Smaller species experience mild adversity under shading in an old-field plant community.

Plant competition experiments commonly suggest that larger species have an advantage, primarily in terms of light acquisition. However, within crowded natural vegetation, where competition evidently impacts fitness, most resident species are relatively small. It remains unclear, therefore, whether the size advantage observed in controlled experiments is normally realized in habitats where competition is most intense. We characterized the light environment and tested for evidence of a size advantage in competition for light in an old-field plant community composed of perennial herbaceous species. We investigated whether larger species contributed to reduced light penetration (i.e., greater shading), and examined the impact of shade on smaller species by testing whether their abundance and richness were lower in plots with less light penetration. Light penetration in plots ranged from 0.3% to 72.4%. Significant effects were more common when analyses focused on small plants that reached reproduction (i.e., flowering rooted units); focusing on only flowering plants (i.e., excluding nonflowering rooted units) can clarify community patterns. Plots with a greater mean species height had significantly lower light penetration, and plots with lower light penetration had significantly lower flowering abundance and richness of small species. However, the impact of shade on the flowering abundance and richness of small species was relatively small (R 2 values between 8% and 15%) and depended on how we defined "small species." Synthesis: Our results confirm that light penetration in herbaceous vegetation can be comparable to levels seen in forests, that plots with taller species cast more shade, and that flowering smaller species are less abundant and diverse in plots where light penetration is low. However, variation in mean plot height explained less than 10% of variation in light penetration, and light penetration explained between 5 and 15% of variation in the flowering abundance and richness of small species. Coupled with the fact that flowering small species were present even within the most heavily shaded plots, our results suggest that any advantage in light competition by large species is limited. One explanation is that at least some small species in these communities are shade-tolerant. Shade tolerance in predominantly herbaceous communities, particularly among small plant species, requires further research.

What does body size mean, from the "plant's eye view"?

Alternative metrics exist for representing variation in plant body size, but the vast majority of previous research for herbaceous plants has focused on dry mass. Dry mass provides a reasonably accurate and easily measured estimate for comparing relative capacity to convert solar energy into stored carbon. However, from a "plant's eye view", its experience of its local biotic environment of immediate neighbors (especially when crowded) may be more accurately represented by measures of "space occupancy" (S-O) recorded in situ-rather than dry mass measured after storage in a drying oven. This study investigated relationships between dry mass and alternative metrics of S-O body size for resident plants sampled from natural populations of herbaceous species found in Eastern Ontario. Plant height, maximum lateral canopy extent, and estimated canopy area and volume were recorded in situ (in the field)-and both fresh and dry mass were recorded in the laboratory-for 138 species ranging widely in body size and for 20 plants ranging widely in body size within each of 10 focal species. Dry mass and fresh mass were highly correlated (r2 > .95) and isometric, suggesting that for some studies, between-species (or between-plant) variation in water content may be unimportant and fresh mass can therefore substitute for dry mass. However, several relationships between dry mass and other S-O body size metrics showed allometry-that is, plants with smaller S-O body size had disproportionately less dry mass. In other words, they have higher "body mass density" (BMD) - more dry mass per unit S-O body size. These results have practical importance for experimental design and methodology as well as implications for the interpretation of "reproductive economy"-the capacity to produce offspring at small body sizes-because fecundity and dry mass (produced in the same growing season) typically have a positive, isometric relationship. Accordingly, the allometry between dry mass and S-O body size reported here suggests that plants with smaller S-O body size-because of higher BMD-may produce fewer offspring, but less than proportionately so; in other words, they may produce more offspring per unit of body size space occupancy.

A global meta-analysis of the relative extent of intraspecific trait variation in plant communities.

Recent studies have shown that accounting for intraspecific trait variation (ITV) may better address major questions in community ecology. However, a general picture of the relative extent of ITV compared to interspecific trait variation in plant communities is still missing. Here, we conducted a meta-analysis of the relative extent of ITV within and among plant communities worldwide, using a data set encompassing 629 communities (plots) and 36 functional traits. Overall, ITV accounted for 25% of the total trait variation within communities and 32% of the total trait variation among communities on average. The relative extent of ITV tended to be greater for whole-plant (e.g. plant height) vs. organ-level traits and for leaf chemical (e.g. leaf N and P concentration) vs. leaf morphological (e.g. leaf area and thickness) traits. The relative amount of ITV decreased with increasing species richness and spatial extent, but did not vary with plant growth form or climate. These results highlight global patterns in the relative importance of ITV in plant communities, providing practical guidelines for when researchers should include ITV in trait-based community and ecosystem studies.

