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Seed size is regarded as a functional trait with very important consequences for the fitness of plant species. Seedlings emerging from larger seeds are more competitive but are more costly to produce than seedlings from smaller seeds. Seed size is also a trait with transgenerational effects, affecting both the fitness of the parent as well as that of the offspring. Theory on the evolution of offspring size predicts an optimum balance between size and number, seen from the parent's perspective; while empirical studies often show selection for larger seeds, seen from the offspring's perspective. Seed size selection arising from post germination traits is, however, often not unidirectional, nor operating with the same strength in all life history stages of the plant. Seed size selection is also environmentally dependent. Even environmental influence might not operate with the same consistency and strength uniformly through the plant's life cycle. This dissertation is intended to study these questions concerning the dynamics of seed size selection in the wild. This work is to my knowledge, the first to document how seed size selection operates through the whole life cycle, with naturally germinated annual plants from the Sonoran Desert. In my first chapter I explored the offspring fitness consequences of seed size in a multiyear observational study using plant demography and relating vital rates (germination, survival, and fecundity) to the size of the seeds that originate individual plants and the environmental variables of precipitation and competition. I detected positive directional selection operating both through survival and fecundity. Water availability increased both survival and fecundity but also strengthened survival selection and had no effect on fecundity selection. Competition detrimental effects were only observed in fecundity but not in plant survival. In my second chapter I ask whether seed size-specific germination could influence seed size selection

Within each leaf is an intricate network of veins. The geometry of this network shows large variation across species and environments, paralleling variation in species' functioning and geographic distributions. Here I develop theory that links leaf venation networks to 1) the worldwide leaf economics spectrum, enabling better understandings of the resource tradeoffs that are central to the terrestrial carbon and water cycles, and 2) atmospheric temperature and carbon dioxide concentrations, enabling better use of leaf fossils for paleoclimate reconstruction. I successfully test these theories across contemporary temperate and tropical sites, and apply them to paleo-sites spanning a 2Myr interval across the Cretaceous-Paleogene boundary. These theoretical and empirical results demonstrate that leaf venation networks can provide an integrative framework for understanding relationships between plant form, function, and environment.

Mutualism is an often-complex positive interaction between species, each of which responds independently to varying biotic and abiotic conditions. Temperature is an important factor that can affect species both directly (e.g., physiologically) and indirectly (e.g., via its effects on interactions with consumers, competitors, and mutualists). Although much research has investigated the consequences of temperature for individual organisms, the effects of temperature on the formation, dissolution, and success of species interactions remain minimally understood. The unique ways in which species respond to temperature likely play a role in structuring communities. Environmental heterogeneity, including the thermal environment, can promote coexistence when species exploit resources in different ways, such as by occupying different thermal niches. This dissertation examines the consequences of temperature for participants in an ant-plant protection mutualism, and investigates how the thermal ecology of individual species affects the interaction. Many mutualisms involve multiple species, or interacting guilds. In these mutualisms, species interact with partner species that vary in multiple characteristics. Mutualists are quite sensitive to both partner quantity and partner quality (e.g., their effectiveness at performing a beneficial task). Mutualisms between ants and plants are common across a variety of habitats worldwide, which differ in thermal range, fluctuation, and seasonality. In light of ants’ well-studied and predictable responses to temperature, ant-plant interaction networks provide excellent systems for studying the thermal ecology of mutualisms. In ant-plant protection mutualisms, each of the participants (ants, plants, and enemies) likely differs in its response to temperature. In addition to the direct effects of temperature on ant species, temperature may affect the magnitude of mutualistic interactions among species by affecting the quantity and quality of

