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Dynamic energy budget (DEB) models provide a mechanistic description of life-histories in terms of fluxes of energy through biological processes. In these models, life-histories are a function of environmental conditions and of fundamental traits of the organism relating to the acquisition, allocation, and use of energy.These traits are described by the parameters of the DEB model, which have been estimated for over 2500 species. Recent work has aimed to compare species on the basis of differences in DEB parameters.We show that caution is required in such analyses, because (i) parameter estimates vary considerably as an artefact of the types of data used to fit the models, and (ii) there is substantial intraspecific variation in parameter values, reflecting biological differences among populations.We show that similar patterns of growth and reproduction can be reproduced with very different parameter sets. Our results imply that direct comparison of DEB parameters across populations or species may be invalid. However, valid comparisons are possible if differences in the types of data used to fit the models are taken into account.We estimated DEB parameters for 16 populations of Trinidadian guppy, identifying differences in resource allocation and metabolic rate consistent with evolved life-history differences among these populations.Variation in parameter values was substantial: if intraspecific variation in DEB parameters is greater than currently measured levels of interspecific variation, the detection of broad-scale patterns in energy budgets across species will be challenging.

In many animal species, individuals engage in fights with conspecifics over access to limited resources (e.g. mates, food, or shelter). Most theory about these intraspecific fights assumes that damage has an important role in determining the contest winner. Thus, defensive structures that reduce the amount of damage an individual accrues during intraspecific competition should provide a fighting advantage.Examples of such damage-reducing structures include the dermal shields of goats, the dorsal osteoderms of crocodiles, and the armored telsons of mantis shrimps. Although numerous studies have identified these defensive structures, no study has investigated whether they influence the outcomes of intraspecific fights.Here, we investigated whether inhibiting damage by enhancing an individual’s armor influenced fighting behavior and success in the giant mesquite bug,Thasus neocalifornicus(Insecta: Hemiptera: Coreidae).We found that experimentally manipulated individuals (i.e. those provided with additional armor) were 1.6 times more likely to win a fight when compared to the control. These results demonstrate that damage, and damage-reducing structures, can influence fighting success.The implications of these results are twofold. First, our results experimentally support a fundamental assumption of most theoretical fighting models: that damage is a fighting cost that can influence contest outcomes. Second, these results highlight the importance of an individual’s defensive capacity, and why defense should not be ignored.

Species contributing high proportions to community biomass strongly influence ecosystem processes within the community. Studies have shown that dominant species may serve as nurse plants, helping to ensure biomass stability of the subordinate species under stress conditions. The question is widely debated as to whether either niche differentiation or neutral processes drive the net outcome of plant interactions within a subordinate plant community. To answer this question, requires precise estimates of individual variation in functional traits and performance. In a 5‐year mesocosm experiment, the functional responses of a subordinate plant community to the removal of the dominant species were evaluated across two drought‐stress scenarios. Small‐scale (i.e. large pots) wetland communities were constructed comprising one dominant species Carex elata and three subordinate species. Removal of the dominant species allowed evaluation of the net effects of drought and interspecific interactions. We estimated the functional divergences for three traits (SLA, leaf dry matter content and allocation to height growth) and compared these with performance differences quantified individually. This enabled distinctions to be made between deterministic (i.e. niche differentiation) and neutral processes driving the drought response of the subordinate community. We showed that the dominant species decreased relative performance differences within the subordinate plant community under conditions of permanent drought stress. These changes were associated with the convergence of traits related to resource acquisition and growth. The dominant species equalized species performance differences by supressing relatively drought‐tolerant species with low competitive ability and by supporting the less drought‐tolerant species with relatively high competitive ability. Meanwhile, under conditions of inter‐annual drought, the subordinate species likely coexisted due to differentiation in resource‐use strategies and the interaction with the dominant species. Inclusion of individual variation in performance with a functional trait approach provides valuable insights into the processes structuring plant communities. Ours is the first study to provide evidence that subordinate species exposed to drought may coexist via neutral processes arising from their interactions with the dominant species, leading to functional convergence of traits associated with the trade‐off between stress tolerance and competitive ability. A free Plain Language Summary can be found within the Supporting Information of this article.

Plant phenology was assessed as one of the most key Essential Biodiversity Variables on species functional traits, which underpins a variety of aspects of ecosystem function and is a vital determinant of species distribution. Flowering phenology is one of the most sensitive indicators of the effects of recent climatic change. Understanding of intraspecific variation in phenological responses to climate change can contribute to the accurate prediction of phenology and the assessment of potential impacts of climate change on organisms. However, this intraspecific variation in phenological sensitivity in natural systems is still unclear, especially at large spatial scales. Using the widespread perennial herb Spiranthes sinensis, we quantified intraspecific variation in phenological responses to warming among groups located along climatic gradients. Phenological data were derived from 1,681 collections of herbarium specimens and photographs across eastern Asia and southeastern Australia over the period 1902–2017 and spans eight humid and eight non‐humid regions. We found that peak flowering dates of Spiranthe sinensis (FLss) showed substantial intraspecific variation in temperature sensitivity, ranging from −5.16 ± 1.13 to 4.16 ± 0.90 days/°C. Across all 16 regions, FLss exhibited the strongest delayed response to warming in southeastern Australia. Temperature sensitivity of the FLss was significantly correlated with annual precipitation and aridity index of a region, suggesting a possible role of declined water availability in reducing temperature sensitivity. Notably, contrasting phenological responses to warming were found between collections with different humidity regimes: humid regions generally flowered earlier (M: −3.57 days/°C) and non‐humid regions exhibited slightly delayed flowering (M: 1.25 days/°C). These findings provide empirical evidence of substantial intraspecific variation in phenological sensitivity across large spatial scales, and highlights that soil water availability may play a key role in a warming planet, especially in non‐humid regions.

