Arthropod responses to the experimental isolation of Amazonian forest fragments

Arthropods are the most diverse and abundant group of animals found in tropical lowland forests, and in light of ongoing global change phenomena, it is essential to better understand their responses to anthropogenic disturbances. Here we present a review of arthropod responses to forest deforestation and fragmentation based on studies conducted at the Biological Dynamics of Forest Fragments Project (BDFFP), located in central Amazonia. These studies involved a wide range of arthropod groups. All but one of the studies evaluated changes in total species number or species density in relation to fragment size, (i.e. area effects), and one-third also evaluated edge effects. Our review indicates that almost every arthropod group studied showed some kind of response to reduction in forest area, including altered abundances, species richness or composition in comparisons of different-sized fragments, fragmented and non-fragmented areas, or comparisons of forest edges and forest interiors. These responses tended to be idiosyncratic, with some groups showing predicted declines in abundance or diversity in the fragments while others show no response or even increases. However, some of the observed effects on arthropods, or on the ecological processes in which they are involved, were transient. The most likely explanation for this was the rapid development of secondary growth around fragments, which greatly increased the connectivity between fragments and the remaining forest. Although the BDFFP has provided many insights regarding the effects of forest fragmentation on arthropod assemblages, many diverse groups, such as canopy arthropods, have received scant attention. For those that have been studied, much remains to be learned regarding the long-term dynamics of these assemblages and how landscape context influences local biodiversity. The BDFFP remains an exceptional site in which to investigate how the ecological interactions in which arthropods are engaged are altered in fragmented landscapes.

Arthropod responses to the experimental isolation of Amazonian forest fragments

Arthropods are the most diverse and abundant group of animals found in tropical lowland forests, and in light of ongoing global change phenomena, it is essential to better understand their responses to anthropogenic disturbances. Here we present a review of arthropod responses to forest deforestation and fragmentation based on studies conducted at the Biological Dynamics of Forest Fragments Project (BDFFP), located in central Amazonia. These studies involved a wide range of arthropod groups. All but one of the studies evaluated changes in total species number or species density in relation to fragment size, (i.e. area effects), and one-third also evaluated edge effects. Our review indicates that almost every arthropod group studied showed some kind of response to reduction in forest area, including altered abundances, species richness or composition in comparisons of different-sized fragments, fragmented and non-fragmented areas, or comparisons of forest edges and forest interiors. These responses tended to be idiosyncratic, with some groups showing predicted declines in abundance or diversity in the fragments while others show no response or even increases. However, some of the observed effects on arthropods, or on the ecological processes in which they are involved, were transient. The most likely explanation for this was the rapid development of secondary growth around fragments, which greatly increased the connectivity between fragments and the remaining forest. Although the BDFFP has provided many insights regarding the effects of forest fragmentation on arthropod assemblages, many diverse groups, such as canopy arthropods, have received scant attention. For those that have been studied, much remains to be learned regarding the long-term dynamics of these assemblages and how landscape context influences local biodiversity. The BDFFP remains an exceptional site in which to investigate how the ecological interactions in which arthropods are engaged are altered in fragmented landscapes.

Arthropod responses to the experimental isolation of Amazonian forest fragments

Arthropods are the most diverse and abundant group of animals found in tropical lowland forests, and in light of ongoing global change phenomena, it is essential to better understand their responses to anthropogenic disturbances. Here we present a review of arthropod responses to forest deforestation and fragmentation based on studies conducted at the Biological Dynamics of Forest Fragments Project (BDFFP), located in central Amazonia. These studies involved a wide range of arthropod groups. All but one of the studies evaluated changes in total species number or species density in relation to fragment size, (i.e. area effects), and one-third also evaluated edge effects. Our review indicates that almost every arthropod group studied showed some kind of response to reduction in forest area, including altered abundances, species richness or composition in comparisons of different-sized fragments, fragmented and non-fragmented areas, or comparisons of forest edges and forest interiors. These responses tended to be idiosyncratic, with some groups showing predicted declines in abundance or diversity in the fragments while others show no response or even increases. However, some of the observed effects on arthropods, or on the ecological processes in which they are involved, were transient. The most likely explanation for this was the rapid development of secondary growth around fragments, which greatly increased the connectivity between fragments and the remaining forest. Although the BDFFP has provided many insights regarding the effects of forest fragmentation on arthropod assemblages, many diverse groups, such as canopy arthropods, have received scant attention. For those that have been studied, much remains to be learned regarding the long-term dynamics of these assemblages and how landscape context influences local biodiversity. The BDFFP remains an exceptional site in which to investigate how the ecological interactions in which arthropods are engaged are altered in fragmented landscapes.