Revising traditional theory on the link between plant body size and fitness under competition: evidence from old-field vegetation.

The selection consequences of competition in plants have been traditionally interpreted based on a "size-advantage" hypothesis - that is, under intense crowding/competition from neighbors, natural selection generally favors capacity for a relatively large plant body size. However, this conflicts with abundant data, showing that resident species body size distributions are usually strongly right-skewed at virtually all scales within vegetation. Using surveys within sample plots and a neighbor-removal experiment, we tested: (1) whether resident species that have a larger maximum potential body size (MAX) generally have more successful local individual recruitment, and thus greater local abundance/density (as predicted by the traditional size-advantage hypothesis); and (2) whether there is a general between-species trade-off relationship between MAX and capacity to produce offspring when body size is severely suppressed by crowding/competition - that is, whether resident species with a larger MAX generally also need to reach a larger minimum reproductive threshold size (MIN) before they can reproduce at all. The results showed that MIN had a positive relationship with MAX across resident species, and local density - as well as local density of just reproductive individuals - was generally greater for species with smaller MIN (and hence smaller MAX). In addition, the cleared neighborhoods of larger target species (which had relatively large MIN) generally had - in the following growing season - a lower ratio of conspecific recruitment within these neighborhoods relative to recruitment of other (i.e., smaller) species (which had generally smaller MIN). These data are consistent with an alternative hypothesis based on a 'reproductive-economy-advantage' - that is, superior fitness under competition in plants generally requires not larger potential body size, but rather superior capacity to recruit offspring that are in turn capable of producing grand-offspring - and hence transmitting genes to future generations - despite intense and persistent (cross-generational) crowding/competition from near neighbors. Selection for the latter is expected to favor relatively small minimum reproductive threshold size and hence - as a tradeoff - relatively small (not large) potential body size.

Effects of 'target' plant species body size on neighbourhood species richness and composition in old-field vegetation.

Competition is generally regarded as an important force in organizing the structure of vegetation, and evidence from several experimental studies of species mixtures suggests that larger mature plant size elicits a competitive advantage. However, these findings are at odds with the fact that large and small plant species generally coexist, and relatively smaller species are more common in virtually all plant communities. Here, we use replicates of ten relatively large old-field plant species to explore the competitive impact of target individual size on their surrounding neighbourhoods compared to nearby neighbourhoods of the same size that are not centred by a large target individual. While target individuals of the largest of our test species, Centaurea jacea L., had a strong impact on neighbouring species, in general, target species size was a weak predictor of the number of other resident species growing within its immediate neighbourhood, as well as the number of resident species that were reproductive. Thus, the presence of a large competitor did not restrict the ability of neighbouring species to reproduce. Lastly, target species size did not have any impact on the species size structure of neighbouring species; i.e. they did not restrict smaller, supposedly poorer competitors, from growing and reproducing close by. Taken together, these results provide no support for a size-advantage in competition restricting local species richness or the ability of small species to coexist and successfully reproduce in the immediate neighbourhood of a large species.

Do citations and impact factors relate to the real numbers in publications? A case study of citation rates, impact, and effect sizes in ecology and evolutionary biology.

Metrics of success or impact in academia may do more harm than good. To explore the value of citations, the reported efficacy of treatments in ecology and evolution from close to 1,500 publications was examined. If citation behavior is rationale, i.e. studies that successfully applied a treatment and detected greater biological effects are cited more frequently, then we predict that larger effect sizes increases study relative citation rates. This prediction was not supported. Citations are likely thus a poor proxy for the quantitative merit of a given treatment in ecology and evolutionary biology-unlike evidence-based medicine wherein the success of a drug or treatment on human health is one of the critical attributes. Impact factor of the journal is a broader metric, as one would expect, but it also unrelated to the mean effect sizes for the respective populations of publications. The interpretation by the authors of the treatment effects within each study differed depending on whether the hypothesis was supported or rejected. Significantly larger effect sizes were associated with rejection of a hypothesis. This suggests that only the most rigorous studies reporting negative results are published or that authors set a higher burden of proof in rejecting a hypothesis. The former is likely true to a major extent since only 29 % of the studies rejected the hypotheses tested. These findings indicate that the use of citations to identify important papers in this specific discipline-at least in terms of designing a new experiment or contrasting treatments-is of limited value.

Size distributions and dispersions along a 485-year chronosequence for sand dune vegetation.