By providing food, living space and reducing biotic and abiotic stress, habitat‐formers support diverse communities. The characteristics that mediate their value to associated organisms, however, are not always clear. Traditional assessments of habitat‐former and associated community relationships use broad‐scale classifications to group habitat‐formers according to identity or gross morphology. Trait‐based approaches instead measure traits at the level of the individual and in doing so allow for predictions that are not constrained to taxonomic groups. By measuring at the level of the individual, trait‐based approaches have the potential to capture not only interspecific variability, but also the intraspecific variation that is often ignored by traditional approaches. Trait‐based approaches may therefore provide a novel way to link the characteristics of habitat‐formers to ecosystem functioning. Here we quantified inter‐ and intraspecific variation in six macroalgal species and tested the relative importance of morphological traits and species identity in predicting the abundance of associated fauna. All species of macroalgae displayed substantial intraspecific morphological variation. Morphological traits were more important than species identity in predicting epifaunal abundances. Total surface area was the most important trait when predicting total abundance per thallus, whilst mean frond length was the most important predictor for abundance per gram algal biomass. This suggests that traits that describe the quantity of available habitat were the most important predictors of community abundance, but the shape or architecture of the habitat also contributes to structuring these communities. Our results indicate that trait‐based approaches may be more useful than traditional species‐based approaches when predicting the abundances of fauna associated with habitat‐forming species. A free Plain Language Summary can be found within the Supporting Information of this article.

1. Despite broad recognition that water is a major limiting factor in arid ecosystems, we lack an empirical understanding of how this resource is shared and distributed among neighbouring plants. Intraspecific variability can further contribute to this variation via divergent life-history traits, including root architecture. We investigated these questions in the shrub Artemisia tridentata and hypothesized that the ability to access and utilize surface water varies among subspecies and cytotypes. 2. We used an isotope tracer to quantify below-ground zone of influence in A. tridentata, and tested whether spatial neighbourhood characteristics can alter plant water uptake. We introduced deuterium-enriched water to the soil in plant interspaces in a common garden experiment and measured deuterium composition of plant stems. We then applied spatially explicit models to test for differential water uptake by A. tridentata, including intermingled populations of three subspecies and two ploidy levels. 3. The results suggest that lateral root functioning in A. tridentata is associated with intraspecific identity and ploidy level. Subspecies adapted to habitats with deep soils generally had a smaller horizontal reach, and polyploid cytotypes were associated with greater water uptake compared to their diploid variants. We also found that plant crown volume was a weak predictor of water uptake, and that neighbourhood crowding had no discernable effect on water uptake. 4. Intraspecific variation in lateral root functioning can lead to differential patterns of resource acquisition, an essential process in arid ecosystems in the contexts of changing climate and seasonal patterns of precipitation. Altogether, we found that lateral root development and activity are more strongly related to genetic variability within A. tridentata than to plant size. Our study highlights how intraspecific variation in life strategies is linked to mechanisms of resource acquisition.

The widespread observation that heat tolerance is less variable than cold tolerance (‘cold-tolerance asymmetry’) leads to the prediction that species exposed to temperatures near their thermal maxima should have reduced evolutionary potential for adapting to climate warming. However, the prediction is largely supported by species-level global studies based on single estimates of both physiological metrics per taxon. We ask whether cold-tolerance asymmetry holds for Iberian lizards after accounting for intraspecific variation in critical thermal maxima (CT) and minima (CT). To do so, we quantified CT and CT for 58 populations of 15 Iberian lizard species (299 individuals). Then, we randomly selected one population from each study species (population sample = 15 CT and CT values), tested for differences between the variance of both thermal metrics across species, and repeated the test for thousands of population samples as if we had undertaken the same study thousands of times, each time sampling one different population per species (as implemented in global studies). The ratio of variances in CT to CT across species varied up to 16-fold depending on the populations chosen. Variance ratios show how much CT departs from the cross-species mean compared to CT, with a unitary ratio indicating equal variance of both thermal limits. Sampling one population per species was six times more likely to result in the observation of greater CT variance (‘heat-tolerance asymmetry’) than cold-tolerance asymmetry. The probability of obtaining the data (given the null hypothesis of equal variance being true) was twice as likely for cases of cold-tolerance asymmetry than for the opposite scenario. Range-wide, population-level studies that quantify heat and cold tolerance of individual species are urgently needed to ascertain the global prevalence of cold-tolerance asymmetry. While broad latitudinal clines of cold tolerance have been strongly supported, heat tolerance might respond to smaller-scale climatic and habitat factors hence go unnoticed in global studies. Studies investigating physiological responses to climate change should incorporate the extent to which thermal traits are characteristic of individuals, populations and/or species. A free Plain Language Summary can be found within the Supporting Information of this article., Funding from British Ecological Society ‘Research Grant’ 4496-5470 to S.H.-P., and CGL2011-26852 project of the Spanish Ministry of Economy and Competitiveness and IC&DT 1/SAESCTN/ALENT-07-0224-FEDER-001755 to M.B.A.

