Diversity of Gall-Inducing Insects Associated With a Widely Distributed Tropical Tree Species: Testing the Environmental Stress Hypothesis.

Abiotic factors can affect plant performance and cause stress, which in turn affects plant-herbivore interactions. The Environmental Stress Hypothesis (ESH) predicts that gall-inducing insect diversity will be greater on host plants that grow in stressful habitats. We tested this hypothesis, considering both historical and ecological scales, using the plant Copaifera langsdorffii Desf. (Fabaceae) as a model because it has a wide geographic distribution and is a super-host of gall-inducing insects. According to the ESH, we predicted that 1) on a historical scale, the diversity of gall-inducing insects will be higher in habitats with greater environmental stress and 2) on an ecological scale, gall-inducing insect diversity will be greater on plants that possess greater levels of foliar sclerophylly. We sampled gall-inducing insects on plants of C. langsdorffii in five sites with different levels of water and soil nutrient availability and separated from each other by a distance of up to 470 km. The composition, richness, and abundance of gall-inducing insects varied among study sites. Plants located in more stressful habitats had higher levels of foliar sclerophylly; but richness and abundance of gall-inducing insects were not affected by host plant sclerophylly. Habitat stress was a good predictor of gall-inducing insect diversity on a regional scale, thus corroborating the first prediction of the ESH. No relationship was found between plant sclerophylly and gall-inducing insect diversity within habitats. Therefore, on a local scale, we did not find support for our second prediction related to the ESH.

Superhost Plants Alter the Structure of Plant-Galling Insect Networks in Neotropical Savannas.

Host plants may harbor a variable number of galling insect species, with some species being able to harbor a high diversity of these insects, being therefore called superhost plants. In the present study, we tested the hypothesis that the occurrence of superhost plant species of genus Qualea (Vochysiaceae) affects the structure of plant-galling insect ecological networks in Brazilian Cerrado. We sampled a total of 1882 plants grouped in 131 species and 43 families, of which 64 species and 31 families of host plants hosted 112 galling insect species. Our results showed that occurrence of superhosts of genus Qualea increased the linkage density of plant species, number of observed interactions, and the size of plant-galling insect networks and negatively affected the network connectance (but had no effect on the residual connectance). Although the occurrence of Qualea species did not affect the plant species richness, these superhosts increased the species richness and the number of interactions of galling insects. Our study represents a step forward in relation to previous studies that investigated the effects of plant diversity on the plant-insect networks, showing that few superhost plant species alter the structure of plant-herbivore networks, even without having a significant effect on plant diversity.

Soil fertility and rainfall during specific phenological phases affect seed trait variation in a widely distributed Neotropical tree, Copaifera langsdorffii.

PREMISE: Knowledge of intra-specific variation in seed traits and its environmental determinants is important for predicting plant responses to environmental changes. Here, we tested the hypothesis that differences in soil fertility and rainfall during specific phenological phases drive variation in seed traits in a widely distributed tree, Copaifera langsdorffii. We also tested the hypothesis that climatic heterogeneity increases within-plant variation in seed traits. METHODS: Inter- and intra-population and within-plant variation in seed mass, number, and seed size/seed number were evaluated for 50 individuals from five populations distributed along a rainfall gradient and occurring on varying soil types. Using multivariate approaches, we tested the effects of soil fertility characteristics and rainfall in five reproductive phenological phases on seed traits. RESULTS: The seed traits varied greatly both among populations and within plants. Inter-population variation in seed mass was driven by total rainfall during fruit development, and variation in seed number was influenced by total rainfall during the dry season before the reproductive phase. Phosphorus levels and potential acidity of the soil also explained the variations in seed mass and seed mass/seed number, respectively. A positive association between intra-annual variation in rainfall and within-plant variation in seed mass and seed number was found. CONCLUSION: Both rainfall during specific reproductive phases and soil conditions shape the variation in the seed mass and number of C. langsdorffii. Environment-driven seed trait variation may contribute to this species' broad niche breadth, which in turn may determine the species' persistence under future climatic conditions.

Variation in community structure of gall-inducing insects associated with a tropical plant supports the hypothesis of competition in stressful habitats.

