Rules of the seed size game: contests between large‐seeded and small‐seeded species

The coexistence of multiple seed size strategies within plant communities have been considered puzzling, based on a theoretical expectation of the existence of an optimal seed size under each set of specific environmental conditions. A model aimed at explaining the coexistence of different seed sizes has been suggested, where a seed size – seed number tradeoff is connected to a tradeoff between competition and colonization, leading to a competitive advantage in larger‐seeded species and a colonization advantage in smaller‐seeded species. Recently an alternative model has been suggested, based on a tradeoff between stress tolerance and fecundity, associated with the variation from large to small seeds. Here, we examine the role of seed size for recruitment in two‐species contests subjected to various treatments. In a garden experiment seeds of 14 plant species were combined pair‐wise into seven pairs, each with one larger‐seeded species and one smaller‐seeded species. Each species‐pair was sown with sparse and dense seed densities and subjected to different treatments of shading and litter. Recruitment was recorded during two years. Our results showed a general advantage of larger‐seeded species over smaller‐seeded species. This seed size advantage increased in treatments with litter, whereas there were minor effects of shade, and no effect of seed density was found. We thus found little support for a density dependent seed size game as assumed in models of a competition‐colonization tradeoff, whereas our results fit well with a model based on a tradeoff between stress tolerance and fecundity. Our experiment provides novel empirical data to theoretical models on co‐existence between multiple seed size strategies.     

Exotic grasses and feces deposition by an exotic herbivore combine to reduce the relative abundance of native forbs

Increased resource availability can facilitate establishment of exotic plant species, especially when coincident with propagule supply. Following establishment, increased resource availability may also facilitate the spread of exotic plant species if it enhances their competitive abilities relative to native species. Exotic Canada geese (Branta canadensis) introduce both exotic grass seed and nutrients to an endangered plant community on the Gulf Islands of southwestern British Columbia, Canada. I used greenhouse experiments to assess the competitive advantage of the exotic grasses relative to native and exotic forbs in this community and to test the impacts of nutrient addition from goose feces on competitive outcomes. I grew experimental communities varying in their proportion of forbs versus exotic grasses, and added goose feces as a nutrient source. I found that both native and exotic forbs produced significantly more biomass in competition with conspecifics than in competition with the grasses, and that the proportional abundance of two out of three native forbs was lowest in the combined presence of exotic grasses and nutrient addition. In a second experiment, I found that in monoculture all species of forbs and grasses showed equal growth responses to nutrients. The exotic species did not convert additional nutrients into additional biomass at a higher rate, but did germinate earlier and grow larger than the native species regardless of nutrient availability. This suggests that the exotic species may have achieved their competitive advantage partly by pre-empting resources in community mixtures. Small and late-germinating native forbs may be particularly vulnerable to competitive suppression from exotic grasses and forbs and may be at an even greater disadvantage if their competitors are benefiting from early access to additional nutrients. In combination, the input of exotic propagules and additional nutrients by nesting geese may compromise efforts to maintain native community composition in this system.     

More plant biomass results in more offspring production in annuals,or does it?

Competitive ability in plants has been previously measured almost exclusively in terms of traits related to growth (biomass) or plant size. In this study, however, we used a multi‐species competition experiment with six annuals to measure relative competitive ability in terms of reproductive output, i.e. the number of offspring produced for the next generation. Under greenhouse conditions, plants of each species were started in pots from germinating seeds and were grown singly (free of competition) and at high density in both monocultures and in mixtures with all study species. Several traits traditionally regarded as determinants of competitive ability in plants were recorded for each species grown singly, including: seed mass, germination time, early growth rate and potential plant size (biomass and height). Under competition, several traits were recorded as indicators of relative performance in both monocultures and mixtures, including: biomass of survivors, total number of survivors, number of reproductive survivors, and reproductive output (total seed production) of the survivors. As expected, species that grew to a larger biomass in isolation had higher seed production in isolation. However, none of the traditional plant growth/size‐related traits, measured either in isolation or under competition, could predict between species variation in reproductive output under competition in either monocultures or mixtures. In mixtures, 97% of this variation in reproductive output could be explained by between‐species variation in the number of reproductive survivors. The results indicate that traits measured on plants grown singly may be poor predictors of reproductive output under competition, and that species’ rank order of competitive ability in terms of the biomass of survivors may bear no relationship to their rank order in terms of the number of offspring produced by these survivors. This has important implications for the interpretation of mechanisms of species coexistence and community assembly within vegetation.     

