Mechanistic links between physiology and spectral reflectance enable previsual detection of oak wilt and drought stress.

Tree mortality due to global change-including range expansion of invasive pests and pathogens-is a paramount threat to forest ecosystems. Oak forests are among the most prevalent and valuable ecosystems both ecologically and economically in the United States. There is increasing interest in monitoring oak decline and death due to both drought and the oak wilt pathogen (Bretziella fagacearum). We combined anatomical and ecophysiological measurements with spectroscopy at leaf, canopy, and airborne levels to enable differentiation of oak wilt and drought, and detection prior to visible symptom appearance. We performed an outdoor potted experiment with Quercus rubra saplings subjected to drought stress and/or artificially inoculated with the pathogen. Models developed from spectral reflectance accurately predicted ecophysiological indicators of oak wilt and drought decline in both potted and field experiments with naturally grown saplings. Both oak wilt and drought resulted in blocked water transport through xylem conduits. However, oak wilt impaired conduits in localized regions of the xylem due to formation of tyloses instead of emboli. The localized tylose formation resulted in more variable canopy photosynthesis and water content in diseased trees than drought-stressed ones. Reflectance signatures of plant photosynthesis, water content, and cellular damage detected oak wilt and drought 12 d before visual symptoms appeared. Our results show that leaf spectral reflectance models predict ecophysiological processes relevant to detection and differentiation of disease and drought. Coupling spectral models that detect physiological change with spatial information enhances capacity to differentiate plant stress types such as oak wilt and drought.

Non-native plants tend to be phylogenetically distant but functionally similar to native plants under intense disturbance at the Three Gorges Reservoir Area.

Darwin's two opposing hypotheses, proposing that non-native species closely or distantly related to native species are more likely to succeed, are known as 'Darwin's Naturalization Conundrum'. Recently, invasion ecologists have sought to unravel these hypotheses. Studies that incorporate rich observational data in disturbed ecosystems that integrate phylogenetic and functional perspectives have potential to shed light on the conundrum. Using 313 invaded plant communities including 46 invasive plant species and 531 native plant species across the Three Gorges Reservoir Area in China, we aim to evaluate the coexistence mechanisms of invasive and native plants by integrating phylogenetic and functional dimensions at spatial and temporal scales. Our findings revealed that invasive plants tended to co-occur more frequently with native plant species that were phylogenetically distant but functionally similar in the reservoir riparian zone. Furthermore, our study demonstrated that the filtering of flood-dry-flood cycles played a significant role in deepening functional similarities of native communities and invasive-native species over time. Our study highlights the contrasting effects of phylogenetic relatedness and functional similarity between invasive and native species in highly flood-disturbed habitats, providing new sights into Darwin's Naturalization Conundrum.

Phylogenetic insights into the Salicaceae: The evolution of willows and beyond.

The Salicaceae includes approximately 54 genera and over 1,400 species with a cosmopolitan distribution. Members of the family are well-known for their diverse secondary plant metabolites, and they play crucial roles in tropical and temperate forest ecosystems. Phylogenetic reconstruction of the Salicaceae has been historically challenging due to the limitations of molecular markers and the extensive history of hybridization and polyploidy within the family. Our study employs whole-genome sequencing of 74 species to generate an extensive phylogeny of the Salicaceae. We generated two RAD-Seq enriched whole-genome sequence datasets and extracted two additional gene sets corresponding to the universal Angiosperms353 and Salicaceae-specific targeted-capture arrays. We reconstructed maximum likelihood-based molecular phylogenies using supermatrix and coalescent-based supertree approaches. Our fossil-calibrated phylogeny estimates that the Salicaceae originated around 128 million years ago and unravels the complex taxonomic relationships within the family. Our findings confirm the non-monophyly of the subgenus Salix s.l. and further support the merging of subgenera Chamaetia and Vetrix, both of which exhibit intricate patterns within and among different sections. Overall, our study not only enhances our understanding of the evolution of the Salicaceae, but also provides valuable insights into the complex relationships within the family.

Consistent physiological, ecological and evolutionary effects of fire regime on conservative leaf economics strategies in plant communities.