Environmental factors act as drivers of species coexistence or competition. Mesic environments favor the action of parasites and predators on gall communities, while the factors that determine the structure of gall communities in xeric environments remain unknown. We evaluated the structure of gall communities along an environmental gradient defined by intrinsic plant characteristics, soil fertility, and aridity, and investigated the role of competition as a structuring force of gall communities in xeric environments. We created null models to compare observed and simulated patterns of co-occurrence of galls and used the C-score index to assess community aggregation or segregation. We used the NES C-score (standardized C-score) to compare patterns of co-occurrence with parameters of environmental quality. Xeric environments had poorer and more arid soils and more sclerophyllous plants than mesic environments, which was reflected in the distribution patterns of gall communities. Values of the C-score index revealed a segregated distribution of gall morphospecies in xeric environments, but a random distribution in mesic environments. The low availability of resources for oviposition and the high density of gallers in xeric environments reinforce interspecific competition as an important structuring force for gall communities in these environments.

Plant phenological asynchrony and community structure of gall-inducing insects associated with a tropical tree species.

The dynamics of occurrence of target organs in plant populations produces windows of opportunity that directly and indirectly affect the structure of herbivore communities. However, mechanisms that drive herbivore specialization between resource patches are still poorly known. In this study, we tested three hypotheses related to variation in host plant phenology and community structure (i.e., composition, richness, and abundance) of gall-forming species: (a) plants with early leaf-flushing in the season will have greater vegetative growth and high contents of secondary chemical compounds; (b) gall-inducing insect community structure changes among temporary resource patches of the host; and (c) interspecific competition is a probable mechanism that drives gall-inducing insect community structure on Copaifera langsdorffii. We monitored daily a total of 102 individuals of the super-host C. langsdorffii from August 2012 to May 2013, to characterize the leaf flushing time of each host plant. The leaf flushing time had a positive relationship with the number of folioles per branch and a negative relationship with branch growth. We sampled a total of 4,906 galls belonging to 24 gall-inducing insect species from 102 individuals of C. langsdorffii. In spite of some gall-inducing species presented high abundance on early leaf-flushing plants, direct and indirect effects of plant phenology on galling insect abundance was species dependent. At the community level, our study revealed that the quality and quantity of plant resources did not affect the richness and abundance of gall-inducing insects associated with C. langsdorffii. However, the richness and composition of gall-inducing species varied according to the variation in leaf flushing time of the host plant. The results of null model analysis showed that galls co-occurrence on C. langsdorffii trees differ more than expected by chance and that interspecific competition can be one potential mechanism structuring this gall-inducing insect community.

Climatic factors shaping intraspecific leaf trait variation of a neotropical tree along a rainfall gradient.

Intraspecific trait variation has been singled out as an important mechanism by which individuals can cope with environmental variations and avoid local extinctions. Here we evaluate variation in metamer traits (i.e., traits associated with internodes, petioles and their corresponding leaves) and parameters of chlorophyll fluorescence within and among populations of a neotropical tree, Copaifera langsdorffii. We also evaluated phenotypic plasticity in natural settings comparing traits between shade and sun-exposed metamers. We selected six populations along a climatic gradient ranging from semi-arid to humid and representing three different biomes (Caatinga, Cerrado, and Atlantic Forest). Local climatic conditions significantly affected the morphological and physiological traits of populations. Trait variation among populations was explained mainly by aridity index and evapotranspiration. Individuals from drier regions had lower specific leaf area (SLA), lower investment in leaf area per total dry mass of metamer (LARm), lower specific petiole length (SPL) and lower potential quantum yield (Fv/Fm, only for sun-exposed metamers). Populations from locations with greater environmental heterogeneity (interannual variation) had greater plasticity in response to light for Fv/Fm and electron transport rate (ETR) and morphological traits related to the hydraulic and biomechanical aspects of the leaves (petiole length, internode length and SPL). High intraspecific variation in metamer traits in C. langsdorffii coupled with its ability to modify these traits in response to different climate conditions can explain the success of the species over a range of different habitats and represent important factors for the persistence of this species in the face of climate change.

