Extreme drought impacts have been underestimated in grasslands and shrublands globally.

Climate change is increasing the frequency and severity of short-term (~1 y) drought events-the most common duration of drought-globally. Yet the impact of this intensification of drought on ecosystem functioning remains poorly resolved. This is due in part to the widely disparate approaches ecologists have employed to study drought, variation in the severity and duration of drought studied, and differences among ecosystems in vegetation, edaphic and climatic attributes that can mediate drought impacts. To overcome these problems and better identify the factors that modulate drought responses, we used a coordinated distributed experiment to quantify the impact of short-term drought on grassland and shrubland ecosystems. With a standardized approach, we imposed ~a single year of drought at 100 sites on six continents. Here we show that loss of a foundational ecosystem function-aboveground net primary production (ANPP)-was 60% greater at sites that experienced statistically extreme drought (1-in-100-y event) vs. those sites where drought was nominal (historically more common) in magnitude (35% vs. 21%, respectively). This reduction in a key carbon cycle process with a single year of extreme drought greatly exceeds previously reported losses for grasslands and shrublands. Our global experiment also revealed high variability in drought response but that relative reductions in ANPP were greater in drier ecosystems and those with fewer plant species. Overall, our results demonstrate with unprecedented rigor that the global impacts of projected increases in drought severity have been significantly underestimated and that drier and less diverse sites are likely to be most vulnerable to extreme drought.

Adjustments in physiological and morphological traits suggest drought-induced competitive release of some California plants.

Drought and competition affect how morphological and physiological traits are expressed in plants. California plants were previously found to respond less negatively to resource limitation compared to invasive counterparts. In a glasshouse in Santa Cruz, CA, USA, we exposed five native California C3 grassland species to episodic drought and competition (via five locally invasive species). We hypothesized that leaf morphology would be more affected by competition, and leaf photosynthetic gas exchange more so by drought, consistent with optimal partitioning and environmental filter theories. We expected that traits would exhibit trade-offs along a spectrum for resource conservatism versus acquisition. Bromus carinatus had greater photosynthetic recovery, while Diplacus aurantiacus had lower percent loss of net assimilation (PLA) and intrinsic water-use efficiency (iWUE) during drought and competition simultaneously compared to just drought. Stipa pulchra and Sidalcea malviflora gas exchange was unaffected by drought, and leaf morphology exhibited drought-related adjustments. Lupinus nanus exhibited trait adjustments for competition but not drought. Functional traits sorted onto two principal components related to trade-offs for resource conservatism versus acquisition, and for above- versus belowground allocation. In summary, morphological traits were affected by competition and drought, whereas physiological traits, like leaf gas exchange, were primarily affected by drought. The grassland plants we studied showed diverse responses to drought and competition with trait trade-offs related to resource conservatism versus acquisition, and for above- versus belowground allocation consistent with optimal partitioning and environmental filter theories. Diplacus aurantiacus experienced competitive release based on greater iWUE and lower PLA when facing drought and competition.

Photosynthetic sensitivity to historic meteorological variability for conifers in the eastern Sierra Nevada.

Increased climatic variability can impact tree physiological processes beyond what is predicted from changes in mean conditions. We assessed the sensitivity of conifer saplings to spatial and temporal variability in meteorological conditions, taking advantage of the end of California's historic drought and the exceedingly wet winter of 2017. We sought to understand how very dry and very wet conditions constrain photosynthesis and growth in four regionally dominant conifers and whether sensitivity in these processes changes across a 500 m gradient in elevation. All species demonstrated phenotypic plasticity in response to temporal differences in precipitation on both inter-annual and seasonal timescales. Net photosynthesis in Pinus contorta decreased from an early season 2016 average of 12.4 to 6.89 µmol CO2 m-2 s-1 later in the summer, but increased 14.1% between seasons in the wet year. By contrast, elevation had almost no effect on instantaneous photosynthetic gas exchange, CO2 response curve parameters, or stem water potential in any of the years for any of the species. Effects of the heavy snow year (2017) on needle growth differed between elevations. Pinus contorta showed a 38.9% increase in average needle length at the lower two elevations but a 31.6% decrease at the highest site compared to the height of the drought. Despite these differences, biological variation was dampened compared to the physical variation between years, suggesting these trees can effectively withstand substantial meteorological variability. Our results show that these species demonstrated considerable ability to tolerate and recover from an extreme drought event.

