Climate change-related growth improvements in a wide niche-breadth tree species across contrasting environments.

BACKGROUND AND AIMS: The vulnerability and responsiveness of forests to drought are immensely variable across biomes. Intraspecific tree responses to drought in species with wide niche breadths that grow across contrasting climatically environments might provide key information regarding forest resistance and changes in species distribution under climate change. Using a species with an exceptionally wide niche breath, we tested the hypothesis that tree populations thriving in dry environments are more resistant to drought than those growing in moist locations. METHODS: We determined temporal trends in tree radial growth of 12 tree populations of Nothofagus antarctica (Nothofagaceae) located across a sharp precipitation gradient (annual precipitation of 500-2000 mm) in Chile and Argentina. Using dendrochronological methods, we fitted generalized additive mixed-effect models to predict the annual basal area increment as a function of year and dryness (De Martonne aridity index). We also measured carbon and oxygen isotope signals (and estimated intrinsic water-use efficiency) to provide potential physiological causes for tree growth responses to drought. KEY RESULTS: We found unexpected improvements in growth during 1980-1998 in moist sites, while growth responses in dry sites were mixed. All populations, independent of site moisture, showed an increase in their intrinsic water-use efficiency in recent decades, a tendency that seemed to be explained by an increase in the photosynthetic rate instead of drought-induced stomatal closure, given that δ18O did not change with time. CONCLUSIONS: The absence of drought-induced negative effects on tree growth in a tree species with a wide niche breadth is promising because it might relate to the causal mechanisms tree species possess to face ongoing drought events. We suggest that the drought resistance of N. antarctica might be attributable to its low stature and relatively low growth rate.

Temporal dynamics in alpine snowpatch plants along a snowmelt gradient explained by functional traits and strategies.

Alpine snowpatches are characterised by persistent snow cover, short growing seasons and periglacial processes, which has resulted in highly specialised plant communities. Hence, these snowpatch communities are among the most threatened from climate change. However, temporal dynamics in snowpatch microclimate and plant composition are rarely explored, especially in the marginal alpine environments of Australia. Seven snowpatches were categorised into early, mid and late snowmelt zones based on growing season length, with soil temperatures recorded from 2003 to 2020 and plant composition surveyed in 84 1 m2 quadrats in 2007, 2013 and 2020. Microclimate, species diversity, plant cover and composition, along with community-weighted trait means and plant strategies were assessed to understand snowpatch dynamics in response to climate change. We found that growing season length and temperatures have increased in late melt zones, while changes were less consistent in early and mid melt zones. There were few changes in species diversity, but increases in graminoids and declines in snowpatch specialists in mid and late melt zones. Community-weighted plant height, leaf area and leaf weight also increased, particularly in mid and late melt zones, while plant strategies shifted from compositions of ruderal-tolerant to stress-tolerant. Here, we show that snowpatch communities are rapidly changing in response to longer growing seasons and warmer temperatures, with the greatest changes occurring where snow persists the longest. The results highlight the climate-induced loss of defining biotic and abiotic characteristics of snowpatches, as temporal convergence of compositions along snowmelt gradients threatens the distinctiveness of snowpatch plant communities.

Hurricanes increase tropical forest vulnerability to drought.

Rapid changes in climate and disturbance regimes, including droughts and hurricanes, are likely to influence tropical forests, but our understanding of the compound effects of disturbances on forest ecosystems is extremely limited. Filling this knowledge gap is necessary to elucidate the future of these ecosystems under a changing climate. We examined the relationship between hurricane response (damage, mortality, and resilience) and four hydraulic traits of 13 dominant woody species in a wet tropical forest subject to periodic hurricanes. Species with high resistance to embolisms (low P50 values) and higher safety margins ( SMP50 ) were more resistant to immediate hurricane mortality and breakage, whereas species with higher hurricane resilience (rapid post-hurricane growth) had high capacitance and P50 values and low SMP50 . During 26 yr of post-hurricane recovery, we found a decrease in community-weighted mean values for traits associated with greater drought resistance (leaf turgor loss point, P50 , SMP50 ) and an increase in capacitance, which has been linked with lower drought resistance. Hurricane damage favors slow-growing, drought-tolerant species, whereas post-hurricane high resource conditions favor acquisitive, fast-growing but drought-vulnerable species, increasing forest productivity at the expense of drought tolerance and leading to higher overall forest vulnerability to drought.

Climate warming leads to advanced fruit development period of temperate woody species but divergent changes in its length.

