Tobacco seed performance is traditionally measured as percentage germination at 14 d under controlled laboratory conditions. However, under greenhouse conditions, seed lots with equal 14-d germination may exhibit substantial differences in uniformity of early seedling growth and spiral root incidence that impact the number of usable transplants. A seedling performance index (SPI) was developed to quantitatively describe greenhouse tobacco seedling performance. The 14-d emergence, relative leaf area uniformity, and seedling leaf area determined by computer image analysis were used to calculate the index. Greenhouse tobacco seed trials demonstrated that seed with the Rickard pellet had higher emergence, but the higher spiral root incidence associated with the Rickard pellet lowered the SPI compared to the Cross Creek pellet. Primed seed lots of flue-cured cultivars (NC 72 and NC 71) had a significantly higher SPI than the nonprimed seed lots at one location but not at a second location. Seed lots sown in Premier Pro-Mix TA commercial medium had a higher spiral root incidence, which resulted in a lower SPI compared to Carolina Choice, Carolina Gold, and Sunshine LP5 commercial media. The index quantitatively determined differences in seedling performance under greenhouse conditions not reflected by standard germination tests. Significant differences in the SPI were observed among seed lots with certified 90% germination. The SPI is a simple method to describe seedling performance because the data used to calculate the SPI is obtained from one seed tray image. In contrast, frequent counting and seedling evaluations are involved with standard germination and vigor tests. / Master of Science

The overwhelming majority of plant species introduced into a new range never become invasive. Consequently, identification of factors allowing the small fraction of successful invaders to naturalize, increase in abundance, and displace resident species continues to be a key area of research in invasion biology. Of the considerable number of hypotheses that have been proposed to resolve why some plant species become noxious pests, the enemy release hypothesis (ERH) is one of the most commonly cited. The ERH maintains that invasive plants succeed in a new range because they are no longer regulated by their coevolved natural enemies, and this reduction in enemy pressure imparts a competitive advantage over native species, which continue to be negatively impacted by top-down processes. Alternatively, the ability of invasive plant species to outperform their counterparts, rather than escape from enemies, may be key in conferring invasion success. The importance of preadapted traits and release from natural enemies in successful invasion remains unclear, likely owing to a lack of empirical studies comparing their effects on relative performance and population growth of closely related species that differ in origin and invasiveness. A system of co-occurring native, introduced non-invasive, and invasive Eugenia congeners exists in south Florida, providing an opportunity to address deficiencies in our understanding of plant invasions by investigating the factors leading to invasion success for Eugenia uniflora. This approach is novel because very few studies have simultaneously incorporated both native and introduced non-invasive congeners into tests of these hypotheses, and no others have done so using this system of Eugenia congeners. The first study in this dissertation tested the ERH using an insect herbivore exclusion experiment in the field to compare the effects of natural enemies on the performance and population growth of Eugenia uniflora and its native congeners. The results showed that E. uniflora sustained more herbivore damage than its native counterparts, and that the effects of herbivores were sufficient to have negative impacts on performance and population growth. In sum, these findings contradict the ERH. Surprisingly, the vast majority of damage to E. uniflora was caused by the recently introduced Sri Lankan weevil (Myllocerus undatus), with which it shares no coevolutionary history. The second study compared seedling performance among native, introduced non-invasive, and invasive Eugenia congeners to determine if the success of E. uniflora can be attributed to superior performance traits. Invasive E. uniflora was found to outperform its native and introduced non-invasive counterparts in a number of seedling traits, including emergence, growth, and survival, in spite of sustaining higher levels of herbivore damage in the field. This result was consistent across years and sites, suggesting that superior performance may be an important factor in invasion success by E. uniflora. The final experiment investigated the role of enemy release on performance of native, introduced non-invasive, and introduced invasive Eugenia seedlings using an insect herbivore exclusion experiment in the field. In this study, the invasive E. uniflora was again found to sustain more damage by foliar herbivores compared to its native and introduced non-invasive counterparts. However, in spite of higher levels of herbivore damage, E. uniflora continued to outperform its congeners in terms of stem growth, and its congeners did not outperform E. uniflora in any attribute. Insect herbivores negatively affected survival of all species, but were found to have little effect on growth. In combination, the results of these studies indicate that the ability of E. uniflora to outperform its native and introduced congeners at the seedling stage, and not release from insect herbivores, may contribute to its success as an invader. Additionally, E. uniflora exhibits relatively low resistance to herbivory in the new range, and instead may possess an ability to tolerate moderate levels of damage. The implications of this study are that enemy release may not be important in determining invasion success in some systems, and that the accumulation of new enemies may mitigate the effects of invasive plants over time. The paucity of studies investigating interactions among invasive plants and herbivores that share no coevolutionary history warrants further research. Finally, this system of Eugenia congeners provides valuable opportunities to test additional hypotheses and to further explore factors leading to invasion success.

