure examined. Although impact is strongly context dependent, some species traits, especially life form, stature and pollination syndrome, may provide a means to predict impact, regardless of the particular habitat and geographical region invaded.

Summary Fine roots acquire essential soil resources and mediate biogeochemical cycling in terrestrial ecosystems. Estimates of carbon and nutrient allocation to build and maintain these structures remain uncertain because of the challenges of consistently measuring and interpreting fine‐root systems. Traditionally, fine roots have been defined as all roots ≤ 2 mm in diameter, yet it is now recognized that this approach fails to capture the diversity of form and function observed among fine‐root orders. Here, we demonstrate how order‐based and functional classification frameworks improve our understanding of dynamic root processes in ecosystems dominated by perennial plants. In these frameworks, fine roots are either separated into individual root orders or functionally defined into a shorter‐lived absorptive pool and a longer‐lived transport fine‐root pool. Using these frameworks, we estimate that fine‐root production and turnover represent 22% of terrestrial net primary production globally – a c . 30% reduction from previous estimates assuming a single fine‐root pool. Future work developing tools to rapidly differentiate functional fine‐root classes, explicit incorporation of mycorrhizal fungi into fine‐root studies, and wider adoption of a two‐pool approach to model fine roots provide opportunities to better understand below‐ground processes in the terrestrial biosphere. Contents Summary 505 I. Introduction 506 II. Ordered variation in fine‐root traits – different functions of different roots 506 III. Pitfalls and platforms – understanding bias and improving estimates of root processes 508 IV. Moving forward using root orders and functional classifications 512 V. Can we integrate mycorrhizal fungi with root classifications? 513 VI. Conclusions and recommendations 514 Acknowledgements 515 References 515

Abstract Losses of nitrogen from the soil/plant system not only reduce soil fertility and plant yield but can also create adverse impacts on the environment. Ammonia emissions into the atmosphere contribute to acid rain and represent an indirect source of nitrous oxide greenhouse gas emissions. Nitrate leaching losses into rivers and lakes can cause eutrophication resulting in excessive growth of aquatic weeds and algae, which can reduce fish populations and the recreational value of the water. Nitrate contamination of drinking water supplies can cause health risks. Legislation that is designed to limit nitrate leaching losses from land has become a constraint on agricultural land use in many countries. Nitrous oxide emissions into the atmosphere contribute to the depletion of the ozone layer and also make a significant contribution to climate change. This review describes the nitrogen cycle in temperate soil/plant systems, the processes involved in each of the individual nitrogen loss pathways, the factors affecting the amounts of losses and the methods that are available to reduce these losses. The review has shown that careful management of temperate soil/plant systems using best management practices and newly developed technologies can increase the sustainability of agriculture and reduce its impact on the environment.

