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Long-term persistence of a neotropical ant-plant population in the absence of obligate plant-ants.

Interactions between ants and ant-plants are considered classic examples of obligate mutualisms. Previous studies have indicated that for many ant-plants the loss of ant colonies results in severe defoliation or mortality. Although individual plants can persist for some period of time without their mutualistic partners, to date populations of ant-free plants have only been recorded at high altitudes or on remote islands where herbivores are also scarce. We studied the interaction between ants, herbivores, and the ant-plant Tococa guianensis in the Cerrado region of central Brazil. Using a survey conducted across a large geographic region, we show that there is interpopulation variation in ant occupancy across sites and habitats. At most sites surveyed, plants were inhabited by Allomerus octoarticulatus, an obligate plant-ant. Plants with obligate ants had significantly lower standing levels of herbivore damage than plants with opportunistic ants and plants with no ant occupants. Furthermore, experimental removal of A. octoarticulatus resulted in increased levels of damage in both young and mature leaves. Despite the protection provided by obligate ants, populations of T. guianensis were found to persist without these ants in some areas. Plants without A. octoarticulatus had significantly greater leaf toughness and trichome density than those with A. octoarticulatus. Furthermore, trichome density in plants with A. octoarticulatus increased after ants were removed, probably as a response induced by increased levels of herbivore damage. To our knowledge, this is the first record of the occurrence of native myrmecophyte populations without their mutualistic ants in mainland low-elevation sites. Several factors may help to explain the long-term persistence of T. guianensis populations without plant-ants in some areas of the Brazilian Cerrado, including its potential for induced morphological defenses against insect herbivores and selection for increased levels of constitutive defenses. In addition, T. guianensis may be under lower herbivore pressure in the sites we studied than in other parts of its range because existing populations were small, isolated, and some were very close to the edge of this species' distribution.