Modeling long-distance seed dispersal of the invasive tree Spathodea campanulata in the Society Islands.

Long-distance dispersal is a key factor explaining the success of invasive alien species, particularly across oceanic islands. However, it is often not feasible to reliably measure long-distance seed dispersal (LDD) over many kilometers in the field. Here, we used a three-dimensional kinematic trajectory model (Computing Atmospheric Trajectory tool [CAT]) initiated on the basis of regional wind field data to assess the potential for LDD of a wind-dispersed invasive tree, Spathodea campanulata (African tulip tree), across the Society Islands (French Polynesia, South Pacific Ocean) following its initial planting and spread on the island of Tahiti. The main objective of our study was to determine whether S. campanulata could be expected to spread naturally among islands. Atmospheric dynamics, seed terminal velocity, precipitation, and temperature of air masses were considered to assess the potential for LDD between oceanic islands, with the island of Tahiti serving as the island source for multiple, geographically distant invasions. Aerial trajectories of modeled S. campanulata seeds indicated that wind-dispersed seeds originating from trees on the island of Tahiti could reach most of the Society Islands and disperse as far as 1364 km. This result suggests that Spathodea can be expected to spread naturally among the Society Islands. When rainfall events were modeled as causal agents of seed settlement, fewer seeds reached distant islands, but more seeds settled on the closest island (20 km away). Including effects of island topography ("barrier effects") also resulted in more seeds settling on the closest island and fewer seeds reaching the most distant islands. Overall, our findings suggest that recent atmospheric models can provide valuable insights into LDD and invasion patterns of wind-dispersed invasive species.

Pre-damage biomass allocation and not invasiveness predicts tolerance to damage in seedlings of woody species in Hawaii.

Plant-herbivore interactions have been predicted to play a fundamental role in plant invasions, although support for this assertion from previous research is mixed. While plants may escape from specialist herbivores in their introduced ranges, herbivory from generalists is common. Tolerance traits may allow non-native plants to mitigate the negative consequences of generalist herbivory that they cannot avoid in their introduced range. Here we address whether tolerance to herbivory, quantified as survival and compensatory growth, is associated with plant invasion success in Hawaii and investigate traits that may enhance tolerance in seedlings, the life stage most susceptible to herbivory. In a greenhouse experiment, we measured seedling tolerance to simulated herbivory through mechanical damage (50% leaf removal) of 16 non-native woody plant species differing in invasion status (invasive vs. non-invasive). Seedlings were grown for 2 weeks following damage and analyzed for biomass to determine whether damaged plants could fully compensate for the lost leaf tissue. Over 99% of all seedlings survived defoliation. Although species varied significantly in their levels of compensation, there was no consistent difference between invasive and non-invasive species. Seedlings of 11 species undercompensated and remained substantially smaller than control seedlings 2 weeks after damage; four species were close to compensating, while one species overcompensated. Across species, compensation was positively associated with an increased investment in potential storage reserves, specifically cotyledons and roots, suggesting that these organs provide resources that help seedlings re-grow following damage. Our results add to a growing consensus that pre-damage growth patterns determine tolerance to damage, even in young seedlings which have relatively low biomass. The lack of higher tolerance in highly invasive species may suggest that invaders overcome herbivory barriers to invasion in other ways, such as resistance traits, or that herbivory does not play an important role in the seedling invasion dynamics of these woody species in Hawaii.

Assembly of nonnative floras along elevational gradients explained by directional ecological filtering.

Nonnative species richness typically declines along environmental gradients such as elevation. It is usually assumed that this is because few invaders possess the necessary adaptations to succeed under extreme environmental conditions. Here, we show that nonnative plants reaching high elevations around the world are not highly specialized stress tolerators but species with broad climatic tolerances capable of growing across a wide elevational range. These results contrast with patterns for native species, and they can be explained by the unidirectional expansion of nonnative species from anthropogenic sources at low elevations and the progressive dropping out of species with narrow elevational amplitudes--a process that we call directional ecological filtering. Independent data confirm that climatic generalists have succeeded in colonizing the more extreme environments at higher elevations. These results suggest that invasion resistance is not conferred by extreme conditions at a particular site but determined by pathways of introduction of nonnative species. In the future, increased direct introduction of nonnative species with specialized ecophysiological adaptations to mountain environments could increase the risk of invasion. As well as providing a general explanation for gradients of nonnative species richness and the importance of traits such as phenotypic plasticity for many invasive species, the concept of directional ecological filtering is useful for understanding the initial assembly of some native floras at high elevations and latitudes.

Meta-analytic evidence that allelopathy may increase the success and impact of invasive grasses.

Background: In the grass family, a disproportionate number of species have been designated as being invasive. Various growth traits have been proposed to explain the invasiveness of grasses; however, the possibility that allelopathy gives invasive grasses a competitive advantage has attracted relatively little attention. Recent research has isolated plant allelochemicals that are mostly specific to the grass family that can breakdown into relatively stable, toxic byproducts. Methods: We conducted a meta-analysis of studies on grass allelopathy to test three prominent hypotheses from invasion biology and competition theory: (1) on native recipients, non-native grasses will have a significantly more negative effect compared to native grasses (Novel Weapons Hypothesis); (2) among native grasses, their effect on non-native recipients will be significantly more negative compared to their effect on native recipients (Biotic Resistance Hypothesis); and (3) allelopathic impacts will increase with phylogenetic distance (Phylogenetic Distance Hypothesis). From 23 studies, we gathered a dataset of 524 observed effect sizes (delta log response ratios) measuring the allelopathic impact of grasses on growth and germination of recipient species, and we used non-linear mixed-effects Bayesian modeling to test the hypotheses. Results: We found support for the Novel Weapons Hypothesis: on native recipients, non-native grasses were twice as suppressive as native grasses (22% vs 11%, respectively). The Phylogenetic Distance Hypothesis was supported by our finding of a significant correlation between phylogenetic distance and allelopathic impact. The Biotic Resistance Hypothesis was not supported. Overall, this meta-analysis adds to the evidence that allelochemicals may commonly contribute to successful or high impact invasions in the grass family. Increased awareness of the role of allelopathy in soil legacy effects associated with grass invasions may improve restoration outcomes through implementation of allelopathy-informed restoration practices. Examples of allelopathy-informed practices, and the knowledge needed to utilize them effectively, are discussed, including the use of activated carbon to neutralize allelochemicals and modify the soil microbial community.