Estratificação vertical de formigas em cerrado strictu sensu no Parque Estadual da Serra de Caldas Novas, Goiás, Brasil

Formicidae is an abundant group in the soil and is also well distributed in all vegetation strata, representing a good model for studies on vertical stratification of the fauna. The aim of this study was to verify a possible stratification of ant assemblages inhabiting the soil, the lower and the higher vegetation strata. Data was collected in the Serra de Caldas Novas State Park, in an area of cerrado strictu sensu. Ant collection was performed using pitfall traps. A total of 11 traps were placed in the soil, 17 in the lower vegetation stratum (dominated by shrubs and small trees) and 23 in the higher vegetation stratum (dominated by taller, mature trees). Forty-nine species of ants from 15 genera and five subfamilies were collected. Accumulation curves indicated that there is 37.5% more species in the soil than in mature trees and 35% more species in mature trees than in shrubs/young trees. There was not a clear vertical stratification between the soil and the two vegetation strata. Therefore, the species present in the vegetation tended to represent a nested subset of those found in the soil. Even without a clear vertical stratification, the diversity of ants in the cerrado vegetation is high, and part of this diversity appears to be explained by the fact that some species are specialized in nesting and/or foraging in the soil, while others in the vegetation.

Formicidae é um grupo abundante no solo e também largamente distribuído em todos os estratos da vegetação, constituindo-se em um bom modelo biológico para estudos de estratificação vertical. O objetivo geral do trabalho foi determinar os padrões de estratificação da assembléia de formigas em três estratos verticais (solo, estrato arbustivo e estrato arbóreo). Os dados foram coletados no Parque Estadual da Serra de Caldas Novas, em uma área de cerrado strictu sensu. A coleta de formigas foi realizada utilizando-se armadilhas do tipo pitfall. Foram instaladas 11 armadilhas no solo, 17 em plantas do estrato arbustivo e 23 em plantas do estrato arbóreo (uma armadilha por planta). Foi coletado um total de 49 espécies de formigas, pertencentes a 15 gêneros e cinco subfamílias. Através de curvas de acumulação de espécies, pôde-se verificar que existem 37,5% a mais de espécies no solo do que nas árvores e 35% a mais de espécies nas árvores do que nos arbustos. Uma análise de agrupamento indicou uma similaridade mediana entre as espécies de formigas do solo e vegetação (estrato arbustivo e arbóreo). Uma análise de aninhamento demonstrou que as espécies presentes no solo são um subconjunto das espécies presentes nos dois estratos de vegetação. Mesmo sem uma estratificação vertical muito clara, a alta diversidade de formigas encontrada no cerrado pode ser explicada em parte pelo fato de algumas espécies serem especializadas em forragear e/ou nidificar no solo e outras na vegetação.

Estratificação vertical de formigas em cerrado strictu sensu no Parque Estadual da Serra de Caldas Novas, Goiás, Brasil

Formicidae is an abundant group in the soil and is also well distributed in all vegetation strata, representing a good model for studies on vertical stratification of the fauna. The aim of this study was to verify a possible stratification of ant assemblages inhabiting the soil, the lower and the higher vegetation strata. Data was collected in the Serra de Caldas Novas State Park, in an area of cerrado strictu sensu. Ant collection was performed using pitfall traps. A total of 11 traps were placed in the soil, 17 in the lower vegetation stratum (dominated by shrubs and small trees) and 23 in the higher vegetation stratum (dominated by taller, mature trees). Forty-nine species of ants from 15 genera and five subfamilies were collected. Accumulation curves indicated that there is 37.5% more species in the soil than in mature trees and 35% more species in mature trees than in shrubs/young trees. There was not a clear vertical stratification between the soil and the two vegetation strata. Therefore, the species present in the vegetation tended to represent a nested subset of those found in the soil. Even without a clear vertical stratification, the diversity of ants in the cerrado vegetation is high, and part of this diversity appears to be explained by the fact that some species are specialized in nesting and/or foraging in the soil, while others in the vegetation.

