Using high-resolution LiDAR-derived canopy structure and topography to characterise hibernaculum locations of the hazel dormouse.

The hazel dormouse is predominantly an arboreal species that moves down to the ground to hibernate in the autumn in temperate parts of its distributional ranges at locations not yet well understood. The main objective of this study is to test whether environmental characteristics surrounding hazel dormouse hibernacula can be identified using high-resolution remote sensing and data collected in situ. To achieve this, remotely sensed variables, including canopy height and cover, topographic slope, sky view, solar radiation and cold air drainage, were modelled around 83 dormouse hibernacula in England (n = 62) and the Netherlands (n = 21), and environmental characteristics that may be favoured by pre-hibernating dormice were identified. Data on leaf litter depth, temperature, canopy cover and distance to the nearest tree were collected in situ and analysed at hibernaculum locations in England. The findings indicated that remotely sensed data were effective in identifying attributes surrounding the locations of dormouse hibernacula and when compared to in situ information, provided more conclusive results. This study suggests that remotely sensed topographic slope, canopy height and sky view have an influence on hazel dormice choosing suitable locations to hibernate; whilst in situ data suggested that average daily mean temperature at the hibernaculum may also have an effect. Remote sensing proved capable of identifying localised environmental characteristics in the wider landscape that may be important for hibernating dormice. This study proposes that this method can provide a novel progression from habitat modelling to conservation management for the hazel dormouse, as well as other species using habitats where topography and vegetation structure influence fine-resolution favourability.

Directional turnover towards larger-ranged plants over time and across habitats.

Species turnover is ubiquitous. However, it remains unknown whether certain types of species are consistently gained or lost across different habitats. Here, we analysed the trajectories of 1827 plant species over time intervals of up to 78 years at 141 sites across mountain summits, forests, and lowland grasslands in Europe. We found, albeit with relatively small effect sizes, displacements of smaller- by larger-ranged species across habitats. Communities shifted in parallel towards more nutrient-demanding species, with species from nutrient-rich habitats having larger ranges. Because these species are typically strong competitors, declines of smaller-ranged species could reflect not only abiotic drivers of global change, but also biotic pressure from increased competition. The ubiquitous component of turnover based on species range size we found here may partially reconcile findings of no net loss in local diversity with global species loss, and link community-scale turnover to macroecological processes such as biotic homogenisation.

The problem of scale in predicting biological responses to climate.

Many analyses of biological responses to climate rely on gridded climate data derived from weather stations, which differ from the conditions experienced by organisms in at least two respects. First, the microclimate recorded by a weather station is often quite different to that near the ground surface, where many organisms live. Second, the temporal and spatial resolutions of gridded climate datasets derived from weather stations are often too coarse to capture the conditions experienced by organisms. Temporally and spatially coarse data have clear benefits in terms of reduced model size and complexity, but here we argue that coarse-grained data introduce errors that, in biological studies, are too often ignored. However, in contrast to common perception, these errors are not necessarily caused directly by a spatial mismatch between the size of organisms and the scale at which climate data are collected. Rather, errors and biases are primarily due to (a) systematic discrepancies between the climate used in analysis and that experienced by organisms under study; and (b) the non-linearity of most biological responses in combination with differences in climate variance between locations and time periods for which models are fitted and those for which projections are made. We discuss when exactly problems of scale can be expected to arise and highlight the potential to circumvent these by spatially and temporally down-scaling climate. We also suggest ways in which adjustments to deal with issues of scale could be made without the need to run high-resolution models over wide extents.

Quantifying resistance and resilience to local extinction for conservation prioritization.

Species-focused conservation planning is often based on reducing local extinction risk at key sites. However, with increasing levels of habitat fragmentation and pressures from climate change and overexploitation, surrounding landscapes also influence the persistence of species populations, and their effects are increasingly incorporated in conservation planning and management for both species and communities. Here, we present a framework based on metapopulation dynamics in fragmented landscapes, for quantifying the survival (resistance) and reestablishment of species populations following localized extinction events (resilience). We explore the application of this framework to guide the conservation of a group of threatened bird species endemic to papyrus (Cyperus papyrus) swamps in East and Central Africa. Using occupancy data for five species collected over two years from a network of wetlands in Uganda, we determine the local and landscape factors that influence local extinction and colonization, and map expected rates of population turnover across the network to draw inferences about the locations that contribute most to regional resistance and resilience for all species combined. Slight variation in the factors driving extinction and colonization between individual papyrus birds led to species-specific differences in the spatial patterns of site-level resistance and resilience. However, despite this, locations with the highest resistance and/or resilience overlapped for most species and reveal where resources could be invested for multispecies persistence. This novel simplified framework can aid decision making associated with conservation planning and prioritization for multiple species residing in overlapping, fragmented habitats; helping to identify key sites that warrant urgent conservation protection, with consideration of the need to adapt and respond to future change.