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.

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

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.

Reproductive strategy of Copaifera langsdorffii (Fabaceae): more seeds or better seeds?

The trade-off seed size/number is a recognized phenomenon capable of shaping ecological processes of colonization and establishment of plant species. Studies that describe trade-offs size/number of seeds in supra-annual fruiting species are still rare. In this study, two predictions for trade-off size/number hypothesis were tested: (i) on a population scale, seeds produced during years of greater reproductive investment showed reduced size, and (ii) on individual scale there is an inverse relationship between size and number of seeds produced by individual plant. To test these predictions, 102 Copaifera langsdorffii adult plants were monitored monthly from January to September during four consecutive years (2008 to 2011) in order to study the reproductive investment of plants. C. langsdorffii plants exhibited reproductive activity only during years 2008 and 2011. The mean number of seeds per branch was 26.4 % greater in 2008 comparatively to 2011. It was also observed that seed size was greater in the year of 2008, when plants produced greater number of fruits. Thus, the data did not support the first prediction of seed size/number hypothesis. In both reproductive years, there was a negative relationship between seed size and number of seeds, supporting the second prediction of the seed size/number hypothesis. The interaction term with the reproductive year suggests that the trade-off size/number of seeds was indeed stronger in 2011, when the plants produced lower seed number. Finally, this study calls the attention to supra-annual fruiting pattern in C. langsdoffii, and suggests that the phenological patterns contribute to explain the wide variation in their seed size and geographical distribution.

El compromiso o balance adaptativo de las semillas en tamaño/ número (trade-off seed size/number) es un reconocido fenómeno capaz de dar forma a los procesos ecológicos de la colonización y el establecimiento de especies de plantas. Los estudios que describen el compromiso adaptativo de las semillas en tamaño/número en especies con fructificación supra-anual siguen siendo raros. En este estudio, se probaron dos hipótesis predictivas para compromiso adap-tativo de las semillas en tamaño/número: (i) una a escala poblacional, semillas producidas durante el año de mayor inversión reproductiva mostraran un tamaño reducido, y (ii) otra a escala individual que se dará una relación inversa entre el tamaño y número de semillas producidas por la planta individual. Para probar estas predicciones, 102 plantas adultas de Copaifera langsdorffii fueron monitoreadas mensualmente entre enero y septiembre, durante cuatro años consecutivos (2008-2011) con el fin de estudiar la inversión reproductiva de las plantas. Plantas langsdorffii C. exhibieron actividad reproductiva sólo durante el 2008 y 2011. El número promedio de semillas por rama fue 26.4 % mayor en 2008 en comparación con 2011. También se observó que el tamaño de la semilla fue mayor en 2008, cuando las plantas producen mayor número de frutos. Por lo tanto, los datos no apoyan la primera hipótesis de predicción de tamaño/número. En ambos años reproductivos, existía una relación negativa entre el tamaño de la semilla y el número de semillas, comprobando la segunda hipótesis de predicción del tamaño/número de semillas. El periodo de interacción con el año reproductivo sugiere que el compromiso adaptativo de las semillas en tamaño/número fue de hecho más fuerte en 2011, cuando las plantas produjeron baja cantidad de semillas. Por último, este estudio llama la atención por el patrón de fructificación supra-anual en C. langsdoffii, y sugiere que los patrones fenológicos contribuyen a explicar la amplia variación en el tamaño de la semilla y la distribución geográfica.

Ecological interactions shape the dynamics of seed predation in Acrocomia aculeata (Arecaceae).