There is increasing evidence that herbivore–plant interactions on a focal plant species are influenced by interspecific neighbourhood effects via neighbouring plants (i.e. an associational effect). However, intraspecific neighborhood effects imposed by plant traits have been less appreciated. Specifically, the significance of intraspecific neighbourhood effects in population‐level consequences of plants has been totally overlooked. Using two varieties of Nicotiana tabacum (high‐ and low‐nicotine), we evaluated the neighbourhood effects based on patch‐level interactions in a split‐plot 3 × 3 factorial experiment that manipulated number of plants (4, 9 and 16 plants) and culture type (monoculture plots with high‐ and low‐nicotine plants, and polyculture plot) in an experimental garden. We found that herbivore visits on plants varied depending on the number of plants per patch and culture type. Presence of more high‐nicotine plants decreased herbivore visits in the four plant plots, and presence of high‐nicotine plants in the nine plant plots decreased herbivore visits on both high‐ and low‐nicotine plants. In contrast, in the 16 plant plots, herbivore visits on high‐nicotine plants in polyculture plots were lower than others, including those on high‐nicotine plants in monoculture plots. Our findings clearly demonstrated that the intraspecific neighbourhood effect could occur depending on the aggregation of highly defended plants (i.e. high density and/or plant‐spacing). This study suggests that multiple mechanisms for the neighbourhood effect simultaneously worked, depending on the patch size and composition of defensive traits of individual plants, and that intraspecific neighbourhood effects may influence population‐level consequences for plant–herbivore interaction. A free Plain Language Summary can be found within the Supporting Information of this article.

1. In biodiversity and ecosystem functioning research, the consideration of intraspecific trait variation and multiple trophic levels - both largely overlooked so far - has been proposed as a promising step towards a consensus about causal relationships and mechanisms.2. Soil invertebrates contribute crucially to leaf litter decomposition and thus to the functioning of one of the most diverse ecosystem components on Earth. On the one hand, a positive effect of leaf litter diversity on decomposition was shown by several studies. On the other hand, at the interspecific level, differences in body size and feeding traits of consumers influence this process by interacting with leaf litter traits that define decomposability, such as nutrient content and physical properties. It is also known that the feeding niche of consumers can vary during their ontogeny as it depends on individual body size, metabolism and physiology. However, whether and how intraspecific trait variation of macrodetritivores contributes through niche partitioning to the positive effect of litter combinations on decomposition was never investigated.3. Here, we tested experimentally the effect of intraspecific body size differences in the common terrestrial isopod species Oniscus asellus on the decomposition of the two co-occurring litter species Acer platanoides and Betula pendula. We hypothesized complementary resource use by two distinct isopod body size classes, leading to decreased competition and higher overall litter decomposition.4. In a full factorial design with all possible combinations of litter species and isopod body size classes, litter mixtures enhanced decomposition beyond the additive effect of each single species, especially in isopod mixed-body size treatments. Our results suggest an interaction between litter and consumer diversity driving intraspecific feeding niche partitioning. Although the focus on a single detritivore species does not allow for generalizations, our results should stimulate the investigation of the mechanisms of biotic interactions among individual organisms.

Plasticity of root traits plays an important role in plant growth and survival under changing climate. Shift in precipitation is one of the most pertinent global change factors driving changes in structure and function of grasslands. However, few studies have investigated intraspecific variation of root traits in response to precipitation change under field conditions. We conducted a 10‐year simulated increased precipitation experiment in a temperate grassland and a 700‐km regional scale transect along a precipitation gradient ranging from 144.23 to 412.29 mm in northern China. The morphological, chemical and anatomical traits of the first two‐order roots were determined on 15 common herbaceous species in the manipulation experiment and two regionally common species (Leymus chinensis, Artemisia frigida) along a precipitation gradient. We found that most of the root traits of the herbaceous species exhibited no significant responses to water addition. The two regionally common species adjusted their root traits at sites with the annual precipitation lower than certain value, that is 250 and 160 mm for L. chinensis and A. frigida, respectively. These results indicate that root traits of the herbaceous species exhibit little plasticity in response to precipitation change and that the adjustment of root traits occurs when the range of annual precipitation exceeds a certain threshold. Root traits of L. chinensis and A. frigida varied differently both in manipulation experiment and along the precipitation gradient. Root traits of L. chinensis were relatively constant, while A. frigida adjusted their morphological root traits in response to water addition. Moreover, L. chinensis showed higher specific root length (SRL) and area, and root N contents at sites with annual precipitation lower than c. 250 mm. In contrast, A. frigida displayed thicker roots with lower SRL and area at sites with annual precipitation lower than c. 160 mm. Our results showed that root traits of herbaceous species in temperate grasslands exhibited little plasticity and that different species have evolved diverse adaptive strategies in response to precipitation change. These novel findings provide valuable information to predict responses of temperate grasslands to future climate change.

Plant performance is driven by nutrient availability. So far, studies on plant nutrient relationships mostly focused on nitrogen (N) and phosphorus (P), whereas potassium (K) received less attention. We evaluated whether plant responses to variation in nutrient availability are similar or different for the nutrients N, P and K and whether the same plant traits are decisive for the competitive ability of species along gradients of N, P and K availabilities. We studied plant trait responses of three temperate perennial grass species (Alopecurus pratensis, Agrostis capillaris and Anthoxanthum odoratum, each dominant under N limitation, P limitation and K limitation, respectively). Each gradient consisted of eight treatments (N1–N8, P1–P8, K1–K8), with N:P:K supply rates being 1:0.8:6 (N1), 128:0.8:6 (N8), 12:0.075:6 (P1), 12:9.6:6 (P8), 12:0.8:0.56 (K1) and 12:0.8:72 (K8) (total supply per plant individual in mg). Plant individuals were kept either in single‐species pots (intraspecific competition) or in mixed‐species pots (interspecific competition). After 2 months, plant biomass, leaf and root traits and root enzyme activity were measured. Most below‐ground traits (e.g. total root length, root diameter) responded most strongly to competition as predictor variable and only secondly to the type of nutrient or their supply rates (unlike RGRTₒₜₐₗ and SLA). Leaf chlorophyll content and root enzyme activity responded most strongly to supply rate of nutrients and the interaction of nutrient type and supply rate, respectively. In line with our prediction, A. pratensis was the superior competitor under N limitation and A. odoratum was the superior competitor under K limitation (assessed by relative dominance [RD] and relative competitive strength [RCS]). Both species showed high investment in total root length. Opposite to our expectation, A. capillaris was a bad competitor under P limitation, with low root length but higher root phosphatase activity. Our results indicate that below‐ground plant traits are decisive in nutrient‐related competition between plant species. Further, competitive strength is determined by a high total root length per soil volume both under N and K limitation. Under P limitation, the model species A. capillaris showed a weak competitive performance either because it was lacking colonization with mycorrhiza and/or the high investment in phosphatase activity posed a trade‐off with higher N investment in phosphatase production. A free Plain Language Summary can be found within the Supporting Information of this article.