Multiple night-time light-emitting diode lighting strategies impact grassland invertebrate assemblages.

White light-emitting diodes (LEDs) are rapidly replacing conventional outdoor lighting technologies around the world. Despite rising concerns over their impact on the environment and human health, the flexibility of LEDs has been advocated as a means of mitigating the ecological impacts of globally widespread outdoor night-time lighting through spectral manipulation, dimming and switching lights off during periods of low demand. We conducted a three-year field experiment in which each of these lighting strategies was simulated in a previously artificial light naïve grassland ecosystem. White LEDs both increased the total abundance and changed the assemblage composition of adult spiders and beetles. Dimming LEDs by 50% or manipulating their spectra to reduce ecologically damaging wavelengths partially reduced the number of commoner species affected from seven to four. A combination of dimming by 50% and switching lights off between midnight and 04:00 am showed the most promise for reducing the ecological costs of LEDs, but the abundances of two otherwise common species were still affected. The environmental consequences of using alternative lighting technologies are increasingly well established. These results suggest that while management strategies using LEDs can be an effective means of reducing the number of taxa affected, averting the ecological impacts of night-time lighting may ultimately require avoiding its use altogether.

Fine-scale climate change: modelling spatial variation in biologically meaningful rates of warming.

The existence of fine-grain climate heterogeneity has prompted suggestions that species may be able to survive future climate change in pockets of suitable microclimate, termed 'microrefugia'. However, evidence for microrefugia is hindered by lack of understanding of how rates of warming vary across a landscape. Here, we present a model that is applied to provide fine-grained, multidecadal estimates of temperature change based on the underlying physical processes that influence microclimate. Weather station and remotely derived environmental data were used to construct physical variables that capture the effects of terrain, sea surface temperatures, altitude and surface albedo on local temperatures, which were then calibrated statistically to derive gridded estimates of temperature. We apply the model to the Lizard Peninsula, United Kingdom, to provide accurate (mean error = 1.21 °C; RMS error = 1.63 °C) hourly estimates of temperature at a resolution of 100 m for the period 1977-2014. We show that rates of warming vary across a landscape primarily due to long-term trends in weather conditions. Total warming varied from 0.87 to 1.16 °C, with the slowest rates of warming evident on north-east-facing slopes. This variation contributed to substantial spatial heterogeneity in trends in bioclimatic variables: for example, the change in the length of the frost-free season varied from +11 to -54 days and the increase in annual growing degree-days from 51 to 267 °C days. Spatial variation in warming was caused primarily by a decrease in daytime cloud cover with a resulting increase in received solar radiation, and secondarily by a decrease in the strength of westerly winds, which has amplified the effects on temperature of solar radiation on west-facing slopes. We emphasize the importance of multidecadal trends in weather conditions in determining spatial variation in rates of warming, suggesting that locations experiencing least warming may not remain consistent under future climate change.

Biogeography of time partitioning in mammals.

Many animals regulate their activity over a 24-h sleep-wake cycle, concentrating their peak periods of activity to coincide with the hours of daylight, darkness, or twilight, or using different periods of light and darkness in more complex ways. These behavioral differences, which are in themselves functional traits, are associated with suites of physiological and morphological adaptations with implications for the ecological roles of species. The biogeography of diel time partitioning is, however, poorly understood. Here, we document basic biogeographic patterns of time partitioning by mammals and ecologically relevant large-scale patterns of natural variation in "illuminated activity time" constrained by temperature, and we determine how well the first of these are predicted by the second. Although the majority of mammals are nocturnal, the distributions of diurnal and crepuscular species richness are strongly associated with the availability of biologically useful daylight and twilight, respectively. Cathemerality is associated with relatively long hours of daylight and twilight in the northern Holarctic region, whereas the proportion of nocturnal species is highest in arid regions and lowest at extreme high altitudes. Although thermal constraints on activity have been identified as key to the distributions of organisms, constraints due to functional adaptation to the light environment are less well studied. Global patterns in diversity are constrained by the availability of the temporal niche; disruption of these constraints by the spread of artificial lighting and anthropogenic climate change, and the potential effects on time partitioning, are likely to be critical influences on species' future distributions.