Tropicalization of temperate ecosystems in North America: The northward range expansion of tropical organisms in response to warming winter temperatures.

Tropicalization is a term used to describe the transformation of temperate ecosystems by poleward-moving tropical organisms in response to warming temperatures. In North America, decreases in the frequency and intensity of extreme winter cold events are expected to allow the poleward range expansion of many cold-sensitive tropical organisms, sometimes at the expense of temperate organisms. Although ecologists have long noted the critical ecological role of winter cold temperature extremes in tropical-temperate transition zones, the ecological effects of extreme cold events have been understudied, and the influence of warming winter temperatures has too often been left out of climate change vulnerability assessments. Here, we examine the influence of extreme cold events on the northward range limits of a diverse group of tropical organisms, including terrestrial plants, coastal wetland plants, coastal fishes, sea turtles, terrestrial reptiles, amphibians, manatees, and insects. For these organisms, extreme cold events can lead to major physiological damage or landscape-scale mass mortality. Conversely, the absence of extreme cold events can foster population growth, range expansion, and ecological regime shifts. We discuss the effects of warming winters on species and ecosystems in tropical-temperate transition zones. In the 21st century, climate change-induced decreases in the frequency and intensity of extreme cold events are expected to facilitate the poleward range expansion of many tropical species. Our review highlights critical knowledge gaps for advancing understanding of the ecological implications of the tropicalization of temperate ecosystems in North America.

Nitrogen Addition Increases the Sensitivity of Photosynthesis to Drought and Re-watering Differentially in C3 Versus C4 Grass Species.

Global change factors, such as variation in precipitation regimes and nitrogen (N) deposition, are likely to occur simultaneously and may have profound impacts on the relative abundance of grasses differing in functional traits, such as C3 and C4 species. We conducted an extreme drought and re-watering experiment to understand differences in the resistance and recovery abilities of C3 and C4 grasses under different N deposition scenarios. A C3 perennial grass (Leymus chinensis) and two C4 grasses (annual species Chloris virgata and perennial species Hemarthria altissima) that co-occur in Northeast China were selected as experimental plants. For both C3 and C4 grasses, N addition caused a strong increase in biomass and resulted in more severe drought stress, leading to a change in the dominant photosynthetic limitation during the drought periods. Although N addition increased antioxidant enzyme activities and protective solute concentrations, the carbon fixing capacity did not fully recover to pre-drought levels by the end of the re-watering period. N addition resulted in lower resilience under the drought conditions and lower resistance at the end of the re-watering. However, N addition led to faster recovery of photosynthesis, especially in the C3 grass, which indicate that the effect of N addition on photosynthesis during drought was asymmetric, especially in the plants with different photosynthetic nitrogen use efficiency (PNUE). These findings demonstrated that nitrogen deposition may significant alter the susceptibility of C3 and C4 grass species to drought stress and re-watering, highlighting the asymmetry between resistance and resilience and to improve our understanding about plant responses to climate change.

Photosynthesis and carbon allocation are both important predictors of genotype productivity responses to elevated CO2 in Eucalyptus camaldulensis.