A shift from exploitation to interference competition with increasing density affects population and community dynamics

Intraspecific competition influences population and community dynamics and occurs via two mechanisms. Exploitative competition is an indirect effect that occurs through use of a shared resource and depends on resource availability. Interference competition occurs by obstructing access to a resource and may not depend on resource availability. Our study tested whether the strength of interference competition changes with protozoa population density. We grew experimental microcosms of protozoa and bacteria under different combinations of protozoan density and basal resource availability. We then solved a dynamic predator–prey model for parameters of the functional response using population growth rates measured in our experiment. As population density increased, competition shifted from exploitation to interference, and competition was less dependent on resource levels. Surprisingly, the effect of resources was weakest when competition was the most intense. We found that at low population densities, competition was largely exploitative and resource availability had a large effect on population growth rates, but the effect of resources was much weaker at high densities. This shift in competitive mechanism could have implications for interspecific competition, trophic interactions, community diversity, and natural selection. We also tested whether this shift in the mechanism of competition with protozoa density affected the structure of the bacterial prey community. We found that both resources and protozoa density affected the structure of the bacterial prey community, suggesting that competitive mechanism may also affect trophic interactions.     

An experimental test of the relationship between seed size and competitive ability in annual plants

Competition–colonization tradeoff models explain the coexistence of competing species in terms of a tradeoff between competitive ability and colonization ability. One class of such models is based on the idea that seed size determines competitive ability, seed number determines colonization ability, and the two traits are negatively correlated such that higher competitive ability of large‐seeded species compensates for their smaller seed number. According to such models, species inhabiting the same community should show a distinct ranking of competitive ability and this ranking should be correlated with seed size. We tested these predictions using a greenhouse competition experiment focusing on 25 annual species that coexist in sandy habitats of the Mediterranean region in Israel. Rankings of species based on their competitive effects on two independent phytometers were positively correlated. Corresponding rankings based on competitive responses were also correlated. Rankings based on competitive effects were correlated with those based on competitive responses. Yet, in spite of the clear hierarchy in all measures of competitive ability, none of the measures was correlated with seed size. While lack of correlation between seed size and competitive ability has been documented in some systems, our study is the first time that absence of such correlation is documented in a system where the existence of competition is well established and the component species show a clear hierarchy of competitive ability.     

Niches, rather than neutrality, structure a grassland pioneer guild

Pioneer species are fast-growing, short-lived gap exploiters. They are prime candidates for neutral dynamics because they contain ecologically similar species whose low adult density is likely to cause widespread recruitment limitation, which slows competitive dynamics. However, many pioneer guilds appear to be differentiated according to seed size. In this paper, we compare predictions from a neutral model of community structure with three niche-based models in which trade-offs involving seed size form the basis of niche differentiation. We test these predictions using sowing experiments with a guild of seven pioneer species from chalk grassland. We find strong evidence for niche structure based on seed size: specifically large-seeded species produce fewer seeds but have a greater chance of establishing on a per-seed basis. Their advantage in establishment arises because there are more microsites suitable for their germination and early establishment and not directly through competition with other seedlings. In fact, seedling densities of all species were equally suppressed by the addition of competitors' seeds. By the adult stage, despite using very high sowing densities, there were no detectable effects of interspecific competition on any species. The lack of interspecific effects indicates that niche differentiation, rather than neutrality, prevails.     