When more than one closely related plant species share the same pollination niche, reproductive interference via interspecific pollen transfer should limit their coexistence. However, some studies have reported the sympatric coexistence of two native close relatives pollinated by the same pollinators under reproductive interference, even without niche partitioning. We examined the frequency dependency of reproductive interference between close relatives in natural conditions and the potential roles of autonomous selfing in mitigating the negative reproductive interference effects between congeneric species. We investigated sympatrically growing Commelina communis (Cc) and C. c. f. ciliata (Ccfc) populations. These species exhibit very large overlaps in habitat preference, flowering phenology and pollination niche, but seldom produce hybrids. First, we conducted a hand‐pollination experiment to examine the negative effects of heterospecific pollen deposition on seed production and the potential of self‐pollination to mitigate the effects in both species. Then, we examined the effects of reproductive interference on reproductive success and the potential for autonomous selfing in the field. We found significant negative effects of heterospecific pollen deposition on seed production and the mitigation effects of prior and competing self‐pollination, in both Cc and Ccfc. For both species in the field, intraspecific pollinator movements and reproductive success significantly decreased with an increase in the relative floral abundance of competing species, although the negative reproductive interference effect on reproductive success was lower in Cc than in Ccfc. We also found greater potential for prior autonomous selfing in Cc than in Ccfc. Our findings suggest that Cc flowers were less affected by reproductive interference from competing species, which was likely due to a higher prior selfing ability compared to Ccfc flowers. The asymmetry in susceptibility to reproductive interference may explain the Cc‐biased distribution in the study area. The study improves our understanding of how prior autonomous selfing can reduce the negative reproductive interference effect from competing species in mixed‐mating species with frequent pollinator visits. A plain language summary is available for this article.

There is remarkable variation in life histories of coexisting plant species. These “alternative designs” for the given set of environmental conditions are likely to play a role in species niche differences and thus may underlie species coexistence, although there is no clear demonstration of it. Currently available data on within‐community differentiation concern primarily easy and measurable traits that are not fully informative of life history variation. Their relevance for functional differentiation of species and species coexistence is far from clear. Here, we examined differentiation of coexisting species in demographic parameters and how it determines species’ response to neighbours. We determined these parameters for a set of 21 co‐occurring species by fitting a process‐based model to a long‐term (30 years) data series of shoot counts. We examined the functional relevance of these parameters using species responses in a field removal experiment. We further asked with which functional traits they are correlated. Species were differentiated along two largely independent axes: (a) slow vs. fast, separating species according to instantaneous growth rate, competitive response and intraspecific density dependence; and (b) dispersal vs. local dynamics, which separated species with strong dispersal (by seeds or vegetative) from species that tended to stay in the occupied spot. While the slow‐fast axis was associated with commonly used leaf and seed traits, the dispersal axis was best predicted by lateral spreading distance. Each of these two axes predicted different components of species’ responses to neighbour competition: the slow‐fast axis was a good predictor of the short‐term response, whereas dispersal axis was a good predictor of the long‐term response. Synthesis. Demographic differentiation of coexisting species resembles to an important degree demographic differentiation known from large‐scale comparisons. This differentiation is functionally meaningful also at the fine scale; its role in species’ responses to competition implies it is involved in species niche differentiation and coexistence. While seed and leaf traits are important correlates of demographic differentiation, a hitherto underappreciated trait, viz. lateral spreading distance, is an important predictor of the dispersal axis at the fine scale and should be more widely used. A plain language summary is available for this article.

Growth in body size is accompanied by changes in foraging capacity and metabolic costs, which lead to changes in competitive ability during ontogeny. The resulting size‐dependent competitive asymmetry influences population dynamics and community structure, but it is not clear whether natural selection leads to asymmetry in intraspecific competition.We address this question by using a size‐structured consumer–resource model, in which the strength and direction of competitive asymmetry between different consumer individuals depends on the scaling of maximum ingestion and maintenance metabolism with consumer body size. We use adaptive dynamics to study selection on the scaling exponents of these processes.Selection leads to an identical scaling of maximum ingestion and maintenance metabolism with consumer body size. Equal scaling exponents neutralize strong competitive differences within the consumer population, because all consumer individuals require the same amount of resources to cover maintenance requirements. Furthermore, the scaling exponents respond adaptively to changes in mortality such that biomass production through growth or reproduction increases in the life stage that is subject to increased mortality. Also, decreasing size at birth leads to increased investment in juvenile growth, while increasing maximum size leads to increased investment in post‐maturation growth and reproduction.These results provide an explanation for observed variation in the ontogenetic scaling of metabolic rate with body size. Data of teleost fish are presented that support these predictions. However, selection towards equal scaling exponents is contradicted by empirical findings, which suggests that additional ecological complexity beyond this basic consumer–resource interaction is required to understand the evolution of size‐dependent asymmetry in intraspecific competition. A plain language summary is available for this article.