Light pollution is associated with earlier tree budburst across the United Kingdom.

The ecological impact of night-time lighting is of concern because of its well-demonstrated effects on animal behaviour. However, the potential of light pollution to change plant phenology and its corresponding knock-on effects on associated herbivores are less clear. Here, we test if artificial lighting can advance the timing of budburst in trees. We took a UK-wide 13 year dataset of spatially referenced budburst data from four deciduous tree species and matched it with both satellite imagery of night-time lighting and average spring temperature. We find that budburst occurs up to 7.5 days earlier in brighter areas, with the relationship being more pronounced for later-budding species. Excluding large urban areas from the analysis showed an even more pronounced advance of budburst, confirming that the urban 'heat-island' effect is not the sole cause of earlier urban budburst. Similarly, the advance in budburst across all sites is too large to be explained by increases in temperature alone. This dramatic advance of budburst illustrates the need for further experimental investigation into the impact of artificial night-time lighting on plant phenology and subsequent species interactions. As light pollution is a growing global phenomenon, the findings of this study are likely to be applicable to a wide range of species interactions across the world.

Quantifying the erosion of natural darkness in the global protected area system.

The nighttime light environment of much of the earth has been transformed by the introduction of electric lighting. This impact continues to spread with growth in the human population and extent of urbanization. This has profound consequences for organismal physiology and behavior and affects abundances and distributions of species, community structure, and likely ecosystem functions and processes. Protected areas play key roles in buffering biodiversity from a wide range of anthropogenic pressures. We used a calibration of a global satellite data set of nighttime lights to determine how well they are fulfilling this role with regard to artificial nighttime lighting. Globally, areas that are protected tend to be darker at night than those that are not, and, with the exception of Europe, recent regional declines in the proportion of the area that is protected and remains dark have been small. However, much of these effects result from the major contribution to overall protected area coverage by the small proportion of individual protected areas that are very large. Thus, in Europe and North America high proportions of individual protected areas (>17%) have exhibited high levels of nighttime lighting in all recent years, and in several regions (Europe, Asia, South and Central America) high proportions of protected areas (32-42%) have had recent significant increases in nighttime lighting. Limiting and reversing the erosion of nighttime darkness in protected areas will require routine consideration of nighttime conditions when designating and establishing new protected areas; establishment of appropriate buffer zones around protected areas where lighting is prohibited; and landscape level reductions in artificial nighttime lighting, which is being called for in general to reduce energy use and economic costs.

Cuantificación de la Erosión de la Oscuridad Natural en el Sistema Global de Áreas Protegidas Resumen El ambiente de luz nocturna de la mayor parte de la Tierra se ha transformado por la introducción de la luz eléctrica. Este impacto continúa esparciéndose con el crecimiento de la población humana y de la extensión de la urbanización. Esto tiene consecuencias profundas sobre la fisiología y el comportamiento de los organismos y afecta a la abundancia y la distribución de especies, a la estructura de la comunidad y probablemente a los procesos y funciones de los ecosistemas. Las áreas protegidas juegan un papel importante en el amortiguamiento de una amplia gama de presiones antropogénicas para la biodiversidad. Usamos una calibración de un conjunto de datos satelitales globales de luces nocturnas para determinar que tan bien desempeñan este papel con respecto a la iluminación nocturna artificial. En un nivel global, las áreas que están protegidas tienden a ser más oscuras en la noche que aquellas que no lo están, y además, con la excepción de Europa, las declinaciones regionales recientes en la proporción del área que está protegida y permanece oscura han sido menores. Sin embargo, muchos de estos efectos resultan de una mayor contribución a la cobertura total del área protegida por parte de la pequeña proporción de áreas protegidas individuales que son muy grandes. Por esto en Europa y América del Norte, una alta proporción de áreas protegidas individuales (>17%) ha exhibido niveles altos de iluminación nocturna en todos los años recientes y en varias regiones (Europa, Asia, América Central y América del Sur) proporciones altas de áreas protegidas (32-42%) han tenido incrementos significativos de iluminación nocturna. Limitar y revertir la erosión de la oscuridad nocturna en las áreas protegidas requerirá una consideración rutinaria de las condiciones nocturnas cuando se designen y establezcan nuevas áreas protegidas; un establecimiento de zonas de amortiguamiento apropiadas donde se prohíba la iluminación alrededor de las áreas protegidas; y una reducción de los niveles de paisaje en la iluminación nocturna artificial, la cual se pide en general para reducir el uso de energía y los costos económicos.