Climate warming has significantly altered the phenology of plants in recent decades. However, in contrast to the widely reported warming-induced extension of vegetative growing season, the response of fruit development period (FDP) from flowering to fruiting remains largely unexplored, particularly for woody plants. Analyzing >560,000 in situ observations of both flowering and fruiting dates for six temperate woody species across 2958 European phenological observations sites during 1980-2013, we found that in all species both flowering and fruiting phenology, that is, the FDP, advanced with climate warming. However, the advancing rates of the two events were not necessarily equal for any given species, resulting in divergent changes in the length of FDP among species with climate warming. During 1980-2013, not only the temperature during FDP but also the forcing requirement for fruit development increased, both affecting the length of FDP. The shortened FDP was mainly due to elevated temperature, thus accelerating the accumulation of forcing, whereas the prolonged FDP was primarily caused by the substantial increase of the forcing requirement of fruiting, which could be fulfilled only in a longer time and thus slowed down the advance of fruiting. This study provides large-scale empirical evidence of warming-induced advances of FDP but divergent changes in its length in temperate woody species. Our findings demonstrate the contrasting reproductive phenological strategies among temperate woody species under the pressure of warming climate, contrary to the lengthening of vegetative growing season, which is by and largely similar with different woody species.

Long-term alfalfa (Medicago sativa L.) establishment could alleviate phosphorus limitation induced by nitrogen deposition in the carbonate soil.

Phosphorus (P) limitation is a widespread problem of primary production in dryland submitted to persistent nitrogen (N) deposition. The legume alfalfa (Medicago sativa L.), which can fix N2, might potentially strengthen P limitation in dryland ecosystems and is widely distributed as forage. However, there is still unclear how alfalfa grassland mobilizes the soil P to meet its demand. In this experiment, alfalfa introduction was used for long-term revegetation to evaluate the P uptake of plants from deep soil and assess the P limitation induced by N deposition compared with fallow. Our results showed that alfalfa introduction increased the soil P storage significantly at 0-2.4 m soil depth (+0.74 Mg ha-1), whereas it decreased at 2.4-4.8 m soil depth (-0.21 Mg ha-1) after 15-year establishment. Alfalfa establishment increased soil organic P concentration (180.9 mg kg-1 vs. 67.2 mg kg-1) and its relative contribution to total P (19.64% vs. 8.08%) at 0-4.8 m. Alfalfa establishment also increased the concentration and proportion of labile and intermediate P fractions at 0-4.8 m (9.12 mg kg-1 vs. 6.87 mg kg-1, 1.12% vs. 0.98%; 16.06 mg kg-1 vs. 8.39 mg kg-1, 1.69% vs. 1.17%). Alfalfa introduction decreased the concentrated HCl-Pi (250.66 mg kg-1 vs. 229.32 mg kg-1, 36.81% vs. 28.91%) in 2.4-4.8 m soil depth. These results indicated that the deep root system of alfalfa grassland could promote the P mobilization from deep to shallow soil. The concentrated HCl-Pi may be the main potential P source of alfalfa from 2.4-4.8 m to 0-2.4 m of soil depth, and long-term establishment of alfalfa can alleviate P limitation caused by N deposition in carbonate soil. Our results suggested that species with deep roots (such as alfalfa) could be selected as an economical way to mitigate nitrogen deposition in drylands.

Are the interaction effects of warming and drought on nutritional status and biomass production in a tropical forage legume greater than their individual effects?

MAIN CONCLUSION: Drought alone and drought plus warming will change the nutrient requirements and biomass distributions of Stylosanthes capitata, while warming will be advantageous only under well-watered condition for the next decades. Climate change effects on natural and managed ecosystems are difficult to predict due to its multi-factor nature. However, most studies that investigate the impacts of climate change factors on plants, such as warming or drought, were conducted under one single stress and controlled environments. In this study, we evaluated the effects of elevated temperature (+ 2 °C) (T) under different conditions of soil water availability (W) to understand the interactive effects of both factors on leaf, stem, and inflorescence macro and micronutrients concentration and biomass allocation of a tropical forage species, Stylosanthes capitata Vogel under field conditions. Temperature control was performed by a temperature free-air controlled enhancement (T-FACE) system. We observed that warming changed nutrient concentrations and plant growth depending on soil moisture levels, but the responses were specific for each plant organ. In general, we found that warming under well-watered conditions greatly improved nutrient concentration and biomass production, whilst the opposite effect was observed under non-irrigated and non-warmed conditions. However, under warmed and non-irrigated conditions, leaf biomass and leaf nutrient concentration were greatly reduced when compared to non-warmed and irrigated plants. Our findings suggest that warming (2 °C above ambient temperature) and drought, as well as both combined stresses, will change the nutrient requirements and biomass distributions between plant aerial organs of S. capitata in tropical ecosystems, which may impact animal feeding in the future.