Intraspecific variation in biomass production responses to elevated atmospheric carbon dioxide (eCO2) could influence tree species' ecological and evolutionary responses to climate change. However, the physiological mechanisms underlying genotypic variation in responsiveness to eCO2 remain poorly understood. In this study, we grew 17 Eucalyptus camaldulensis Dehnh. subsp. camaldulensis genotypes (representing provenances from four different climates) under ambient atmospheric CO2 and eCO2. We tested whether genotype leaf-scale photosynthetic and whole-tree carbon (C) allocation responses to eCO2 were predictive of genotype biomass production responses to eCO2. Averaged across genotypes, growth at eCO2 increased in situ leaf net photosynthesis (Anet) (29%) and leaf starch concentrations (37%). Growth at eCO2 reduced the maximum carboxylation capacity of Rubisco (-4%) and leaf nitrogen per unit area (Narea, -6%), but Narea calculated on a total non-structural carbohydrate-free basis was similar between treatments. Growth at eCO2 also increased biomass production and altered C allocation by reducing leaf area ratio (-11%) and stem mass fraction (SMF, -9%), and increasing leaf mass area (18%) and leaf mass fraction (5%). Overall, we found few significant CO2 × provenance or CO2 × genotype (within provenance) interactions. However, genotypes that showed the largest increases in total dry mass at eCO2 had larger increases in root mass fraction (with larger decreases in SMF) and photosynthetic nitrogen-use efficiency (PNUE) with CO2 enrichment. These results indicate that genetic differences in PNUE and carbon sink utilization (in roots) are both important predictors of tree productivity responsiveness to eCO2.

Relationships between climate of origin and photosynthetic responses to an episodic heatwave depend on growth CO2 concentration for Eucalyptus camaldulensis var. camaldulensis.

Stressful episodic weather is likely to affect the C balance of trees as the climate changes, potentially altering survival. However, the role of elevated CO2 concentration ([CO2]) in tolerating off-season episodic extremes is not clear. We tested for interactive effects of elevated CO2 and springtime heat stress on photosynthesis for seven genotypes of Eucalyptus camaldulensis Dehnh. var. camaldulensis, representing its widespread distribution across south-eastern Australia. We grew clonal material under glasshouse conditions of ambient (aCO2; 400 parts per million (ppm)) or elevated (eCO2; 640ppm) [CO2], and air temperatures of 25:17°C (day:night), and measured the electron transport rate in PSII (ETR), stomatal conductance to water vapour (gs) and net CO2 assimilation (A). Measurements were made before, during and after a four-day temperature excursion of 35:27°C. ETR and A were ~17% higher for plants grown in eCO2 than in aCO2. Photosynthesis remained stable for plants in eCO2 during the heatwave. Based on the effect size ratio (eCO2:aCO2), gs and ETR were temporarily affected more by the heatwave than A. A reduction in ETR in eCO2 was the only lasting effect of the heatwave. There were no significant differences among genotypes. Correlations between photosynthesis and climate of origin differed for plants grown in aCO2 compared with eCO2, suggesting potential complex and multiple control points on photosynthesis.

Pushing precipitation to the extremes in distributed experiments: recommendations for simulating wet and dry years.

Intensification of the global hydrological cycle, ranging from larger individual precipitation events to more extreme multiyear droughts, has the potential to cause widespread alterations in ecosystem structure and function. With evidence that the incidence of extreme precipitation years (defined statistically from historical precipitation records) is increasing, there is a clear need to identify ecosystems that are most vulnerable to these changes and understand why some ecosystems are more sensitive to extremes than others. To date, opportunistic studies of naturally occurring extreme precipitation years, combined with results from a relatively small number of experiments, have provided limited mechanistic understanding of differences in ecosystem sensitivity, suggesting that new approaches are needed. Coordinated distributed experiments (CDEs) arrayed across multiple ecosystem types and focused on water can enhance our understanding of differential ecosystem sensitivity to precipitation extremes, but there are many design challenges to overcome (e.g., cost, comparability, standardization). Here, we evaluate contemporary experimental approaches for manipulating precipitation under field conditions to inform the design of 'Drought-Net', a relatively low-cost CDE that simulates extreme precipitation years. A common method for imposing both dry and wet years is to alter each ambient precipitation event. We endorse this approach for imposing extreme precipitation years because it simultaneously alters other precipitation characteristics (i.e., event size) consistent with natural precipitation patterns. However, we do not advocate applying identical treatment levels at all sites - a common approach to standardization in CDEs. This is because precipitation variability varies >fivefold globally resulting in a wide range of ecosystem-specific thresholds for defining extreme precipitation years. For CDEs focused on precipitation extremes, treatments should be based on each site's past climatic characteristics. This approach, though not often used by ecologists, allows ecological responses to be directly compared across disparate ecosystems and climates, facilitating process-level understanding of ecosystem sensitivity to precipitation extremes.

