Resource allocation in Copaifera langsdorffii (Fabaceae): how supra-annual fruiting affects plant traits and herbivory?

AbstractPlants have limited resources to invest in reproduction, vegetative growth and defense against herbivorous. Trade-off in resources allocation promotes changes in plant traits that may affect higher trophic levels. In this study, we evaluated the trade-off effect between years of high and low fruiting on the investment of resources for growth and defense, and their indirect effects on herbivory in Copaifera langsdorffii. Our questions were: (i) does the resource investment on reproduction causes a depletion in vegetative growth as predicted by the Carbon/Nutrient Balance hypothesis (CNBH), resulting in more availability of resources to be allocated for defense?, (ii) does the variation in resource allocation for growth and defense between years of high and low fruiting leads to indirect changes in herbivory? Thirty-five trees located in a Cerrado area were monitored during 2008 (year of high fruiting) and 2009 (year of no fruiting) to evaluate the differential investment in vegetative traits (biomass, growth and number of ramifications), plant defense (tannin concentration and plant hypersensitivity) and herbivory (galling attack and folivory). According to our first question, we observed that in the fruiting year, woody biomass negatively affected tannin concentration, indicating that fruit production restricted the resources that could be invested both in growth as in defense. In the same way, we observed an inter-annual variation in herbivorous attack, and found that plants with higher leaf biomass and tannin concentration, experienced higher galling attack and hypersensitive reaction, regardless years. These findings suggested that plants’ resistance to herbivory is a good proxy of plant defense and an effective defense strategy for C. langsdorffii, besides the evidence of indirect responses of the third trophic level, as postulated by the second question. In summary, the supra-annual fruiting pattern promoted several changes on plant development, demonstrating the importance of evaluating different plant traits when characterizing the vegetative investment. As expected by theory, the trade-off in resource allocation favored changes in defense compounds production and patterns of herbivory. The understanding of this important element of insect-plant interactions will be fundamental to decipher coevolutionary life histories and interactions between plant species reproduction and herbivory. Besides that, only through long-term studies we will be able to build models and develop more accurate forecasts about the factors that trigger the bottom-up effect on herbivory performance, as well the top-down effect of herbivores on plant trait evolution. Rev. Biol. Trop. 64 (2): 507-520. Epub 2016 June 01.

ResumenLas plantas tienen recursos limitados para invertir en reproducción, crecimiento vegetativo y defensa contra herbívoros. El cambio en la distribución de recursos promueve variaciones en rasgos vegetales, que pueden afectar los niveles tróficos superiores. Durante dos años consecutivos de alta y baja inversión reproductiva se evaluó el cambio de recursos entre crecimiento vegetativo y defensa, y su efecto indirecto sobre la herbivoría en Copaifera langsdorffii. Nos preguntamos: i) ¿La inversión de recursos para la reproducción causa reducción del crecimiento vegetativo, como predice la hipótesis de equilibrio carbono/nutrientes, haciendo posible gastar más recursos en defensa? ii) ¿La variación en distribución de recursos para crecimiento y defensa entre años de alta y baja fructificación modifica indirectamente la herbivoría? Se monitorearon treinta y cinco árboles durante 2008 (gran fructificación) y 2009 (sin fructificación) en un área de vegetación de cerrado (Brasil), para evaluar la inversión diferencial en rasgos vegetativos (biomasa, crecimiento y No. de ramificaciones), defensa (concentración de taninos e hipersensibilidad vegetal) y herbivoría. De acuerdo a nuestra primera pregunta, se observó que en el año de fructificación la biomasa leñosa afectó negativamente la concentración de taninos, indicando que la producción de frutos redujo los recursos que podían invertirse en crecimiento y defensa. Además, la resistencia de las plantas y el ataque de agallas fueron influidos positivamente por la concentración de taninos y la biomasa foliar, lo que sugiere que la resistencia de los árboles a la herbivoría es un buen indicador de defensa vegetal y una estrategia efectiva de defensa de C. langsdorffii, además hay evidencia de respuesta trófica indirecta, como se postula en la segunda pregunta. En resumen, el patrón de fructificación supra-anual provoca varios efectos en el desarrollo de las plantas, mostrando la importancia de evaluar diversos rasgos vegetales al caracterizar la inversión de recursos de una especie. Como se esperaba, el cambio en la distribución de recursos modifica la producción de compuestos de defensa y los patrones de herbivoría. El entendimiento de este elemento importante de las interacciones insecto-planta será fundamental para descifrar la historia natural coevolutiva y las interacciones entre reproducción vegetal y ataque herbívoro. Además de eso, solo a través de estudios a largo plazo vamos a ser capaces de construir modelos y desarrollar pronósticos más precisos acerca de los factores que desencadenan el efecto de abajo hacia arriba en el rendimiento de la herbivoría, así el efecto de arriba hacia abajo de los herbívoros sobre la evolución de las plantas.

Desenvolvimento de um modelo para previsão de ocorrência do bicho-furão-dos-citros, Ecdytolopha aurantiana (Lima) (Lepidoptera: Tortricidae / Development of a model to predict the occurrence of citrus fruit borer, Ecdytolopha aurantiana (Lima) (Lepidoptera: Tortricidae)

O presente trabalho teve como objetivo desenvolver um modelo para previsão de ocorrência do bicho-furão-dos-citros, Ecdytolopha aurantiana (Lima). Para tanto, 553 áreas localizadas em 17 fazendas do estado de São Paulo foram monitoradas por meio de armadilhas com feromônio sexual de E. aurantiana durante um ano. O tipo de solo, temperatura mensal média do local, variedade de citros, idade das plantas e uso de agroquímicos para o controle de E. aurantiana, foram utilizados como variáveis para cada área mencionada. A maior influência sobre a flutuação populacional do bicho-furão-dos-citros foi exercida pelo tipo de solo, seguido pela temperatura mensal média do local, variedade de citros, idade das plantas e uso de agroquímicos. A ocorrência de E. aurantiana em função da temperatura foi diferente para cada combinação de tipo de solo, variedade de citros, idade das plantas e uso de agroquímicos. O modelo desenvolvido pode prever o potencial de ocorrência de E. aurantiana em função da temperatura ou dos meses do ano, levando-se em consideração o tipo de solo, variedade de citros, idade das plantas e aplicação de agroquímicos. O programa (BF) elaborado na linguagem R conta com equações para simular as diversas situações de ocorrência de E. aurantiana. O modelo de previsão de ocorrência de bicho-furão pode ser aperfeiçoado com a coleta de dados mais regulares e de forma contínua.

The goal of this work was to develop a model to predict the occurrence of Ecdytolopha aurantiana (Lima), based on monitoring data collected through sexual pheromone traps. Soil type, site temperature, citrus variety, age of plants and use of chemicals to control E. aurantiana influenced the population dynamics of the insect. The highest influence was exerted by the soil type, followed by site temperature, citrus variety, age of plants and the use of chemicals for E. aurantiana control. The occurrence of E. aurantiana according to temperature is different for each combination of soil type, citrus variety, age of plants and use of chemicals. The model developed can predict the occurrence potential of E. aurantiana according to temperature or months of the year, taking into account soil type, citrus variety, age of plants and chemicals spraying. The elaborated software (BF), designed in R language, includes equations that simulate the various situations of E. aurantiana occurrence. The predicting model of citrus fruit borer occurrence can be improved with more frequent and continuous data collecting.