Fertilization affects the establishment ability of species differing in seed mass via direct nutrient addition and indirect competition effects

Fertilization causes species loss and species dominance changes in plant communities worldwide. However, it still remains unclear how fertilization acts upon species functional traits, e.g. seed mass. Seed mass is a key trait of the regeneration strategy of plants, which influences a range of processes during the seedling establishment phase. Fertilization may select upon seed mass, either directly by increased nutrient availability or indirectly by increased competition. Since previous research has mainly analyzed the indirect effects of fertilization, we disentangled the direct and indirect effects to examine how nutrient availability and competition influence the seed mass relationships on four key components during seedling establishment: seedling emergence, time of seedling emergence, seedling survival and seedling growth. We conducted a common garden experiment with 22 dry grassland species with a two‐way full factorial design that simulated additional nutrient supply and increased competition. While we found no evidence that fertilization either directly by additional nutrient supply or indirectly by increased competition alters the relationship between seed mass and (time of) seedling emergence, we revealed that large seed mass is beneficial under nutrient‐poor conditions (seedlings have greater chances of survival, particularly in nutrient‐poor soils) as well as under competition (large‐seeded species produced larger seedlings, which suffered less from competition than small‐seeded species). Based on these findings, we argue that both factors, i.e. nutrient availability and competition intensity, ought to be considered to understand how fertilization influences seedling establishment and species composition with respect to seed mass in natural communities. We propose a simple conceptual model, in which seed mass in natural communities is determined by competition intensity and nutrient availability. Here, we hypothesize that seed mass shows a U‐shaped pattern along gradients of soil fertility, which may explain the contrasting soil fertility‐seed mass relationships found in the recent literature.     

Enhancing Native Forb Establishment and Persistence Using a Rich Seed Mixture

Establishing native forbs is crucial for invasive plant management and restoring a desirable plant community. Our objectives were to determine (1) if increasing forb seed density results in increased forb establishment; (2) if a species‐rich mixture of forbs has greater establishment and survivorship than a single species; and (3) if mixtures of forbs are more competitive with Spotted knapweed (Centaurea maculosa) than a forb monoculture. To test our first two objectives, we seeded monocultures of Purple coneflower (Echinacea angustifolia), Arrowleaf balsamroot (Balsamorhiza sagittata), Annual sunflower (Helianthus annuus), Dotted gayfeather (Liatris punctata), Western white yarrow (Achillea millefolium), Sticky geranium (Geranium viscosissimum), as well as a mixture of all forbs. Pots were seeded at 800 or 2,000 seeds/m² and watered twice or thrice weekly. The highest seed density produced the highest plant density, which averaged 4.35 plants/pot. The density of the mixture was similar to the mean density seen for individual species, and it doubled in response to the highest seed density. To test our third objective, Spotted knapweed and Purple coneflower were arranged in an addition series matrix with a maximum total density of 4,000 seeds/pot. We found that the forb mixture was seven times more competitive with Spotted knapweed than Purple coneflower alone. Using a mixture of forbs rather than a single species enhances forb establishment in various and unpredictable environments because the mixture possesses a variety of traits that may match year–year and site–site conditions. Once established, the mixture may have a greater chance of persisting than a monoculture.     

Seed Density Significantly Affects Species Richness and Composition in Experimental Plant Communities

Studies on the importance of seed arrival for community richness and composition have not considered the number of seeds arriving and its effect on species richness and composition of natural communities is thus unknown. A series of experimental dry grassland communities were established. All communities were composed of the same 44 species in exactly the same proportions on two substrates using three different seed densities.The results showed that seed density had an effect on species richness only at the beginning of the experiment. In contrast, the effects on species composition persisted across the entire study period. The results do not support the prediction that due to higher competition for light in nutrient-rich soil, species richness will be the highest in the treatment with the lowest seed density. However, the prevalence of small plants in the lowest seed density supported the expectation that low seed density guarantees low competition under high soil nutrients. In the nutrient-poor soil, species richness was the highest at the medium seed density, indicating that species richness reflects the balance between competition and limitations caused by the availability of propagules or their ability to establish themselves. This medium seed density treatment also contained the smallest plants.The results demonstrate that future seed addition experiments need to consider the amount of seed added so that it reflects the amount of seed that is naturally found in the field. Differences in seed density, mimicking different intensity of the seed rain may also explain differences in the composition of natural communities that cannot be attributed to habitat conditions. The results also have important implications for studies regarding the consequences of habitat fragmentation suggesting that increasing fragmentation may change species compositions not only due to different dispersal abilities but also due to differential response of plants to overall seed density.     