Water relations and photosynthesis along an elevation gradient for Artemisia tridentata during an historic drought.

Quantifying the variation in plant-water relations and photosynthesis over environmental gradients and during unique events can provide a better understanding of vegetation patterns in a future climate. We evaluated the hypotheses that photosynthesis and plant water potential would correspond to gradients in precipitation and soil moisture during a lengthy drought, and that experimental water additions would increase photosynthesis for the widespread evergreen shrub Artemisia tridentata ssp. vaseyana. We quantified abiotic conditions and physiological characteristics for control and watered plants at 2135, 2315, and 2835 m near Mammoth Lakes, CA, USA, at the ecotone of the Sierra Nevada and Great Basin ecoregions. Snowfall, total precipitation, and soil moisture increased with elevation, but air temperature and soil N content did not. Plant water potential (Ψ), stomatal conductance (g s), maximum photosynthetic rate (A max), carboxylation rate (V cmax), and electron transport rate (J max) all significantly increased with elevations. Addition of water increased Ψ, g s, J max, and A max only at the lowest elevation; g s contributed about 30 % of the constraints on photosynthesis at the lowest elevation and 23 % at the other two elevations. The physiology of this foundational shrub species was quite resilient to this 1-in-1200 year drought. However, plant water potential and photosynthesis corresponded to differences in soil moisture across the gradient. Soil re-wetting in early summer increased water potential and photosynthesis at the lowest elevation. Effects on water relations and photosynthesis of this widespread, cold desert shrub species may be disproportionate at lower elevations as drought length increases in a future climate.

Genetic variation in circadian regulation of nocturnal stomatal conductance enhances carbon assimilation and growth.

Circadian resonance, whereby a plant's endogenous rhythms are tuned to match environmental cues, has been repeatedly shown to be adaptive, although the underlying mechanisms remain elusive. Concomitantly, the adaptive value of nocturnal transpiration in C3 plants remains unknown because it occurs without carbon assimilation. These seemingly unrelated processes are interconnected because circadian regulation drives temporal patterns in nocturnal stomatal conductance, with maximum values occurring immediately before dawn for many species. We grew individuals of six Eucalyptus camaldulensis genotypes in naturally lit glasshouses and measured sunset, predawn and midday leaf gas exchange and whole-plant biomass production. We tested whether sunrise anticipation by the circadian clock and subsequent increases in genotype predawn stomatal conductance led to rapid stomatal opening upon illumination, ultimately affecting genotype differences in carbon assimilation and growth. We observed faster stomatal responses to light inputs at sunrise in genotypes with higher predawn stomatal conductance. Moreover, early morning and midday stomatal conductance and carbon assimilation, leaf area and total plant biomass were all positively correlated with predawn stomatal conductance across genotypes. Our results lead to the novel hypothesis that genotypic variation in the circadian-regulated capacity to anticipate sunrise could be an important factor underlying intraspecific variation in tree growth.

Characterizing differences in precipitation regimes of extreme wet and dry years: implications for climate change experiments.

Climate change is intensifying the hydrologic cycle and is expected to increase the frequency of extreme wet and dry years. Beyond precipitation amount, extreme wet and dry years may differ in other ways, such as the number of precipitation events, event size, and the time between events. We assessed 1614 long-term (100 year) precipitation records from around the world to identify key attributes of precipitation regimes, besides amount, that distinguish statistically extreme wet from extreme dry years. In general, in regions where mean annual precipitation (MAP) exceeded 1000 mm, precipitation amounts in extreme wet and dry years differed from average years by ~40% and 30%, respectively. The magnitude of these deviations increased to >60% for dry years and to >150% for wet years in arid regions (MAP<500 mm). Extreme wet years were primarily distinguished from average and extreme dry years by the presence of multiple extreme (large) daily precipitation events (events >99th percentile of all events); these occurred twice as often in extreme wet years compared to average years. In contrast, these large precipitation events were rare in extreme dry years. Less important for distinguishing extreme wet from dry years were mean event size and frequency, or the number of dry days between events. However, extreme dry years were distinguished from average years by an increase in the number of dry days between events. These precipitation regime attributes consistently differed between extreme wet and dry years across 12 major terrestrial ecoregions from around the world, from deserts to the tropics. Thus, we recommend that climate change experiments and model simulations incorporate these differences in key precipitation regime attributes, as well as amount into treatments. This will allow experiments to more realistically simulate extreme precipitation years and more accurately assess the ecological consequences.