Formicidae é um grupo abundante no solo e também largamente distribuído em todos os estratos da vegetação, constituindo-se em um bom modelo biológico para estudos de estratificação vertical. O objetivo geral do trabalho foi determinar os padrões de estratificação da assembléia de formigas em três estratos verticais (solo, estrato arbustivo e estrato arbóreo). Os dados foram coletados no Parque Estadual da Serra de Caldas Novas, em uma área de cerrado strictu sensu. A coleta de formigas foi realizada utilizando-se armadilhas do tipo pitfall. Foram instaladas 11 armadilhas no solo, 17 em plantas do estrato arbustivo e 23 em plantas do estrato arbóreo (uma armadilha por planta). Foi coletado um total de 49 espécies de formigas, pertencentes a 15 gêneros e cinco subfamílias. Através de curvas de acumulação de espécies, pôde-se verificar que existem 37,5% a mais de espécies no solo do que nas árvores e 35% a mais de espécies nas árvores do que nos arbustos. Uma análise de agrupamento indicou uma similaridade mediana entre as espécies de formigas do solo e vegetação (estrato arbustivo e arbóreo). Uma análise de aninhamento demonstrou que as espécies presentes no solo são um subconjunto das espécies presentes nos dois estratos de vegetação. Mesmo sem uma estratificação vertical muito clara, a alta diversidade de formigas encontrada no cerrado pode ser explicada em parte pelo fato de algumas espécies serem especializadas em forragear e/ou nidificar no solo e outras na vegetação.

Levels of leaf Herbivory in Amazonian trees from different stages In forest regeneration

Leaves from 120 canopy trees and 60 understory tree saplings growing in primary and secondary forests near Manaus, Brazil, were collected for determination of standing levels of herbivory (percent leaf area lost). Overall, levels of herbivory on leaves of central Amazonian trees were low. About one quarter of the leaves examined (n = 855) had no damage at all. In most other Neotropical sites studied the mean percentage of herbivory was found to vary between 5.7 and 13.1%, whereas in Manaus it was only 3.1%. The data presented here support the contention that levels of herbivore damage are positively related to soil fertility. No significant difference was found in herbivory levels between canopy trees and understory saplings. Also, there was no difference in damage between leaves from pioneer and late successional trees. Field assays of preference, however, revealed that leaves from pioneer trees are more palatable to leaf-cutting ants (Atta laevigata). This effect was dependent upon leaf age, being observed in mature leaves, but not in young leaves. The greater rate of leaf production in secondary forests may be a factor accounting for the greater abundance of leaf-cutting ants in secondary compared to primary forests.

Folhas de 120 árvores do dossel e de 60 arvoretas do sub-bosque foram coletadas em florestas primárias e secundárias de terra-firme próximas a Manaus, para medição dos níveis de herbivoria (porcentagem da área foliar danificada por herbívoros). Os níveis de herbivoria foram baixos. Cerca de 25% das folhas examinadas (n = 855) não tinham dano algum. Na maioria das outras florestas Neotropicais estudadas a porcentagem média de herbivoria ficou entre 5,7 e 13,1%, enquanto em Manaus este valor foi de apenas 3,1%. Os dados apresentados aqui dão apoio a hipótese de que os níveis dc herbivoria são positivamente relacionados a fertilidade do solo. Não houve diferença no nível de herbivoria entre árvores do dossel e arvoretas do sub-bosque. Também, não encontrou-se diferença entre folhas de plantas pioneiras da mata secundária e aquelas de final de sucessão encontradas na mata primária. Testes de preferência, entretanto, mostraram que folhas das plantas pioneiras são mais palatáveis as saúvas (Atta laevigata). Este efeito foi dependente da idade foliar, tendo sido observado em folhas maduras mas não em folhas jovens. A maior taxa de produção de folhas na floresta secundária pode ser um dos fatores responsáveis pela maior abundância de saúvas na mata secundária do que na mata primária.

Variação espaço-temporal na atividade forrageira da Saúva (Atta laevigata)