Functional traits determine tree growth and ecosystem productivity of a tropical montane forest: Insights from a long‐term nutrient manipulation experiment

Trait‐response effects are critical to forecast community structure and biomass production in highly diverse tropical forests. Ecological theory and few observation studies indicate that trees with acquisitive functional traits would respond more strongly to higher resource availability than those with conservative traits. We assessed how long‐term tree growth in experimental nutrient addition plots (N, P, and N + P) varied as a function of morphological traits, tree size, and species identity. We also evaluated how trait‐based responses affected stand scale biomass production considering the community structure. We found that tree growth depended on interactions between functional traits and the type or combination of nutrients added. Common species with acquisitive functional traits responded more strongly to nutrient addition, mainly to N + P. Phosphorous enhanced the growth rates of species with acquisitive and conservative traits, had mostly positive effects on common species and neutral or negative effects in rare species. Moreover, trees receiving N + P grew faster irrespective of their initial size relative to trees in control or to trees in other treatment plots. Finally, species responses were highly idiosyncratic suggesting that community processes including competition and niche dimensionality may be altered under increased resource availability. We found no statistically significant effects of nutrient additions on aboveground biomass productivity because acquisitive species had a limited potential to increase their biomass, possibly due to their generally lower wood density. In contrast, P addition increased the growth rates of species characterized by more conservative resource strategies (with higher wood density) that were poorly represented in the plant community. We provide the first long‐term experimental evidence that trait‐based responses, community structure, and community processes modulate the effects of increased nutrient availability on biomass productivity in a tropical forest.     

Species richness and evenness respond in a different manner to propagule density in developing prairie microcosm communities

Diversity has two basic components: richness, or number of species in a given area, and evenness, or how relative abundance or biomass is distributed among species. Previously, we found that richness and evenness can be negatively related across plant communities and that evenness can account for more variation in Shannon’s diversity index (H′) than richness, which suggests that relationships among diversity components can be complex. Non-positive relationships between evenness and richness could arise due to the effects of migration rate or local species interactions, and relationships could vary depending on how these two processes structure local communities. Here we test whether diversity components are equally or differentially affected over time by changes in seed density (and associated effects on established plant density and competition) in greenhouse communities during the very early stages of community establishment. In our greenhouse experiment, we seeded prairie microcosms filled with bare field soil at three densities with draws from a mix of 22 grass and forb species to test if increased competition intensity or seedling density would affect the relationships among diversity components during early community establishment. Increased seed density treatments caused diversity components to respond in a different manner and to have different relationships with time. Richness increased linearly with seed density early in the experiment when seedling emergence was high, but was unrelated to density later in the experiment. Evenness decreased log-linearly with seed densities on all sampling dates due to a greater dominance by Rudbeckia hirta with higher densities. Early in the experiment, diversity indices weakly reflected differences in richness, but later, after the competitive effects of Rudbeckia hirta became more intense, diversity indices more strongly reflected differences in evenness. This suggests that species evenness and diversity indices do not always positively covary with richness. Based on these results, we suggest that evenness and richness can be influenced by different processes, with richness being more influenced by the number of emerging seedlings and evenness more by species interactions like competition. These results suggest that both diversity components should be measured in plant diversity studies whenever it is possible.     