Trait intraspecific variability determines community dynamics and species coexistence. In response to competition, plants can display intraspecific variability to enhance their competitive ability or stabilise their niche differences with competitors. This response is multidimensional because it involves changes along different functional axes and inevitable trade‐offs between traits. Here, we transposed the recent concept of the multidimensional trait space to the analysis of intraspecific plant response to competition. We specifically tested the following: (a) in the absence of competitors, the plant multidimensional trait space will be packed towards strategies promoting plant colonisation; and (b) with competitors, the plant multidimensional trait space will be directed towards competition with its size and shaping characteristics dependent on competitor species richness. We studied trait intraspecific variability of two clonal species, Brachypodium pinnatum (L.) P. Beauv. and Elytrigia repens (L.) Gould, in response to competition. We analysed plant response in the absence of competitors and in competition. Competition treatments included intraspecific and interspecific experimental mixtures with increasing species richness. For each target species and each treatment, we built an hypervolume based on six traits involved in the three‐dimensional competition (i.e., ramet and connection traits). We measured these hypervolumes for their size, similarity and the contribution of traits in their shaping. In the absence of competitors and for both species, we demonstrated a multidimensional trait space packing towards a colonisation strategy. Under competition, the multidimensional trait spaces of the two target species were the widest at the extremes of the richness gradient, that is, intraspecific and interspecific high richness competition treatments. High intraspecific variability either promoted niche differentiation from individuals of similar species or reflected the large range of competitive responses deployed when plants were faced with many different competitor identities. The multidimensional response process was based on fine adjustments of various traits depending on the surrounding neighbourhood composition and more specifically, on the competitor functional similarity with the target species. This study emphasises the multidimensionality of species competitive response, and also underlines the so far neglected importance of competitor species richness for trait intraspecific variability and subsequently community assembly. A plain language summary is available for this article.

The impact of plant–plant interactions on species diversity patterns has been broadly addressed in stressful environments, such as alpine ecosystems, where foundation species promote species richness by creating habitat for other species. However, foundation species with contrasting phenotypes might modify the microhabitat differently, which would alter the subordinate community composition, and coincide with distinct feedback effects of those subordinate species on the foundation species. However, the precise interaction mechanisms that facilitate species are not fully understood, especially the relative contribution of above- and below-ground compartments of foundation species to subordinate species and the potential feedbacks they receive. We explored whether two contrasted canopy phenotypes (tight and loose) of the shrub Cytisus galianoi differed in their effects on the microhabitat and on subordinate plant community composition in a dry subalpine system. We also experimentally distinguished the relative contribution of above- (canopy) and below-ground (soil) effects of C. galianoi on the most frequent subordinate species, Festuca indigesta, as well as the reciprocal effects of F. indigesta on C. galianoi. We performed observational and manipulative experiments to assess the influence of phenotypic differences of the shrub on understorey microhabitat and subordinate plant community composition. Reciprocal effects were assessed by removing either F. indigesta from the understorey of the two shrub phenotypes or the C. galianoi canopy from the immediate vicinity of F. indigesta. The two C. galianoi phenotypes differed in mean values of functional traits (like stem density or plant height), modified their understorey microhabitats differently and hosted distinct subordinate communities. Loose phenotypes had more positive effects on community composition and diversity than tight phenotypes. In addition, tight phenotypes simultaneously showed both more positive above-ground and more negative below-ground effects on F. indigesta than loose phenotypes. There were no significant feedback effects of F. indigesta on C. galianoi. The two phenotypes of the foundation species C. galianoi showed contrasting effects on the subordinate plant community: Compared to the tight phenotype, the loose phenotype had higher associated species diversity and reduced reciprocal interaction intensities above- and below-ground with the subordinate species F. indigesta. This highlights the impact of phenotypic variation for plant interactions and community-level diversity. A plain language summary is available for this article., This work was funded by the Organismo Autónomo Parques Nacionales (Grant Nos. 0002/9 and 1578/2015) and AEI- MINECO (CGL2014-59010-R and CGL2017-84515-R). N.P. was supported by a JAEPredoc CSIC and a Brazilian CAPES PVE Grant Number 88881.068053/2014; C.S. by the Swiss National Science Foundation (PZ00P3_148261, PP00P3_170645); P.M. by LINCGlobal, GACR 17-20839S and by a CzechPolar Grant Number MSMT LM2015083; and C.A. by a Spanish Government “Ramón y Cajal” contract (RYC-2012-12277).

A frequent question in invasion ecology is whether the traits of alien plant species differ from natives. This question is increasingly addressed at the plant community scale to assess the role trait differences play in community assembly. Nevertheless, there remains considerable debate as to the value of native vs. alien trait comparisons. Recommendations are made to address seven conceptual and methodological limitations in current approaches used to assess trait differences between native and alien plants: account for spatial scale dependence, make an informed choice of traits, evaluate multiple trait dimensions, incorporate intraspecific trait variation, use appropriate null models, examine environmental gradients, and assess temporal variation in trait spectra. Tackling these limitations will dramatically improve the value of native vs. alien trait comparisons. Traits should be selected based on expectations of abiotic and/or biotic constraints. Phylogenetic distances can provide valuable insights but are no substitute for the measurement of individual traits when contrasting alien and native species. An ideal set of traits would include one or more leaf, stem, seed and root traits as well as flowering phenology and plant height at maturity. Comparisons of intraspecific trait variation among native and alien species within the same plant community are rare but could improve predictions of interspecific competition and the response of communities to plant invasions. The full plant community should be examined rather than selected species subsets and comparisons between native and alien species traits are better undertaken at a neighbourhood scale. The frequency distribution of alien and native species traits may change over time through competitive exclusion as well as contemporary evolution. Repeated surveys of species abundances and traits are required to understand the consequences of trait differences between aliens and natives for plant community assembly. Analyses of the role of species traits in plant invasions need to quantify differences in a priori identified traits within communities along suitable environmental gradients and test hypothetical trait patterns against appropriate null expectations. The infrequent application of such approaches may explain the limited generalisations regarding the role of trait differences between native and alien species in the invasion of plant communities. A plain language summary is available for this article.

