Co-occurrence patterns in a diverse arboreal ant community are explained more by competition than habitat requirements.

A major goal of community ecology is to identify the patterns of species associations and the processes that shape them. Arboreal ants are extremely diverse and abundant, making them an interesting and valuable group for tackling this issue. Numerous studies have used observational data of species co-occurrence patterns to infer underlying assembly processes, but the complexity of these communities has resulted in few solid conclusions. This study takes advantage of an observational dataset that is unusually well-structured with respect to habitat attributes (tree species, tree sizes, and vegetation structure), to disentangle different factors influencing community organization. In particular, this study assesses the potential role of interspecific competition and habitat selection on the distribution patterns of an arboreal ant community by incorporating habitat attributes into the co-occurrence analyses. These findings are then contrasted against species traits, to explore functional explanations for the identified community patterns. We ran a suite of null models, first accounting only for the species incidence in the community and later incorporating habitat attributes in the null models. We performed analyses with all the species in the community and then with only the most common species using both a matrix-level approach and a pairwise-level approach. The co-occurrence patterns did not differ from randomness in the matrix-level approach accounting for all ant species in the community. However, a segregated pattern was detected for the most common ant species. Moreover, with the pairwise approach, we found a significant number of negative and positive pairs of species associations. Most of the segregated associations appear to be explained by competitive interactions between species, not habitat affiliations. This was supported by comparisons of species traits for significantly associated pairs. These results suggest that competition is the most important influence on the distribution patterns of arboreal ants within the focal community. Habitat attributes, in contrast, showed no significant influence on the matrix-wide results and affected only a few associations. In addition, the segregated pairs shared more biological characteristic in common than the aggregated and random ones.

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.