Water availability affects tree species performance and distributions in tropical forests. However, there are no studies that have measured detailed spatial variation in soil water availability within a tropical forest. This limits our understanding of how water availability shapes the demography and distributions of tree species within tropical forests. In this dissertation, I measured detailed spatial variation in soil water potential (SWP), the relevant measure of water availability for plant performance, in the seasonal tropical moist forest of the 50-ha Forest Dynamics Plot on Barro Colorado Island, Panama. In Paper 1, I mapped spatial variation in SWP across the 50-ha plot in various stages of the dry season using information on topography, soil type, dry season intensity and more. In Paper 2, I quantified the soil moisture niches of species in terms of demographic responses (growth and mortality) and species distributions. I related seedling growth and mortality responses to SWP of 62 species to their distributional centre along the SWP gradient, using data from 20 years of annual seedling censuses across 200 seedling census sites. I found that species that grew faster (slow) with increasing SWP were more common on wetter (drier) parts of the SWP gradient. Moreover, wet-distributed species grew faster on the wet side of the SWP gradient than dry-distributed species. Mortality was unrelated to species distributions but decreased strongly with seedling height. These findings indicate that species with a growth advantage with respect to SWP grow faster out of the vulnerable small size ranges, reducing their mortality in later seedling stages and thus shaping species distributions indirectly. This mechanism is a form of niche differentiation that contributes to species coexistence. In Paper 3, I related seedling growth and mortality responses to spatiotemporal variation in water availability with responses to light availability, another highly limiting resource in tropical forests. I found an interspecific trade-off in responses to shade versus inter-annual drought (dry season intensity): species that performed relatively well in the shade performed worse during more severe dry seasons and vice versa. This trade-off enables coexistence, because species are adapted to perform well under either shade or drought. In sum, water availability contributes to the maintenance of the high diversity of tropical forests through hydrological niche differentiation and a trade-off between performance in shade versus drought. Future work can use my SWP maps and species responses to SWP to identify the functional traits that underlie the species responses and improve Dynamic Global Vegetation Models. Finally, my work facilitates the prediction of future species composition, diversity and ecosystem functioning of tropical forests with shifts in rainfall patterns caused by climate change.

Interações interespecíficas são consideradas importantes forças estruturadoras da vegetação. Enquanto a competição aparentemente prevalece em ambientes com menor estresse abiótico, a facilitação tende a aumentar em importância em ambientes de maior estresse. Consequentemente, é esperado que a facilitação desempenhe um papel importante em áreas tropicais degradadas, nas quais as condições abióticas, envolvendo altos níveis de irradiação, temperatura e evapotranspiração, são muito distintas das condições ótimas para a maioria das espécies florestais. Além disso, face à complexidade dos processos estruturadores de comunidades florestais tropicais altamente diversas, os ecólogos têm percebido a necessidade de simplificação. Um caminho promissor para isso é por meio do uso de características funcionais, que podem ser medidas para qualquer espécie e comparadas em diferentes sistemas e locais. No entanto, em áreas degradadas ainda é incerta a relação entre as características funcionais e o desempenho das espécies. Neste contexto, planejamos um projeto de restauração na planície costeira do sudeste brasileiro para comparar o crescimento e a sobrevivência de mudas de árvores plantadas em diferentes densidades (sistema isolado ou sistema agregado) em um experimento fatorial com adição de nutrientes. No primeiro capítulo analisamos a sobrevivência, altura, diâmetro ao nível do solo e projeção da copa de 4.132 mudas de 19 espécies de diferentes classificações sucessionais, durante um período de 18 meses. Usamos modelos de efeitos mistos para analisar a relação entre o desempenho de espécies e os tratamentos e selecionamos os melhores modelos utilizando o critério de informação de Akaike (AIC). Verificamos uma maior sobrevivência nos sistemas agregados para as espécies não-pioneiras, indicando o predomínio do processo de facilitação. Em contrapartida, constatamos um menor crescimento em diâmetro nos sistema de plantio agregado, indicando o predomínio de competição. A adição de nutrientes não afetou a sobrevivência nos sistemas agregados, mas, surpreendentemente, teve um efeito negativo sobre as plantas isoladas. Além disso, a adição de nutrientes teve um efeito positivo sobre crescimento, diâmetro e projeção da copa nos sistemas de plantio agregado (indicando a redução da competição), especialmente para espécies pioneiras. Uma vez que o estabelecimento de espécies não-pioneiras em áreas degradadas pode ser bastante difícil, a facilitação para a sobrevivência dessas espécies em sistemas agregados pode ser utilizada como instrumento para melhorar os modelos de restauração. No segundo capítulo utilizamos algumas características funcionais que são facilmente mensuráveis para investigar sua relação com o crescimento e sobrevivência das mudas. Além disso, testamos o efeito das