Human alteration of natural light cycles: causes and ecological consequences.

Artificial light at night is profoundly altering natural light cycles, particularly as perceived by many organisms, over extensive areas of the globe. This alteration comprises the introduction of light at night at places and times at which it has not previously occurred, and with different spectral signatures. Given the long geological periods for which light cycles have previously been consistent, this constitutes a novel environmental pressure, and one for which there is evidence for biological effects that span from molecular to community level. Here we provide a synthesis of understanding of the form and extent of this alteration, some of the key consequences for terrestrial and aquatic ecosystems, interactions and synergies with other anthropogenic pressures on the environment, major uncertainties, and future prospects and management options. This constitutes a compelling example of the need for a thoroughly interdisciplinary approach to understanding and managing the impact of one particular anthropogenic pressure. The former requires insights that span molecular biology to ecosystem ecology, and the latter contributions of biologists, policy makers and engineers.

Eyespot variation and field temperature in the Meadow Brown butterfly.

Since the classic work of E.B. Ford, explanations for eyespot variation in the Meadow Brown butterfly have focused on the role of genetic polymorphism. The potential role of thermal plasticity in this classic example of natural selection has therefore been overlooked. Here, we use large daily field collections of butterflies from three sites, over multiple years, to examine whether field temperature is correlated with eyespot variation, using the same presence/absence scoring as Ford. We show that higher developmental temperature in the field leads to the disappearance of the spots visible while the butterfly is at rest, explaining the historical observation that hindwing spotting declines across the season. Strikingly, females developing at 11°C have a median of six spots and those developing at 15°C only have three. In contrast, the large forewing eyespot is always present and scales with forewing length. Furthermore, in contrast to the smaller spots, the size of the large forewing spot is best explained by calendar date (days since 1st March) rather than the temperature at pupation. As this large forewing spot is involved in startling predators and/or sexual selection, its constant presence is therefore likely required for defence, whereas the disappearance of the smaller spots over the season may help with female crypsis. We model annual total spot variation with phenological data from the UK and derive predictions as to how spot patterns will continue to change, predicting that female spotting will decrease year on year as our climate warms.

Penetrance interactions of colour pattern loci in the African Monarch and their implications for the evolution of dominance.

Scoring the penetrance of heterozygotes in complex phenotypes, like colour pattern, is difficult and complicates the analysis of systems in which dominance is incomplete or evolving. The African Monarch (Danaus chrysippus) represents an example where colour pattern heterozygotes, formed in the contact zone between the different subspecies, show such intermediate dominance. Colour pattern in this aposematic butterfly is controlled by three loci A, B and C. The B and C loci are closely linked in a B/C supergene and significant interaction of B and C phenotypes is therefore expected via linkage alone. The A locus, however, is not linked to B/C and is found on a different chromosome. To study interactions between these loci we generated colour pattern heterozygotes by crossing males and females bearing different A and B/C genotypes, collected from different parts of Africa. We derived a novel scoring system for the expressivity of the heterozygotes and, as predicted, we found significant interactions between the genotypes of the closely linked B and C loci. Surprisingly, however, we also found highly significant interactions between C and the unlinked A locus, modifications that generally increased the resemblance of heterozygotes to homozygous ancestors. In contrast, we found no difference in the penetrance of any of the corresponding heterozygotes from crosses conducted either in allopatry or sympatry, in reciprocal crosses of males and females, or in the presence or absence of endosymbiont mediated male-killing or its associated neoW mediated sex-linkage of colour pattern. Together, this data supports the idea that the different colour morphs of the African Monarch meet transiently in the East African contact zone and that genetic modifiers act to mask inappropriate expression of colour patterns in the incorrect environments.

Changing streetlighting schemes and the ecological availability of darkness.