Forest regeneration in abandoned lands of Amazonia can be affected by leaf-cutting ants, since these ants are important predators of the seeds and seedlings of tree species which eventually arrive or emerge in these areas. However, little is known about how rates of seed and seedling predation vary in space and time. With this objective, I established a plot of 80 by 120 m, divided into 10 10 m quadrants, in an abandoned field near Manaus, in Brazilian Amazonia. There were three leaf-cutting ant (Atta laevigata) nests close to this plot. For one year, I determined which plants (tree species only) were attacked by the ants. I also determined the temporal and spatial variation in seed predation. For this, "seed baits" (20 Phaseolus vulgaris seeds in each) were distributed along the plot for one year. Finally, I determined if seedlings (Bellucia imperialis) transplanted into bare soil had a greater chance of being attacked than those transplanted into areas with a dense grass cover. Results show that seed predation activity was spatially aggregated. More predation was observed in the south-east and north-west portions of the plot, generally close to where nests were located. Strong temporal variation in seed predation rates was also observed. The greatest rates were observed around July, while the lowest ones around January. The temporal distribution of attacks on tree seedlings and saplings followed a similar pattern. There was a clear decrease in the number of plants attacked between November and January. The proportion of plants attacked per quadrant varied between 6 and 89%. The number of B. imperialis seedlings attacked did not vary in response to vegetation cover (with or without grass cover), presumably due to the large spatial heterogeneity in A. laevigata foraging activity, which obscured any eventual effect of vegetation cover.

Na Amazônia, a regeneração natural da floresta em pastos e cultivos abandonados pode ser afetada pelas saúvas, pois estas são importantes predadoras das plântulas e das sementes das espécies arbóreas que eventualmente ali se estabelecem. Entretanto, pouco se sabe sobre como as chances de uma dada planta ser atacada ou de uma semente ser removida pelas saúvas varia no tempo e no espaço. Com este intuito, estabeleci uma parcela de 80 120 m, subdividida em quadrantes de 10 10m, em um campo abandonado próximo a Manaus. Na periferia desta parcela haviam 3 ninhos da saúva Atta laevigata. Ao longo de 1 ano, determinei quais plantas (das espécies arbóreas somente) foram atacadas pelas formigas. Determinei também como variou a atividade de coleta de sementes por A. laevigata. Para isto utilizei grãos de feijão (Phaseolus vulgaris). Iscas, com cerca de 20 feijões cada, foram distribuídas pela parcela ao longo do ano. Finalmente, determinei se plantas (mudas de Bellucia imperialis) estabelecidas em solo desnudo tinham uma maior chance de serem atacadas do que aquelas estabelecidas em solo com cobertura de gramíneas. Os resultados mostram que a atividade de coleta de sementes foi espacialmente agregada. A maior atividade foi observada nas regiões sudeste e noroeste da parcela, em geral, próximo aos ninhos. Forte variação temporal na coleta de sementes também foi observada. O pico máximo de atividade ocorreu em meados de julho e o de menor atividade em meados de janeiro. A atividade de corte seguiu um padrão similar. Houve uma clara diminuição no número de plantas atacadas entre novembro e janeiro. A proporção de plantas atacadas por parcela variou entre 6 e 89%. A incidência de ataques por saúvas sobre mudas de B. imperialis não variou em função da cobertura vegetal (com ou sem gramíneas). Isto possivelmente em função da forte heterogeneidade espacial na atividade de A. laevigata, que encobriu qualquer efeito devido a cobertura vegetal.

Variação espaço-temporal na atividade forrageira da Saúva (Atta laevigata)

Forest regeneration in abandoned lands of Amazonia can be affected by leaf-cutting ants, since these ants are important predators of the seeds and seedlings of tree species which eventually arrive or emerge in these areas. However, little is known about how rates of seed and seedling predation vary in space and time. With this objective, I established a plot of 80 by 120 m, divided into 10 10 m quadrants, in an abandoned field near Manaus, in Brazilian Amazonia. There were three leaf-cutting ant (Atta laevigata) nests close to this plot. For one year, I determined which plants (tree species only) were attacked by the ants. I also determined the temporal and spatial variation in seed predation. For this, "seed baits" (20 Phaseolus vulgaris seeds in each) were distributed along the plot for one year. Finally, I determined if seedlings (Bellucia imperialis) transplanted into bare soil had a greater chance of being attacked than those transplanted into areas with a dense grass cover. Results show that seed predation activity was spatially aggregated. More predation was observed in the south-east and north-west portions of the plot, generally close to where nests were located. Strong temporal variation in seed predation rates was also observed. The greatest rates were observed around July, while the lowest ones around January. The temporal distribution of attacks on tree seedlings and saplings followed a similar pattern. There was a clear decrease in the number of plants attacked between November and January. The proportion of plants attacked per quadrant varied between 6 and 89%. The number of B. imperialis seedlings attacked did not vary in response to vegetation cover (with or without grass cover), presumably due to the large spatial heterogeneity in A. laevigata foraging activity, which obscured any eventual effect of vegetation cover.

