Polyploidy on Islands: Its Emergence and Importance for Diversification.

Whole genome duplication or polyploidy is widespread among floras globally, but traditionally has been thought to have played a minor role in the evolution of island biodiversity, based on the low proportion of polyploid taxa present. We investigate five island systems (Juan Fernández, Galápagos, Canary Islands, Hawaiian Islands, and New Zealand) to test whether polyploidy (i) enhances or hinders diversification on islands and (ii) is an intrinsic feature of a lineage or an attribute that emerges in island environments. These island systems are diverse in their origins, geographic and latitudinal distributions, levels of plant species endemism (37% in the Galapagos to 88% in the Hawaiian Islands), and ploidy levels, and taken together are representative of islands more generally. We compiled data for vascular plants and summarized information for each genus on each island system, including the total number of species (native and endemic), generic endemicity, chromosome numbers, genome size, and ploidy levels. Dated phylogenies were used to infer lineage age, number of colonization events, and change in ploidy level relative to the non-island sister lineage. Using phylogenetic path analysis, we then tested how the diversification of endemic lineages varied with the direct and indirect effects of polyploidy (presence of polyploidy, time on island, polyploidization near colonization, colonizer pool size) and other lineage traits not associated with polyploidy (time on island, colonizer pool size, repeat colonization). Diploid and tetraploid were the most common ploidy levels across all islands, with the highest ploidy levels (>8x) recorded for the Canary Islands (12x) and New Zealand (20x). Overall, we found that endemic diversification of our focal island floras was shaped by polyploidy in many cases and certainly others still to be detected considering the lack of data in many lineages. Polyploid speciation on the islands was enhanced by a larger source of potential congeneric colonists and a change in ploidy level compared to overseas sister taxa.

Evolutionary priority effects persist in anthropogenically created habitats, but not through nonnative plant invasion.

Evolutionary priority effects, where early-arriving lineages occupy niche space via diversification and preclude dominance of later arrivals, have been observed in alpine and forest communities. However, the potential for evolutionary priority effects to persist in an era of rapid global change remains unclear. Here, we use a natural experiment of historical disturbance in New Zealand to test whether anthropogenic changes in available habitat and nonnative invasion eliminate the role of evolutionary priority effects in community assembly. We also test whether evolutionary priority effects diminish with decreasing resource availability. Older plant clades, as estimated by clade crown age, were relatively more abundant in both primary and secondary grassland. Relative abundance in primary grassland decreased with clade stem age, but only weakly. However, for both clade age estimates, relative abundance decreased with age when nonnative biomass was high and soil moisture was low. Our data show that patterns in community structure consistent with evolutionary priority effects can occur in both primary and secondary grasslands, the latter created by anthropogenic disturbance. However, nonnative invasion may overwhelm the effect of immigration timing on community dominance, possibly as a result of high immigration rates and preadaptation to anthropogenically modified environments.

Spatial heterogeneity of plant-soil feedbacks increases per capita reproductive biomass of species at an establishment disadvantage.

Plant-soil feedbacks have been widely implicated as a driver of plant community diversity, and the coexistence prediction generated by a negative plant-soil feedback can be tested using the mutual invasibility criterion: if two populations are able to invade one another, this result is consistent with stable coexistence. We previously showed that two co-occurring Rumex species exhibit negative pairwise plant-soil feedbacks, predicting that plant-soil feedbacks could lead to their coexistence. However, whether plants are able to reproduce when at an establishment disadvantage ("invasibility"), or what drivers in the soil might correlate with this pattern, are unknown. To address these questions, we created experimental plots with heterogeneous and homogeneous soils using field-collected conditioned soils from each of these Rumex species. We then allowed resident plants of each species to establish and added invader seeds of the congener to evaluate invasibility. Rumex congeners were mutually invasible, in that both species were able to establish and reproduce in the other's resident population. Invaders of both species had twice as much reproduction in heterogeneous compared to homogeneous soils; thus the spatial arrangement of plant-soil feedbacks may influence coexistence. Soil mixing had a non-additive effect on the soil bacterial and fungal communities, soil moisture, and phosphorous availability, suggesting that disturbance could dramatically alter soil legacy effects. Because the spatial arrangement of soil patches has coexistence implications, plant-soil feedback studies should move beyond studies of mean effects of single patch types, to consider how the spatial arrangement of patches in the field influences plant communities.