BACKGROUND As global climate change accelerates, one of the most urgent tasks for the coming decades is to develop accurate predictions about biological responses to guide the effective protection of biodiversity. Predictive models in biology provide a means for scientists to project changes to species and ecosystems in response to disturbances such as climate change. Most current predictive models, however, exclude important biological mechanisms such as demography, dispersal, evolution, and species interactions. These biological mechanisms have been shown to be important in mediating past and present responses to climate change. Thus, current modeling efforts do not provide sufficiently accurate predictions. Despite the many complexities involved, biologists are rapidly developing tools that include the key biological processes needed to improve predictive accuracy. The biggest obstacle to applying these more realistic models is that the data needed to inform them are almost always missing. We suggest ways to fill this growing gap between model sophistication and information to predict and prevent the most damaging aspects of climate change for life on Earth. ADVANCES On the basis of empirical and theoretical evidence, we identify six biological mechanisms that commonly shape responses to climate change yet are too often missing from current predictive models: physiology; demography, life history, and phenology; species interactions; evolutionary potential and population differentiation; dispersal, colonization, and range dynamics; and responses to environmental variation. We prioritize the types of information needed to inform each of these mechanisms and suggest proxies for data that are missing or difficult to collect. We show that even for well-studied species, we often lack critical information that would be necessary to apply more realistic, mechanistic models. Consequently, data limitations likely override the potential gains in accuracy of more realistic models. Given the enormous challenge of collecting this detailed information on millions of species around the world, we highlight practical methods that promote the greatest gains in predictive accuracy. Trait-based approaches leverage sparse data to make more general inferences about unstudied species. Targeting species with high climate sensitivity and disproportionate ecological impact can yield important insights about future ecosystem change. Adaptive modeling schemes provide a means to target the most important data while simultaneously improving predictive accuracy. OUTLOOK Strategic collections of essential biological information will allow us to build generalizable insights that inform our broader ability to anticipate species’ responses to climate change and other human-caused disturbances. By increasing accuracy and making uncertainties explicit, scientists can deliver improved projections for biodiversity under climate change together with characterizations of uncertainty to support more informed decisions by policymakers and land managers. Toward this end, a globally coordinated effort to fill data gaps in advance of the growing climate-fueled biodiversity crisis offers substantial advantages in efficiency, coverage, and accuracy. Biologists can take advantage of the lessons learned from the Intergovernmental Panel on Climate Change’s development, coordination, and integration of climate change projections. Climate and weather projections were greatly improved by incorporating important mechanisms and testing predictions against global weather station data. Biology can do the same. We need to adopt this meteorological approach to predicting biological responses to climate change to enhance our ability to mitigate future changes to global biodiversity and the services it provides to humans. Emerging models are beginning to incorporate six key biological mechanisms that can improve predictions of biological responses to climate change. Models that include biological mechanisms have been used to project (clockwise from top) the evolution of disease-harboring mosquitoes, future environments and land use, physiological responses of invasive species such as cane toads, demographic responses of penguins to future climates, climate-dependent dispersal behavior in butterflies, and mismatched interactions between butterflies and their host plants. Despite these modeling advances, we seldom have the detailed data needed to build these models, necessitating new efforts to collect the relevant data to parameterize more biologically realistic predictive models.

Treelines are temperature sensitive transition zones that are expected to respond to climate warming by advancing beyond their current position. Response to climate warming over the last century, however, has been mixed, with some treelines showing evidence of recruitment at higher altitudes and/or latitudes (advance) whereas others reveal no marked change in the upper limit of tree establishment. To explore this variation, we analysed a global dataset of 166 sites for which treeline dynamics had been recorded since 1900 AD. Advance was recorded at 52% of sites with only 1% reporting treeline recession. Treelines that experienced strong winter warming were more likely to have advanced, and treelines with a diffuse form were more likely to have advanced than those with an abrupt or krummholz form. Diffuse treelines may be more responsive to warming because they are more strongly growth limited, whereas other treeline forms may be subject to additional constraints.

World production of wine has steadily risen over recent years and consumption has not kept pace with this increase, thus many countries have surpluses of wine which pose problems in international trade. Despite these problems, there is not, generally, a surplus of high quality wines. Quality is not easy to define, but ideally, it should be related to intrinsic visual, taste, or aroma characters which are perceived as above average for that type of wine. Usually this is reflected in the price paid for that wine — although price is not necessarily a reliable indicator since it can be influenced also by fashion, tradition, availability and personal preferences. Unfortunately, despite the many references to quality and the amount of work which directly or indirectly refers to it, there is still confusion over what contribution climates, sites, and viticultural practices really make. This paper is a review of the effects of these environmental and management practices which may change grape composition and wine quality. Its scope is limited to table wines rather than fortified wines.

Recent comprehensive data provided through the DAISIE project ( www.europe‐aliens.org ) have facilitated the development of the first pan‐European assessment of the impacts of alien plants, vertebrates, and invertebrates – in terrestrial, freshwater, and marine environments – on ecosystem services. There are 1094 species with documented ecological impacts and 1347 with economic impacts. The two taxonomic groups with the most species causing impacts are terrestrial invertebrates and terrestrial plants. The North Sea is the maritime region that suffers the most impacts. Across taxa and regions, ecological and economic impacts are highly correlated. Terrestrial invertebrates create greater economic impacts than ecological impacts, while the reverse is true for terrestrial plants. Alien species from all taxonomic groups affect “supporting”, “provisioning”, “regulating”, and “cultural” services and interfere with human well‐being. Terrestrial vertebrates are responsible for the greatest range of impacts, and these are widely distributed across Europe. Here, we present a review of the financial costs, as the first step toward calculating an estimate of the economic consequences of alien species in Europe.