Simulated climate change decreases nutrient resorption from senescing leaves.

Nutrient resorption is the process whereby plants recover nutrients from senescing leaves and reallocate them to storage structures or newer tissues. Elemental resorption of foliar N and P has been shown to respond to temperature and precipitation, but we know remarkably little about the influence of warming and drought on the resorption of these and other essential plant macro- and micronutrients, which could alter the ability of species to recycle their nutrients. We conducted a 5 year manipulative field study to simulate predicted climate change conditions and studied the effects of warming (W), rainfall reduction (RR), and their combination (W+RR) on nutrient resorption efficiency in five coexisting shrub species in a semiarid shrubland. Both mature and senesced leaves showed significant reductions in their nutrient contents and an altered stoichiometry in response to climate change conditions. Warming (W, W+RR) reduced mature leaf N, K, Ca, S, Fe, and Zn and senesced leaf N, Ca, Mg, S, Fe, and Zn contents relative to ambient temperature conditions. Warming increased mature leaf C/N ratios and decreased N/P and C/P ratios and increased senesced leaf C/N and C/P ratios. Furthermore, W and W+RR reduced nutrient resorption efficiencies for N (6.3%), K (19.8%), S (70.9%) and increased Ca and Fe accumulation in senesced leaves (440% and 35.7%, respectively) relative to the control treatment. Rainfall reduction decreased the resorption efficiencies of N (6.7%), S (51%), and Zn (46%). Reductions in nutrient resorption efficiencies with warming and/or rainfall reduction were rather uniform and consistent across species. The negative impacts of warming and rainfall reduction on foliar nutrient resorption efficiency will likely cause an impairment of plant nutrient budgets and fitness across coexisting native shrubs in this nutrient-poor habitat, with probable implications for key ecosystem functions such as reductions in nutrient retention in vegetation, litter decomposition, and nutrient cycling rates.

Effects of moisture dynamics on bryophyte carbon fluxes in a tropical cloud forest.

Bryophytes play key roles in the ecological function of a number of major world biomes but remain understudied compared with vascular plants. Little is known about bryophyte responses to different aspects of predicted changes in moisture dynamics with climate change. In this study, CO2 fluxes and photosynthetic light responses were measured within bryophyte mesocosms, being subjected to different amounts, frequencies, and types (mist or rainfall) of water addition, both before and after different periods of complete desiccation. Bryophyte carbon fluxes and photosynthetic light response were generally affected by the magnitude and type, but not frequency, of watering events. Desiccation suppressed bryophyte carbon uptake even after rehydration, and the degree of uptake suppression progressively increased with desiccation duration. Estimated ecosystem-level bryophyte respiration and net carbon uptake were c. 58% and c. 3%, respectively, of corresponding fluxes from tree foliage at the site. Our results suggest that a simplified representation of precipitation processes may be sufficient to accurately model bryophyte carbon cycling under future climate scenarios. Further, we find that projected increases in drought could have strong negative impacts on bryophyte and ecosystem carbon storage, with major consequences for a wide range of ecosystem processes.

Ongoing seasonally uneven climate warming leads to earlier autumn growth cessation in deciduous trees.

Ongoing global warming is causing phenological shifts that affect photosynthesis and growth rates in temperate woody species. However, the effects of seasonally uneven climate warming-as is occurring in much of Europe, where the winter/spring months are warming twice as fast than the summer/autumn months-on autumn growth cessation (completion of overwintering buds) and leaf senescence, and possible carry-over effects between phenophases, remain under-investigated. We conducted experiments in which we exposed saplings of canopy and understory species to 4 °C warming in winter/spring, summer/autumn, or all year to disentangle how the timing of bud break, bud set completion, and leaf senescence is affected by seasonally uneven warming. All-year warming led to significantly delayed leaf senescence, but advanced bud set completion; summer/autumn warming only delayed leaf senescence; and winter/spring warming advanced both bud set and senescence. The non-parallel effects of warming on bud completion and leaf senescence show that leaf senescence alone is an inadequate proxy for autumn growth cessation in trees and counterintuitively suggest that continued uneven seasonal warming will advance cessation of primary growth in autumn, even when leaf senescence is delayed. Phenological responses to warming treatments (earlier spring onset, later autumn senescence) were more than twice as high in understory species than in canopy species, which can partly be explained by the absence of carry-over effects among phenophases in the former group. This underscores the need to consider differences among plant functional types when forecasting the future behaviour of ecosystems.

