ABSTRACT
Shade tolerance (the capacity to survive and grow over long periods under shade) is a key component of plant fitness and the foundation of current theories of forest succession in tropical rain forests. It serves as a paradigm to understand the optimal allocation of limited resources under dynamic light regimes. I analyze how tropical rain forest succession influences the expression of ecophysiological mechanisms leading to shade tolerance, and identify future areas that will increase our understanding of the ecological and evolutionary consequences of this phenomenon. Shade tolerance is a multivariate, continuous functional trait reflecting the growth-mortality trade-off of investing resources under limited light vs. exploiting high light conditions. I propose the life cycle successional trajectory model of Gómez-Pompa & Vázquez-Yanes as an integrative tool to understand tropical rain forest succession. This model shows how species distribute along the successional environmental gradient based on their degree of shade tolerance and represents a more integrative paradigm to understand the interface between different aspects of species diversity (ontogenetic variation and functional diversity) throughout succession. It proposes that different trait combinations determining shade tolerance are expressed at different stages of the life cycle, which affects how and when plants enter the successional trajectory. Models explaining the expression of shade tolerance (resource availability, carbon gain, CSR, resource competition) are based on whole-plant economics and are not mutually exclusive. The analysis of shade tolerance is biased towards tree seedlings in the understory of mature forests. Other life stages (juvenile and adult trees), life forms, and microhabitats throughout the forest profile are almost always excluded from these analyses. More integrative explanations based on the distribution of functional traits among species, ontogenetic stages, and the nature of the environmental gradient are being developed based on long-term data and chronosequence comparisons. In summary, shade-tolerance is a complex phenomenon, is determined by multiple characters that change ontogenetically over space and time and entails considerable plasticity. Current methods do not account for this plasticity. Understanding the nature of shade tolerance and its functional basis is critical to comprehending plant performance and improving the management, restoration and conservation of tropical rain forests given the combined threats of global warming and habitat loss.
La tolerancia a la sombra (la capacidad de sobrevivir y crecer durante largos períodos bajo sombra profunda) es un componente clave del valor adaptativo de la planta y la base de las teorías actuales de la sucesión forestal de la selva tropical. Sirve como un paradigma para entender la asignación óptima de recursos limitados bajo regímenes dinámicos de luz. En esta revisión analizo cómo la sucesión de los bosques tropicales lluviosos influye en la expresión de los mecanismos ecofisiológicos que conducen a la tolerancia a la sombra, e identifico áreas futuras que pueden aumentar nuestra comprensión de las consecuencias ecológicas y evolutivas de este fenómeno. La tolerancia a la sombra es un rasgo funcional continuo y multivariable que refleja el balance de invertir recursos bajo condiciones de luz limitada versus crecer más rápidamente en condiciones de luz intensa. Propongo el modelo de ciclo de vida a lo largo de la trayectoria de sucesión de Gómez-Pompa y Vázquez-Yanes como una herramienta integradora para entender la sucesión de la selva tropical. Este modelo muestra cómo las especies se distribuyen a lo largo del gradiente ambiental en función de su grado de tolerancia a la sombra, y representa un paradigma más integrador para comprender la interacción entre los diferentes componentes de la diversidad de especies (diversidad taxonómica y funcional y variación ontogenética) a lo largo de la sucesión. El modelo propone que las diferentes combinaciones de caracteres funcionales que determinan la tolerancia a la sombra se expresan en diferentes etapas del ciclo de vida, y afectan cómo y cuándo las plantas ingresan en el proceso de sucesión. Los modelos que explican la expresión de tolerancia a la sombra (disponibilidad de recursos, ganancia de carbono, CSR, competencia de recursos) se basan en la economía de toda la planta y no son mutuamente excluyentes. Se están desarrollando explicaciones más integradoras basadas en la distribución de caracteres funcionales entre especies, etapas ontogenéticas, y micrositios, mediante el uso de estudios de cronosecuencia y metadatos colectados a largo plazo. El análisis de la tolerancia a la sombra está sesgado hacia las plántulas de árboles y el sotobosque. Otras formas de vida y microhábitats dentro del perfil del bosque están casi excluidas de estos análisis. En resumen, la tolerancia a la sombra es un fenómeno complejo, está determinada por múltiples caracteres funcionales que cambian ontogenéticamente en el espacio y el tiempo, e implica una considerable plasticidad. Los métodos actuales no toman en cuenta esta plasticidad. Comprender la naturaleza de la tolerancia a la sombra y su base funcional es fundamental para entender el crecimiento de la planta y mejorar la gestión, restauración, y conservación de los bosques tropicales, los cuales enfrentan las amenazas combinadas del calentamiento global y la pérdida de hábitat.
ABSTRACT
This study described the feeding habits of the characin Astyanax aff. fasciatus. The diet compositions of specimens from two sites (A and B) on a river in Southern Brazil were compared according to the size of individuals and seasonal period. The collections were performed monthly from March 2005 to February 2006, where the stomach contents of 290 specimens were assessed. Food items for A. aff. fasciatus were basically composed of plants and insects, especially leaf fragments, seeds, fruits, filamentous algae, aquatic and terrestrial insects and insect fragments. At site A, the most common items were insect and plant fragments. Conversely at site B, plant fragments were more representative. In general, all items of animal origin showed the highest feeding index values at site A, whereas at site B detritus and grass items were more abundant. The composition of items varied seasonally, with higher diversity of items being recorded during the spring at both sites. Smaller individuals preferred items of animal origin, while the larger ones consumed mainly items of plant origin. According to its size, A. aff. fasciatus in this study may be considered a species with insectivorous tendencies when immature or herbivorous tendencies when adult. Nevertheless, its feeding habits may be flexible according to resource availability, showing wide ontogenetic, besides spatial and temporal variation.
Este estudo descreveu os hábitos alimentares do lambari Astyanax aff. fasciatus. Foram comparadas as composições alimentares de espécimes de dois sítios (A e B) de um rio no sul do Brasil de acordo com o tamanho dos indivíduos e do período sazonal. As coletas foram realizadas mensalmente de março de 2005 a fevereiro de 2006, sendo o conteúdo estomacal de 290 exemplares analisado. A dieta de A. aff. fasciatus foi composta basicamente por plantas e insetos, especialmente fragmentos de folhas, sementes, frutos, algas filamentosas, insetos aquáticos e terrestres e fragmentos de insetos. No sítio A, os itens de maior importância foram fragmentos de insetos e plantas. Ao contrário no sítio B, fragmentos de plantas foram mais representativos. Em geral, todos os itens de origem animal apresentaram os maiores valores do índice alimentar no sítio A, enquanto que no sítio B detritos e gramíneas foram mais abundantes. A composição dos itens ingeridos variou sazonalmente, sendo a maior diversidade registrada durante o período da primavera para ambos os sítios. Indivíduos pequenos preferiram itens de origem animal enquanto que os maiores consumiram principalmente itens de origem vegetal. De acordo com seu tamanho A. aff. fasciatus neste estudo pode ser considerada uma espécie com tendências à insetivoria quando imaturo, ou à herbivoria quando adulto. No entanto, seu hábito alimentar pode ser flexível de acordo com a disponibilidade de recursos, mostrando ampla variação tanto ontogenética quanto espacial e temporal.