Abstract. 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 ¼ 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.
Temporal priority can affect individual performance and reproduction, as well as community assembly, but whether these effects persist over time remains unclear, and their demographic mechanisms have been little explored. The continued dominance of exotic annual grasses in California has been commonly attributed to their demonstrated early germination and rapid early growth relative to native perennial grasses. This advantage may play a crucial role in the structure of California exotic annual grasslands, as well as in the practice of native grassland restoration. We tested whether a two-week planting advantage under field conditions increased individual survival, growth, and reproduction for four native perennial grass species and whether these effects persisted over three years. We show that short-term priority significantly increased the establishment success of' native perennial grasses. Increased density of native grass seedlings presaged later large increases in cover that were not evident in the first year after planting. Although priority effects at the individual level may diminish over time, short differences in emergence timing can have long-lasting effects on community structure. Earlier germination and faster initial growth of exotic annual species may help explain their unprecedented invasion and continued dominance of California grasslands. Finally, these results highlight the importance of priority effects for effective exotic annual control during native grassland restoration in California: initial control can increase the establishment of native perennial seedlings, which then results in long-term control by mature native individuals.
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.
Interannual variation in experimental field conditions produce variability in the results of experiments monitored over multiple years, termed here "year effects." When experimental treatments are replicated in separate years, interannual variation may influence treatment effects and produce significant treatment by initiation-year interactions. Understanding the frequency and strength of these effects requires initiating identical experiments across years. We conducted a review of literature covering more than 500 experimental articles published in 7 journals between 1966 and 2008. Only 5% of the 276 general ecological field studies initiated experiments in multiple years. This rarity was even more evident in the journal Restoration Ecology, in which none of the 173 surveyed experimental studies initiated experiments in multiple years. In contrast, 48% of the 58 field experiments published in an agronomy journal were replicated across years. We found only 17 studies that tested treatment by initiation-year interactions. Despite their rarity, 76% of these studies found significant interactions between treatment and initiation year. We conclude that the results of many ecological field experiments are likely to be contingent on the year in which they are implemented. We discuss the importance of treatment by initiation-year interactions in ecology and restoration, factors that have hindered the inclusion of temporal replication in the past, and some suggestions for the appropriate design and analysis of temporally replicated experiments. We argue for more deliberate investigation of temporal contingency in ecological experimentation, especially in the field of restoration ecology, which may be particularly sensitive to treatment by initiation-year interactions.
Throughout the western United States, native perennial grasses are being supplanted by aggressive non-native annuals. We show that giving native grasses just a two-week germination 'head start' over exotic invasive grasses shifts the competitive edge strongly in their favour. We also show that the strength of this advantage differs strikingly depending on the site where the experiment is carried out, and the weather in the initial weeks of the experiment. These results a) give insight into the reasons for the competitive advantage that annuals usually demonstrate, and b) are an example of the likelihood that ecological experiments often produce results that are limited to a particular time and place, and less general than we might wish to believe.
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.
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.
In the Central Valley of California, native perennial grass species have been largely replaced by Eurasian annual species, while in many parts of the Mediterranean Basin native perennial grasses continue to dominate, even on disturbed or degraded sites. We assessed whether differences in summer rainfall patterns have lead to the development of different plant-water strategies between grasses from these two regions. We compared six measures of plant-water physiology for three guilds of grasses: California perennial grasses, Mediterranean perennial grasses, and Mediterranean annual grasses. Discriminant analysis distinguished between the three guilds; Mediterranean perennial grasses were characterized by a more conservative water-relations physiology than Mediterranean annual grasses, whereas California perennial grasses were in some ways intermediate between the two Mediterranean grass guilds. For individual traits, California perennial grasses were either intermediate or more like Mediterranean annuals than Mediterranean perennials. Our results suggest California perennials are more drought tolerant than Mediterranean annuals but less drought tolerant than Mediterranean perennials, despite the fact that California's Central Valley has a more intense summer drought than the Mediterranean Basin. These patterns may help explain why Mediterranean annuals, but not Mediterranean perennials, have been more successful invaders of interior California grasslands.
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