Impact of intra- versus inter-annual snow depth variation on water relations and photosynthesis for two Great Basin Desert shrubs.

Snowfall provides the majority of soil water in certain ecosystems of North America. We tested the hypothesis that snow depth variation affects soil water content, which in turn drives water potential (Ψ) and photosynthesis, over 10 years for two widespread shrubs of the western USA. Stem Ψ (Ψ stem) and photosynthetic gas exchange [stomatal conductance to water vapor (g s), and CO2 assimilation (A)] were measured in mid-June each year from 2004 to 2013 for Artemisia tridentata var. vaseyana (Asteraceae) and Purshia tridentata (Rosaceae). Snow fences were used to create increased or decreased snow depth plots. Snow depth on +snow plots was about twice that of ambient plots in most years, and 20 % lower on -snow plots, consistent with several down-scaled climate model projections. Maximal soil water content at 40- and 100-cm depths was correlated with February snow depth. For both species, multivariate ANOVA (MANOVA) showed that Ψ stem, g s, and A were significantly affected by intra-annual variation in snow depth. Within years, MANOVA showed that only A was significantly affected by spatial snow depth treatments for A. tridentata, and Ψ stem was significantly affected by snow depth for P. tridentata. Results show that stem water relations and photosynthetic gas exchange for these two cold desert shrub species in mid-June were more affected by inter-annual variation in snow depth by comparison to within-year spatial variation in snow depth. The results highlight the potential importance of changes in inter-annual variation in snowfall for future shrub photosynthesis in the western Great Basin Desert.

Utilizing intraspecific variation in phenotypic plasticity to bolster agricultural and forest productivity under climate change.

Climate change threatens the ability of agriculture and forestry to meet growing global demands for food, fibre and wood products. Information gathered from genotype-by-environment interactions (G × E), which demonstrate intraspecific variation in phenotypic plasticity (the ability of a genotype to alter its phenotype in response to environmental change), may prove important for bolstering agricultural and forest productivity under climate change. Nonetheless, very few studies have explicitly quantified genotype plasticity-productivity relationships in agriculture or forestry. Here, we conceptualize the importance of intraspecific variation in agricultural and forest species plasticity, and discuss the physiological and genetic factors contributing to intraspecific variation in phenotypic plasticity. Our discussion highlights the need for an integrated understanding of the mechanisms of G × E, more extensive assessments of genotypic responses to climate change under field conditions, and explicit testing of genotype plasticity-productivity relationships. Ultimately, further investigation of intraspecific variation in phenotypic plasticity in agriculture and forestry may prove important for identifying genotypes capable of increasing or sustaining productivity under more extreme climatic conditions.

Quantifying ecological memory in plant and ecosystem processes.

The role of time in ecology has a long history of investigation, but ecologists have largely restricted their attention to the influence of concurrent abiotic conditions on rates and magnitudes of important ecological processes. Recently, however, ecologists have improved their understanding of ecological processes by explicitly considering the effects of antecedent conditions. To broadly help in studying the role of time, we evaluate the length, temporal pattern, and strength of memory with respect to the influence of antecedent conditions on current ecological dynamics. We developed the stochastic antecedent modelling (SAM) framework as a flexible analytic approach for evaluating exogenous and endogenous process components of memory in a system of interest. We designed SAM to be useful in revealing novel insights promoting further study, illustrated in four examples with different degrees of complexity and varying time scales: stomatal conductance, soil respiration, ecosystem productivity, and tree growth. Models with antecedent effects explained an additional 18-28% of response variation compared to models without antecedent effects. Moreover, SAM also enabled identification of potential mechanisms that underlie components of memory, thus revealing temporal properties that are not apparent from traditional treatments of ecological time-series data and facilitating new hypothesis generation and additional research.