When do plant radiations influence community assembly? The importance of historical contingency in the race for niche space.

Plant radiations are widespread but their influence on community assembly has rarely been investigated. Theory and some evidence suggest that radiations can allow lineages to monopolize niche space when founding species arrive early into new bioclimatic regions and exploit ecological opportunities. These early radiations may subsequently reduce niche availability and dampen diversification of later arrivals. We tested this hypothesis of time-dependent lineage diversification and community dominance using the alpine flora of New Zealand. We estimated ages of 16 genera from published phylogenies and determined their relative occurrence across climatic and physical gradients in the alpine zone. We used these data to reconstruct occupancy of environmental space through time, integrating palaeoclimatic and palaeogeological changes. Our analysis suggested that earlier-colonizing lineages encountered a greater availability of environmental space, which promoted greater species diversity and occupancy of niche space. Genera that occupied broader niches were subsequently more dominant in local communities. An earlier time of arrival also contributed to greater diversity independently of its influence in accessing niche space. We suggest that plant radiations influence community assembly when they arise early in the occupancy of environmental space, allowing them to exclude later-arriving colonists from ecological communities by niche preemption.

Plant trait expression responds to establishment timing.

Trait divergence between co-occurring individuals could decrease the strength of competition between these individuals, thus promoting their coexistence. To test this hypothesis, we manipulated establishment timing for four congeneric pairs of perennial plants and assessed trait plasticity. Because soil conditions can affect trait expression and competition, we grew the plants in field-collected soil from each congener. Competition was generally weak across species, but the order of establishment affected divergence in biomass between potmates for three congeneric pairs. The type of plastic response differed among genera, with trait means of early-establishing individuals of Rumex and Solanum spp. differing from late-establishing individuals, and trait divergence between potmates of Plantago and Trifolium spp. depending on which species established first. Consistent with adaptive trait plasticity, higher specific leaf area (SLA) and root-shoot ratio in Rumex spp. established later suggest that these individuals were maximizing their ability to capture light and soil resources. Greater divergence in SLA correlated with increased summed biomass of competitors, which is consistent with trait divergence moderating the strength of competition for some species. Species did not consistently perform better in conspecific or congener soil, but soil type influenced the effect of establishment order. For example, biomass divergence between Rumex potmates was greater in R. obtusifolius soil regardless of which species established first. These results suggest that plant responses to establishment timing act in a species-specific fashion, potentially enhancing coexistence in plant communities.

Experimental protocol for manipulating plant-induced soil heterogeneity.

Coexistence theory has often treated environmental heterogeneity as being independent of the community composition; however biotic feedbacks such as plant-soil feedbacks (PSF) have large effects on plant performance, and create environmental heterogeneity that depends on the community composition. Understanding the importance of PSF for plant community assembly necessitates understanding of the role of heterogeneity in PSF, in addition to mean PSF effects. Here, we describe a protocol for manipulating plant-induced soil heterogeneity. Two example experiments are presented: (1) a field experiment with a 6-patch grid of soils to measure plant population responses and (2) a greenhouse experiment with 2-patch soils to measure individual plant responses. Soils can be collected from the zone of root influence (soils from the rhizosphere and directly adjacent to the rhizosphere) of plants in the field from conspecific and heterospecific plant species. Replicate collections are used to avoid pseudoreplicating soil samples. These soils are then placed into separate patches for heterogeneous treatments or mixed for a homogenized treatment. Care should be taken to ensure that heterogeneous and homogenized treatments experience the same degree of soil disturbance. Plants can then be placed in these soil treatments to determine the effect of plant-induced soil heterogeneity on plant performance. We demonstrate that plant-induced heterogeneity results in different outcomes than predicted by traditional coexistence models, perhaps because of the dynamic nature of these feedbacks. Theory that incorporates environmental heterogeneity influenced by the assembling community and additional empirical work is needed to determine when heterogeneity intrinsic to the assembling community will result in different assembly outcomes compared with heterogeneity extrinsic to the community composition.