Predicting the interactions between climate change and ecosystems remains a core problem in global change research; tropical forest ecosystems are of particular importance because of their disproportionate role in global carbon and water cycling. Amazonia is unique among tropical forest ecosystems, exhibiting a high degree of coupling with its regional hydrometeorology, such that the stability of the entire forest-climate system is dependent on the functioning of its component parts. Belowground ecohydrological interactions between soil moisture environments and the roots which permeate them initiate the water transport pathway to leaf stomata, yet despite the disproportionate role they play in vegetation-atmosphere coupling in Amazonian forest ecosystems, the impacts of climate variability on the belowground environment remain understudied. The research which follows is designed to address critical knowledge gaps in our understanding of root functioning in Amazonian tropical forests as it relates to seasonality and extremes in belowground moisture regime as well as discerning which ecohydrological mechanisms govern ecosystem-level processes of carbon and water flux. A secondary research theme is the evaluation and use of models of ecosystem function as applied to Amazonia - these models are the "knowledge boxes" which build in the ecohydrological hypotheses (some testable than others) deemed to be most important for the forest ecosystems of Amazonia. In what follows, I investigate (i) which mechanisms of water supply (from the soil environment) and water demand (by vegetation) regulate the magnitude and seasonality of evapotranspiration across broad environmental gradients of Amazonia, (ii) how specific hypotheses of root function are or are not corroborated by soil moisture measurements conducted under normal seasonal and experimentally-induced extreme drought conditions, and (iii) the linkage between an extreme drought event with associated impacts on root zone s

Canopy structure has long been thought to influence the productivity and ecological dynamics of tropical forests by altering the availability of light to leaves. Theories and methods that can connect detailed quantitative observations of canopy structure with forest dynamics, however, have been lacking. There is urgent need to resolve this uncertainty because human-caused climate change may alter canopy structure and function in the Amazon. This work addresses this problem by, first, developing methods based on LiDAR remote sensing of fine-scale structural variation to predict the spatial structure of leaf area and light in forest canopies of the central Amazon (Appendices B&C). I show that LiDAR-based leaf area and light estimates can be used to predict the productivity of tree size groups and one-hectare forest plots--as well as differences between 2 sites separated by 500km (App. B). Sites also differed in canopy structure and the distribution of tree frequencies over size (size or diameter distribution). A model based on tree architecture, however, was able to connect observed differences in canopy architecture with size distributions to predict plot and site differences (App. D). This model showed that tree architecture is plastic in different light environments. While plasticity may increase light absorption, the smallest size groups appeared light limited. Absorption over size groups in one site, but not the other, agreed with the hypothesis of energetic equivalence across size structure. Ultimately, the performance of individual trees of different sizes in different canopy environments links forest demography with canopy structure and ecosystem function--I present a study aimed at improving tests of individual level theories for the role of light dependence in tree growth (App. A). Together, this work quantitatively connects canopy structure with forest carbon dynamics and demographic structure and further develops LiDAR as premier tool for studying forest e

Drought-induced tree mortality is an emerging global phenomenon that appears related to climate change and rising temperatures in particular, and may be an early indication of vegetation change. However, vegetation response to climate change is uncertain, particularly for future novel climates. Notably, no current models of vegetation change attempt to mechanistically predict plant mortality, and in particular, mortality of trees, which exerts strong influences on ecological function. Resolving uncertainties surrounding the physiological mechanism and temperatures sensitivity of tree mortality is a current challenge in global change ecology. The objectives of this dissertation were to 1) consider tree mortality consequences for earth system processes related to carbon, water, and energy exchange that include climate regulation; 2) explore tree mortality effects on the water cycle by developing hypotheses and research needs; 3) quantify the temperature sensitivity of drought-induced tree mortality and gain insight into the physiological mechanism of mortality; 4) quantify the relationships among temperature, stored carbohydrate resources, and gas exchange to further elucidate physiological tree mortality mechanisms; and 5) quantify the sensitivity of two species of pine seedlings to progressively elevated temperatures and relate mortality to the effect of temperature on carbon metabolism. Major findings of this dissertation relate to the temperature sensitivity, physiological mechanism, and implications of tree mortality. Assessment of the potential consequences of tree mortality for earth system processes documented the contrasting influences of tree mortality on the terrestrial C cycle and land-surface energy exchange, the balance of which will determine the net effects on climate regulation (Appendix A). Following a survey of the ecohydrology literature, thresholds for tree mortality to cause watershed changes were hypothesized at ~20% loss of canopy cover, ~500 m

