Additive genetic effects in interacting species jointly determine the outcome of caterpillar herbivory.

Plant-insect interactions are common and important in basic and applied biology. Trait and genetic variation can affect the outcome and evolution of these interactions, but the relative contributions of plant and insect genetic variation and how these interact remain unclear and are rarely subject to assessment in the same experimental context. Here, we address this knowledge gap using a recent host-range expansion onto alfalfa by the Melissa blue butterfly. Common garden rearing experiments and genomic data show that caterpillar performance depends on plant and insect genetic variation, with insect genetics contributing to performance earlier in development and plant genetics later. Our models of performance based on caterpillar genetics retained predictive power when applied to a second common garden. Much of the plant genetic effect could be explained by heritable variation in plant phytochemicals, especially saponins, peptides, and phosphatidyl cholines, providing a possible mechanistic understanding of variation in the species interaction. We find evidence of polygenic, mostly additive effects within and between species, with consistent effects of plant genotype on growth and development across multiple butterfly species. Our results inform theories of plant-insect coevolution and the evolution of diet breadth in herbivorous insects and other host-specific parasites.

Biochemistry and genetics of floral scent: a historical perspective.

Floral scent plays a crucial role in the reproductive process of many plants. Humans have been fascinated by floral scents throughout history, and have transported and traded floral scent products for which they have found multiple uses, such as in food additives, hygiene and perfume products, and medicines. Yet the scientific study of how plants synthesize floral scent compounds began later than studies on most other major plant metabolites, and the first report of the characterization of an enzyme responsible for the synthesis of a floral scent compound, namely linalool in Clarkia breweri, a California annual, appeared in 1994. In the almost 30 years since, enzymes and genes involved in the synthesis of hundreds of scent compounds from multiple plant species have been described. This review recapitulates this history and describes the major findings relating to the various aspects of floral scent biosynthesis and emission, including genes and enzymes and their evolution, storage and emission of scent volatiles, and the regulation of the biochemical processes.

Nitrogen-mediated volatilisation of defensive metabolites in tomato confers resistance to herbivores.

Plants synthesise a vast array of volatile organic compounds (VOCs), which serve as chemical defence and communication agents in their interactions with insect herbivores. Although nitrogen (N) is a critical resource in the production of plant metabolites, its regulatory effects on defensive VOCs remain largely unknown. Here, we investigated the effect of N content in tomato (Solanum lycopersicum) on the tobacco cutworm (Spodoptera litura), a notorious agricultural pest, using biochemical and molecular experiments in combination with insect behavioural and performance analyses. We observed that on tomato leaves with different N contents, S. litura showed distinct feeding preference and growth and developmental performance. Particularly, metabolomics profiling revealed that limited N availability conferred resistance upon tomato plants to S. litura is likely associated with the biosynthesis and emission of the volatile metabolite α-humulene as a repellent. Moreover, exogenous application of α-humulene on tomato leaves elicited a significant repellent response against herbivores. Thus, our findings unravel the key factors involved in N-mediated plant defence against insect herbivores and pave the way for innovation of N management to improve the plant defence responses to facilitate pest control strategies within agroecosystems.

Insect herbivory on woody broadleaf seedlings along a subtropical elevational gradient supports the resource concentration hypothesis.

PREMISE: Theories of plant-herbivore interactions hold that seedlings are more vulnerable to herbivory in warmer and more stable climates at lower elevations. Hypotheses of plant apparency, resource concentration, and resource availability have been proposed to explain variability in leaf herbivory. However, seasonal differences in the effects of these hypotheses on leaf herbivory on seedlings remain unclear. METHODS: We evaluated the three herbivory hypotheses by comparing the percentage and frequency of leaf herbivory in understory broadleaf seedlings in a subtropical forest in May (spring) and October (autumn) along an elevational gradient (290-1370 m a.s.l.). In total, we measured 2890 leaves across 696 seedlings belonging to 95 species and used beta regressions to test the effects of plant apparency (e.g., leaf area, seedling height), resource concentration (e.g., plant species diversity), and resource availability (e.g., canopy openness, soil available N and P) on leaf herbivory. RESULTS: Seedlings exhibited unimodal patterns of leaf herbivory along elevation, with drivers of leaf herbivory varying by the month. Variation in the frequency of leaf herbivory was best explained by the resource concentration hypothesis (e.g., plant species diversity) in both months, and herbivory was lower on seedlings in sites with higher plant diversity. Plant apparency hypothesis (e.g., leaf area, seedling height) was weakly supported only in spring, and the evidence for resource availability hypothesis (e.g., canopy openness, soil nutrients) was mixed. CONCLUSIONS: This study supports the resource concentration hypothesis and reveals the importance of seasonal difference on understanding leaf herbivory patterns and the drivers of plant diversity in subtropical forests.