Summary Pathways describe the processes that result in the introduction of alien species from one location to another. A framework is proposed to facilitate the comparative analysis of invasion pathways by a wide range of taxa in both terrestrial and aquatic ecosystems. Comparisons with a range of data helped identify existing gaps in current knowledge of pathways and highlight the limitations of existing legislation to manage introductions of alien species. The scheme aims for universality but uses the European Union as a case study for the regulatory perspectives. Alien species may arrive and enter a new region through three broad mechanisms: importation of a commodity, arrival of a transport vector, and/or natural spread from a neighbouring region where the species is itself alien. These three mechanisms result in six principal pathways: release, escape, contaminant, stowaway, corridor and unaided. Alien species transported as commodities may be introduced as a deliberate release or as an escape from captivity. Many species are not intentionally transported but arrive as a contaminant of a commodity, for example pathogens and pests. Stowaways are directly associated with human transport but arrive independently of a specific commodity, for example organisms transported in ballast water, cargo and airfreight. The corridor pathway highlights the role transport infrastructures play in the introduction of alien species. The unaided pathway describes situations where natural spread results in alien species arriving into a new region from a donor region where it is also alien. Vertebrate pathways tend to be characterized as deliberate releases, invertebrates as contaminants and plants as escapes. Pathogenic micro‐organisms and fungi are generally introduced as contaminants of their hosts. The corridor and unaided pathways are often ignored in pathway assessments but warrant further detailed consideration. Synthesis and applications. Intentional releases and escapes should be straightforward to monitor and regulate but, in practice, developing legislation has proved difficult. New introductions continue to occur through contaminant, stowaway, corridor and unaided pathways. These pathways represent special challenges for management and legislation. The present framework should enable these trends to be monitored more clearly and hopefully lead to the development of appropriate regulations or codes of practice to stem the number of future introductions.

In this article the authors focus on the use of a modal preferences questionnaire as a catalyst to empower students to reflect on their own sensory preferences and modify their study methods accordingly. The authors discuss the development and use of the questionnaire, strategies for students to use in modifying their learning behavior, responses of students and faculty to the technique, and directions for further investigation of modal preferences. Over the last four decades the literature from both psychology and educa-tion has supported the proposition that learners of all ages have different yet consistent ways of responding in learning situations. These behaviors or predispositions to behave in a particular fashion have been termed learning

The abundance and composition of soil ammonia-oxidizing bacteria (AOB) and ammonia-oxidizing archaea (AOA) were investigated by using quantitative real-time polymerase chain reaction, cloning and sequencing approaches based on amoA genes. The soil, classified as agri-udic ferrosols with pH (H(2)O) ranging from 3.7 to 6.0, was sampled in summer and winter from long-term field experimental plots which had received 16 years continuous fertilization treatments, including fallow (CK0), control without fertilizers (CK) and those with combinations of fertilizer nitrogen (N), phosphorus (P) and potassium (K): N, NP, NK, PK, NPK and NPK plus organic manure (OM). Population sizes of AOB and AOA changed greatly in response to the different fertilization treatments. The NPK + OM treatment had the highest copy numbers of AOB and AOA amoA genes among the treatments that received mineral fertilizers, whereas the lowest copy numbers were recorded in the N treatment. Ammonia-oxidizing archaea were more abundant than AOB in all the corresponding treatments, with AOA to AOB ratios ranging from 1.02 to 12.36. Significant positive correlations were observed among the population sizes of AOB and AOA, soil pH and potential nitrification rates, indicating that both AOB and AOA played an important role in ammonia oxidation in the soil. Phylogenetic analyses of the amoA gene fragments showed that all AOB sequences from different treatments were affiliated with Nitrosospira or Nitrosospira-like species and grouped into cluster 3, and little difference in AOB community composition was recorded among different treatments. All AOA sequences fell within cluster S (soil origin) and cluster M (marine and sediment origin). Cluster M dominated exclusively in the N, NP, NK and PK treatments, indicating a pronounced difference in the community composition of AOA in response to the long-term fertilization treatments. These findings could be fundamental to improve our understanding of the importance of both AOB and AOA in the cycling of nitrogen and other nutrients in terrestrial ecosystems.