Evapotranspiration dominates the water budget in drylands, exerting important controls not only on the dynamics of water, but also on the amount and distribution of vegetation on a landscape. The spatial and temporal variability of vegetation cover imposes constraints on key ecohydrological processes that feedback to the dynamics of evapotranspiration and, most importantly, its partitioning between direct evaporation and transpiration from plants, one of the most significant ecohydrological challenges. Yet, lacking are systematic evaluations of how variations in woody plant cover--a fundamental vegetation attribute of landscapes that can vary spatially with amount of cover and temporally with leaf phenology-- influence the dynamics of soil microclimate and ultimately the partitioning of evapotranspiration into its components. This study presents the results of field experiments that systematically evaluated the effects of amount of canopy cover and its seasonality in both surface microclimate and soil evaporation. These field observations are complemented by controlled experiments that directly evaluate the relationship between amount of canopy cover and the partitioning of evapotranspiration, with an assessment of its larger-scale implications using a regional land surface-atmosphere model. Finally, this study presents a classroom-adaptation of the evapotranspiration partitioning experiment that was used to effectively translate new scientific concepts and information into k-12 classrooms. Overall, the results from this study provide a comprehensive understanding about the interactive ways in which canopy cover, canopy structure attributes and plant phenology influence soil surface microclimate--characterized by near-ground solar radiation and soil temperature--and soil evaporation. More specifically, the results illustrate how the main control of deciduous-woody vegetation on soil evaporation is the addition of litter to the surface. However, in absence of litter

Wind erosion and associated dust emissions play a fundamental role in many ecological processes, yet most ecological studies do not explicitly consider dust-driven processes despite the growing body of evidence suggesting that wind erosion is a key driver of land surface dynamics and many other environmentally relevant processes such as desertification. This study provides explicit support for a pervasive underlying but untested desertification hypothesis by showing that at the vegetation patch scale shrubs are significantly more efficient at capturing wind-blown sediment and other resources such as nutrients than grasses and that this difference is amplified following disturbance. At the landscape scale, the spacing and shape of woody plants were found to be a major determinant of dryland aeolian sediment transport processes in grasslands, shrublands, woodlands and forests, particularly following disturbance. This study also found that disturbance such as fire can have a significant influence on background dust emissions, which can have important consequences for many basic ecological and hydrological processes. Potential interactions between aeolian and fluvial processes were also evaluated in this study, and a new conceptual framework was developed that highlights important differences and similarities between the two processes as a function of scale-dependencies, mean annual precipitation, and disturbance. This study also explicitly evaluates the effect of climatic extremes and alternate land management practices on the absolute and relative magnitudes of wind and water erosion. Notably, results indicate that wet/dry climatic extremes and grazing can increase the wind-to-water erosion ratio, whereas burning disproportionally increases water erosion relative to wind erosion.

Diversity within a habitat is determined largely by ecology and species interactions. Studies to date, however, rarely examined the role of intraspecific aggression in promoting coexistence and diversity. This is especially true in cities, where community ecology is poorly understood. This knowledge is important for basic understanding of how ecological principals come into play in our newly created habitats, as well as for reconciling human-dominated areas for wildlife.I studied the effect of human-made habitats on hummingbird abundance and diversity in Tucson, Arizona. To do that, I examined community organization and competitive interactions among four hummingbird species. I answer the questions: What is the community organization of hummingbirds in Tucson? How do characteristics of human habitats (e.g., landscaping and artificial resources) affect diversity? What mechanism underlies this pattern? And how can we apply this knowledge to conservation?To perform this study, I established a citizen science project - the Tucson Hummingbird Project (http://hummingbirds.arizona.edu). Trained participants reported abundance and behavior of hummingbirds in their backyards. Landscaping and resources (feeders and nectar plants) varied between yards.Results show that the distribution of hummingbirds in Tucson varies by species. Diversity, rather than merely abundance, increased with higher habitat heterogeneity and with more resources. Competitive interactions differ between species. Notably, intraspecific competition takes precedence over interspecific competition in the dominant and most common species, Anna's hummingbird.Based on the data, I suggest that Aggressive Resource Neglect (ARN) promotes coexistence and results in higher diversity when resources are augmented. When there are more feeders, they are distributed over a larger area. This reduces the ability of a territory-owner to defend these resources. While the territory-owner chases intruders, other individuals g