GC-MS-based Metabolomics Unravels Metabolites across Larval Development and Diapause of a Specialist Insect.

Plant-insect interactions are a driving force into ecosystem evolution and community dynamics. Many insect herbivores enter diapause, a developmental arrest stage in anticipation of adverse conditions, to survive and thrive through seasonal changes. Herein, we investigated the roles of medium- to non-polar metabolites during larval development and diapause in a specialist insect herbivore, Chlosyne lacinia, reared on Aldama robusta leaves. Varying metabolites were determined using gas chromatography-mass spectrometry (GC-MS)-based metabolomics. Sesquiterpenes and steroids were the main metabolites putatively identified in A. robusta leaves, whereas C. lacinia caterpillars were characterized by triterpenes, steroids, fatty acids, and long-chain alkanes. We found out that C. lacinia caterpillars biosynthesized most of the identified steroids and fatty acids from plant-derived ingested metabolites, as well as all triterpenes and long-chain alkanes. Steroids, fatty acids, and long-chain alkanes were detected across all C. lacinia instars and in diapausing caterpillars. Sesquiterpenes and triterpenes were also detected across larval development, yet they were not detected in diapausing caterpillars, which suggested that these metabolites were converted to other molecules prior to the diapause stage. Our findings shed light on the chemical content variation across C. lacinia development and diapause, providing insights into the roles of metabolites in plant-insect interactions.

A semiochemical view of the ecology of the seed beetle Acanthoscelides obtectus Say (Coleoptera: Chrysomelidae, Bruchinae).

The dried bean beetle, Acanthoscelides obtectus, is an economically important pest of stored legumes worldwide. Tracking the human-aided dispersion of its primary hosts, the Phaseolus vulgaris beans, it is now widespread in most bean-growing areas of the tropics and subtropics. In temperate regions where it can only occasionally overwinter in the field, A. obtectus proliferates in granaries, having multiple generations a year. Despite its negative impact on food production, no sensitive detection or monitoring tools exist, and the reduction of local populations still relies primarily on inorganic insecticides as fumigating agents. However, in the quest to produce more nutritious food more sustainably and healthily, the development of environmentally benign crop protection methods is vital against A. obtectus. For this, knowledge of the biology and chemistry of both the host plant and its herbivore will underpin the development of, among others, chemical ecology-based approaches to form an essential part of the toolkit of integrated bruchid management. We review the semiochemistry of the mate- and host-finding behaviour of A. obtectus and provide new information about the effect of seed chemistry on the sensory and behavioural ecology of host acceptance and larval development.

Transcriptomic and metabolomic analyses reveals keys genes and metabolic pathways in tea (Camellia sinensis) against six-spotted spider mite (Eotetranychus Sexmaculatus).

BACKGROUND: Six-spotted spider mite (Eotetranychus sexmaculatus) is one of the most damaging pests of tea (Camellia sinensis). E. sexmaculatus causes great economic loss and affects tea quality adversely. In response to pests, such as spider mites, tea plants have evolved resistance mechanisms, such as expression of defense-related genes and defense-related metabolites. RESULTS: To evaluate the biochemical and molecular mechanisms of resistance in C. sinensis against spider mites, "Tianfu-5" (resistant to E. sexmaculatus) and "Fuding Dabai" (susceptible to E. sexmaculatus) were inoculated with spider mites. Transcriptomics and metabolomics based on RNA-Seq and liquid chromatography coupled to tandem mass spectrometry (LC-MS/MS) technology were used to analyze changes in gene expression and metabolite content, respectively. RNA-Seq data analysis revealed that 246 to 3,986 differentially expressed genes (DEGs) were identified in multiple compared groups, and these DEGs were significantly enriched in various pathways, such as phenylpropanoid and flavonoid biosynthesis, plant-pathogen interactions, MAPK signaling, and plant hormone signaling. Additionally, the metabolome data detected 2,220 metabolites, with 194 to 260 differentially abundant metabolites (DAMs) identified in multiple compared groups, including phenylalanine, lignin, salicylic acid, and jasmonic acid. The combined analysis of RNA-Seq and metabolomic data indicated that phenylpropanoid and flavonoid biosynthesis, MAPK signaling, and Ca2+-mediated PR-1 signaling pathways may contribute to spider mite resistance. CONCLUSIONS: Our findings provide insights for identifying insect-induced genes and metabolites and form a basis for studies on mechanisms of host defense against spider mites in C. sinensis. The candidate genes and metabolites identified will be a valuable resource for tea breeding in response to biotic stress.