Aggregated seed arrival alters plant diversity in grassland communities

Aims Species aggregation is commonly seen in plant communities and may increase diversity by causing intraspecific competition to exceed interspecific competition. One potential source of this spatial aggregation is seed dispersal but it is unclear to what extent aggregated seed distributions affect plant diversity in real communities. Using a field experiment, I tested whether uniform or aggregated seed arrival alters community structure and whether these effects vary with sowing density.Methods The experiment consisted of two spatial seeding treatments (uniform and aggregated) that were fully crossed with three seed density treatments. Sixty, 3 × 4-m plots were arrayed in a low-diversity grassland located in Kansas, USA. Each plot was divided into forty-eight, 0.5 × 0.5-m patches. For aggregated seeding treatments, each of the 15 species was sown into three randomly selected patches within the plot (3×15 = 45). To create a uniform species arrival but control for the seed addition method, all 15 species were sown into 45 individual patches (with three patches remaining unsown) within each plot. Seed mass for each species was held constant at the plot scale between uniform or aggregated treatments within a given level of the sowing density treatment. After two growing seasons, plant density was quantified for all sown species in 15 randomly selected patches from each plot.Important findings I found evidence for shifts in community structure in response to the different spatial seeding patterns. The evenness of added species was higher under aggregated than uniform sowing patterns. There was no detectable effect of aggregated seed sowing on species richness at 3.75 m 2 scale. However, when species richness was extrapolated to larger scales (11.25 m 2), aggregated sowing was predicted to have greater richness than uniform sowing. Effects of seed aggregation on community structure were apparent only at moderate to high sowing rates, yet the latter are within the range of measured seed dispersal in similar grasslands. Additionally, as sowing density increased, seed mass became an increasingly effective predictor of relative abundances for added species, but only under uniform sowing patterns supporting the idea that aggregated dispersal may buffer weaker (smaller seeded) species from competition during colonization. This is the first experiment to show that aggregated seed dispersal patterns can increase at least some components of plant diversity in undisturbed grasslands and suggests that previous seed dispersal experiments, which utilize uniform seed sowing, may underestimate the potential effect of dispersal on plant community structure.     

Does the seed size/number trade‐off model determine plant community structure? An assessment of the model mechanisms and their generality

This paper examines four key mechanisms of the seed size/number trade-off (SSNT) models to assess their relevance to a general understanding of plant community structure. Mechanism 1 is that large seeds have a greater probability of winning in competition against smaller seeds. I provide interspecific experimental evidence that there is a competitive hierarchy among seedlings based on seed size. Mechanism 2 is that a trade-off exists between the number and size of seeds produced for a given reproductive allocation. Negative correlations between seed size and number were found consistently across a range of species from a range of habitats, from published literature. Mechanism 3, that seedling-seedling competition is an important influence on species composition, was found to exist potentially in a range of environments, including annual-dominated, post-fire and gap-dynamic communities. However, there is little quantitative evidence available and this is likely to be a restrictive mechanism. Mechanism 4, that small seeds are superior colonists due to their greater number, was tested in a field experiment in a calcareous grassland community. No supporting evidence was found, suggesting that the SSNT is not an important determinant of structure in this community. Thus two of the four mechanisms can be considered to hold true generally, while the third mechanism may be valid in particular environments. The fourth mechanism did not apply in the community tested, but could be tested in a wider range of communities.     

Influence of seed size on performance of non-native annual plant species in a novel community at two planting densities

Climate warming enables plant species to migrate to higher latitudes and altitudes. Within Europe, the Mediterranean harbours many species that might expand their ranges towards Western Europe. Small seed size may facilitate dispersal, however, it may impair establishment of the range-expanding plant species in the novel vegetation. In a greenhouse experiment, we examined effects of average seed size of Mediterranean plant species on their establishment in a mixed community of Western European plant species. Applying two levels of densities of the natives and a herbivory treatment, we tested how seed size is linked to response in plant growth and fitness in novel vegetation. While all non-native plant species showed a negative response to increased planting density, species with small seeds showed a less negative response. This effect persisted under herbivory. Our data suggest that small-seeded non-native plant species may tolerate competitive pressure from novel plant communities better than large-seeded species, so that small seed size may confer a higher probability of establishment of non-native species in novel communities.     