interações interespecíficas na relação das características funcionais com o desempenho das plantas. Para espécies pioneiras, uma maior área foliar específica (SLA - Specific leaf area), foi relacionada com uma maior taxa de sobrevivência. Entretanto, ao considerarmos as relações interespecíficas (sistemas agregados), maior SLA foi relacionada com uma menor taxa de sobrevivência. Em geral, as características funcionais explicaram apenas parcialmente o desempenho de mudas no contexto de restauração de áreas degradadas. Nosso trabalho contribuiu para a discussão sobre o potencial da aplicação prática da abordagem funcional na escolha das espécies para a recuperação de áreas degradadas. O caráter manipulativo da restauração ecológica baseada no plantio de mudas amplia as perspectivas para implementar experimentos que visem testar novas hipóteses em ecologia e refinar modelos de restauração / Interspecific interactions are considered to be important structuring forces in early successional vegetation. Whereas competition seems to prevail in less severe environments, facilitation tends to increase in importance in harsh environments. Hence, facilitation is expected to play an important role in degraded tropical areas with high irradiance, heat loads, and evapotranspiration, where conditions are far from optimum for most forest species. Moreover, in order to understand complex structuring process of high diverse tropical forests, ecologists have realized the need for simplification. A promising way to do that is through the use of functional traits, which can be measured for any species and compared in different systems and different locations around the world. However, in these conditions, little is known about the role of species interaction on the relation of functional traits and species performance. In this scenario, we planned a restoration project on the southeastern Brazilian coastal plain to compare growth and survival of tree seedlings planted at two densities (isolated or aggregated) in a factorial experiment with nutrient addition. In the first chapter we analyzed survival, height, ground level diameter, and crown projection of 4,132 saplings from 19 species that ranged along a successional gradient, over an 18-month period. We used mixed-effect models to analyze the relationship between species performance and treatments, and Akaike\'s information criterion (AIC) to select the models. The best model showed higher survival in aggregated plantations (indicating facilitation) for non-pioneer species. In contrast, we found lower diameter growth in aggregated plantation (indicating competition). Fertilizer addition did not affect survival in clusters, but, surprisingly, it had a negative effect on isolated plants of both pioneer and non-pioneer species. On the other hand, fertilizer addition had a positive effect on diameter and crown projection growth in aggregated plantations (reducing competition), especially for pioneer species. Thus, whether facilitation or competition was the predominant interaction depended on the response analyzed. We concluded that, as establishment of non-pioneer species in disturbed sites can be challenging, restoration designs could take advantage of higher survival rates in clusters and use resource addition to ameliorate growth and decrease competition for limited resources. In the second chapter we used the core simple-measurable traits to investigate whether functional traits were related to growth and survival and whether there was a difference in the effect of the functional traits on plant performance of isolated individuals or of individuals subjected to interaction with neighbors. Our main findings were that for pioneer species, the greater the specific leaf areas (SLA), the greater the survival rates, but, under aggregated conditions, the greater the SLA the lower the survival rates. However, functional traits only partially explained saplings performance in a restoration context and more research is needed to understand its role in predicting seedlings outcomes, especially considering the potential applicability of this methodological approach. The manipulative character of ecological restoration broadens perspectives to use experiments to generate and test new hypotheses in ecology and to refine restoration models

Interações interespecíficas são consideradas importantes forças estruturadoras da vegetação. Enquanto a competição aparentemente prevalece em ambientes com menor estresse abiótico, a facilitação tende a aumentar em importância em ambientes de maior estresse. Consequentemente, é esperado que a facilitação desempenhe um papel importante em áreas tropicais degradadas, nas quais as condições abióticas, envolvendo altos níveis de irradiação, temperatura e evapotranspiração, são muito distintas das condições ótimas para a maioria das espécies florestais. Além disso, face à complexidade dos processos estruturadores de comunidades florestais tropicais altamente diversas, os ecólogos têm percebido a necessidade de simplificação. Um caminho promissor para isso é por meio do uso de características funcionais, que podem ser medidas para qualquer espécie e comparadas em diferentes sistemas e locais. No entanto, em áreas degradadas ainda é incerta a relação entre as características funcionais e o desempenho das espécies. Neste contexto, planejamos um projeto de restauração na planície costeira do sudeste brasileiro para comparar o crescimento e a sobrevivência de mudas de árvores plantadas em diferentes densidades (sistema isolado ou sistema agregado) em um experimento fatorial com adição de nutrientes. No primeiro capítulo analisamos a sobrevivência, altura, diâmetro ao nível do solo e projeção