Transcriptome features of stone cell development in weevil-resistant and susceptible Sitka spruce.

Stone cells are a specialized, highly lignified cell type found in both angiosperms and gymnosperms. In conifers, abundance of stone cells in the cortex provides a robust constitutive physical defense against stem feeding insects. Stone cells are a major insect-resistance trait in Sitka spruce (Picea sitchensis), occurring in dense clusters in apical shoots of trees resistant (R) to spruce weevil (Pissodes strobi) but being rare in susceptible (S) trees. To learn more about molecular mechanisms of stone cell formation in conifers, we used laser microdissection and RNA sequencing to develop cell-type-specific transcriptomes of developing stone cells from R and S trees. Using light, immunohistochemical, and fluorescence microscopy, we also visualized the deposition of cellulose, xylan, and lignin associated with stone cell development. A total of 1293 genes were differentially expressed at higher levels in developing stone cells relative to cortical parenchyma. Genes with potential roles in stone cell secondary cell wall formation (SCW) were identified and their expression evaluated over a time course of stone cell formation in R and S trees. The expression of several transcriptional regulators was associated with stone cell formation, including a NAC family transcription factor and several genes annotated as MYB transcription factors with known roles in SCW formation.

Enhanced ß-glucosidase in Western flower thrips affects its interaction with the redox-based strategies of kidney beans under elevated CO2.

ß-Glucosidase is validated as an elicitor for early immune responses in plants and it was detected in the salivary glands of Frankliniella occidentalis in previous research. Seven differentially expressed genes encoding ß-glucosidase were obtained by comparing the transcriptomes of F. occidentalis adults grown under two different CO2 concentrations (800 vs. 400 ppm), which might be associated with the differences in the interaction between F. occidentalis adults and its host plant, Phaseolus vulgaris under different CO2 levels. To verify this speculation, changes in defense responses based on the production and elimination of reactive oxygen species (ROS) in P. vulgaris leaves treated with three levels of ß-glucosidase activity under ambient CO2 (aCO2 ) and elevated CO2 (eCO2 ) were measured in this study. According to the results, significantly higher levels of ROS were noticed under eCO2 compared to aCO2 , which was caused by the increased ß-glucosidase activity in thrips due to increased cellulose content in P. vulgaris leaves under eCO2 . Together with the lower activities of superoxide dismutase (SOD), peroxidase (POD) and catalase (CAT) in injured leaves under eCO2 , P. vulgaris leaves would be negatively affected on redox-based defense by eCO2 , thus facilitating thrips damage under climate change.

Effects of plant diversity on species-specific herbivory: patterns and mechanisms.

Invertebrate herbivory can shape plant communities when impacting growth and fitness of some plant species more than other species. Previous studies showed that herbivory varies among plant species and that species-specific herbivory is affected by the diversity of the surrounding plant community. However, mechanisms underlying this variation are still poorly understood. In this study, we investigate how plant traits and plant apparency explain differences in herbivory among plant species and we explore the effect of plant community diversity on these species-specific relationships. We found that species differed in the herbivory they experienced. Forbs were three times more damaged by herbivores than grasses. Variability within grasses was caused by differences in leaf dry matter content (LDMC). Furthermore, higher plant diversity increased herbivory on 15 plant species and decreased herbivory on nine species. Variation within forb and grass species in their response to changing plant diversity was best explained by species' physical resistance (LDMC, forbs) and biomass (grasses). Overall, our results show that herbivory and diversity effects on herbivory differ among species, and that, depending on the plant functional group, either species-specific traits or apparency are driving those differences. Thus, herbivores might selectively consume palatable forbs or abundant grasses with contrasting consequences for plant community composition in grasslands dominated by either forbs or grasses.