The functional response of plant communities to disturbance is hypothesised to be controlled by changes in environmental conditions and evolutionary history of species within the community. However, separating these influences using direct manipulations of repeated disturbances within ecosystems is rare. We evaluated how 41 years of manipulated fire affected plant leaf economics by sampling 89 plant species across a savanna-forest ecotone. Greater fire frequencies created a high-light and low-nitrogen environment, with more diverse communities that contained denser leaves and lower foliar nitrogen content. Strong trait-fire coupling resulted from the combination of significant intraspecific trait-fire correlations being in the same direction as interspecific trait differences arising through the turnover in functional composition along the fire-frequency gradient. Turnover among specific clades helped explain trait-fire trends, but traits were relatively labile. Overall, repeated burning led to reinforcing selective pressures that produced diverse plant communities dominated by conservative resource-use strategies and slow soil nitrogen cycling.

Drought response strategies are coupled with leaf habit in 35 evergreen and deciduous oak (Quercus) species across a climatic gradient in the Americas.

Distinct survival strategies can result from trade-offs in plant function under contrasting environments. Investment in drought resistance mechanisms can enhance survivorship but result in conservative growth. We tested the hypothesis that the widespread oaks (Quercus spp.) of the Americas exhibit an interspecific trade-off between drought resistance and growth capacity. Using experimental water treatments, we isolated adaptive trait associations among species in relation to their broad climates of origin and tested for correlated evolution between plant functional responses to water availability and habitat. Across all lineages, oaks displayed plastic drought responses - typically acclimating through osmolyte accumulation in leaves and/or employing conservative growth. Oaks from xeric climates had higher osmolytes and reduced stomatal pore area index, which allows for moderated gas exchange and limits tissue loss. Patterns suggest drought resistance strategies are convergent and under strong adaptive pressure. Leaf habit, however, mediates the growth and drought resistance strategies of oaks. Deciduous species, and evergreen species from xeric climates, have increased drought tolerance through osmoregulation, which allows for continuous, conservative growth. Evergreen mesic species show limited drought resistance but could enhance growth under well-watered conditions. Consequently, evergreen species from mesic environments are especially vulnerable to chronic drought and climate change.

Disentangling leaf structural and material properties in relationship to their anatomical and chemical compositional traits in oaks (Quercus L.).

BACKGROUND AND AIMS: The existence of sclerophyllous plants has been considered an adaptive strategy against different environmental stresses. Given that it literally means 'hard-leaved', it is essential to quantify the leaf mechanical properties to understand sclerophylly. However, the relative importance of each leaf trait for mechanical properties is not yet well established. METHODS: Genus Quercus is an excellent system to shed light on this because it minimizes phylogenetic variation while having a wide variation in sclerophylly. We measured leaf anatomical traits and cell wall composition, analysing their relationship with leaf mass per area and leaf mechanical properties in a set of 25 oak species. KEY RESULTS: The upper epidermis outer wall makes a strong and direct contribution to the leaf mechanical strength. Moreover, cellulose plays a crucial role in increasing leaf strength and toughness. The principal component analysis plot based on leaf trait values clearly separates Quercus species into two groups corresponding to evergreen and deciduous species. CONCLUSIONS: Sclerophyllous Quercus species are tougher and stronger owing to their thicker epidermis outer wall and/or higher cellulose concentration. Furthermore, section Ilex species share common traits, although they occupy different climates. In addition, evergreen species living in mediterranean-type climates share common leaf traits irrespective of their different phylogenetic origin.

Remotely detected aboveground plant function predicts belowground processes in two prairie diversity experiments.