Na Amazônia, a regeneração natural da floresta em pastos e cultivos abandonados pode ser afetada pelas saúvas, pois estas são importantes predadoras das plântulas e das sementes das espécies arbóreas que eventualmente ali se estabelecem. Entretanto, pouco se sabe sobre como as chances de uma dada planta ser atacada ou de uma semente ser removida pelas saúvas varia no tempo e no espaço. Com este intuito, estabeleci uma parcela de 80 120 m, subdividida em quadrantes de 10 10m, em um campo abandonado próximo a Manaus. Na periferia desta parcela haviam 3 ninhos da saúva Atta laevigata. Ao longo de 1 ano, determinei quais plantas (das espécies arbóreas somente) foram atacadas pelas formigas. Determinei também como variou a atividade de coleta de sementes por A. laevigata. Para isto utilizei grãos de feijão (Phaseolus vulgaris). Iscas, com cerca de 20 feijões cada, foram distribuídas pela parcela ao longo do ano. Finalmente, determinei se plantas (mudas de Bellucia imperialis) estabelecidas em solo desnudo tinham uma maior chance de serem atacadas do que aquelas estabelecidas em solo com cobertura de gramíneas. Os resultados mostram que a atividade de coleta de sementes foi espacialmente agregada. A maior atividade foi observada nas regiões sudeste e noroeste da parcela, em geral, próximo aos ninhos. Forte variação temporal na coleta de sementes também foi observada. O pico máximo de atividade ocorreu em meados de julho e o de menor atividade em meados de janeiro. A atividade de corte seguiu um padrão similar. Houve uma clara diminuição no número de plantas atacadas entre novembro e janeiro. A proporção de plantas atacadas por parcela variou entre 6 e 89%. A incidência de ataques por saúvas sobre mudas de B. imperialis não variou em função da cobertura vegetal (com ou sem gramíneas). Isto possivelmente em função da forte heterogeneidade espacial na atividade de A. laevigata, que encobriu qualquer efeito devido a cobertura vegetal.

Levels of leaf Herbivory in Amazonian trees from different stages In forest regeneration

Leaves from 120 canopy trees and 60 understory tree saplings growing in primary and secondary forests near Manaus, Brazil, were collected for determination of standing levels of herbivory (percent leaf area lost). Overall, levels of herbivory on leaves of central Amazonian trees were low. About one quarter of the leaves examined (n = 855) had no damage at all. In most other Neotropical sites studied the mean percentage of herbivory was found to vary between 5.7 and 13.1%, whereas in Manaus it was only 3.1%. The data presented here support the contention that levels of herbivore damage are positively related to soil fertility. No significant difference was found in herbivory levels between canopy trees and understory saplings. Also, there was no difference in damage between leaves from pioneer and late successional trees. Field assays of preference, however, revealed that leaves from pioneer trees are more palatable to leaf-cutting ants (Atta laevigata). This effect was dependent upon leaf age, being observed in mature leaves, but not in young leaves. The greater rate of leaf production in secondary forests may be a factor accounting for the greater abundance of leaf-cutting ants in secondary compared to primary forests.

Folhas de 120 árvores do dossel e de 60 arvoretas do sub-bosque foram coletadas em florestas primárias e secundárias de terra-firme próximas a Manaus, para medição dos níveis de herbivoria (porcentagem da área foliar danificada por herbívoros). Os níveis de herbivoria foram baixos. Cerca de 25% das folhas examinadas (n = 855) não tinham dano algum. Na maioria das outras florestas Neotropicais estudadas a porcentagem média de herbivoria ficou entre 5,7 e 13,1%, enquanto em Manaus este valor foi de apenas 3,1%. Os dados apresentados aqui dão apoio a hipótese de que os níveis dc herbivoria são positivamente relacionados a fertilidade do solo. Não houve diferença no nível de herbivoria entre árvores do dossel e arvoretas do sub-bosque. Também, não encontrou-se diferença entre folhas de plantas pioneiras da mata secundária e aquelas de final de sucessão encontradas na mata primária. Testes de preferência, entretanto, mostraram que folhas das plantas pioneiras são mais palatáveis as saúvas (Atta laevigata). Este efeito foi dependente da idade foliar, tendo sido observado em folhas maduras mas não em folhas jovens. A maior taxa de produção de folhas na floresta secundária pode ser um dos fatores responsáveis pela maior abundância de saúvas na mata secundária do que na mata primária.