Occupancy of cold habitats by evergreen species requires structural modification of photosynthetic organs for stress resistance and longevity. Such modifications have been described at interspecific level, while intraspecific variation has been underexplored. To identify structural and anatomical traits that may be adaptive in cold environments, we studied intraspecific variability of needles of Scots pine (Pinus sylvestris L.), a wide-ranging tree, along a 1,900 km temperate-boreal transect in Europe. Needles from 20 sites representing mean minimum winter temperatures between -4.0 and -19.9°C and mean annual temperatures between 8.3 and -1.7°C were sampled for measurements of leaf mass per area (LMA, g/m2), leaf density (LD, g/cm3) and 30 other morpho-anatomical traits. Needles from cold sites lived longer, were shorter, showed higher LMA and LD, had narrower and more collapse-resistant tracheids, thicker epidermal cells with thicker cell walls and wider resin ducts occupying larger fraction of needle volume in comparison to needles from warmer sites. Along the steep climatic gradient, needles presented a coordinated phenotypic spectrum of external and internal traits that are largely interpretable in functional, adaptive terms. This intraspecific pattern of covarying traits provides insight into the adaptive syndrome associated with stress tolerance and extended needle longevity under cold conditions of high latitudes. A plain language summary is available for this article. (Less)

Summary Interspecific variation in elemental composition is well known and often leads to predictable differences in ecosystem interactions, but little is known about the extent, causes and importance of intraspecific variation in elemental composition. If intraspecific variation is substantial and has a genetic basis, it may underlie an important mechanism of evolutionary interplay with ecology as individuals compensate for evolutionary changes in elemental demand. To investigate the extent and causes of intraspecific elemental variation in vertebrates, we sampled evolutionary model species Gasterosteus aculeatus (Threespine Stickleback) from 12 locations in British Columbia, Canada. Fish were phenotyped, genotyped for Eda alleles underlying lateral plate variation and assayed for elemental content (C, N, P). We found stickleback vary widely in elemental composition (2.2 – 6.5%P; 3.0 – 9.4:1 N:P). Phenotypic models explained the majority of this variation using bony armour traits (pelvis length, lateral plate count), bone mineralization, body size and condition. Subsequent genetic models found allelic variation at Eda generates a 7-14% change in whole organism N:P. As Eda allele frequencies are commonly changed through strong natural selection in freshwater habitats, we infer that stickleback elemental composition can evolve rapidly. Further, as genetics are known to drive variation in many of the other influential traits, we conclude that genetic variation constitutes a major source of variation in the elemental composition of Gasterosteus aculeatus. As such, we find that elemental composition has a large evolutionary potential which may underlie important evo-eco interactions. This article is protected by copyright. All rights reserved.

Summary Ecological theory has long recognized the importance of positive and negative species interactions as drivers of food web structure, yet many studies have only focused on competition. Because competitive and facilitative mechanisms operate simultaneously, but through different food web pathways, the balance of their combined effects can produce complex and variable responses. We used a response-surface experimental design to assess the roles of negative (e.g., intra-, interspecific competition) and positive (e.g., facilitation) interactions between native and invasive juvenile fishes. We tested whether these interactions alter the densities of planktonic and benthic invertebrates to evaluate the magnitude and mechanism(s) influencing the acceptance or resistance of biological invaders. Interactions between bighead carp (Hypophthalmichthys nobilis) and bluegill (Lepomis macrochirus) or common carp (Cyprinus carpio) were evaluated in mesocosms. Intraspecific interactions were 1.5 to 2.4 times stronger than interspecific interactions between carp species. The only instance of interspecific competition resulted in bighead carp reducing the daily growth of bluegill, whereas the reciprocal interaction resulted in facilitation. Facilitation occurred when bluegill increased the daily growth of low density bighead carp treatments, despite increased numbers of fishes. Bighead carp also increased densities of benthic Chironomidae larvae, which were subsequently consumed by bluegill, but did not result in enhanced bluegill growth. These suites of interactions were not observed between common and bighead carp. Our response-surface design proved useful for comparing the relative magnitude of intra- vs. interspecific competition, identifying facilitation among species, and tracing attendant effects on invertebrate communities. By accounting for the directionality of interactions within our experimental framework and tracking responses of prey at lower trophic levels, we provide a clearer understanding of how competitive effects and stressed consumers alter prey communities and influence facilitation. This article is protected by copyright. All rights reserved.

Summary 1.Trait-based models of ecological communities typically assume intraspecific variation in functional traits is not important, though such variation can change species trait rankings along gradients in resources and environmental conditions, and thus influence community structure and function. 2.We examined the degree of intraspecific relative to interspecific variation, and reaction norms of 11 functional traits for 57 forest understory plant species, including: intrinsic water-use efficiency (iWUE), Δ15N, 5 leaf traits, 2 stem traits and 2 root traits along gradients in light, nitrogen, moisture and understory cover. 3.Our results indicate that interspecific trait variation exceeded intraspecific variation by at least 50% for most, but not all traits. Intraspecific variation in Δ15N, iWUE, leaf nitrogen content and root traits was high (47-70%) compared with most leaf traits and stem traits (13-38%). 4.Δ15N varied primarily along gradients in abiotic conditions, while light and understory cover were relatively less important. iWUE was related primarily to light transmission, reflecting increases in photosynthesis relative to stomatal conductance. Leaf traits varied mainly as a function of light availability, with some reaction norms depending on understory cover. Plant height increased with understory cover, while stem specific density was related primarily to light. Resources, environmental conditions and understory cover did not contribute strongly to the observed variation in root traits. 5.Gradients in resources, environmental conditions and competition all appear to control intraspecific variability in most traits to some extent. However, our results suggest that species cross-over (i.e., trait rank reversals) along the gradients measured here are generally not a concern. 6.Intraspecific variability in understory plant species traits can be considerable. However, trait data collected under a narrow range of environmental conditions appears sufficient to establish species rankings and scale between community and ecosystem levels using trait-based models. Investigators may therefore focus on obtaining a sufficient sample size within a single set of conditions rather than characterizing trait variation across entire gradients in order to optimize sampling efforts. This article is protected by copyright. All rights reserved.