Maritime climate influence on chaparral composition and diversity in the coast range of central California.

We investigated the hypothesis that maritime climatic factors associated with summer fog and low cloud stratus (summer marine layer) help explain the compositional diversity of chaparral in the coast range of central California. We randomly sampled chaparral species composition in 0.1-hectare plots along a coast-to-interior gradient. For each plot, climatic variables were estimated and soil samples were analyzed. We used Cluster Analysis and Principle Components Analysis to objectively categorize plots into climate zone groups. Climate variables, vegetation composition and various diversity measures were compared across climate zone groups using ANOVA and nonmetric multidimensional scaling. Differences in climatic variables that relate to summer moisture availability and winter freeze events explained the majority of variance in measured conditions and coincided with three chaparral assemblages: maritime (lowland coast where the summer marine layer was strongest), transition (upland coast with mild summer marine layer influence and greater winter precipitation), and interior sites that generally lacked late summer water availability from either source. Species turnover (ß-diversity) was higher among maritime and transition sites than interior sites. Coastal chaparral differs from interior chaparral in having a higher obligate seeder to facultative seeder (resprouter) ratio and by being dominated by various Arctostaphylos species as opposed to the interior dominant, Adenostoma fasciculatum. The maritime climate influence along the California central coast is associated with patterns of woody plant composition and ß-diversity among sites. Summer fog in coastal lowlands and higher winter precipitation in coastal uplands combine to lower late dry season water deficit in coastal chaparral and contribute to longer fire return intervals that are associated with obligate seeders and more local endemism. Soil nutrients are comparatively less important in explaining plant community composition, but heterogeneous azonal soils contribute to local endemism and promote isolated chaparral patches within the dominant forest vegetation along the coast.

Soil microbial and nutrient responses to 7 years of seasonally altered precipitation in a Chihuahuan Desert grassland.

Soil microbial communities in Chihuahuan Desert grasslands generally experience highly variable spatiotemporal rainfall patterns. Changes in precipitation regimes can affect belowground ecosystem processes such as decomposition and nutrient cycling by altering soil microbial community structure and function. The objective of this study was to determine if increased seasonal precipitation frequency and magnitude over a 7-year period would generate a persistent shift in microbial community characteristics and soil nutrient availability. We supplemented natural rainfall with large events (one/winter and three/summer) to simulate increased precipitation based on climate model predictions for this region. We observed a 2-year delay in microbial responses to supplemental precipitation treatments. In years 3-5, higher microbial biomass, arbuscular mycorrhizae abundance, and soil enzyme C and P acquisition activities were observed in the supplemental water plots even during extended drought periods. In years 5-7, available soil P was consistently lower in the watered plots compared to control plots. Shifts in soil P corresponded to higher fungal abundances, microbial C utilization activity, and soil pH. This study demonstrated that 25% shifts in seasonal rainfall can significantly influence soil microbial and nutrient properties, which in turn may have long-term effects on nutrient cycling and plant P uptake in this desert grassland.

Photosynthesis of seedlings of Otoba novogranatensis (Myristicaceae) and Ruagea glabra (Meliaceae) in abandoned pasture, secondary forest and plantation habitats in Costa Rica.