Artificial light at night (ALAN), from streetlights and other sources, has a wide variety of impacts on the natural environment. A significant challenge remains, however, to predict at intermediate spatial extents (e.g. across a city) the ALAN that organisms experience under different lighting regimes. Here we use Monte Carlo radiative Transfer to model the three-dimensional lighting environment at, and just above, ground level, on the spatial scales at which animals and humans experience it. We show how this technique can be used to model a suite of both real and hypothetical lighting environments, mimicking the transition of public infrastructure between different lighting technologies. We then demonstrate how the behaviour of animals experiencing these simulated lighting environments can be emulated to probe the availability of darkness, and dark corridors, within them. Our simulations show that no single lighting technology provides an unmitigated alleviation of negative impacts within urban environments, and that holistic treatments of entire lighting environments should be employed when understanding how animals use and traverse them.

Range expansion through fragmented landscapes under a variable climate.

Ecological responses to climate change may depend on complex patterns of variability in weather and local microclimate that overlay global increases in mean temperature. Here, we show that high-resolution temporal and spatial variability in temperature drives the dynamics of range expansion for an exemplar species, the butterfly Hesperia comma. Using fine-resolution (5 m) models of vegetation surface microclimate, we estimate the thermal suitability of 906 habitat patches at the species' range margin for 27 years. Population and metapopulation models that incorporate this dynamic microclimate surface improve predictions of observed annual changes to population density and patch occupancy dynamics during the species' range expansion from 1982 to 2009. Our findings reveal how fine-scale, short-term environmental variability drives rates and patterns of range expansion through spatially localised, intermittent episodes of expansion and contraction. Incorporating dynamic microclimates can thus improve models of species range shifts at spatial and temporal scales relevant to conservation interventions.

Street lighting changes the composition of invertebrate communities.

Artificial lighting has been used to illuminate the nocturnal environment for centuries and continues to expand with urbanization and economic development. Yet, the potential ecological impact of the resultant light pollution has only recently emerged as a major cause for concern. While investigations have demonstrated that artificial lighting can influence organism behaviour, reproductive success and survivorship, none have addressed whether it is altering the composition of communities. We show, for the first time, that invertebrate community composition is affected by proximity to street lighting independently of the time of day. Five major invertebrate groups contributed to compositional differences, resulting in an increase in the number of predatory and scavenging individuals in brightly lit communities. Our results indicate that street lighting changes the environment at higher levels of biological organization than previously recognized, raising the potential that it can alter the structure and function of ecosystems.

Long-term effects of artificial nighttime lighting and trophic complexity on plant biomass and foliar carbon and nitrogen in a grassland community.

The introduction of artificial nighttime lighting due to human settlements and transport networks is increasingly altering the timing, intensity, and spectra of natural light regimes worldwide. Much of the research on the impacts of nighttime light pollution on organisms has focused on animal species. Little is known about the impacts of daylength extension due to outdoor lighting technologies on wild plant communities, despite the fact that plant growth and development are under photoperiodic control. In a five-year field experiment, artificial ecosystems ("mesocosms") of grassland communities both alone or in combination with invertebrate herbivores and predators were exposed to light treatments that simulated street lighting technologies (low-pressure sodium, and light-emitting diode [LED]-based white lighting), at ground-level illuminance. Most of the plant species in the mesocosms did not exhibit changes in biomass accumulation after 5 years of exposure to the light treatments. However, the white LED treatment had a significant negative effect on biomass production in the herbaceous species Lotus pedunculatus. Likewise, the interaction between the white LED treatment and the presence of herbivores significantly reduced the mean shoot/root ratio of the grass species Holcus lanatus. Artificial nighttime lighting had no effect on the foliar carbon or nitrogen in most of the grassland species. Nevertheless, the white LED treatment significantly increased the leaf nitrogen content in Lotus corniculatus in the presence of herbivores. Long-term exposure to artificial light at night had no general effects on plant biomass responses in experimental grassland communities. However, species-specific and negative effects of cool white LED lighting at ground-level illuminance on biomass production and allocation in mixed plant communities are suggested by our findings. Further studies on the impacts of light pollution on biomass accumulation in plant communities are required as these effects could be mediated by different factors, including herbivory, competition, and soil nutrient availability.

Recent advances in the remote sensing of insects.