Volatiles as Attractants of Mahogany Shoot Borer, Hypsipyla grandella Zeller (Lepidoptera: Pyralidae).

The mahogany shoot borer, Hypsipyla grandella Zeller (Lepidoptera: Pyralidae), is one of the most economically important pests in all American tropical forests because it prevents the establishment of monoculture plantations of the family Meliaceae, such as Spanish cedar, Cedrela odorata L. Various studies have focussed on the bioecological aspects and the chemical and silvicultural control of this pest. However, relatively little is known about the biological interactions between this insect and its host plant. In this study, the shoot borer's behavior and attraction response to cedar host plants was evaluated in field cages. We also identified the volatiles emitted by healthy C. odorata plants that were attractive to H. grandella adults. The attraction to headspace volatiles from cedar plants and a synthetic blend were evaluated in a Y-glass tube olfactometer. We observed that virgin and mated females exhibited low activity at night, frequent movement of the antennae, sporadic flight activity, and short (< 10 s) and long (> 30 s) wing-fanning. Virgin females assumed a calling position, whereas mated females exhibited three periods of oviposition. The results showed that all evaluated categories - virgin females, virgin males, and mated females - were attracted to cedar plants. We identified the following volatile compounds: α-pinene, (E)-ß-ocimene, 2-ethyl-1-hexanol, limonene, nonanal, (E)-4,8-dimethyl-1,3,7-nonatriene, α-copaene, ß-caryophyllene, and germacrene D. A synthetic blend significantly attracted virgin male and mated female shoot borers. Our results suggested that C. odorata volatiles compounds are responsible for the attraction of H. grandella.

Structural patterns of Lepidoptera galls and the case of Andescecidium parrai (Cecidosidae) galls on Schinus polygama (Anacardiaceae).

Gall anatomical and metabolic peculiarities are determined by the feeding habit of the gall inducer, but develop under the constraints of the host plants. The chewing habit of the Lepidoptera larvae imposes a high impact on the host plant cells, and supposedly drives peculiar structural and histochemical patterns. So, our starting point was the search of such patterns in literature, and the test of these traits on the Andescecidium parrai (Cecidosidae)-Schinus polygama (Anacardiaceae) system, as a case study in Chilean flora. The literature on the structure of lepidopteran galls in the temperate and tropical regions comprises 13 works, describing stems as the most frequent host organs, followed by leaves, buds, and flowers. As common structural traits of Lepidoptera galls, the literature converge in describing the processes of cell hypertrophy and hyperplasia, resulting in a variable number of common storage parenchyma layers, interspersed by the redifferentiated sclerenchyma, vascular, and typical nutritive cells around the larval chamber. These nutritive cells accumulate lipids and proteins, which support the lepidopteran larvae nutrition. As expected, the A. parrai galls follow the patterns herein described for the lepidoptera-induced galls, but with peculiarities associated with its host organ. Even though the Lepidoptera galls have destructive mouthparts and can induce large and complex galls, they cannot alter important conservative features of their hosts' organs.

Insect herbivory antagonizes leaf cooling responses to elevated temperature in tomato.

As global climate change brings elevated average temperatures and more frequent and extreme weather events, pressure from biotic stresses will become increasingly compounded by harsh abiotic stress conditions. The plant hormone jasmonate (JA) promotes resilience to many environmental stresses, including attack by arthropod herbivores whose feeding activity is often stimulated by rising temperatures. How wound-induced JA signaling affects plant adaptive responses to elevated temperature (ET), however, remains largely unknown. In this study, we used the commercially important crop plant Solanum lycopersicum (cultivated tomato) to investigate the interaction between simulated heat waves and wound-inducible JA responses. We provide evidence that the heat shock protein HSP90 enhances wound responses at ET by increasing the accumulation of the JA receptor, COI1. Wound-induced JA responses directly interfered with short-term adaptation to ET by blocking leaf hyponasty and evaporative cooling. Specifically, leaf damage inflicted by insect herbivory or mechanical wounding at ET resulted in COI1-dependent stomatal closure, leading to increased leaf temperature, lower photosynthetic carbon assimilation rate, and growth inhibition. Pharmacological inhibition of HSP90 reversed these effects to recapitulate the phenotype of a JA-insensitive mutant lacking the COI1 receptor. As climate change is predicted to compound biotic stress with larger and more voracious arthropod pest populations, our results suggest that antagonistic responses resulting from a combination of insect herbivory and moderate heat stress may exacerbate crop losses.

