Outside the envelope: rare events disrupt the relationshipbetween climate factors and species interactions.

The order in which species arrive during community assembly can be an important driver of community composition and function. However, the strength of these priority effects can be variable, in part because of strong site and year effects. To understand how priority effects vary in importance with abiotic conditions, we initiated identical community assembly experiments in which we varied the timing of arrival of native and exotic grass species in each of 4 yr across three grassland sites in northern California. This uniquely replicated experiment tested the power of priority to determine initial community structure in a restoration context across a natural range of conditions. There were large and significant differences in both total seeded cover and the strength of priority across sites and years of initiation, confirming the suspicion that most ecological experiments may lack spatial and temporal generality. On the other hand, much of the variation in strength of priority could be related to climate. Strikingly, however, the model fit across the three sites and the first 3 yr of the study (the first nine experiments) was radically altered when we included the fourth year, which was characterized by an unusual weather pattern with higher temporal variability in rainfall (a rainfall pattern predicted to increase with climate change). This year produced relatively low strength of priority, supporting the suggestion that highly variable climates may be associated with lower strength of priority effects. Experiments that examine community assembly over a range of naturally occurring abiotic conditions enhance our ability to predict when priority effects will be important, allowing us to explore shifting patterns of community assembly in the face of climate change and optimize restoration strategies based on environmental conditions.

Persistent asymmetrical priority effects in a California grassland restoration experiment.

The order of species arrival can dramatically alter the trajectory of community development. While there is experimental evidence that priority effects can be important drivers of community structure early on, the persistence and duration of these effects is unclear. Here we report on a community assembly experiment in which a mix of four native grasses and a mix of four native forbs were planted on their own, together, or with one-year priority over the other guild. We found positive effects of priority for both grasses and forbs in the initial years of the experiment. However, 6-8 yr after planting, the effectiveness of priority treatments were mixed. Some species became rare, persisting only in treatments in which they had been given priority; others continued to maintain high cover and exhibit a strong positive signal of priority effects; still others remained common but no longer showed a signature of the initial priority effects; and finally, some species became locally extinct across all experimental plots. Grass priority over forbs was strong and persistent, but not forb priority over grasses. Our results demonstrate that the long-term benefits of temporal priority can persist for at least 8 yr for some, but not all species, and these continued effects result in distinct community composition. Manipulating the trajectory of community assembly through priority in seeding has potential as a useful tool for restoration.

Do invasive species perform better in their new ranges?

A fundamental assumption in invasion biology is that most invasive species exhibit enhanced performance in their introduced range relative to their home ranges. This idea has given rise to numerous hypotheses explaining "invasion success" by virtue of altered ecological and evolutionary pressures. There are surprisingly few data, however, testing the underlying assumption that the performance of introduced populations, including organism size, reproductive output, and abundance, is enhanced in their introduced compared to their native range. Here, we combined data from published studies to test this hypothesis for 26 plant and 27 animal species that are considered to be invasive. On average, individuals of these 53 species were indeed larger, more fecund, and more abundant in their introduced ranges. The overall mean, however, belied significant variability among species, as roughly half of the investigated species (N=27) performed similarly when compared to conspecific populations in their native range. Thus, although some invasive species are performing better in their new ranges, the pattern is not universal, and just as many are performing largely the same across ranges.

Can initial intraspecific spatial aggregation increase multi-year coexistence by creating temporal priority?

Both intraspecific spatial aggregation and temporal priority effects have the potential to increase long-term species coexistence. Theory and models suggest that intraspecific aggregation can facilitate coexistence via limited dispersal or asymmetric interaction distances. During community assembly, intraspecific aggregation may also delay interactions between more and less competitive species, thus creating opportunities for priority effects to facilitate longer-term coexistence. Few empirical studies have tested predictions about aggregation and coexistence, especially in the context of community assembly or ecological restoration. We investigated (1) impacts of intraspecific aggregation on the assembly of eight-species communities over three years, (2) the scale dependence of these impacts, and (3) implications for California prairie restoration. We planted eight native species in each of 19, 5 m wide, octagonal plots. Species were either interspersed throughout the plot or aggregated into eight, 2.2-m(2), wedge-shaped, monospecific sectors. Over three years, species diversity declined more quickly in interspersed plots than in aggregated plots. Two species had higher cover or increased more in interspersed than aggregated plots and were identified as "aggressives." Four species had higher cover or increased more in aggregated than interspersed plots and were identified as "subordinates." Within aggregated plots, aggressive species expanded beyond the sector in which they were originally seeded. Cover of aggressive species increased faster and reached higher values in sectors that were adjacent to the originally planted sector, compared to nonadjacent sectors. Cover of aggressive species also increased more and faster near plot centers, compared to plot edges. Areas near plot centers were representative of smaller aggregation patches since species were planted closer to heterospecific neighbors. Two subordinate species maintained higher cover near plot edges than near plot centers. Moreover, two subordinate species maintained higher cover when seeded in sectors farther away from aggressive species. These results suggest that initial intraspecific aggregation can facilitate species coexistence for at least three years, and larger aggregation patches may be more effective than smaller ones in the face of dispersing dominants. The creation of temporal priority effects may represent an underappreciated pathway by which intraspecific aggregation can increase coexistence. Restorationists may be able to maintain more diverse communities by planting in a mosaic of monospecific patches.

Initial success of native grasses is contingent on multiple interactions among exotic grass competition, temporal priority, rainfall and site effects.

Ecological communities are increasingly being recognized as the products of contemporary drivers and historical legacies that are both biotic and abiotic. In an attempt to unravel multiple layers of ecological contingency, we manipulated (i) competition with exotic annual grasses, (ii) the timing of this competition (temporal priority in arrival/seeding times) and (iii) watering (simulated rainfall) in a restoration-style planting of native perennial grasses. In addition, we replicated this experiment simultaneously at three sites in north-central California. Native perennial grasses had 73-99 % less cover when planted with exotic annuals than when planted alone, but this reduction was greatly ameliorated by planting the natives 2 weeks prior to the exotics. In a drought year, irrigation significantly reduced benefits of early planting so that these benefits resembled those observed in a non-drought year. There were significant differences across the three sites (site effects and interactions) in (i) overall native cover, (ii) the response of natives to competition, (iii) the strength of the temporal priority effect and (iv) the degree to which supplemental watering reduced priority effects. These results reveal the strong multi-layered contingency that underlies even relatively simple communities.