BACKGROUND: The complex network of direct and indirect relationships determines not only the species abundances but also the community characteristics such as diversity and stability. In this context, seed predation is a direct interaction that affects the reproductive success of the plant. For Acrocomia aculeata, the seed predation by Pachymerus cardo and Speciomerus revoili in post-dispersal may destroy more than 70% of the propagules and is influenced by the herbivory of the fruits during pre-dispersal. Fruits of plants with a higher level of herbivory during pre-dispersal are less attacked by predators in post-dispersal. We proposed a hypothesis that describes this interaction as an indirect defense mediated by fungi in a multitrophic interaction. As explanations, we proposed the predictions: i) injuries caused by herbivores in the fruits of A. aculeata favor fungal colonization and ii) the colonization of A. acuelata fruit by decomposing fungi reduces the selection of the egg-laying site by predator. METHODOLOGY/PRINCIPAL FINDINGS: For prediction (i), differences in the fungal colonization in fruits with an intact or damaged epicarp were evaluated in fruits exposed in the field. For prediction (ii), we performed fruit observations in the field to determine the number of eggs of P. cardo and/or S. revoili per fruit and the amount of fungal colonization in the fruits. In another experiment, in the laboratory, we use P. cardo females in a triple-choice protocol. Each insect to choose one of the three options: healthy fruits, fruits with fungus, or an empty pot. The proposed hypothesis was corroborated. Fruits with injuries in the epicarp had a higher fungal colonization, and fruits colonized by fungi were less attractive for egg-laying by seed predators. CONCLUSION/SIGNIFICANCE: This study emphasizes the importance of exploring the networks of interactions between multitrophic systems to understand the dynamics and maintenance of natural populations.

Differential effects of land use on ant and herbivore insect communities associated with Caryocar brasiliense (Caryocaraceae).

Simplification of natural habitats leads to a modification of the community associated with a host plant. Pequi trees (Caryocar brasiliense) are common to find in central Brazil, especially in the middle of monocultures, such as soy, corn, pasturelands or Eucalyptus plantations. On this scenario we hypothesized that habitat modification differentially affects the diversity of ants and herbivore insects associated with this species. The aim of the work was to test if C. brasiliense trees located in human modified habitats, support a lower species richness and abundance of ants, and a greater species richness and abundance of insect herbivores, compared to preserved cerrado habitats. The study was conducted in a Cerrado area located in Northern Minas Gerais State, Brazil. Ants and herbivore insects were collected monthly during 2005 using beating technique. The results showed that ant species richness was higher in pequi trees located in preserved Cerrado, followed by trees in pastureland and Eucalyptus plantation, respectively. The ant abundance was lower in the Eucalyptus plantation but no difference in ant abundance was observed between trees in pastureland and the preserved Cerrado. Moreover, herbivore insects exhibited lower number of species and individuals in trees located in the preserved Cerrado than in the pastureland and Eucalyptus plantation. We concluded that habitats simplified by human activities may result in diversity loss and may change species interactions.

Differential effects of land use on ant and herbivore insect communities associated with Caryocar brasiliense (Caryocaraceae)

Simplification of natural habitats leads to a modification of the community associated with a host plant. Pequi trees (Caryocar brasiliense) are common to find in central Brazil, especially in the middle of monocultures, such as soy, corn, pasturelands or Eucalyptus plantations. On this scenario we hypothesized that habitat modification differentially affects the diversity of ants and herbivore insects associated with this species. The aim of the work was to test if C. brasiliense trees located in human modified habitats, support a lower species richness and abundance of ants, and a greater species richness and abundance of insect herbivores, compared to preserved cerrado habitats. The study was conducted in a Cerrado area located in Northern Minas Gerais State, Brazil. Ants and herbivore insects were collected monthly during 2005 using beating technique. The results showed that ant species richness was higher in pequi trees located in preserved Cerrado, followed by trees in pastureland and Eucalyptus plantation, respectively. The ant abundance was lower in the Eucalyptus plantation but no difference in ant abundance was observed between trees in pastureland and the preserved Cerrado. Moreover, herbivore insects exhibited lower number of species and individuals in trees located in the preserved Cerrado than in the pastureland and Eucalyptus plantation. We concluded that habitats simplified by human activities may result in diversity loss and may change species interactions.