Imaging spectroscopy provides the opportunity to incorporate leaf and canopy optical data into ecological studies, but the extent to which remote sensing of vegetation can enhance the study of belowground processes is not well understood. In terrestrial systems, aboveground and belowground vegetation quantity and quality are coupled, and both influence belowground microbial processes and nutrient cycling. We hypothesized that ecosystem productivity, and the chemical, structural and phylogenetic-functional composition of plant communities would be detectable with remote sensing and could be used to predict belowground plant and soil processes in two grassland biodiversity experiments: the BioDIV experiment at Cedar Creek Ecosystem Science Reserve in Minnesota and the Wood River Nature Conservancy experiment in Nebraska. We tested whether aboveground vegetation chemistry and productivity, as detected from airborne sensors, predict soil properties, microbial processes and community composition. Imaging spectroscopy data were used to map aboveground biomass, green vegetation cover, functional traits and phylogenetic-functional community composition of vegetation. We examined the relationships between the image-derived variables and soil carbon and nitrogen concentration, microbial community composition, biomass and extracellular enzyme activity, and soil processes, including net nitrogen mineralization. In the BioDIV experiment-which has low overall diversity and productivity despite high variation in each-belowground processes were driven mainly by variation in the amount of organic matter inputs to soils. As a consequence, soil respiration, microbial biomass and enzyme activity, and fungal and bacterial composition and diversity were significantly predicted by remotely sensed vegetation cover and biomass. In contrast, at Wood River-where plant diversity and productivity were consistently higher-belowground processes were driven mainly by variation in the quality of aboveground inputs to soils. Consequently, remotely sensed functional, chemical and phylogenetic composition of vegetation predicted belowground extracellular enzyme activity, microbial biomass, and net nitrogen mineralization rates but aboveground biomass (or cover) did not. The contrasting associations between the quantity (productivity) and quality (composition) of aboveground inputs with belowground soil attributes provide a basis for using imaging spectroscopy to understand belowground processes across productivity gradients in grassland systems. However, a mechanistic understanding of how above and belowground components interact among different ecosystems remains critical to extending these results broadly.

Representing plant diversity in land models: An evolutionary approach to make "Functional Types" more functional.

Plants are critical mediators of terrestrial mass and energy fluxes, and their structural and functional traits have profound impacts on local and global climate, biogeochemistry, biodiversity, and hydrology. Yet, Earth System Models (ESMs), our most powerful tools for predicting the effects of humans on the coupled biosphere-atmosphere system, simplify the incredible diversity of land plants into a handful of coarse categories of "Plant Functional Types" (PFTs) that often fail to capture ecological dynamics such as biome distributions. The inclusion of more realistic functional diversity is a recognized goal for ESMs, yet there is currently no consistent, widely accepted way to add diversity to models, that is, to determine what new PFTs to add and with what data to constrain their parameters. We review approaches to representing plant diversity in ESMs and draw on recent ecological and evolutionary findings to present an evolution-based functional type approach for further disaggregating functional diversity. Specifically, the prevalence of niche conservatism, or the tendency of closely related taxa to retain similar ecological and functional attributes through evolutionary time, reveals that evolutionary relatedness is a powerful framework for summarizing functional similarities and differences among plant types. We advocate that Plant Functional Types based on dominant evolutionary lineages ("Lineage Functional Types") will provide an ecologically defensible, tractable, and scalable framework for representing plant diversity in next-generation ESMs, with the potential to improve parameterization, process representation, and model benchmarking. We highlight how the importance of evolutionary history for plant function can unify the work of disparate fields to improve predictive modeling of the Earth system.

Enhanced light interception and light use efficiency explain overyielding in young tree communities.

Diverse plant communities are often more productive than mono-specific ones. Several possible mechanisms underlie this phenomenon but their relative importance remains unknown. Here we investigated whether light interception alone or in combination with light use efficiency (LUE) of dominant and subordinate species explained greater productivity of mixtures relative to monocultures (i.e. overyielding) in 108 young experimental tree communities. We found mixed-species communities that intercepted more light than their corresponding monocultures had 84% probability of overyielding. Enhanced LUE, which arose via several pathways, also mattered: the probability of overyielding was 71% when, in a mixture, species with higher 'inherent' LUE (i.e. LUE in monoculture) intercepted more light than species with lower LUE; 94% when dominant species increased their LUE in mixture; and 79% when subordinate species increased their LUE. Our results suggest that greater light interception and greater LUE, generated by inter and intraspecific variation, together drive overyielding in mixed-species forests.

Disease and fire interact to influence transitions between savanna-forest ecosystems over a multi-decadal experiment.

Global change is shifting disturbance regimes that may rapidly change ecosystems, sometimes causing ecosystems to shift between states. Interactions between disturbances such as fire and disease could have especially severe effects, but experimental tests of multi-decadal changes in disturbance regimes are rare. Here, we surveyed vegetation for 35 years in a 54-year fire frequency experiment in a temperate oak savanna-forest ecotone that experienced a recent outbreak of oak wilt. Different fire regimes determined whether plots were savanna or forest by regulating tree abundance (r2  = 0.70), but disease rapidly reversed the effect of fire exclusion, increasing mortality by 765% in unburned forests, but causing relatively minor changes in frequently burned savannas. Model simulations demonstrated that disease caused unburned forests to transition towards a unique woodland that was prone to transition to savanna if fire was reintroduced. Consequently, disease-fire interactions could shift ecosystem resilience and biome boundaries as pathogen distributions change.