Abiotic and biotic resistance to grass invasion in serpentine annual plant communities

Biological invasions severely impact native plant communities, causing dramatic shifts in species composition and the restriction of native species to spatially isolated refuges. Competition from resident species and the interaction between resource limitation and competition have been overlooked as mechanisms of community resistance in refugia habitats. We examined the importance of these factors in determining the resistance of California serpentine plant communities to invasion by three common European grasses, Avena barbata, Bromus diandrus, and Hordeum murinum. We added seeds of each of these grasses to plots subjected to six levels of resource addition (N, P, Ca, H₂O, all resources together, and a no-addition control) and two levels of competition (with resident community present or removed). Resource limitation and competition had strong effects on the biomass and reproduction of the three invaders. The addition of all resources together combined with the removal of the resident community yielded individual plants that were fourfold to 20-fold larger and sixfold to 20-fold more fecund than plants from control plots. Competitor removal alone yielded invaders that were twofold to sevenfold larger and twofold to ninefold more fecund. N addition alone or in combination with other resources led to a twofold to ninefold increase in the biomass and fecundity of the invaders. No other resource alone significantly affected native or invader performance, suggesting that N was the key limiting resource during our experiment. We found a significant interaction between abiotic and biotic resistance for Bromus, which experienced increased competitive suppression in fertilized plots. The threefold increase in resident biomass with N addition was likely responsible for this result. Our results confirm that serpentine plant communities are severely N limited, which, in combination with competition from resident species, promotes the resistance of these systems to invasions. Our work suggests that better understanding the relative sensitivities of invaders and residents to the physical environment is critical to predicting how abiotic and biotic factors interact to determine community resistance.     

A trade off in the absolute magnitudes of seed and microsite limitations and their effects on population regulation

Microsite limitation (ML) occurs when the number of suitable sites for development determines population size. Seed limitation (SL) arises when seeds cannot occupy all microsites. Both limitations have been measured on relative scales, but absolute values would allow analyzing the relationship between them and their contributions to population regulation. A tradeoff is expected because small‐seeded species are prolific and thus could have low SL, whereas large seeds perform better under different conditions, lowering ML. We assessed whether there is such tradeoff, and if it is mediated by seed size. We measured the absolute magnitudes of limitations through seed addition experiments, and tested whether they determine population density in 14 herbs with different seed sizes in a semiarid grassland. All species were limited by microsites and seeds. The tradeoff occurred only during germination, but was seemingly not mediated by seed size. Annual species were more limited by microsites than perennials, which suffered more SL. Population densities were smaller for species with greater ML measured over the life cycle, but marginally so when ML was measured during germination. SL only reduced annual‐plant density. These negative correlations indicate that our measurements correspond to actual limitations. Microsites limited populations depending on the process analyzed (germination, survival and whole life‐cycle) and the life history of species, suggesting that an integral approach to ML is required rather than focusing only on germination. Comparing annuals and perennials, we found that the strongest limitation is the main determinant of population size. The tradeoff was consistent with seed size and competition–colonization tradeoff, but our results were inconclusive. The SL–ML tradeoff seems to arise from still‐unidentified evolutionary constraints, and may contribute to the maintenance of species diversity, preventing any single species from becoming dominant and excluding its neighbors.     

Fruit size preference in the New Zealand pigeon (Hemiphaga novaeseelandiae)

Abstract We investigated whether the New Zealand pigeon Hemiphaga novaeseelandiae (Columbidae) exhibits size‐based preferences for fruits. We tested the hypothesis that in small‐fruited species, pigeons would prefer larger fruits, but in larger‐fruited species, this preference would reverse as the pigeons become increasingly limited by their gape size. We collected undispersed fruits and bird‐dispersed seeds of 10 plant species, some over several sites or years (13 datasets in total). We estimated the fruit size of dispersed seeds by fitting regressions of fruit diameter to seed diameter in intact fruits. We were able to predict fruit diameter from seed diameter in 12 of the 13 populations, although the relationship was stronger in single‐seeded species than in multi‐seeded species. Seven of the 12 populations tested showed a significant difference in seed diameter among undispersed and dispersed seeds. However, our results showed no consistent pattern in fruit size preference by the New Zealand pigeon and did not support our hypothesis. The large‐bodied New Zealand pigeon is generally not gape limited and fruit size preferences appear to be independent of mean fruit size.     