Using a sand dune chronosequence that spans 485 years of primary succession, we collected nearest-neighbor vegetation data to test two predictions associated with the traditional "size-advantage" hypothesis for plant competitive ability: (1) the relative representation of larger species should increase in later stages of succession; and (2) resident species that are near neighbors should, over successional time, become more similar in plant body size and/or seed size than expected by random assembly. The first prediction was supported over the time period between mid to later succession, but the second prediction was not; that is, there was no temporal pattern across the chronosequence indicating that either larger resident species, or larger seeded resident species, increasingly exclude smaller ones from local neighborhoods over time. Rather, neighboring species were generally more different from each other in seed sizes than expected by random assembly. As larger species accumulate over time, some relatively small species are lost from later stages of succession, but species size distributions nevertheless remain strongly right-skewed-even in late succession-and species of disparate sizes are just as likely as in early succession to coexist as immediate neighbors. This local-scale coexistence of disparate sized neighbors might be accounted for-as in traditional interpretations-in terms of species differences in "physical-space-niches" (e.g., involving different rooting depths), combined with possible facilitation effects. We propose, however, that this coexistence may also occur because competitive ability involves more than just a size advantage, with traits associated with survival (tolerance of intense competition) and fecundity (offspring production despite intense competition) being at least equally important.

The role of plant species size in invasibility: a field experiment.

Large plant species self-thin to disproportionately lower densities than smaller plant species, and therefore may leave more patches of unused space suitable for invasion. Using experimental monocultures of 11 old-field perennial plant species differing in maximum size, as well as mixtures composed of all monoculture species, we tested our primary hypothesis that monocultures of larger species will be more susceptible to natural invasion. After 3 years, monocultures of larger species were invaded by a significantly greater number of species, and more ramets, from the surrounding vegetation. Invading plant species were significantly smaller than the monoculture species being invaded, suggesting that smaller plant species may be better invaders. Thus, we quantified a trade-off between species size, which is frequently associated with increased competitive ability for light, and invasibility, suggesting one reason why large and small species coexist in virtually all plant communities. Although we expected that invasion would enhance biomass production by more fully capturing available resources, we found that the most highly invaded plots of each species produced significantly less biomass. This suggests that increased diversity resulting from invasion did not result in complementary resource use. Mixture plots containing all experimental species did not admit a significantly different number of invading ramets or species than most monocultures, indicating no obvious role for diversity in resistance to invasion, or complementary resource use. Our results suggest that relatively large species may be limited in their capacity to competitively exclude other, smaller species from communities because pure stands of the former are more susceptible to invasion by the latter.

Does publication in top-tier journals affect reviewer behavior?

We show that when ecologists act as reviewers their reported rejection rates recommended for manuscripts increases with their publication frequency in high impact factor journals. Rejection rate however does not relate to reviewer age. These results indicate that the likelihood of getting a paper accepted for publication may depend upon factors in addition to scientific merit. Multiple reviewer selection for a given manuscript therefore should consider not only appropriate expertise, but also reviewers that have variable publication experience with a range of different journals to ensure balanced treatment. Interestingly since age did not relate to rejection rates, more senior scientists are not necessarily more jaded in reviewing practices.

Systematic variation in reviewer practice according to country and gender in the field of ecology and evolution.

The characteristics of referees and the potential subsequent effects on the peer-review process are an important consideration for science since the integrity of the system depends on the appropriate evaluation of merit. In 2006, we conducted an online survey of 1334 ecologists and evolutionary biologists pertaining to the review process. Respondents were from Europe, North America and other regions of the world, with the majority from English first language countries. Women comprised a third of all respondents, consistent with their representation in the scientific academic community. Among respondents we found no correlation between the time typically taken over a review and the reported average rejection rate. On average, Europeans took longer over reviewing a manuscript than North Americans, and females took longer than males, but reviewed fewer manuscripts. Males recommended rejection of manuscripts more frequently than females, regardless of region. Hence, editors and potential authors should consider alternative sets of criteria, to what exists now, when selecting a panel of referees to potentially balance different tendencies by gender or region.

Double-blind review favours increased representation of female authors.

Double-blind peer review, in which neither author nor reviewer identity are revealed, is rarely practised in ecology or evolution journals. However, in 2001, double-blind review was introduced by the journal Behavioral Ecology. Following this policy change, there was a significant increase in female first-authored papers, a pattern not observed in a very similar journal that provides reviewers with author information. No negative effects could be identified, suggesting that double-blind review should be considered by other journals.

Life history traits in selfing versus outcrossing annuals: exploring the 'time-limitation' hypothesis for the fitness benefit of self-pollination.