Coupling spectral and resource-use complementarity in experimental grassland and forest communities.

Reflectance spectra provide integrative measures of plant phenotypes by capturing chemical, morphological, anatomical and architectural trait information. Here, we investigate the linkages between plant spectral variation, and spectral and resource-use complementarity that contribute to ecosystem productivity. In both a forest and prairie grassland diversity experiment, we delineated n-dimensional hypervolumes using wavelength bands of reflectance spectra to test the association between the spectral space occupied by individual plants and their growth, as well as between the spectral space occupied by plant communities and ecosystem productivity. We show that the spectral space occupied by individuals increased with their growth, and the spectral space occupied by plant communities increased with ecosystem productivity. Furthermore, ecosystem productivity was better explained by inter-individual spectral complementarity than by the large spectral space occupied by productive individuals. Our results indicate that spectral hypervolumes of plants can reflect ecological strategies that shape community composition and ecosystem function, and that spectral complementarity can reveal resource-use complementarity.

Deciduous and evergreen oaks show contrasting adaptive responses in leaf mass per area across environments.

Increases in leaf mass per area (LMA) are commonly observed in response to environmental stresses and are achieved through increases in leaf thickness and/or leaf density. Here, we investigated how the two underlying components of LMA differ in relation to species native climates and phylogeny, across deciduous and evergreen species. Using a phylogenetic approach, we quantified anatomical, compositional and climatic variables from 40 deciduous and 45 evergreen Quercus species from across the Northern Hemisphere growing in a common garden. Deciduous species from shorter growing seasons tended to have leaves with lower LMA and leaf thickness than those from longer growing seasons, while the opposite pattern was found for evergreens. For both habits, LMA and thickness increased in arid environments. However, this shift was associated with increased leaf density in evergreens but reduced density in deciduous species. Deciduous and evergreen oaks showed fundamental leaf morphological differences that revealed a diverse adaptive response. While LMA in deciduous species may have diversified in tight coordination with thickness mainly modulated by aridity, diversification of LMA within evergreens appears to be dependent on the infrageneric group, with diversification in leaf thickness modulated by both aridity and cold, while diversification in leaf density is only modulated by aridity.

Climate and phylogenetic history structure morphological and architectural trait variation among fine-root orders.

Fine roots mediate below-ground resource acquisition, yet understanding of how fine-root functional traits vary along environmental gradients, within branching orders and across phylogenetic scales remains limited. Morphological and architectural fine-root traits were measured on individual root orders of 20 oak species (genus Quercus) from divergent climates of origin that were harvested after three growing seasons in a glasshouse. These were then compared with similar measurements obtained from a phylogenetically diverse dataset of woody species from the Fine-Root Ecology Database (FRED). For the oaks, only precipitation seasonality and growing season moisture availability were correlated to aspects of root diameter and branching. Strong correlations among root diameters and architecture of different branch orders were common, while correlations between diameter and length were weakly negative. By contrast, the FRED dataset showed strong positive correlations between diameter and length and fewer correlations between root diameter and architectural traits. Our findings suggest that seasonal patterns of water availability are more important drivers of root adaptation in oaks than annual averages in precipitation and temperature. Furthermore, contrasting patterns of trait relationships between the oak and FRED datasets suggest that branching patterns are differentially constrained at narrow vs broad phylogenetic scales.

Foliar functional traits from imaging spectroscopy across biomes in eastern North America.

Foliar functional traits are widely used to characterize leaf and canopy properties that drive ecosystem processes and to infer physiological processes in Earth system models. Imaging spectroscopy provides great potential to map foliar traits to characterize continuous functional variation and diversity, but few studies have demonstrated consistent methods for mapping multiple traits across biomes. With airborne imaging spectroscopy data and field data from 19 sites, we developed trait models using partial least squares regression, and mapped 26 foliar traits in seven NEON (National Ecological Observatory Network) ecoregions (domains) including temperate and subtropical forests and grasslands of eastern North America. Model validation accuracy varied among traits (normalized root mean squared error, 9.1-19.4%; coefficient of determination, 0.28-0.82), with phenolic concentration, leaf mass per area and equivalent water thickness performing best across domains. Across all trait maps, 90% of vegetated pixels had reasonable values for one trait, and 28-81% provided high confidence for multiple traits concurrently. Maps of 26 traits and their uncertainties for eastern US NEON sites are available for download, and are being expanded to the western United States and tundra/boreal zone. These data enable better understanding of trait variations and relationships over large areas, calibration of ecosystem models, and assessment of continental-scale functional diversity.