Remote sensing has revolutionised many aspects of ecological research, enabling spatiotemporal data to be collected in an efficient and highly automated manner. The last two decades have seen phenomenal growth in capabilities for high-resolution remote sensing that increasingly offers opportunities to study small, but ecologically important organisms, such as insects. Here we review current applications for using remote sensing within entomological research, highlighting the emerging opportunities that now arise through advances in spatial, temporal and spectral resolution. Remote sensing can be used to map environmental variables, such as habitat, microclimate and light pollution, capturing data on topography, vegetation structure and composition, and luminosity at spatial scales appropriate to insects. Such data can also be used to detect insects indirectly from the influences that they have on the environment, such as feeding damage or nest structures, whilst opportunities for directly detecting insects are also increasingly available. Entomological radar and light detection and ranging (LiDAR), for example, are transforming our understanding of aerial insect abundance and movement ecology, whilst ultra-high spatial resolution drone imagery presents tantalising new opportunities for direct observation. Remote sensing is rapidly developing into a powerful toolkit for entomologists, that we envisage will soon become an integral part of insect science.

Environmental risks from artificial nighttime lighting widespread and increasing across Europe.

The nighttime environment of much of Earth is being changed rapidly by the introduction of artificial lighting. While data on spatial and temporal variation in the intensity of artificial lighting have been available at a regional and global scale, data on variation in its spectral composition have only been collected for a few locations, preventing variation in associated environmental and human health risks from being mapped. Here, we use imagery obtained using digital cameras by astronauts on the International Space Station to map variation in the spectral composition of lighting across Europe for 2012-2013 and 2014-2020. These show a regionally widespread spectral shift, from that associated principally with high-pressure sodium lighting to that associated with broad white light-emitting diodes and with greater blue emissions. Reexpressing the color maps in terms of spectral indicators of environmental pressures, we find that this trend is widely increasing the risk of harmful effects to ecosystems.

Artificial nighttime lighting impacts visual ecology links between flowers, pollinators and predators.

The nighttime environment is being altered rapidly over large areas worldwide through introduction of artificial lighting, from streetlights and other sources. This is predicted to impact the visual ecology of many organisms, affecting both their intra- and interspecific interactions. Here, we show the effects of different artificial light sources on multiple aspects of hawkmoth visual ecology, including their perception of floral signals for pollination, the potential for intraspecific sexual signalling, and the effectiveness of their visual defences against avian predators. Light sources fall into three broad categories: some that prevent use of chromatic signals for these behaviours, others that more closely mimic natural lighting conditions, and, finally, types whose effects vary with light intensity and signal colour. We find that Phosphor Converted (PC) amber LED lighting - often suggested to be less harmful to nocturnal insects - falls into this third disruptive group, with unpredictable consequences for insect visual ecology depending on distance from the light source and the colour of the objects viewed. The diversity of impacts of artificial lighting on hawkmoth visual ecology alone argues for a nuanced approach to outdoor lighting in environmentally sensitive areas, employing intensities and spectra designed to limit those effects of most significant concern.

Pervasiveness of Biological Impacts of Artificial Light at Night.

Artificial light at night (ALAN) and its associated biological impacts have regularly been characterized as predominantly urban issues. Although far from trivial, this would imply that these impacts only affect ecosystems that are already heavily modified by humans and are relatively limited in their spatial extent, at least as compared with some key anthropogenic pressures on the environment that attract much more scientific and public attention, such as climate change or plastic pollution. However, there are a number of reasons to believe that ALAN and its impacts are more pervasive, and therefore need to be viewed from a broader geographic perspective rather than an essentially urban one. Here we address, in turn, 11 key issues when considering the degree of spatial pervasiveness of the biological impacts of ALAN. First, the global extent of ALAN is likely itself commonly underestimated, as a consequence of limitations of available remote sensing data sources and how these are processed. Second and third, more isolated (rural) and mobile (e.g., vehicle headlight) sources of ALAN may have both very widespread and important biological influences. Fourth and fifth, the occurrence and impacts of ALAN in marine systems and other remote settings, need much greater consideration. Sixth, seventh, and eighth, there is growing evidence for important biological impacts of ALAN at low light levels, from skyglow, and over long distances (because of the altitudes from which it may be viewed by some organisms), all of which would increase the areas over which impacts are occurring. Ninth and tenth, ALAN may exert indirect biological effects that may further expand these areas, because it has a landscape ecology (modifying movement and dispersal and so hence with effects beyond the direct extent of ALAN), and because ALAN interacts with other anthropogenic pressures on the environment. Finally, ALAN is not stable, but increasing rapidly in global extent, and shifting toward wavelengths of light that often have greater biological impacts.