Water is a critical resource for which plants compete in many terrestrial communities. In arid communities where water most limits plant growth, rainfall events occur in discrete, pulsed events. These pulses of water create highly variable soil moisture availabilities. Plant species respond differently to variation in soil water availabilities throughout a season and between years. How species vary in their responses to a range of water availabilities is thought to influence community and ecosystem properties. Many previously proposed hypotheses are not suitable to explain rapidly fluctuating resource availabilities or numerous input events throughout the growing season. This dissertation uses variation in water availability as a model resource to examine how species characteristics influence the process of exploitation competition within plant communities.Experiments were conducted to examine variation in growing season, exploitation competition between several pairs of co-occurring species in the Sonoran desert. Three separate studies evaluated several components of community dynamics thought to be influenced by exploitation competition. Spatial attributes of exploitation competition were assessed by measuring the performance of a deep-rooted species across the boundary of a natural expansion of a shallowly rooted species. Then, neighborhood composition was varied for species of similar growth-form to address the affects of species characteristics to shifts in abundances under field conditions. Lastly, species from the neighborhood composition study were placed under controlled, manipulated water availabilities to measure their fundamental operational conditions.Performances of plant species in all experiments were assessed using a combination of physiological and vegetative measurements, capturing the responses of the plants to both the dynamic growth conditions during the growing season, and integrated measures of plant performance post growth season. A shared p

Precipitation, more than any other environmental factor, controls patterns of ecosystem production and biogeochemical cycling in arid and semiarid environments. Growing-season rains in these regions are highly unpredictable as they come in intermittent pulses varying in size, frequency and spatial extent, thereby producing unique hydrological patterns that constrain the location and residence time of soil water available for biological activity. In order to understand how arid and semiarid ecosystems respond to inputs of precipitation within the context of ecosystem science and global change studies, knowledge is needed on how plants and other organisms respond as an integrated system to such environmental control. The focus of my research was to understand how the distribution of precipitation events influences the dynamics of carbon cycling in semiarid ecosystems. At a semiarid riparian woodland, measurements of CO2 exchange and evapotranspiration revealed that following precipitation events occurring soon after prolonged dry periods the efficiency of rain-use (amount of carbon gain per unit of precipitation over a specific period time) was low. Precipitation did not readily stimulate primary productivity, water was mainly lost as soil evaporation and large respiratory CO2 effluxes were observed. This commonly observed features in seasonally dry ecosystems might have profound consequences for the seasonal and annual carbon balance. In this woodland, 47% of the precipitation within a single growing season (May-October) was returned to atmosphere as soil evaporation and the CO2 efflux observed just during the first rainy month (July) was equivalent to almost 50% of the net carbon gain observed over the six-month growing season. Results from experimental irrigations in understory plots of riparian mesquite woodland revealed that the magnitude and duration of the large CO2 fluxes occurring soon after rainfall was higher in plots located under tree canopies where, rela

Biological activity in desert soils is driven by water availability. The nature of individual precipitation events is critical to understanding soil moisture availability. Rain falls as discrete events (pulses) that vary in size and sequencing, resulting in soil "wet-dry cycles". Soil organisms are responsive to wet-dry cycles with rapid changes in activity. How soil activity is driven by changes in water content associated with individual pulses is poorly understood. The effects of precipitation on soil processes likely depend on ecosystem structure, which influences the soil environment. The goal of this dissertation was to determine how soil carbon cycling responds to precipitation in the context of ecosystem structure (plant composition, geomorphology) and climate.I used differences in stable carbon isotopic composition of soil organisms and plants to understand how positioning in the soil profile influences biological responses to different sized pulses. I evaluated how soil texture and grass species composition affect soil process response to rainfall in different seasons. I manipulated rainfall sequence to understand the interaction between closely spaced rainfall events of different sizes on soil processes. I evaluated the role of plant functional types in influencing soil microclimate and litter deposition and the response of soil processes to seasonal rainfall.Chamber measurements of soil and plant CO2 flux were used to understand their response to rainfall. I found that surface organisms are more responsive to small rainfall events due to the relationship between pulse size and infiltration. While soil texture and season of rainfall are important, the best predictor of the response of soil respiration to rainfall was initial activity levels. Grass species was not important. Grass roots and soil microbes differ in response to sequences of precipitation. Grasses responded less to subsequent large events if they were already 'activated' by a recent rainfall

