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A challenge for studies on the organization of ant assemblages in forest ecosystems is to disentangle the causal effects of species occurrences. The structural and functional attributes of trees can act as environmental filters for ground-dwelling ant species influencing resource availability and the microclimate. The biotic interactions, especially competition, can work together with plant characteristics influencing ant species occurrences. To test the importance of tree traits and species interactions on co-occurrence patterns of ants, we collected grounddwelling ants, with pitfalls and litter sampling, beneath the canopies of four tree species during the rainy and dry seasons in a restored forest. We used five predictors (tree identity, crown size, trunk circumference, litter depth, and leaves density) to model the presence probabilities of ants . Hence, we applied habitat constrained null models in pairwise analyses to disentangle the causal effects of ants co-occurrences. The random pattern predominated in the assemblages, making up 96% of all possible species pairs combinations. Overall, 50% of the species pairs that showed nonrandomness in the ant occurrences were interpreted as resulting from environmental filters, 36% as negative associations and 14% as positive associations. Additionally, we found that the effects of season and the sampling technique on the ant assemblages were also important. We suggest that the ideas of the trees as templates and the paradigm of competition are both useful for understanding pairwise occurrence patterns in ant assemblages, and can be tested using tree traits as predictors in ant species distribution models for running constrained null models. A R T I C L E I N F O Article history: Received 29 March 2020 Accepted 19 January 2021 Available online 01 March 2021 Associate Editor: Ricardo Solar

revista brasileira de entomologia/revista brasileira de entomologia

Revisiting the ideas of trees as templates and the competition paradigm in pairwise analyses of ground-dwelling ant species occurrences in a tropical forest