da copa de 4.132 mudas de 19 espécies de diferentes classificações sucessionais, durante um período de 18 meses. Usamos modelos de efeitos mistos para analisar a relação entre o desempenho de espécies e os tratamentos e selecionamos os melhores modelos utilizando o critério de informação de Akaike (AIC). Verificamos uma maior sobrevivência nos sistemas agregados para as espécies não-pioneiras, indicando o predomínio do processo de facilitação. Em contrapartida, constatamos um menor crescimento em diâmetro nos sistema de plantio agregado, indicando o predomínio de competição. A adição de nutrientes não afetou a sobrevivência nos sistemas agregados, mas, surpreendentemente, teve um efeito negativo sobre as plantas isoladas. Além disso, a adição de nutrientes teve um efeito positivo sobre crescimento, diâmetro e projeção da copa nos sistemas de plantio agregado (indicando a redução da competição), especialmente para espécies pioneiras. Uma vez que o estabelecimento de espécies não-pioneiras em áreas degradadas pode ser bastante difícil, a facilitação para a sobrevivência dessas espécies em sistemas agregados pode ser utilizada como instrumento para melhorar os modelos de restauração. No segundo capítulo utilizamos algumas características funcionais que são facilmente mensuráveis para investigar sua relação com o crescimento e sobrevivência das mudas. Além disso, testamos o efeito das interações interespecíficas na relação das características funcionais com o desempenho das plantas. Para espécies pioneiras, uma maior área foliar específica (SLA - Specific leaf area), foi relacionada com uma maior taxa de sobrevivência. Entretanto, ao considerarmos as relações interespecíficas (sistemas agregados), maior SLA foi relacionada com uma menor taxa de sobrevivência. Em geral, as características funcionais explicaram apenas parcialmente o desempenho de mudas no contexto de restauração de áreas degradadas. Nosso trabalho contribuiu para a discussão sobre o potencial da aplicação prática da abordagem funcional na escolha das espécies para a recuperação de áreas degradadas. O caráter manipulativo da restauração ecológica baseada no plantio de mudas amplia as perspectivas para implementar experimentos que visem testar novas hipóteses em ecologia e refinar modelos de restauração / Interspecific interactions are considered to be important structuring forces in early successional vegetation. Whereas competition seems to prevail in less severe environments, facilitation tends to increase in importance in harsh environments. Hence, facilitation is expected to play an important role in degraded tropical areas with high irradiance, heat loads, and evapotranspiration, where conditions are far from optimum for most forest species. Moreover, in order to understand complex structuring process of high diverse tropical forests, ecologists have realized the need for simplification. A promising way to do that is through the use of functional traits, which can be measured for any species and compared in different systems and different locations around the world. However, in these conditions, little is known about the role of species interaction on the relation of functional traits and species performance. In this scenario, we planned a restoration project on the southeastern Brazilian coastal plain to compare growth and survival of tree seedlings planted at two densities (isolated or aggregated) in a factorial experiment with nutrient addition. In the first chapter we analyzed survival, height, ground level diameter, and crown projection of 4,132 saplings from 19 species that ranged along a successional gradient, over an 18-month period. We used mixed-effect models to analyze the relationship between species performance and treatments, and Akaike\'s information criterion (AIC) to select the models. The best model showed higher survival in aggregated plantations (indicating facilitation) for non-pioneer species. In contrast, we found lower diameter growth in aggregated plantation (indicating competition). Fertilizer addition did not affect survival in clusters, but, surprisingly, it had a negative effect on isolated plants of both pioneer and non-pioneer species. On the other hand, fertilizer addition had a positive effect on diameter and crown projection growth in aggregated plantations (reducing competition), especially for pioneer species. Thus, whether facilitation or competition was the predominant interaction depended on the response analyzed. We concluded that, as establishment of non-pioneer species in disturbed sites can be challenging, restoration designs could take advantage of higher survival rates in clusters and use resource addition to ameliorate growth and decrease competition for limited resources. In the second chapter we used the core simple-measurable traits to investigate whether functional traits were related to growth and survival and whether there was a difference in the effect of the functional traits on plant performance of isolated individuals or of individuals subjected to interaction with neighbors. Our main findings were that for pioneer species, the greater the specific leaf areas (SLA), the greater the survival rates, but, under aggregated conditions, the greater the SLA the lower the survival rates. However, functional traits only partially explained saplings performance in a restoration context and more research is needed to understand its role in predicting seedlings outcomes, especially considering the potential applicability of this methodological approach. The manipulative character of ecological restoration broadens perspectives to use experiments to generate and test new hypotheses in ecology and to refine restoration models