La simplificación de los hábitats naturales conduce a una modificación de la comunidad asociada con una planta huésped. En la región central de Brasil es muy común encontrar árboles aislados de “pequi” (Caryocar brasiliense) en medio de monocultivos de soya, maíz, pastos o plantaciones de eucalipto. Bajo este escenario, nosotros predecimos que la modificación del hábitat afecta diferencialmente la diversidad de hormigas y herbívoros asociados a árboles de “pequi”. El objetivo de este artículo fue determinar que árboles de “pequi” localizados en hábitats modificados por actividades humanas (i.e. pastizales y plantaciones de Eucalyptus), soportan una menor abundancia y riqueza de especies de hormigas. Por el contrario, se espera una mayor abundancia y riqueza de especies de herbívoros en hábitats modificados. Este estudio fue realizado en un área de Cerrado (sabana brasileña) localizado en el Norte del estado de Minas Gerais, Brasil. Las hormigas y los herbívoros fueron muestreados usando técnicas de golpe. La riqueza de especies de hormigas fue mayor en árboles de “pequi” localizados en áreas preservadas de Cerrado, seguida por árboles de pastizales y finalmente por árboles de plantaciones de Eucalyptus. La abundancia de hormigas fue menor en plantaciones de Eucalyptus, pero no hubo diferencias en la abundancia de hormigas entre árboles de pastizales y áreas preservadas de cerrado. Por otra parte, el número de especies de insectos herbívoros fue menor en árboles de áreas preservadas de Cerrado que en pastizales y plantaciones de Eucalyptus. Concluimos que las actividades humanas reducen la complejidad del hábitat resultando en la pérdida de la diversidad de insectos y procesos ecológicos.

Insect herbivores associated with Baccharis dracunculifolia (Asteraceae): responses of gall-forming and free-feeding insects to latitudinal variation.

The spatial heterogeneity hypothesis has been invoked to explain the increase in species diversity from the poles to the tropics: the tropics may be more diverse because they contain more habitats and micro-habitats. In this paper, the spatial heterogeneity hypothesis prediction was tested by evaluating the variation in richness of two guilds of insect herbivores (gall-formers and free-feeders) associated with Baccharis dracunculifolia (Asteraceae) along a latitudinal variation in Brazil. The seventeen populations of B. dracunculifolia selected for insect herbivores sampling were within structurally similar habitats, along the N-S distributional limit of the host plant, near the Brazilian sea coast. Thirty shrubs were surveyed in each host plant population. A total of 8 201 galls and 864 free-feeding insect herbivores belonging to 28 families and 88 species were sampled. The majority of the insects found on B. dracunculifolia were restricted to a specific site rather than having a geographic distribution mirroring that of the host plant. Species richness of free-feeding insects was not affected by latitudinal variation corroborating the spatial heterogeneity hypothesis. Species richness of gall-forming insects was positively correlated with latitude, probably because galling insect associated with Baccharris genus radiated in Southern Brazil. Other diversity indices and evenness estimated for both gall-forming and free feeding insect herbivores, did not change with latitude, suggesting a general structure for different assemblages of herbivores associated with the host plant B. dracunculifolia. Thus it is probable that, insect fauna sample in each site resulted of large scale events, as speciation, migration and coevolution, while at local level, the population of these insects is regulated by ecological forces which operate in the system.

Insect herbivores associated with Baccharis dracunculifolia (Asteraceae): responses of gall-forming and free-feeding insects to latitudinal variation

The spatial heterogeneity hypothesis has been invoked to explain the increase in species diversity from the poles to the tropics: the tropics may be more diverse because they contain more habitats and microhabitats. In this paper, the spatial heterogeneity hypothesis prediction was tested by evaluating the variation in richness of two guilds of insect herbivores (gall-formers and free-feeders) associated with Baccharis dracunculifolia (Asteraceae) along a latitudinal variation in Brazil. The seventeen populations of B. dracunculifolia selected for insect herbivores sampling were within structurally similar habitats, along the N-S distributional limit of the host plant, near the Brazilian sea coast. Thirty shrubs were surveyed in each host plant population. A total of 8 201 galls and 864 free-feeding insect herbivores belonging to 28 families and 88 species were sampled. The majority of the insects found on B. dracunculifolia were restricted to a specific site rather than having ageographic distribution mirroring that of the host plant. Species richness of free-feeding insects was not affected by latitudinal variation corroborating the spatial heterogeneity hypothesis. Species richness of gall-forming insects was positively correlated with latitude, probably because galling insect associated with Baccharris genus radiated in Southern Brazil. Other diversity indices and evenness estimated for both gall-forming and free feeding insect herbivores, did not change with latitude, suggesting a general structure for different assemblages of herbivores associated with the host plant B. dracunculifolia. Thus it is probable that, insect fauna sample in each site resulted of large scale events, as speciation, migration and coevolution, while at local level, the population of these insects is regulated by ecological forces which operate in the system. Rev. Biol. Trop. 59 (3): 1419-1432. Epub 2011 September 01.