Summary Early postnatal development can have profound effects on life-history traits later in life. One mechanism hypothesized to mediate this relationship is the anabolic hormone, insulin-like growth factor-1 (IGF-1). IGF-1 contributes importantly to postnatal growth, and thus offers a means by which environmental and genetic variation might direct organismal development, reproduction and survival. We tested whether juvenile concentrations of IGF-1 can predict intraspecific variation in life-history traits later in life using longitudinal data from free-living female spotted hyenas (Crocuta crocuta). We found that juvenile concentrations of IGF-1 predicted heavier juvenile mass, which in turn predicted greater survival to reproductive maturity. However, independent of mass, higher juvenile concentrations of IGF-1 predicted earlier age at first parturition and reduced longevity in adulthood. Our results highlight the importance of early postnatal development as a determination period in mammals and suggest that concentrations of IGF-1 during this sensitive period can be used to predict important later-life trade-offs between growth, reproductive fitness and life span in wild, long-lived animals. A lay summary is available for this article.

1.Physiological and energetic mechanisms have been proposed to constrain body sizes of organisms along climatic gradients; however, these provide contrasting predictions. While Bergmann′s rule predicts increases of body sizes in cooler climates resulting from physiological constraints, energy-based community assembly rules suggest declines in the mean body size of species caused by increased extinction probabilities for large-bodied species in low-energy habitats. 2.We tested these contrasting hypotheses by quantifying trait distributions in bee communities along a 3.6 km elevational gradient at Mt. Kilimanjaro. Traditionally, intra- and interspecific trait shifts along environmental gradients have been investigated in isolation. However, a surge of theoretical approaches and studies on plants demonstrated that the explicit integration of trait variation among and within species can be essential for identifying the mechanisms that shape traits and related ecosystem functions along environmental gradients. We therefore studied variation of body size and related morphological traits at both the intra- and interspecific level. 3.We found support for both physiological constraints and energy-based community assembly rules as drivers of trait distribution in bee communities along elevational gradients, which, however, affected different levels of biotic organization, i.e. the population and community level. While the number of bee species with large body sizes declined with elevation, individuals within species became on average larger, resulting in contrasting trends in morphometric parameters at the community versus population level. 4.Furthermore, body size within bee communities became less variable at higher elevations, largely as a result of a non-random, directive loss of species, but paralleled by a decline in intraspecific variance, suggesting intensified filtering effects with increasing elevation. Similar patterns were found for other functional traits related to the foraging ecology of bees (tongue length, relative forewing length). 5.We conclude that along climatic gradients both physiological and energetic constraints shape trait distributions of pollinators, but at different levels of biological organization. This article is protected by copyright. All rights reserved.

Understanding species differences in plant functional traits has been critical in developing a mechanistic understanding of terrestrial ecological processes. Greater attention is now being placed on understanding the extent, causes and consequences of intraspecific trait variation (ITV). ITV is especially important in governing ecological processes in cropping systems, where only a small number of species or genotypes exist in high abundances. However, it remains unclear if key principles of trait-based ecology – namely the leaf economics spectrum (LES) – also describe intraspecific variation in crop functional biology. There also remains a need to understand whether ITV within crops is random, or structured across environmental, management-related or biological levels of organization in agroecosystems. We employed a nested design field survey to evaluate ITV in leaf traits in coffee (Coffea arabica), one of the world's most widespread tropical crops. We evaluated ITV in eight physiological, morphological and chemical leaf traits, across five nested categorical levels (sites, management systems, spatial location, plant identity, branch identity). We compared patterns of LES trait covariation in coffee, to interspecific patterns observed across over 700 wild plant species. Patterns of bivariate and multivariate ITV in coffee were broadly consistent with, but considerably weaker than, interspecific patterns associated with the LES, indicating that crops may systematically diverge from global patterns of trait trade-offs observed in wild plants. Physiological traits varied most widely (coefficient of variation (cv) 42–107%), followed by morphological traits (cv = 15–38%) and chemical traits (cv = 3–11%). Physiological ITV was best explained by the site in which a coffee plant was growing (17–55% explained), while ITV for chemical traits was best explained by management treatments within sites (25–36%); morphological ITV was higher even at the individual tree level or branch level and remained largely unexplained. Our results support the hypothesis that artificial selection and high-resource agricultural environments lead crops to systematically deviate from patterns of leaf trait covariation observed across wild plants species. Coupled with an understanding of how different traits vary systematically across multiple levels of biological organization, these findings help integrate ITV into future analyses of agroecosystem structure and function. (Resume d'auteur)