Plant water limitation and its impact on the oviposition preferences of the monarch butterfly (Lepidoptera: Nymphalidae).

Intensifying drought conditions across the western United States due to global climate change are altering plant-insect interactions. Specialist herbivores must find their host plants within a matrix of nonhosts, and thus often rely upon specific plant secondary chemistry for host location and oviposition cues. Climate-induced alterations to plant chemistry could thus affect female selection of larval food plants. Here, we investigated whether host-plant water limitation influenced oviposition preference in a threatened invertebrate: the monarch butterfly (Danaus plexippus). We found that females deposited more eggs on reduced-water than on well-watered narrowleaf milkweed plants (Asclepias fascicularis), but we could not attribute this change to any specific change in plant chemistry. Specialist herbivores, such as the monarch butterfly, which are tightly linked to specific plant cues, may experience shift in preferences under global-change conditions. Understanding oviposition preferences will be important to directing ongoing habitat restoration activities for this declining insect.

Small Brown Planthopper Nymph Infestation Regulates Plant Defenses by Affecting Secondary Metabolite Biosynthesis in Rice.

The small brown planthopper (SBPH, Laodelphax striatellus) is one of the most destructive insect pests in rice (Oryza sativa), which is the world's major grain crop. The dynamic changes in the rice transcriptome and metabolome in response to planthopper female adult feeding and oviposition have been reported. However, the effects of nymph feeding remain unclear. In this study, we found that pre-infestation with SBPH nymphs increased the susceptibility of rice plants to SBPH infestation. We used a combination of broadly targeted metabolomic and transcriptomic studies to investigate the rice metabolites altered by SBPH feeding. We observed that SBPH feeding induced significant changes in 92 metabolites, including 56 defense-related secondary metabolites (34 flavonoids, 17 alkaloids, and 5 phenolic acids). Notably, there were more downregulated metabolites than upregulated metabolites. Additionally, nymph feeding significantly increased the accumulation of seven phenolamines and three phenolic acids but decreased the levels of most flavonoids. In SBPH-infested groups, 29 differentially accumulated flavonoids were downregulated, and this effect was more pronounced with infestation time. The findings of this study indicate that SBPH nymph feeding suppresses flavonoid biosynthesis in rice, resulting in increased susceptibility to SBPH infestation.

Identification of Potential Gene Targets for Suppressing Oviposition in Holotrichia parallela Using Comparative Transcriptome Analysis.

Holotrichia parallela is an important plant pest. Comparative feeding experiments showed that the egg production, oviposition duration and survival rate of H. parallela beetles were significantly higher when they fed on elm leaves than when they fed on willow or purpus privet leaves. RNA sequencing was used to determine transcriptomic changes associated with oviposition. Comparative transcriptome analysis revealed that the beetles that fed on elm and willow had a total of 171 genes with differential expression. When the beetles fed on elm and purpus privet, 3568 genes had differential expression. The vitellogenesis, ovarian serine protease, odorant-binding proteins, acyl-CoA synthetase and follicle cell proteins were commonly upregulated genes in elm-fed beetles compared with those fed on willow/purpus privet leaves. The involvement of the follicle cell protein 3C gene in the regulation of oviposition was confirmed using RNA interference. The results provide insights into the molecular mechanisms underlying oviposition in H. parallela feeding on different host plants. This study also describes a method for identifying potentially effective genes for pest control.

Exploring spatiotemporal dynamics of flower visitor association pattern on two Avicennia mangroves: a network approach.

Plant-flower visitor interaction is one of the most important relationships regarding the co-existence of the floral and faunal communities. The implication of network approaches is an efficient way to understand the impact of community structure on ecosystem functionality. To understand the association pattern of flower visitors, we performed this study on Avicennia officinalis and Avicennia marina mangroves from the islands of Indian Sundarban over three consecutive years. We found that visiting time and sites (islands) influenced the abundance of visitors. The bipartite networks showed a significant generalized structure for both site-visitor and visiting time-visitor networks where the strength and specialization of visitor species showed a highly and moderately significant positive correlation between both networks respectively. All the site-wise visiting time-visitor networks and year-wise site-visitor networks were significantly modular in structure. For both the plants, most of the visitors showed a generalized association pattern among islands and also among visiting times. Additionally, the study of the foraging behavior of dominant visitors showed Apis dorsata and Apis mellifera as the potential visitors for these plants. Our results showed that flower visitor networks are spatiotemporally dynamic. The interactions of visitors with flowers at different times influence their contribution to the network for becoming a generalist or peripheral species in the context of their visiting time, which may subsequently change over islands. This approach will help to devise more precise plant species-specific conservation strategies by understanding the contribution of visitors through the spatiotemporal context.