Human land use threatens global biodiversity and compromises multiple ecosystem functions critical to food production. Whether crop yield-related ecosystem services can be maintained by a few dominant species or rely on high richness remains unclear. Using a global database from 89 studies (with 1475 locations), we partition the relative importance of species richness, abundance, and dominance for pollination; biological pest control; and final yields in the context of ongoing land-use change. Pollinator and enemy richness directly supported ecosystem services in addition to and independent of abundance and dominance. Up to 50% of the negative effects of landscape simplification on ecosystem services was due to richness losses of service-providing organisms, with negative consequences for crop yields. Maintaining the biodiversity of ecosystem service providers is therefore vital to sustain the flow of key agroecosystem benefits to society.

Interest in biochar stems from its potential agronomic benefits and carbon sequestration ability. Biochar application alters soil nitrogen (N) dynamics. This review establishes emerging trends and gaps in biochar-N research. Biochar adsorption of NO3−, up to 0.6 mg g−1 biochar, occurs at pyrolysis temperatures >600 °C with amounts adsorbed dependent on feedstock and NO3− concentration. Biochar NH4+ adsorption depends on feedstock, but no pyrolysis temperature trend is apparent. Long-term practical effectiveness of inorganic-N adsorption, as a NO3− leaching mitigation option, requires further study. Biochar adsorption of ammonia (NH3) decreases NH3 and NO3− losses during composting and after manure applications, and offers a mechanism for developing slow release fertilisers. Reductions in NH3 loss vary with N source and biochar characteristics. Manure derived biochars have a role as N fertilizers. Increasing pyrolysis temperatures, during biochar manufacture from manures and biosolids, results in biochars with decreasing hydrolysable organic N and increasing aromatic and heterocyclic structures. The short- and long-term implications of biochar on N immobilisation and mineralization are specific to individual soil-biochar combinations and further systematic studies are required to predict agronomic and N cycling responses. Most nitrous oxide (N2O) studies measuring nitrous oxide (N2O) were short-term in nature and found emission reductions, but long-term studies are lacking, as is mechanistic understanding of reductions. Stable N isotopes have a role in elucidating biochar-N-soil dynamics. There remains a dearth of information regarding effects of biochar and soil biota on N cycling. Biochar has potential within agroecosystems to be an N input, and a mitigation agent for environmentally detrimental N losses. Future research needs to systematically understand biochar-N interactions over the long term.

Species moved by human activities beyond the limits of their native geographic ranges into areas in which they do not naturally occur (termed aliens) can cause a broad range of significant changes to recipient ecosystems; however, their impacts vary greatly across species and the ecosystems into which they are introduced. There is therefore a critical need for a standardised method to evaluate, compare, and eventually predict the magnitudes of these different impacts. Here, we propose a straightforward system for classifying alien species according to the magnitude of their environmental impacts, based on the mechanisms of impact used to code species in the International Union for Conservation of Nature (IUCN) Global Invasive Species Database, which are presented here for the first time. The classification system uses five semi-quantitative scenarios describing impacts under each mechanism to assign species to different levels of impact-ranging from Minimal to Massive-with assignment corresponding to the highest level of deleterious impact associated with any of the mechanisms. The scheme also includes categories for species that are Not Evaluated, have No Alien Population, or are Data Deficient, and a method for assigning uncertainty to all the classifications. We show how this classification system is applicable at different levels of ecological complexity and different spatial and temporal scales, and embraces existing impact metrics. In fact, the scheme is analogous to the already widely adopted and accepted Red List approach to categorising extinction risk, and so could conceivably be readily integrated with existing practices and policies in many regions.