Summary Colour signals and colour vision play a pivotal role in intraspecific communication and predator-prey interactions. However, the costs of expressing conspicuous sexual signals at multiple trophic levels have been largely overlooked. Sexual signals can also experience character displacement in sympatric populations of closely-related species, leading to potential changes in conspicuousness. We here investigate a bird-damselfly-fruit fly predator-prey system, where two closely related damselfly species have conspicuous, sexually selected wing coloration. The damselflies can occur in sympatry and allopatry and reproductive character displacement in the coloration size has been previously reported. We quantify the damselfly wing reflectance from replicated sympatric and allopatric populations, and use receptor noise models to investigate the visual discriminability of the wing coloration for the bird, damselfly and fly vision systems, against natural backgrounds. We perform electroretinograms to study damselfly eye sensitivity. We also estimate damselfly predation risk in natural populations. We find that the chromatic component of wing coloration makes males highly discriminable to the predator, but not to the prey. However, female wing coloration is predominantly cryptic for the predator and prey, and interestingly, also for male damselflies. A female being cryptic to conspecifics likely reduces male harassment. The estimates of predation risk partially support the discriminability results. We also show that there is no difference in colour vision sensitivity between the two damselfly species and sexes, and no difference in wing coloration or its discriminability between sympatric and allopatric populations. Our results suggest that sexually selected traits can be antagonistically selected by predators and prey, and that this antagonistic selection can be sex-dependent: males are paying a large cost in terms of conspicuousness, while females remain mostly cryptic. Our study thus emphasizes the need for investigating visual communication at multi-trophic levels since the degree of colour discriminability can differ between predators, prey and the focal species. This article is protected by copyright. All rights reserved.

Summary Ecological and evolutionary processes historically have been assumed to operate on significantly different time-scales. We know now from theory and work in experimental and model systems that these processes can feed back on each other on mutually relevant time-scales. Here, we present evidence of a soil-mediated eco-evolutionary feedback on the population dynamics of an invasive biennial plant, Alliaria petiolata. As populations age, natural selection drives down production of A. petiolata's important antimycorrhizal allelochemical, sinigrin. This occurs due to density-dependent selection on sinigrin, which is favoured under interspecific, but disfavoured under intraspecific, competition. We show that population stochastic growth rates (λS) and plant densities are positively related to sinigrin concentration measured in seedling roots. This interaction is mediated by sinigrin's positive effect on seedling and summer survival, which are important drivers of λS. Together, these illustrate how the evolution of a trait shaped by natural selection can influence the ecology of a species over a period of just years to decades, altering its trajectory of population growth and interactions with the species in the soil and plant communities it invades. Our findings confirm the predictions that eco-evolutionary feedbacks occur in natural populations. Furthermore, they improve our conceptual framework for projecting future population growth by linking the variation in plant demography to a critical competitive trait (sinigrin) whose selective advantages decrease as populations age.

Individuals vary in the extent to which they associate with conspecifics, but little is known about the energetic underpinnings of this variation in sociability. Group-living allows individuals to find food more consistently, but within groups, there can be competition for food items. Individuals with an increased metabolic rate could display decreased sociability to reduce competition. Long-term food deprivation (FD) may alter any links between sociability and metabolic rate by affecting motivation to find food. We examined these issues in juvenile qingbo carp Spinibarbus sinensis, to understand how FD and metabolic rate affect sociability. Like many aquatic ectotherms, this species experiences seasonal bouts of FD. Individuals were either: (i) food-deprived for 21 days; or (ii) fed a maintenance ration (control). Fish from each treatment were measured for standard metabolic rate (SMR) and tested for sociability twice: once in the presence of a control stimulus shoal and once with a food-deprived stimulus shoal. Control individuals ventured further from stimulus shoals over a 30-min trial, while food-deprived fish did not change their distance from stimulus shoals as trials progressed. Control fish with a higher SMR were least sociable. Well-fed controls showed decreased sociability when exposed to food-deprived stimulus shoals, but there was evidence of consistency in relative sociability between exposures to different shoal types. Results contrast with previous findings that several days of fasting causes individuals to decrease associations with conspecifics. Prolonged FD may cause individuals to highly prioritize food acquisition, and the decreased vigilance that would accompany continuous foraging may heighten the need for the antipredator benefits of shoaling. Conversely, decreased sociability in well-fed fish with a high SMR probably minimizes intraspecific competition, allowing them to satisfy an increased energetic demand while foraging. Together, these results suggest that FD – a challenge common for many ectothermic species – can affect individual sociability as well as the attractiveness of groups towards conspecifics. In addition, the lack of a link between SMR and sociability in food-deprived fish suggests that, in situations where group membership is linked to fitness, the extent of correlated selection on metabolic traits may be context-dependent.

Summary Comparative trait analyses of native and invasive plant species have enriched our understanding of the strategies that make plants successful in particular contexts. However, it is generally unknown whether traits associated with invasiveness arise de novo in the introduced range or represent a case of ‘pre-adaptation’ of some species to become invasive. We compared the functional strategies of two invasive species, Prunus serotina, a tree native to Eastern North America (ENA) and invasive in European forests, and Rhamnus cathartica, a shrub native to Europe and invasive in ENA, in both their native and invasive ranges. We measured leaf functional traits related to plant carbon and nitrogen economics in populations across northeastern ENA (New York) and northern France. This reciprocal field approach is unique, comparing in situ physiology within and between each species' shared ranges. For both species, we found striking differences in leaf economic traits and intraspecific trait scaling relationships. Prunus serotina exhibited similar photosynthetic rates in New York (native range) and France (invasive range), but French populations had significantly greater leaf carbon investments in the form of increased leaf respiration, construction costs, and carbon concentrations. Rhamnus cathartica exhibited 50–60% higher photosynthetic rates in New York (invasive range) than France (native range), along with increased leaf nitrogen costs. New York populations also had significantly lower nitrogen resorption efficiency, with autumn leaf nitrogen concentrations 69% lower in France. Intraspecific trait differences between native and invasive ranges indicate shifts in resource-use strategies between the native and invasive range are possible in invasive species. While further investigations with additional species would be needed to determine the role of phenotypic plasticity and prevalence of post-introduction trait shifts, our results question the assumption that functional strategies of invasive species are conserved from the native range and highlight the utility of measuring in situ functional strategies of reciprocal invaders to provide mechanistic understanding of invasions.