Arid and semi-arid ecosystems of the southwestern U.S. are experiencing major changes that have profound impacts for community structure and ecosystem function. First, these ecosystems are experiencing dramatic shifts in vegetation composition as a result of the invasion of non-native species. Second these ecosystems are predicted to undergo substantial shifts in climate regime, which include increases in the variability and frequency of extreme temperature and precipitation events. It is not well understood how these current and predicted changes will affect the physiological performance of different plant types in arid and semi-arid ecosystems. To address the effect of these changes, this dissertation focused on the photosynthetic response of a native and non-native grass species, and dominant shrub species to precipitation across contrasting soil surfaces in southeastern Arizona. The native and non-native grasses were exposed to wet and dry seasonal precipitation and responses to precipitation events ('pulses') were measured over the course of a summer growing season. To gain a mechanistic understanding of these patterns, the biochemical and diffusion limitations to photosynthetic function were measured over the course of a pulse period. Building on this foundation, natural stands of the non-native grass species were exposed to sequences of different sized pulse events. The physiological performance of a dominant shrub species, Larrea tridentata, was measured in order to determine the biochemical and diffusional constraints to photosynthetic function across seasons and contrasting soil surfaces. The results showed that leaf area development of these grass species affects water availability and time lags in photosynthetic response. Initial soil moisture conditions across contrasting soil surfaces influence the magnitude of photosynthetic response in grasses. Large photosynthetic responses of the non-native grass require large and consecutive precipitation pulses.

Key to evaluating the consequences of woody plant encroachment on water and carbon cycling in semiarid ecosystems is a mechanistic understanding of how biological and non-biological processes influence water loss to the atmosphere. To better understand how precipitation is partitioned into the components of evapotranspiration (bare-soil evaporation and plant transpiration) and their relationship to plant uptake of carbon dioxide (CO2) as well as ecosystem respiratory efflux, we measured whole plant transpiration, evapotranspiration, and CO2 fluxes over the course of a growing season at a semiarid Chihuahuan Desert shrubland site in south-eastern Arizona. Whole plant transpiration was measured using the heat balance sap-flow method, while evapotranspiration and net ecosystem exchange (NEE) of CO2 were quantified using the Bowen ratio technique.

Escalating demands on freshwater resources coupled with decreasing supplies dictates the need for improved groundwater modeling techniques and a more direct linkage between groundwater and riparian vegetation. This study describes the development and application of a new methodology and associated software model to simulate riparian and wetland evapotranspiration (RIP-ET). Traditional groundwater modeling, based on a quasi-linear relationship between evapotranspiration (ET) rates and groundwater elevation, has been used to predict changes in regional groundwater levels, and thus riparian vegetation potential. Our new approach uses multiple non-linear, segmented flux curves that more accurately reflect plant eco-physiology, thus simulating riparian and wetland ET in a manner that more realistically reflects the complexity inherent in these ecosystems. By decoupling evaporation and transpiration, both processes are more accurately defined and quantified. Plant functional groups (PFGs) based on water tolerance ranges and rooting depths are used to describe the interactive processes of plant transpiration with groundwater conditions. ET flux rate curves set the extinction and saturation extinction depths and define the group's ET flux rate as a function of water table depth. For each group, the curve simulates transpiration declines that occur both as water levels decline below rooting depths and as waters rise to levels that produce anoxic soil conditions. Each of the multiple transpiration curves reflects a particular plant functional group, thus reflecting the variability in vegetative conditions.The RIP-ET model also improves accuracy by more effectively dealing with spatial issues of plant and water table distribution. ET can now be calculated by delineating the area of all plant assemblages (or habitat types), assigning accurate elevations to the individual polygons and then applying multiple ET curves to a single model cell. The ability to assign a unique surface