Resilience of seed production to a severe El Niño-induced drought across functional groups and dispersal types.

More frequent and severe El Niño Southern Oscillations (ENSO) are causing episodic periods of decreased rainfall. Although the effects of these ENSO-induced droughts on tree growth and mortality have been well studied, the impacts on other demographic rates such as reproduction are less well known. We use a four-year seed rain dataset encompassing the most severe ENSO-induced drought in more than 30 years to assess the resilience (i.e., resistance and recovery) of the seed composition and abundance of three forest types in a tropical dry forest. We found that forest types showed distinct differences in the timing, duration, and intensity of drought during the ENSO event, which likely mediated seed composition shifts and resilience. Drought-deciduous species were particularly sensitive to the drought with overall poor resilience of seed production, whereby seed abundance of this functional group failed to recover to predrought levels even two years after the drought. Liana and wind-dispersed species were able to maintain seed production both during and after drought, suggesting that ENSO events promote early successional species or species with a colonization strategy. Combined, these results suggest that ENSO-induced drought mediates the establishment of functional groups and dispersal types suited for early successional conditions with more open canopies and reduced competition among plants. The effects of the ENSO-induced drought on seed composition and abundance were still evident two years after the event suggesting the recovery of seed production requires multiple years that may lead to shifts in forest composition and structure in the long term, with potential consequences for higher trophic levels like frugivores.

Legacy effects of land-use modulate tree growth responses to climate extremes.

Climate change can impact forest ecosystem processes via individual tree and community responses. While the importance of land-use legacies in modulating these processes have been increasingly recognised, evidence of former land-use mediated climate-growth relationships remain rare. We analysed how differences in former land-use (i.e. forest continuity) affect the growth response of European beech to climate extremes. Here, using dendrochronological and fine root data, we show that ancient forests (forests with a long forest continuity) and recent forests (forests afforested on former farmland) clearly differ with regard to climate-growth relationships. We found that sensitivity to climatic extremes was lower for trees growing in ancient forests, as reflected by significantly lower growth reductions during adverse climatic conditions. Fine root morphology also differed significantly between the former land-use types: on average, trees with high specific root length (SRL) and specific root area (SRA) and low root tissue density (RTD) were associated with recent forests, whereas the opposite traits were characteristic of ancient forests. Moreover, we found that trees of ancient forests hold a larger fine root system than trees of recent forests. Our results demonstrate that land-use legacy-mediated modifications in the size and morphology of the fine root system act as a mechanism in regulating drought resistance of beech, emphasising the need to consider the 'ecological memory' of forests when assessing or predicting the sensitivity of forest ecosystems to global environmental change.

Elevated temperature drives a shift from selfing to outcrossing in the insect-pollinated legume, faba bean (Vicia faba).

Climate change can threaten the reproductive success of plants, both directly, through physiological damage during increasingly extreme weather events, and indirectly, through disruption of plant-pollinator interactions. To explore how plant-pollinator interactions are modified by extreme weather, we exposed faba bean (Vicia faba) plants to elevated temperature for 5 d during flowering, simulating a heatwave. We then moved the plants to flight cages with either bumblebees or no pollinators, or to two field sites, where plants were enclosed in mesh bags or pollinated by wild insect communities. We used a morphological marker to quantify pollen movement between experimental plants. There was a substantial increase in the level of outcrossing by insect pollinators following heat stress. Proportion outcrossed seed increased from 17 % at control temperature, to 33 % following heat stress in the flight cages, and from 31 % to 80 % at one field site, but not at the other (33 % to 32 %). Abiotic stress can dramatically shift the relative contributions of cross- and self-pollination to reproduction in an insect pollinated plant. The resulting increases in gene flow have broad implications for genetic diversity and functioning of ecosystems, and may increase resilience by accelerating the selection of more stress-tolerant genotypes.

Intra-annual plasticity of growth mediates drought resilience over multiple years in tropical seedling communities.