Leaf reflectance spectra capture the evolutionary history of seed plants.

Leaf reflectance spectra have been increasingly used to assess plant diversity. However, we do not yet understand how spectra vary across the tree of life or how the evolution of leaf traits affects the differentiation of spectra among species and lineages. Here we describe a framework that integrates spectra with phylogenies and apply it to a global dataset of over 16 000 leaf-level spectra (400-2400 nm) for 544 seed plant species. We test for phylogenetic signal in spectra, evaluate their ability to classify lineages, and characterize their evolutionary dynamics. We show that phylogenetic signal is present in leaf spectra but that the spectral regions most strongly associated with the phylogeny vary among lineages. Despite among-lineage heterogeneity, broad plant groups, orders, and families can be identified from reflectance spectra. Evolutionary models also reveal that different spectral regions evolve at different rates and under different constraint levels, mirroring the evolution of their underlying traits. Leaf spectra capture the phylogenetic history of seed plants and the evolutionary dynamics of leaf chemistry and structure. Consequently, spectra have the potential to provide breakthrough assessments of leaf evolution and plant phylogenetic diversity at global scales.

Genomic landscape of the global oak phylogeny.

The tree of life is highly reticulate, with the history of population divergence emerging from populations of gene phylogenies that reflect histories of introgression, lineage sorting and divergence. In this study, we investigate global patterns of oak diversity and test the hypothesis that there are regions of the oak genome that are broadly informative about phylogeny. We utilize fossil data and restriction-site associated DNA sequencing (RAD-seq) for 632 individuals representing nearly 250 Quercus species to infer a time-calibrated phylogeny of the world's oaks. We use a reversible-jump Markov chain Monte Carlo method to reconstruct shifts in lineage diversification rates, accounting for among-clade sampling biases. We then map the > 20 000 RAD-seq loci back to an annotated oak genome and investigate genomic distribution of introgression and phylogenetic support across the phylogeny. Oak lineages have diversified among geographic regions, followed by ecological divergence within regions, in the Americas and Eurasia. Roughly 60% of oak diversity traces back to four clades that experienced increases in net diversification, probably in response to climatic transitions or ecological opportunity. The strong support for the phylogeny contrasts with high genomic heterogeneity in phylogenetic signal and introgression. Oaks are phylogenomic mosaics, and their diversity may in fact depend on the gene flow that shapes the oak genome.

Multi-temporal assessment of grassland α- and ß-diversity using hyperspectral imaging.

While more and more studies are exploring the application of remote sensing in assessing biodiversity for different ecosystems, most consider biodiversity at one point in time. Using several remote-sensing-based metrics, we asked how well remote sensing can detect biodiversity (both α- and ß-diversity) in a prairie grassland across time using airborne hyperspectral data collected in two successive years (2017 and 2018) and at different periods in the growing season (2018). The ability to detect biodiversity using "spectral diversity" and "spectral species" types indeed varied significantly over a 2-yr timespan. Toward the end of the growing season in 2018, the relationship between field- and remote-sensing-based α- and ß-diversity weakened compared to data collected from the same season in the previous year. This contrasting pattern between the two years was likely influenced by prescribed fire, altered weather, and the resulting shifting species composition and phenology. These findings indicate that direct detection of α- and ß-diversity in grasslands should be multi-temporal when possible and should consider the effect of disturbances, climate variables, and phenology. We demonstrate an essential role for airborne platforms in developing a global biodiversity monitoring system involving forthcoming space-borne hyperspectral sensors.

Consequences of drought severity for tropical live oak (Quercus oleoides) in Mesoamerica.

