RESUMEN
In this study, we report the first records and morphological characterization of galls in a Cerrado area in western Bahia, Brazil. The data were collected monthly over two hours between March and September 2015. Fifteen gall morphotypes were found in twelve plant species distributed among seven families. The plant family with the greatest richness of galls was Fabaceae (n = 8). The following gall morphologies were found: globoid, lenticular, marginal leaf roll, conical, cylindrical, fusiform, spherical and pocket shaped. Cecidomyiidae induced globoid, lenticular, conical, fusiform, spherical and cylindrical morphotypes. In addition, species of microhymenoptera belonging to the Eulophidae, Eurytomidae and Encyrtidae families were found. Marginal leaf roll and pocket-shaped galls induced by Thysanoptera were also verified. Springtails were also identified as a successor. Undescribed species of Schizomyia and Lopesia were recorded in B. cupulata and Andira humilis, respectively.(AU)
Nesse trabalho são apresentadas informações sobre a ocorrência e a caracterização morfológica de galhas induzidas por insetos em uma área de Cerrado do Oeste da Bahia, Brasil. Os materiais foram coletados mensalmente durante duas horas entre Março e Setembro de 2015. Foram encontrados quinze morfotipos de galhas em doze espécies vegetais, distribuídas em sete famílias. A família de planta com maior riqueza de galhas foi Fabaceae (n = 8). Com relação à morfologia externa, foram encontradas galhas com forma globoide, lenticular, enrolamento marginal, cônica, cilíndrica, fusiforme, esférica e bolso. Quanto à fauna associada às galhas, os morfotipos globoide, cônico, lenticular, fusiforme, esférico e cilíndrico foram induzidos por Cecidomyiidae. Além disso, espécies de microhimenópteros pertencentes às famílias Eulophidae, Eurytomidae e Encyrtidae foram encontrados. A indução de galhas por Thysanoptera, nos morfotipos enrolamento marginal e formato de bolso, também foi detectada. Espécies de colembôlos foram identificadas como sucessoras. Espécies não descritas de Schizomyia e Lopesia foram registradas em Bauhinia cupulata e Andira humilis, respectivamente.(AU)
Asunto(s)
Tumores de Planta , Interacciones Huésped-Patógeno , Herbivoria/clasificación , Dípteros/patogenicidad , Biodiversidad , Pradera , BrasilRESUMEN
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.
Asunto(s)
Animales , Adaptación Fisiológica/fisiología , Herbivoria/fisiología , Insectos/fisiología , Fabaceae/fisiología , Tumores de Planta , Herbivoria/clasificación , Insectos/clasificación , Fabaceae/crecimiento & desarrollo , Fabaceae/parasitologíaRESUMEN
Plants 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.
Asunto(s)
Adaptación Fisiológica/fisiología , Fabaceae/fisiología , Herbivoria/fisiología , Insectos/fisiología , Animales , Fabaceae/crecimiento & desarrollo , Fabaceae/parasitología , Herbivoria/clasificación , Insectos/clasificación , Tumores de PlantaRESUMEN
Species diversity of insect herbivores associated to canopy may vary local and geographically responding to distinct factors at different spatial scales. The aim of this study was to investigate how forest canopy structure affects insect herbivore species richness and abundance depending on feeding guilds' specificities. We tested the hypothesis that habitat structure affects insect herbivore species richness and abundance differently to sap-sucking and chewing herbivore guilds. Two spatial scales were evaluated: inside tree crowns (fine spatial cale) and canopy regions (coarse spatial scale). In three sampling sites we measured 120 tree crowns, grouped n five points with four contiguous tree crowns. Insects were sampled by beating method from each crown and data were summed up for analyzing each canopy region. In crowns (fine spatial scale) we measured habitat tructure: trunk circumference, tree height, canopy depth, number of ramifications and maximum ramification level. In each point, defined as a canopy region (coarse spatial scale), we measured habitat structure using a vertical cylindrical transect: tree species richness, leaf area, sum of strata heights and maximum canopy height. A principal component analysis based on the measured variables for each spatial scale was run to estimate habitat structure parameters. To test the effects of habitat structure upon herbivores, different general linear models were adjusted using the first two principal components as explanatory variables. Sap-sucking insect species richness and all herbivore abundances increased with size of crown at fine spatial scale. On the other hand, chewer species richness and abundance increased with resource quantity at coarse scale. Feeding specialization, resources availability, and agility are discussed as ecological causes of the found pattern.
Asunto(s)
Ecosistema , Herbivoria/clasificación , Insectos/clasificación , Animales , Biodiversidad , Herbivoria/fisiología , Insectos/fisiología , Densidad de Población , ÁrbolesRESUMEN
Species diversity of insect herbivores associated to canopy may vary local and geographically responding to distinct factors at different spatial scales. The aim of this study was to investigate how forest canopy structure affects insect herbivore species richness and abundance depending on feeding guilds´ specificities. We tested the hypothesis that habitat structure affects insect herbivore species richness and abundance differently to sap-sucking and chewing herbivore guilds. Two spatial scales were evaluated: inside tree crowns (fine spatial scale) and canopy regions (coarse spatial scale). In three sampling sites we measured 120 tree crowns, grouped in five points with four contiguous tree crowns. Insects were sampled by beating method from each crown and data were summed up for analyzing each canopy region. In crowns (fine spatial scale) we measured habitat structure: trunk circumference, tree height, canopy depth, number of ramifications and maximum ramification level. In each point, defined as a canopy region (coarse spatial scale), we measured habitat structure using a vertical cylindrical transect: tree species richness, leaf area, sum of strata heights and maximum canopy height. A principal component analysis based on the measured variables for each spatial scale was run to estimate habitat structure parameters. To test the effects of habitat structure upon herbivores, different general linear models were adjusted using the first two principal components as explanatory variables. Sap-sucking insect species richness and all herbivore abundances increased with size of crown at fine spatial scale. On the other hand, chewer species richness and abundance increased with resource quantity at coarse scale. Feeding specialization, resources availability, and agility are discussed as ecological causes of the found pattern.