Enrichment planting in naturally recovering secondary forests or in tree plantations is increasingly being used as strategy to restore later-successional, large-seeded tropical forest trees. We seeded two tree species (Otoba novogranatensis and Ruagea glabra) in three agricultural sites in Southern Costa Rica: abandoned pastures, eight to ten year old secondary forests and three year old tree plantations (containing two N-fixing of four total tree species). We measured micrometeorological conditions, soil water content, plant water potential, leaf area, foliar C and N, and photosynthesis to better understand mechanistic responses of seedlings to conditions in the different successional habitats. Micrometeorological conditions, soil water content, and plant water potential were generally similar across habitats. Certain aspects of leaves (such as Specific Leaf Area and foliar N content), and photosynthesis (e.g. quantum yield and electron transport rate) were highest in the plantations, intermediate in the secondary forests, and lowest in abandoned pastures. Enhanced rates of photosynthetic biochemistry (such as Vxmax and Jmax) and Photosystem II efficiency (e.g. thermal energy dissipation) occurred in leaves from the plantations compared to the abandoned pastures, which may be related to higher leaf %N content. Results suggest that foliar N may be of greater importance than soil water content and micrometeorological factors in driving differences in photosynthetic processes across planting habitats. Planting seeds of these two species in plantations containing three year old trees (including two N-fixing species) enhances certain aspects of their photosynthesis and growth, compared to seedlings in abandoned pastures with non-native grasses, and thus can help increase forest recovery on abandoned agricultural lands.

Photosynthesis of seedlings of Otoba novogranatensis (Myristicaceae) and Ruagea glabra (Meliaceae) in abandoned pasture, secondary forest and plantation habitats in Costa Rica / Fotosíntesis de plántulas de Otoba novogranatensis (Myristicaceae) y Ruagea glabra (Meliaceae) en pastizales abandonados, bosques secundarios y hábitats de plantaciones en Costa Rica

Enrichment planting in naturally recovering secondary forests or in tree plantations is increasingly being used as strategy to restore later-successional, large-seeded tropical forest trees. We seeded two tree species (Otoba novogranatensis and Ruagea glabra) in three agricultural sites in Southern Costa Rica: abandoned pastures, eight to ten year old secondary forests and three year old tree plantations (containing two N-fixing of four total tree species). We measured micrometeorological conditions, soil water content, plant water potential, leaf area, foliar C and N, and photosynthesis to better understand mechanistic responses of seedlings to conditions in the different successional habitats. Micrometeorological conditions, soil water content, and plant water potential were generally similar across habitats. Certain aspects of leaves (such as Specific Leaf Area and foliar N content), and photosynthesis (e.g. quantum yield and electron transport rate) were highest in the plantations, intermediate in the secondary forests, and lowest in abandoned pastures. Enhanced rates of photosynthetic biochemistry (such as Vcmax and Jmax) and Photosystem II efficiency (e.g. thermal energy dissipation) occurred in leaves from the plantations compared to the abandoned pastures, which may be related to higher leaf %N content. Results suggest that foliar N may be of greater importance than soil water content and micrometeorological factors in driving differences in photosynthetic processes across planting habitats. Planting seeds of these two species in plantations containing three year old trees (including two N-fixing species) enhances certain aspects of their photosynthesis and growth, compared to seedlings in abandoned pastures with non-native grasses, and thus can help increase forest recovery on abandoned agricultural lands.

El enriquecimiento de bosques secundarios o plantaciones forestales en proceso de regeneración natural por medio de la siembra de plántulas es una práctica cada vez más utilizada para restaurar bosques tropicales en estado de sucesión tardía. Sembramos dos especies de árboles (Otoba novogranatensis y Ruagea glabra) en pastizales abandonados, bosques secundarios de ocho a diez años de edad y plantaciones forestales de tres años de edad (con dos especies fijadoras de Nitrógeno de un total de cuatro especies) en tres sitios agrícolas en el Sur de Costa Rica. Medimos condiciones micrometeorológicas, contenido de agua del suelo, potencial hídrico de las plantas, área foliar, C y N foliar, y fotosíntesis para entender de una mejor manera las respuestas funcionales de las plántulas ante condiciones de distintos estadíos sucesionales. Las condiciones micrometeorológicas, contenido hídrico del suelo y el potencial hídrico de las plantas fueron mayoritariamente similares entre hábitats. Algunos aspectos de las hojas (como Área Foliar Específica y contenido de N foliar) y fotosíntesis (ej.: rendimiento cuántico y tasa de transporte de electrones) presentaron valores mayores en las plantaciones, intermedios en los bosques secundarios y menores en los pastizales abandonados. Se obtuvo un aumento en las tasas fotosintéticas bioquímicas (como Vcmax, Jmax) y la eficiencia del Fotosistema II (ej.: disipación de energía térmica) en hojas provenientes de las plantaciones comparado a las de los pastizales, posiblemente relacionado a un mayor %N foliar. Los resultados sugieren que el N foliar puede ser más importante que el contenido de agua del suelo y que los factores micrometeorológicos para marcar diferencias en los procesos fotosintéticos entre hábitats. Las plántulas de estas dos especies en las plantaciones con árboles de tres años de edad (incluyendo dos fijadoras de N) incrementaron ciertos aspectos de su fotosíntesis y crecimiento comparado a las plántulas en los pastizales abandonados de especies exóticas, por lo tanto, esta práctica puede ayudar a incrementar la recuperación de los bosques en áreas agrícolas abandonadas.