Pesticide resistance in arthropods: Ecology matters too.

Pesticide resistance development is an example of rapid contemporary evolution that poses immense challenges for agriculture. It typically evolves due to the strong directional selection that pesticide treatments exert on herbivorous arthropods. However, recent research suggests that some species are more prone to evolve pesticide resistance than others due to their evolutionary history and standing genetic variation. Generalist species might develop pesticide resistance especially rapidly due to pre-adaptation to handle a wide array of plant allelochemicals. Moreover, research has shown that adaptation to novel host plants could lead to increased pesticide resistance. Exploring such cross-resistance between host plant range evolution and pesticide resistance development from an ecological perspective is needed to understand its causes and consequences better. Much research has, however, been devoted to the molecular mechanisms underlying pesticide resistance while both the ecological contexts that could facilitate resistance evolution and the ecological consequences of cross-resistance have been under-studied. Here, we take an eco-evolutionary approach and discuss circumstances that may facilitate cross-resistance in arthropods and the consequences cross-resistance may have for plant-arthropod interactions in both target and non-target species and species interactions. Furthermore, we suggest future research avenues and practical implications of an increased ecological understanding of pesticide resistance evolution.

Chemical defense acquired via pharmacophagy can lead to protection from predation for conspecifics in a sawfly.

Chemical defense is a widespread anti-predator strategy exhibited by organisms, with individuals either synthesizing or extrinsically acquiring defensive chemicals. In some species, such defences can also be transferred among conspecifics. Here, we tested the effects of pharmacophagy on the defense capability of the turnip sawfly, Athalia rosae, which can acquire neo-clerodane diterpenoids (clerodanoids) via pharmacophagy when having access to the plant Ajuga reptans. We show that clerodanoid access mediates protection against predation by mantids for the sawflies, both in a no-choice feeding assay and a microcosm setup. Even indirect access to clerodanoids, via nibbling on conspecifics that had access to the plant, resulted in protection against predation albeit to a lower degree than direct access. Furthermore, sawflies that had no direct access to clerodanoids were consumed less frequently by mantids when they were grouped with conspecifics that had direct access. Most, but not all, of such initially undefended sawflies could acquire clerodanoids from conspecifics that had direct access to the plant, although in low quantities. Together our results demonstrate that clerodanoids serve as a chemical defense that can also be transferred by interactions among conspecifics. Moreover, the presence of chemically defended individuals in a group can confer protection onto conspecifics that had no direct access to clerodanoids.

The Matthew effect: Common species become more common and rare ones become more rare in response to artificial light at night.

Artificial light at night (ALAN) has been and still is rapidly spreading and has become an important component of global change. Although numerous studies have tested its potential biological and ecological impacts on animals, very few studies have tested whether it affects alien and native plants differently. Furthermore, common plant species, and particularly common alien species, are often found to benefit more from additional resources than rare native and rare alien species. Whether this is also the case with regard to increasing light due to ALAN is still unknown. Here, we tested how ALAN affected the performance of common and rare alien and native plant species in Germany directly, and indirectly via flying insects. We grew five common alien, six rare alien, five common native, and four rare native plant species under four combinations of two ALAN (no ALAN vs. ALAN) and two insect-exclusion (no exclusion vs. exclusion) treatments, and compared their biomass production. We found that common plant species, irrespective of their origin, produced significantly more biomass than rare species and that this was particularly true under ALAN. Furthermore, alien species tended to show a slightly stronger positive response to ALAN than native species did (p = .079). Our study shows that common plant species benefited more from ALAN than rare ones. This might lead to competitive exclusion of rare species, which could have cascading impacts on other trophic levels and thus have important community-wide consequences when ALAN becomes more widespread. In addition, the slightly more positive response of alien species indicates that ALAN might increase the risk of alien plant invasions.