BACKGROUND: Most self-pollinating plants are annuals. According to the 'time-limitation' hypothesis, this association between selfing and the annual life cycle has evolved as a consequence of strong r-selection, involving severe time-limitation for completing the life cycle. Under this model, selection from frequent density-independent mortality in ephemeral habitats minimizes time to flower maturation, with selfing as a trade-off, and/or selection minimizes the time between flower maturation and ovule fertilization, in which case selfing has a direct fitness benefit. Predictions arising from this hypothesis were evaluated using phylogenetically-independent contrasts of several life history traits in predominantly selfing versus outcrossing annuals from a data base of 118 species distributed across 14 families. Data for life history traits specifically related to maturation and pollination times were obtained by monitoring the start and completion of different stages of reproductive development in a greenhouse study of selfing and outcrossing annuals from an unbiased sample of 25 species involving five pair-wise family comparisons and four pair-wise genus comparisons. RESULTS: Selfing annuals in general had significantly shorter plant heights, smaller flowers, shorter bud development times, shorter flower longevity and smaller seed sizes compared with their outcrossing annual relatives. Age at first flower did not differ significantly between selfing and outcrossing annuals. CONCLUSIONS: This is the first multi-species study to report these general life-history differences between selfers and outcrossers among annuals exclusively. The results are all explained more parsimoniously by selection associated with time-limitation than by selection associated with pollinator/mate limitation. The shorter bud development time reported here for selfing annuals is predicted explicitly by the time-limitation hypothesis for the fitness benefit of selfing (and not by the alternative 'reproductive assurance' hypothesis associated with pollinator/mate limitation). Support for the time-limitation hypothesis is also evident from published surveys: whereas selfers and outcrossers are about equally represented among annual species as a whole, selfers occur in much higher frequencies among the annual species found in two of the most severely time-limited habitats where flowering plants grow--deserts and cultivated habitats.

Allometric gender allocation in Ambrosia Artemisiifolia (Asteraceae) has adaptive plasticity.

Evidence is reported for size-dependent (allometric) gender allocation in the monoecious, wind-pollinated annual Ambrosia artemissifolia. Consistent with established theory, the pattern of allometry displayed adaptive plasticity, depending on the environmental cause of variation in plant size. Plant size gradients were generated in both field and greenhouse experiments using separate and combined gradients of shading, soil nutrient levels, and neighbor proximity. When plant size constraints involved light limitation from shading (e.g., because of close neighbor proximity), decreasing plant size was generally associated with decreasing maleness and increasing femaleness (based on relative male and female flower production, respectively). This is consistent with the "pollen-dispersal" hypothesis in which the consequences of relatively small plant size (among larger neighbors) imposes less severe limitation for female reproductive success than for male reproductive success (because success as an outcrossing donor of wind-dispersed pollen increases with increasing plant height, especially when neighbors are present). However, when size was constrained by soil nutrient limitation alone (i.e., without shading effects), the results had the converse allometric relationship; i.e., decreasing plant size was generally associated with increasing maleness and decreasing femaleness. This is consistent with the "size-advantage" and "time-limitation" hypotheses in which energetic and time limitations (respectively) associated with relatively small plant size impose a less severe limitation for male reproductive success than for female reproductive success.

Allometry and development in herbaceous plants: functional responses of meristem allocation to light and nutrient availability.

We examined the relationship between meristem allocation and plant size for four annual plant species: Arabidopsis thaliana, Arenaria serphyllifolia, Brassica rapa, and Chaenorrhinum minus. Gradients of light and nutrient availability were used to obtain a range of plant sizes for each of these species. Relative allocation to reproductive, inactive, and growth meristems were used to measure reproductive effort, apical dominance, and branching intensity, respectively. We measured allocation to each of these three meristem fates at weekly intervals throughout development and at final developmental stage. At all developmental stages reproductive effort and branching intensity tended to increase with increasing plant size (i.e., due to increasing resource availability) and apical dominance tended to decrease with increasing plant size. We interpret these responses as a strategy for plants to maximize fitness across a range of environments. In addition, significant differences in meristem response among species may be important in defining the range of habitats in which a species can exist and may help explain patterns of species competition and coexistence in habitats with variable resource availability.

The analysis of contact sampling data.

Use of the chi-square statistic for investigating interspecific association has been incorrectly applied to the contact sampling method because of an invalid probability argument. The usual method of calculating expected frequencies leads to a large overestimation and hence bias towards negative association. A correct method of analysis is presented and discussed.