Precipitation patterns are changing across the globe causing more severe and frequent drought for many forest ecosystems. Although research has focused on the resistance of tree populations and communities to these novel precipitation regimes, resilience of forests is also contingent on recovery following drought, which remains poorly understood, especially in aseasonal tropical forests. We used rainfall exclusion shelters to manipulate the interannual frequency of drought for diverse seedling communities in a tropical forest and assessed resistance, recovery and resilience of seedling growth and mortality relative to everwet conditions. We found seedlings exposed to recurrent periods of drought altered their growth rates throughout the year relative to seedlings in everwet conditions. During drought periods, seedlings grew slower than seedlings in everwet conditions (i.e., resistance phase) while compensating with faster growth after drought (i.e., recovery phase). However, the response to frequent drought was species dependent as some species grew significantly slower with frequent drought relative to everwet conditions while others grew faster with frequent drought due to overcompensating growth during the recovery phase. In contrast, mortality was unrelated to rainfall conditions and instead correlated with differences in light. Intra-annual plasticity of growth and increased annual growth of some species led to an overall maintenance of growth rates of tropical seedling communities in response to more frequent drought. These results suggest these communities can potentially adapt to predicted climate change scenarios and that plasticity in the growth of species, and not solely changes in mortality rates among species, may contribute to shifts in community composition under drought.

Responses of lichen communities to 18 years of natural and experimental warming.

Background and Aims: Climate change is expected to have major impacts on high alpine and arctic ecosystems in the future, but empirical data on the impact of long-term warming on lichen diversity and richness are sparse. This study report the effects of 18 years of ambient and experimental warming on lichens and vascular plant cover in two alpine plant communities, a dry heath with sparse canopy cover (54 %) and a mesic meadow with a more developed (67 %) canopy cover, in sub-arctic Sweden. Methods: The effects of long-term passive experimental warming using open top chambers (OTCs) on lichens and total vascular plant cover, and the impact of plant cover on lichen community parameters, were analysed. Key Results: Between 1993 and 2013, mean annual temperature increased about 2 °C. Both site and experimental warming had a significant effect on cover, species richness, effective number of species evenness of lichens, and total plant canopy cover. Lichen cover increased in the heath under ambient conditions, and remained more stable under experimental warming. The negative effect on species richness and effective number of species was driven by a decrease in lichens under experimental warming in the meadow. Lichen cover, species richness, effective number of species evenness were negatively correlated with plant canopy cover. There was a significant negative impact on one species and a non-significant tendency of lower abundance of the most common species in response to experimental warming. Conclusions: The results from the long-term warming study imply that arctic and high alpine lichen communities are likely to be negatively affected by climate change and an increase in plant canopy cover. Both biotic and abiotic factors are thus important for future impacts of climate change on lichens.

The contrasting effects of short-term climate change on the early recruitment of tree species.

Predictions of plant responses to climate change are frequently based on organisms' presence in warmer locations, which are then assumed to reflect future performance in cooler areas. However, as plant life stages may be affected differently by environmental changes, there is little empirical evidence that this approach provides reliable estimates of short-term responses to global warming. Under this premise, we analyzed 8 years of early recruitment data, seed production and seedling establishment and survival, collected for two tree species at two latitudes. We quantified recruitment to a wide range of environmental conditions, temperature, soil moisture and light, and simulated recruitment under two forecasted climatic scenarios. Annual demographic transitions were affected by the particular conditions taking place during their onset, but the effects of similar environmental shifts differed among the recruitment stages; seed production was higher in warmer years, while seedling establishment and survival peaked during cold years. Within a species, these effects also varied between latitudes; increasing temperatures at the southern location will have stronger detrimental effects on recruitment than similar changes at the northern locations. Our simulations illustrate that warmer temperatures may increase seed production, but they will have a negative effect on establishment and survival. When the three early recruitment processes were simultaneously considered, simulations showed little change in recruitment dynamics at the northern site and a slight decrease at the southern site. It is only when we considered these three stages that we were able to assess likely changes in early recruitment under the predicted conditions.

Testing the large genome constraint hypothesis: plant traits, habitat and climate seasonality in Liliaceae.