La hipótesis de heterogeneidad espacial se ha utilizado para explicar el aumento en la diversidad de especies desde los polos a los trópicos: los trópicos pueden ser más diversos ya que están conformados por una mayor cantidad de hábitats y micro-hábitats. En este estudio, la hipótesis de heterogeneidad espacial se puso a prueba evaluando la variación en la riqueza de dos gremios de insectos herbívoros (formadores de agallas y de alimentación libre) asociados con B. dracunculifolia (Asteracea) a lo largo de un gradiente latitudinal en Brasil. Las diecisiete poblaciones de B. dracunculifolia seleccionadas para el muestreo de insectos herbívoros estaban en hábitats con una estructura similar, a lo largo del límite Norte-Sur de distribución de la planta hospedera, cerca de la costa brasileña. De cada población de planta hospedera, se muestrearon treinta arbustos y se obtuvo un total de 8 201 agallas y 864 insectos de alimentación libre pertenecientes a 28 familias y 88 especies. La mayoría de los insectos que se encontraron en B. dracunculifolia estaban restringidos a un sitio específico en lugar de tener una distribución geográfica similar a la de la planta hospedera. La riqueza de especies de insectos de alimentación libre no se vió afectada por el gradiente latitudinal, por lo que se corroboró la hipótesis de heterogeneidad espacial. Mientras que la riqueza de especies de insectos formadores de agallas se correlacionó positivamente con la latitud, probablemente debido a que los insectos asociados al género Baccharis se extendieron hacia el sur de Brasil. Otros índices de diversidad y equidad estimados no variaron con la latitud para ninguno de los dos grupos de insectos herbívoros evaluados, lo que sugirie una estructura general para diferentes conjuntos de herbívoros asociados con la planta hospedera B. dracunculifolia. Por lo tanto, es probable que, la muestra de insectos en cada sitio sea resultado de eventos a gran escala, como la especiación, migración y coevolución; mientras que a nivel local la población de estos insectos está regulada por fuerzas ecológicas que operan dentro del sistema.

Ants and their effects on an insect herbivore community associated with the inflorescences of Byrsonima crassifolia ( Linnaeus ) H B K ( Malpighiaceae )

Os efeitos de formigas na comunidade de insetos em inflorescências de Byrsonima crassifolia (Malpighiaceae) foram testados em um experimento de exclusão em uma vegetacão de cerrado no Sudeste do Brasil. Quarenta e quatro espécies de insetos (23 famílias) e nove espécies de formigas (seis gêneros e três subfamílias) foram encontradas nas inflorescências de B. crassifolia. A exclusão das formigas, principalmente de Camponotus sericeiventris e de Camponotus spp. reduziu a populacão de membracídeos para 20por cento da abundância original. Exclusão das formigas e o tempo influenciaram a abundância de insetos mastigadores (exclusão, P<0,001; tempo, P<0,002) e sugadores (exclusão, P<0,02; tempo, P<0,01). Insetos mastigadores e sugadores foram encontrados duas vezes mais em inflorescências com formigas excluídas quando comparados com inflorescências controle (P<0,001). Insetos sugadores foram encontrados 1,5 vezes mais em inflorescências com formigas excluídas do que no controle. Apenas o tempo influenciou significativamente a riqueza de insetos mastigadores e sugadores associados com as inflorescências de B. crassifolia. Inflorescências em ramos controle foram significativamente menos atacadas por herbívoros do que inflorescências em ramos com formigas excluídas (P<0,001). Portanto, estes resultados sugerem que a presenca das formigas influencia a estrutura da comunidade de insetos herbívoros associados com B. crassifolia.