In two Costa Rican and three Honduran sites that vary in rainfall and soil properties, we used natural isotopes, a soil water balance model, and broad-scale climate-based drought indices to study shifts in water use with ontogeny from seedlings to mature tropical live oak (Quercus oleoides) trees. Water use patterns help to explain persistence of this broadly distributed species in Mesoamerica and to evaluate likely threats of ongoing climate changes. At the end of the dry season, soil δ18 O profiles can be described by one-phase exponential decay curves. Minimum values reflect geographic origins of the last significant rain event, and curvature is inversely related to canopy closure, demonstrating its role in controlling topsoil evaporation. Partitioning of soil water sources for transpiration was analyzed with a mixing model. In the Costa Rican sites, in a relatively dry year, saplings and mature trees took up water from the upper soil. In a relatively wet year in the Honduran sites, we observed deeper water extraction. In all sites, soil storage dampens extreme variation in water availability. The size dependence of water uptake with larger stems exploiting deeper layers is translated into variation in bulk leaf δ13 C-based water use efficiency (WUE) with the exception of mature trees. From 1932 to 2015, drought severity was evaluated with the Standardized Precipitation Evapotranspiration Index (SPEI) concurrently with simulations of the soil water balance model. Drought occurrence increased, regardless of the time period, averaged across 6, 12, or 24 months. All ontogenetic stages in all populations experienced frequent water limitation. We found evidence for linear trends toward aridification with increases of return periods of drought for October SPEI-24 declining from 42 to 6 yr in Costa Rica and from 21 to 7 yr in Honduras and recent occurrence of multiyear droughts from 2013 to 2016. October SPEI-12 and SPEI-24 were significantly related to the Oceanic Niño Indices demonstrating that local inter-annual variations in drought severity in Mesoamerica are modulated by large-scale climate forces. Drought severity in the near-term future depends on the extent to which the Pacific will adopt a more La Niña-like vs. a more El Niño-like state under ongoing climatic changes.

Horticultural availability and homeowner preferences drive plant diversity and composition in urban yards.

Understanding the factors that influence biodiversity in urban areas is important for informing management efforts aimed at enhancing the ecosystem services in urban settings and curbing the spread of invasive introduced species. We determined the ecological and socioeconomic factors that influence patterns of plant richness, phylogenetic diversity, and composition in 133 private household yards in the Minneapolis-Saint Paul Metropolitan area, Minnesota, USA. We compared the composition of spontaneously occurring plant species and those planted by homeowners with composition in natural areas (at the Cedar Creek Ecosystem Science Reserve) and in the horticulture pool of species available from commercial growers. Yard area and fertilizer frequency influenced species richness of the spontaneous species but expressed homeowner values did not. In contrast, the criteria that homeowners articulated as important in their management decisions, including aesthetics, wildlife, neatness and food provision, significantly predicted cultivated species richness. Strikingly, the composition of plant species that people cultivated in their yards resembled the taxonomic and phylogenetic composition of species available commercially. In contrast, the taxonomic and phylogenetic composition of spontaneous species showed high similarity to natural areas. The large fraction of introduced species that homeowners planted was a likely consequence of what was available for them to purchase. The study links the composition and diversity of yard flora to their natural and anthropogenic sources and sheds light on the human factors and values that influence the plant diversity in residential areas of a major urban system. Enhanced understanding of the influences of the sources of plants, both native and introduced, that enter urban systems and the human factors and values that influence their diversity is critical to identifying the levers to manage urban biodiversity and ecosystem services.

Traits linked with species invasiveness and community invasibility vary with time, stage and indicator of invasion in a long-term grassland experiment.

Much uncertainty remains about traits linked with successful invasion - the establishment and spread of non-resident species into existing communities. Using a 20-year experiment, where 50 non-resident (but mostly native) grassland plant species were sown into savannah plots, we ask how traits linked with invasion depend on invasion stage (establishment, spread), indicator of invasion success (occupancy, relative abundance), time, environmental conditions, propagule rain, and traits of invaders and invaded communities. Trait data for 164 taxa showed that invader occupancy was primarily associated with traits of invaders, traits of recipient communities, and invader-community interactions. Invader abundance was more strongly associated with community traits (e.g. proportion legume) and trait differences between invaders and the most similar resident species. Annuals and invaders with high-specific leaf area were only successful early in stand development, whereas invaders with conservative carbon capture strategies persisted long-term. Our results indicate that invasion is context-dependent and long-term experiments are required to comprehensively understand invasions.