Seed and establishment limitation contribute to long-term native forb declines in California grasslands.

The effects of exotic species invasions on biodiversity vary with spatial scale, and documentation of local-scale changes in biodiversity following invasion is generally lacking. Coupling long-term observations of local community dynamics with experiments to determine the role played by exotic species in recruitment limitation of native species would inform both our understanding of exotic impacts on natives at local scales and regional-scale management efforts to promote native persistence. We used field experimentation to quantify propagule and establishment limitation in a suite of native annual forbs in a California reserve, and compared these findings to species abundance trends within the same sites over the past 48 years. Observations at 11 paired sites (inside and outside the reserve) indicated that exotic annual plants have continued to increase in abundance over the past 48 years. This trend suggests the system has not reached equilibrium > 250 years after exotic species began to spread, and 70 years after livestock grazing ceased within the reserve. Long-term monitoring observations also indicated that six native annual forb species went extinct from more local populations than were colonized. To determine the potential role of exotic species in these native plant declines, we added seed of these species into plots adjacent to monitoring sites where plant litter and live grass competition were removed. Experimental results suggest both propagule and establishment limitation have contributed to local declines observed for these native forbs. Recruitment was highest at sites that had current or historical occurrences of the seeded species, and in plots where litter was removed. Grazing history (i.e., location within or outside the reserve) interacted with exotic competition removal, such that removal of live grass competition increased recruitment in more recently grazed sites. Abundance of forbs was positively related to recruitment, while abundance of exotic forbs was negatively related. Thus, exotic competition is likely only one factor contributing to local declines of native species in invaded ecosystems, with a combination of propagule limitation, site quality, and land use history also playing important and interactive roles in native plant recruitment.

Regional and decadal patterns of native and exotic plant coexistence in California grasslands.

Coexistence through a variety of mechanisms is possible for species with differential responses to environmental conditions. Understanding the role of environmental heterogeneity in mediating coexistence of species of different provenance (i.e., native vs. exotic) has important implications for theory and management. We used two California grassland data sets, one spanning seven years at three reserves along a 500-km latitudinal gradient and one spanning 47 years at 11 sites within a single 1000-ha reserve, to determine how environmental heterogeneity in space and time contributes to variability in provenance group abundance and diversity, and whether native and exotic species respond similarly to spatial and temporal variability. We found that temporal environmental heterogeneity is the primary determinant of provenance group abundance, while spatial and temporal environmental heterogeneity both contribute to community diversity. Spatial and temporal heterogeneity must therefore be considered simultaneously when examining community dynamics and species coexistence. Provenance was a poor general predictor of species response; native and exotic species exhibit similar spatiotemporal patterns in some cases but not others. Plant persistence may depend more upon the abiotic environment than competition from the other provenance group as native and exotic diversity were generally positively correlated. Furthermore, mesoscale (10(2)-10(3) m) spatial heterogeneity may be a greater mediator of provenance group coexistence than temporal heterogeneity or spatial heterogeneity at other scales.

Phylogeny and provenance affect plant-soil feedbacks in invaded California grasslands.

Plant-soil feedbacks can affect plant community dynamics by influencing processes of coexistence or invasion, or by maintaining alternate stable states. Darwin's naturalization hypothesis suggests that phylogenetic relatedness should be a critical factor governing such feedbacks in invaded communities but is rarely considered in soil feedback studies. We investigated the effects of soil biota from experimentally established native and invaded California grassland communities on resource capture and allocation of three native and three exotic grass species, comprising three tribes, grown in the laboratory. Phylogeny was the single greatest determinant of grass biomass, root:shoot ratio, and growth rate, with presence of soil biota explaining the second greatest proportion of variance in total grass biomass. Similar trends were observed in soil collected from naturally occurring stands of native perennial and exotic annual grasses. Species of similar life history/provenance exhibited similar biomass responses to the same soil community, while more closely related species exhibited similar root:shoot ratio responses to the same soil community. Relationships between the plant community composition of a field plot and species responses to soil inoculum collected from that field plot were idiosyncratic, with many aspects of plant community structure potentially contributing to soil feedbacks. Thus, future studies should explicitly consider both phylogeny and provenance and evaluate soil feedbacks in a community setting.