Seed size, seedling morphology, and response to deep shade and damage in neotropical rain forest trees

To investigate the existence of coordinated sets of seedling traits adapted to contrasting establishment conditions, we examined evolutionary convergence in seedling traits for 299 French Guianan woody plant species and the stress response in a shadehouse of species representing seed size gradients within five major cotyledon morphology types. The French Guianan woody plant community has larger seeds than other tropical forest communities and the largest proportion of hypogeal cotyledon type (59.2%) reported for tropical forests. Yet the community includes many species with intermediate size seeds that produce seedlings with different cotyledonal morphologies. A split-plot factorial design with two light levels (0.8% and 16.1% PAR) and four damage treatments (control, seed damage, leaf damage, stem damage) was used in the shadehouse experiment. Although larger-seeded species had higher survival and slower growth, these patterns were better explained by cotyledon type than by seed mass. Even larger-seeded species with foliar cotyledons grew faster than species with reserve-type cotyledons, and survival after stem grazing was five times higher in seedlings with hypogeal cotyledons than with epigeal cotyledons. Thus, to predict seedling performance using seed size, seedling morphology must also be considered.     

Natural selection on body size is mediated by multiple interacting factors: a comparison of beetle populations varying naturally and experimentally in body size

Body size varies considerably among species and among populations within species, exhibiting many repeatable patterns. However, which sources of selection generate geographic patterns, and which components of fitness mediate evolution of body size, are not well understood. For many animals, resource quality and intraspecific competition may mediate selection on body size producing large-scale geographic patterns. In two sequential experiments, we examine how variation in larval competition and resource quality (seed size) affects the fitness consequences of variation in body size in a scramble-competing seed-feeding beetle, Stator limbatus. Specifically, we compared fitness components among three natural populations of S. limbatus that vary in body size, and then among three lineages of beetles derived from a single base population artificially selected to vary in size, all reared on three sizes of seeds at variable larval density. The effects of larval competition and seed size on larval survival and development time were similar for larger versus smaller beetles. However, larger-bodied beetles suffered a greater reduction in adult body mass with decreasing seed size and increasing larval density; the relative advantage of being large decreased with decreasing seed size and increasing larval density. There were highly significant interactions between the effects of seed size and larval density on body size, and a significant three-way interaction (population-by-density-by-seed size), indicating that environmental effects on the fitness consequences of being large are nonadditive. Our study demonstrates how multiple ecological variables (resource availability and resource competition) interact to affect organismal fitness components, and that such interactions can mediate natural selection on body size. Studying individual factors influencing selection on body size may lead to misleading results given the potential for nonlinear interactions among selective agents.     

Females of the weevil Zabrotes subfasciatus manipulate the size and number of eggs according to the host seed availability

Abstract. In some insects, the finding of oviposition substrate triggers the uptake into oocytes of yolk proteins that are stored in the fat body during post‐embryonic development. The main host of the bean weevil Zabrotes subfasciatus (Coleoptera; Chrysomelidae; Bruchinae; Amblycerini), in which larval resources are the sole source for future egg maturation, is Phaseolus vulgaris. Despite not feeding as adults, females of this species are able to lay eggs after encountering host seeds but it is not known how females react to changes in the availability of bean seeds. In the present study, the behaviour of Z. subfasciatus facing two very different environments for oviposition is investigated, as well as how this influences offspring fitness. The results obtained show that females of Z. subfasciatus react to variations in the availability of seeds belonging to the same host species by adjusting egg size and number. Females on low bean seed density lay larger and fewer eggs than those on high bean seed density, demonstrating a trade‐off between these reproductive traits. Moreover, females can adjust egg size to changing levels of host availability during the first 4 days of their oviposition period. Although no difference in offspring weight is found, those from small eggs (low competition environment) result in larger adults. No response to selection on these traits after rearing beetles on the same host for 40 generations is observed. This unresponsiveness may indicate that beetle populations behave according to their reaction norm that already allows rapid adaptation to a varying amount of host‐seed availability and better exploitation of the environments of this widespread stored‐seed pest.