In recent years, research in invasion biology has focused increasing attention on understanding the role of phenology in shaping plant invasions. Multiple studies have found non-native species that tend to flower distinctly early or late in the growing season, advance more with warming or have shifted earlier with climate change compared with native species. This growing body of literature has focused on patterns of phenological differences, but there is a need now for mechanistic studies of how phenology contributes to invasions. To do this, however, requires understanding how phenology fits within complex functional trait relationships. Towards this goal, we review recent literature linking phenology with other functional traits, and discuss the role of phenology in mediating how plants experience disturbance and stress-via climate, herbivory and competition-across the growing season. Because climate change may alter the timing and severity of stress and disturbance in many systems, it could provide novel opportunities for invasion-depending upon the dominant climate controller of the system, the projected climate change, and the traits of native and non-native species. Based on our current understanding of plant phenological and growth strategies-especially rapid growing, early-flowering species versus later-flowering species that make slower-return investments in growth-we project optimal periods for invasions across three distinct systems under current climate change scenarios. Research on plant invasions and phenology within this predictive framework would provide a more rigorous test of what drives invader success, while at the same time testing basic plant ecological theory. Additionally, extensions could provide the basis to model how ecosystem processes may shift in the future with continued climate change.

Plant invasions have tremendous potential to alter food webs by changing basal resources. Recent studies document how plant invasions may contribute to increased arthropod abundances in detritus-based food webs. An obvious mechanism for this phenomenon—a bottom-up effect resulting from elevated levels of detritus from the invasive plant litter—has not been explicitly studied. We examined the effects of an annual grass invasion on ground arthropod assemblages in the coastal sage scrub (CSS) of southern California. Bottom-up food web theory predicts that the addition of detritus would increase generalist-feeding arthropods at all trophic levels; accordingly, we expected increases in fungi, Collembola, and common predators such as mites and spiders. For the common ant taxa, habitat alteration may also be important for predicting responses. Thus we expected that Forelius mccooki and Pheidole vistana, the most common ant species, would decline because of changes in soil temperature (F. mccooki) and habitat structure (P. vistana) associated with litter. We studied trends observationally and conducted a 3-year experiment in which we manipulated litter quantity. In contrast to other published studies, most detritus-based arthropod taxa declined in areas of high grass invasion, and, within trophic levels, responses often varied idiosyncratically. For the two most common taxa, a native ant (F. mccooki), and predatory mites in the Anystidae, we experimentally linked declines in abundance to increased levels of invasive grass litter. Such declines, especially those exhibited by the most common ant taxa, could have cascading effects on the CSS ecosystem, where ants are numerically dominant and thus may have broad influences on food web and ecosystem properties. Our results highlight that accurately predicting arthropod responses to invasive plant litter requires careful consideration of the structural and food resources provided by detritus to each particular food web.

Phenological events - defined points in the life cycle of a plant or animal - have been regarded as highly plastic traits, reflecting flexible responses to various environmental cues. The ability of a species to track, via shifts in phenological events, the abiotic environment through time might dictate its vulnerability to future climate change. Understanding the predictors and drivers of phenological change is therefore critical. Here, we evaluated evidence for phylogenetic conservatism - the tendency for closely related species to share similar ecological and biological attributes - in phenological traits across flowering plants. We aggregated published and unpublished data on timing of first flower and first leaf, encompassing ̃4000 species at 23 sites across the Northern Hemisphere. We reconstructed the phylogeny for the set of included species, first, using the software program Phylomatic, and second, from DNA data. We then quantified phylogenetic conservatism in plant phenology within and across sites. We show that more closely related species tend to flower and leaf at similar times. By contrasting mean flowering times within and across sites, however, we illustrate that it is not the time of year that is conserved, but rather the phenological responses to a common set of abiotic cues. Our findings suggest that species cannot be treated as statistically independent when modelling phenological responses. Synthesis. Closely related species tend to resemble each other in the timing of their life-history events, a likely product of evolutionarily conserved responses to environmental cues. The search for the underlying drivers of phenology must therefore account for species' shared evolutionary histories. Closely related species tend to resemble each other in the timing of their life-history events, a likely product of evolutionarily conserved responses to environmental cues. The search for the underlying drivers of phenology must therefore account for species' shared evolutionary histories. © 2013 British Ecological Society.

Exotic species dominate many communities; however the functional significance of species' biogeographic origin remains highly contentious. This debate is fuelled in part by the lack of globally replicated, systematic data assessing the relationship between species provenance, function and response to perturbations. We examined the abundance of native and exotic plant species at 64 grasslands in 13 countries, and at a subset of the sites we experimentally tested native and exotic species responses to two fundamental drivers of invasion, mineral nutrient supplies and vertebrate herbivory. Exotic species are six times more likely to dominate communities than native species. Furthermore, while experimental nutrient addition increases the cover and richness of exotic species, nutrients decrease native diversity and cover. Native and exotic species also differ in their response to vertebrate consumer exclusion. These results suggest that species origin has functional significance, and that eutrophication will lead to increased exotic dominance in grasslands.

Invasions have increased the size of regional species pools, but are typically assumed to reduce native diversity. However, global-scale tests of this assumption have been elusive because of the focus on exotic species richness, rather than relative abundance. This is problematic because low invader richness can indicate invasion resistance by the native community or, alternatively, dominance by a single exotic species. Here, we used a globally replicated study to quantify relationships between exotic richness and abundance in grass-dominated ecosystems in 13 countries on six continents, ranging from salt marshes to alpine tundra. We tested effects of human land use, native community diversity, herbivore pressure, and nutrient limitation on exotic plant dominance. Despite its widespread use, exotic richness was a poor proxy for exotic dominance at low exotic richness, because sites that contained few exotic species ranged from relatively pristine (low exotic richness and cover) to almost completely exotic-dominated ones (low exotic richness but high exotic cover). Both exotic cover and richness were predicted by native plant diversity (native grass richness) and land use (distance to cultivation). Although climate was important for predicting both exotic cover and richness, climatic factors predicting cover (precipitation variability) differed from those predicting richness (maximum temperature and mean temperature in the wettest quarter). Herbivory and nutrient limitation did not predict exotic richness or cover. Exotic dominance was greatest in areas with low native grass richness at the site- or regional-scale. Although this could reflect native grass displacement, a lack of biotic resistance is a more likely explanation, given that grasses comprise the most aggressive invaders. These findings underscore the need to move beyond richness as a surrogate for the extent of invasion, because this metric confounds monodominance with invasion resistance. Monitoring species' relative abundance will more rapidly advance our understanding of invasions.