La diversidad de especies de insectos herbívoros asociados con el dosel puede variar geográficamente y responder a distintos factores a diferentes escalas espaciales. El objetivo de este estudio fue investigar cómo la estructura del dosel afecta la riqueza de especies de insectos herbívoros y la abundancia en función la especialización alimenticia. Se evaluó la hipótesis que propone que la estructura del hábitat afecta en forma diferente la riqueza y abundancia de especies de insectos que se alimentan de savia y la de especies herbívoras masticadoras. Dos escalas espaciales fueron evaluadas: el interior de las copas de árboles (escala fina) y regiones del dosel (escala gruesa). En tres sitios de muestreo medimos 120 copas de árboles, agrupadas en cinco puntos con cuatro copas de árboles contiguas. Los insectos fueron muestreados golpeando las copas y los datos fueron sumados para analizar cada región del dosel. En las copas (escala espacial fina) medimos la estructura del hábitat: circunferencia del tronco, altura del árbol, profundidad del dosel, número de ramificaciones y máximo nivel de ramificación. En cada punto, definiendo una región del dosel (escala gruesa), medimos la estructura del hábitat usando un transecto cilíndrico vertical: riqueza de especies árboles, área foliar, sumatoria de altura de los estratos y máxima altura del dosel. Fue realizado un análisis de componentes principales basado en las variables medidas para cada escala espacial para estimar los parámetros de la estructura del hábitat. Para probar los efectos de la estructura del hábitat sobre los herbívoros, se ajustaron diferentes modelos lineares generales usando estos componentes principales como variables causales. La riqueza de especies chupadoras de savia y la abundancia de todas las especies herbívoras se incrementaron con el tamaño de la copa en la escala espacial final. Por otro lado, la riqueza y abundancia de especies masticadoras incrementaron con la complejidad de la estructura del hábitat en la escala más gruesa. La especialización alimenticia, la disponibilidad de recursos y la movilidad son propuestas como los factores ecológicos que explican los patrones observados.
Asunto(s)
Animales , Ecosistema , Herbivoria/clasificación , Insectos/clasificación , Biodiversidad , Herbivoria/fisiología , Insectos/fisiología , Densidad de Población , ÁrbolesRESUMEN
Our goal was to identify herbivory patterns from two insect guilds associated with Cnidoscolus quercifolius in a tropical deciduous forest in northeastern Brazil. We sampled four different habitats: (1) forest edge, (2) mesic (near to the perennial water source), (3) forest interior and (4) rupestrian fields. Habitat edge had lower leaf damage than rupestrian, mesic and forest interior habitats. Nevertheless, abundance of galls at the edge habitat was higher than at mesic, forest interior and/or rupestrian habitats. There was no difference in gall mortality by natural enemies among the four habitats sampled, demonstrating the absence of any influence of top-down controls related to abundance of galls. Trophic relationships were not related to the patterns of distribution among habitats of two insect herbivorous guilds associated with C. quercifolius. Our results demonstrated that environmental heterogeneity of dry forests can significantly alter important ecological interactions and experimental studies are needed to better understand the mechanisms responsible for differences in herbivory among habitats.
Asunto(s)
Ecosistema , Euphorbiaceae/parasitología , Herbivoria/fisiología , Interacciones Huésped-Parásitos/fisiología , Insectos/fisiología , Animales , Brasil , Herbivoria/clasificación , Insectos/clasificaciónRESUMEN
Our goal was to identify herbivory patterns from two insect guilds associated with Cnidoscolus quercifolius in a tropical deciduous forest in northeastern Brazil. We sampled four different habitats: (1) forest edge, (2) mesic (near to the perennial water source), (3) forest interior and (4) rupestrian fields. Habitat edge had lower leaf damage than rupestrian, mesic and forest interior habitats. Nevertheless, abundance of galls at the edge habitat was higher than at mesic, forest interior and/or rupestrian habitats. There was no difference in gall mortality by natural enemies among the four habitats sampled, demonstrating the absence of any influence of top-down controls related to abundance of galls. Trophic relationships were not related to the patterns of distribution among habitats of two insect herbivorous guilds associated with C. quercifolius. Our results demonstrated that environmental heterogeneity of dry forests can significantly alter important ecological interactions and experimental studies are needed to better understand the mechanisms responsible for differences in herbivory among habitats.
Nosso objetivo foi identificar os padrões de herbivoria de duas guildas de insetos herbívoros associadas à Cnidoscolus quercifolius, em uma floresta estacional seca do nordeste do Brasil. Foram amostrados quatro diferentes habitats: (1) borda florestal; (2) mésico (próximo a uma fonte de água perene); (3) interior florestal, e (4) rupestre. O habitat borda apresentou uma quantidade de dano foliar inferior aos habitats rupestre, mésico e interior florestal. Diferentemente, a abundância de galhas no habitat borda foi superior aos habitats mésico, interior florestal e rupestre. Não houve diferença na mortalidade de galhas por inimigos naturais entre os quatro habitats, o que revela ausência de influência de controles top-down relacionados à abundância de galhas. Relações tróficas não parecem influenciar os padrões de distribuição entre habitats das duas guildas de insetos herbívoros associadas à C. quercifolius. Nossos resultados demonstram que a heterogeneidade ambiental de florestas secas pode alterar significativamente importantes interações ecológicas e que trabalhos experimentais são necessários para uma melhor compreensão dos mecanismos responsáveis pelas diferenças nas taxas de herbivoria entre habitats.