The factors driving genome size evolution in Liliaceae were examined. In particular, we investigated whether species with larger genomes are confined to less stressful environments with a longer vegetative season. We tested our hypotheses by correlating the genome size with other plant traits and environmental variables. To determine the adaptive nature of the genome size, we also compared the performances of Brownian motion (BM) processes with those inferred by Ornstein-Uhlenbeck (OU) models of trait evolution. A positive correlation of genome size with plant size, mean temperature and habitat moisture and a negative correlation with altitude and precipitation seasonality were found. Models of trait evolution revealed a deviation from a drift process or BM. Instead, changes in genome size were significantly associated with precipitation regimes according to an OU process. Specifically, the evolutionary optima towards which the genome size evolves were higher for humid climates and lower for drier ones. Taken together, our results indicate that the genome size increase in Liliaceae is constrained by climate seasonality.

Contrasting nonstructural carbohydrate dynamics of tropical tree seedlings under water deficit and variability.

Drought regimes can be characterized by the variability in the quantity of rainfall and the duration of rainless periods. However, most research on plant response to drought has ignored the impacts of rainfall variation, especially with regard to the influence of nonstructural carbohydrates (NSCs) in promoting drought resistance. To test the hypothesis that these components of drought differentially affect NSC dynamics and seedling resistance, we tracked NSC in plant tissues of tropical tree seedlings in response to manipulations of the volume and frequency of water applied. NSC concentrations decreased in woody tissues under infrequent-high watering but increased under no watering. A faster decline of growth relative to stomatal conductance in the no watering treatment was consistent with NSC accumulation as a result of an uncoupling of growth and photosynthesis, while usage of stored NSCs in woody tissues to maintain function may account for the NSC decline under infrequent-high watering. NSCs, and specifically stem NSCs, contributed to drought resistance under severe water deficits, while NSCs had a less clear role in drought resistance to variability in water availability. The contrasting response of NSCs to water variability and deficit indicates that unique processes support seedling resistance to these components of drought.

Simulating the germination response to diurnally alternating temperatures under climate change scenarios: comparative studies on Carex diandra seeds.

BACKGROUND AND AIMS: Environmental temperature regulates plant regeneration via seed in several superimposed ways, and this complex regulation will be disrupted by climate change. The role of diurnally alternating temperatures (ΔT) in terminating dormancy will be a major factor in this disruption, as its effects on seed germination are immediate. METHODS: The effect of ΔT on seed germination was modelled using two populations of the wetland sedge Carex diandra, one from a montane site and one from a subalpine site. A cardinal-temperature model was fitted to germination results obtained from a thermal gradient plate, and the model was used to simulate changes in germination under two possible future climate scenarios (RCP2·6 and RCP8·5, for representative concentration pathways) as defined by the Intergovernmental Panel on Climate Change. KEY RESULTS: Scenario RCP2·6 projected moderate increases in average temperatures and ΔT, whereas RCP8·5 projected greater warming and higher ΔT. Increasing ΔT decreased the base temperature for seed germination and the thermal time required for germination. The effect of higher ΔT together with the higher temperatures increased germination under both climate scenarios. CONCLUSIONS: Carex diandra germination is highly responsive to potential changes in ΔT, and thus this study highlights the role of ΔT in seed responses to climate change. Comprehensive cardinal-temperature models, encompassing the different effects of temperature on seed germination, are needed to understand how climate change will affect plant regeneration.

Biotic context and soil properties modulate native plant responses to enhanced rainfall.

BACKGROUND AND AIMS: The environmental and biotic context within which plants grow have a great potential to modify responses to climatic changes, yet few studies have addressed both the direct effects of climate and the modulating roles played by variation in the biotic (e.g. competitors) and abiotic (e.g. soils) environment. METHODS: In a grassland with highly heterogeneous soils and community composition, small seedlings of two native plants, Lasthenia californica and Calycadenia pauciflora, were transplanted into factorially watered and fertilized plots. Measurements were made to test how the effect of climatic variability (mimicked by the watering treatment) on the survival, growth and seed production of these species was modulated by above-ground competition and by edaphic variables. KEY RESULTS: Increased competition outweighed the direct positive impacts of enhanced rainfall on most fitness measures for both species, resulting in no net effect of enhanced rainfall. Both species benefitted from enhanced rainfall when the absence of competitors was accompanied by high soil water retention capacity. Fertilization did not amplify the watering effects; rather, plants benefitted from enhanced rainfall or competitor removal only in ambient nutrient conditions with high soil water retention capacity. CONCLUSIONS: The findings show that the direct effects of climatic variability on plant fitness may be reversed or neutralized by competition and, in addition, may be strongly modulated by soil variation. Specifically, coarse soil texture was identified as a factor that may limit plant responsiveness to altered water availability. These results highlight the importance of considering the abiotic as well as biotic context when making future climate change forecasts.