Global change effects on Bromus tectorum L. (Poaceae) at its high-elevation range margin.

Global change is likely to affect invasive species distribution, especially at range margins. In the eastern Sierra Nevada, California, USA, the invasive annual grass, Bromus tectorum, is patchily distributed and its impacts have been minimal compared with other areas of the Intermountain West. We used a series of in situ field manipulations to determine how B. tectorum might respond to changing climatic conditions and increased nitrogen deposition at the high-elevation edge of its invaded range. Over 3 years, we used snow fences to simulate changes in snowpack, irrigation to simulate increased frequency and magnitude of springtime precipitation, and added nitrogen (N) at three levels (0, 5, and 10 g m(-2) ) to natural patches of B. tectorum growing under the two dominant shrubs, Artemisia tridentata and Purshia tridentata, and in intershrub spaces (INTR). We found that B. tectorum seedling density in April was lower following deeper snowpack possibly due to delayed emergence, yet there was no change in spikelet production or biomass accumulation at the time of harvest. Additional spring rain events increased B. tectorum biomass and spikelet production in INTR plots only. Plants were primarily limited by water in 2009, but colimited by N and water in 2011, possibly due to differences in antecedent moisture conditions at the time of treatments. The threshold at which N had an effect varied with magnitude of water additions. Frequency of rain events was more influential than magnitude in driving B. tectorum growth and fecundity responses. Our results suggest that predicted shifts from snow to rain could facilitate expansion of B. tectorum at high elevation depending on timing of rain events and level of N deposition. We found evidence for P-limitation at this site and an increase in P-availability with N additions, suggesting that stoichiometric relationships may also influence B. tectorum spread.

Influence of summer marine fog and low cloud stratus on water relations of evergreen woody shrubs (Arctostaphylos: Ericaceae) in the chaparral of central California.

Mediterranean-type climate (MTC) regions around the world are notable for cool, wet winters and hot, dry summers. A dominant vegetation type in all five MTC regions is evergreen, sclerophyllous shrubland, called chaparral in California. The extreme summer dry season in California is moderated by a persistent low-elevation layer of marine fog and cloud cover along the margin of the Pacific coast. We tested whether late dry season water potentials (Ψ(min)) of chaparral shrubs, such as Arctostaphylos species in central California, are influenced by this coast-to-interior climate gradient. Lowland coastal (maritime) shrubs were found to have significantly less negative Ψ(min) than upland interior shrubs (interior), and stable isotope (δ(13)C) values exhibited greater water use efficiency in the interior. Post-fire resprouter shrubs (resprouters) had significantly less negative Ψ(min) than co-occurring obligate seeder shrubs (seeders) in interior and transitional chaparral, possibly because resprouters have deeper root systems with better access to subsurface water than shallow-rooted seeders. Unexpectedly, maritime resprouters and seeders did not differ significantly in their Ψ(min), possibly reflecting more favorable water availability for shrubs influenced by the summer marine layer. Microclimate and soil data also suggest that maritime habitats have more favorable water availability than the interior. While maritime seeders constitute the majority of local Arctostaphylos endemics, they exhibited significantly greater vulnerability to xylem cavitation than interior seeders. Because rare seeders in maritime chaparral are more vulnerable to xylem cavitation than interior seeders, the potential breakdown of the summer marine layer along the coast is of potential conservation concern.