Herbarium specimens can reveal impacts of climate change on plant phenology; a review of methods and applications.

Studies in plant phenology have provided some of the best evidence for large-scale responses to recent climate change. Over the last decade, more than thirty studies have used herbarium specimens to analyze changes in flowering phenology over time, although studies from tropical environments are thus far generally lacking. In this review, we summarize the approaches and applications used to date. Reproductive plant phenology has primarily been analyzed using two summary statistics, the mean flowering day of year and first-flowering day of year, but mean flowering day has proven to be a more robust statistic. Two types of regression models have been applied to test for associations between flowering, temperature and time: flowering day regressed on year and flowering day regressed on temperature. Most studies analyzed the effect of temperature by averaging temperatures from three months prior to the date of flowering. On average, published studies have used 55 herbarium specimens per species to characterize changes in phenology over time, but in many cases fewer specimens were used. Geospatial grid data are increasingly being used for determining average temperatures at herbarium specimen collection locations, allowing testing for finer scale correspondence between phenology and climate. Multiple studies have shown that inferences from herbarium specimen data are comparable to findings from systematically collected field observations. Understanding phenological responses to climate change is a crucial step towards recognizing implications for higher trophic levels and large-scale ecosystem processes. As herbaria are increasingly being digitized worldwide, more data are becoming available for future studies. As temperatures continue to rise globally, herbarium specimens are expected to become an increasingly important resource for analyzing plant responses to climate change.

Persistence at the final stage of volcanic island ontogeny: Abiotic predictors explain native plant species richness on 111 remote Pacific atolls.

AIM: The final island ontogeny of the general dynamic model (GDM) (i.e., before island submergence) in tropical oceans corresponds to the coral atoll stage. Here, we examined whether the species richness of native vascular plants (indigenous and endemic species) on atolls is controlled by spatial and/or physical processes. We also predicted that atolls strongly affected by anthropogenic disturbance would have lower native species richness than predicted by spatial and physical processes. LOCATION: Marshall Islands, Kiribati Islands, Nauru, Niue, Johnston, Cook Islands, French Polynesia and Pitcairn Islands (Pacific Ocean). TAXON: Native vascular plants. METHODS: We used stepwise regression to test the relative influence of five biogeographic variables on native species richness. Relationships were assessed for the full set of 111 Pacific coral atolls, as well as for atoll subsets ranging from 9 to 45 atolls. An index of human impact was then estimated, and residuals in the regression model predicting species richness from biogeographic variables were compared with the level of human impact. RESULTS: A regression model including atoll area, highest atoll elevation, the stepping stone distances from the nearest raised atoll and volcanic island explained native species richness on the 111 Pacific coral atolls. Regression models for different archipelagos and atoll subsets were also significant. Endemic species richness was significantly linked with highest atoll elevation and the stepping stone distances from the nearest raised atoll. Residuals in the biogeographic regression model were barely related to human impact across the 111 atolls but were significantly related to human impact in the Kiribati atolls. MAIN CONCLUSIONS: Native species richness on atolls is mainly controlled by physical and spatial characteristics. However, anthropogenic disturbances have altered the predicted pattern of native species richness leading to a lower model fit in some atoll subsets.

Short lag times for invasive tropical plants: evidence from experimental plantings in Hawai'i.

BACKGROUND: The lag time of an invasion is the delay between arrival of an introduced species and its successful spread in a new area. To date, most estimates of lag times for plants have been indirect or anecdotal, and these estimates suggest that plant invasions are often characterized by lag times of 50 years or more. No general estimates are available of lag times for tropical plant invasions. Historical plantings and documentation were used to directly estimate lag times for tropical plant invasions in Hawai'i. METHODOLOGY/PRINCIPAL FINDINGS: Historical planting records for the Lyon Arboretum dating back to 1920 were examined to identify plants that have since become invasive pests in the Hawaiian Islands. Annual reports describing escape from plantings were then used to determine the lag times between initial plantings and earliest recorded spread of the successful invaders. Among 23 species that eventually became invasive pests, the average lag time between introduction and first evidence of spread was 14 years for woody plants and 5 years for herbaceous plants. CONCLUSIONS/SIGNIFICANCE: These direct estimates of lag times are as much as an order of magnitude shorter than previous, indirect estimates, which were mainly based on temperate plants. Tropical invaders may have much shorter lag times than temperate species. A lack of direct and deliberate observations may have also inflated many previous lag time estimates. Although there have been documented cases of long lag times due to delayed arrival of a mutualist or environmental changes over time, this study suggests that most successful invasions are likely to begin shortly after arrival of the plant in a suitable habitat, at least in tropical environments. Short lag times suggest that controlled field trials may be a practical element of risk assessment for plant introductions.