1. The seeds of many plant species produce mucilage on their surfaces that when wetted and dried, firmly adheres seeds to surfaces and substrates. Previous studies have demonstrated that seed anchorage to the ground can reduce seed predation, although only a few species have thus far been tested.2. Here we investigated whether binding to the ground reduces seed removal by harvester ants Pogonomyrmex subdentatus, an important granivore, for 53 species with mucilaginous seeds. We also explored functional traits that associate with seed removal risk to understand the ecological and evolutionary context of this granivory resistance trait.3. Using a field cafeteria choice experiment, we compared harvester ant seed removal of wetted ground-bound seeds to dry unbound control seeds for these 53 species. We developed a simple assay to score dislodgement force. We examined whether this force, seed mass and seed mucilage production explained the interspecific variation in protection that we observed in field seed removal. We integrated these experiments with a broad scale test of correlates of seed attachment using a previously published dataset of attachment potential of mucilaginous seeds for 432 species, examining correlations of attachment potential with 13 plant traits and the climate characteristics of the species' range. 4. Binding to the ground reduced seed removal in 42 of 53 species tested. The benefit increased with seed dislodgement force, which itself increased with mucilage production, but not with seed mass. In the larger dataset, shorter plant life span, higher temperature, more solar radiation, higher humidity, fewer wet days per year and higher seed density correlated positively with the odds of seed attachment. We also found that attachment potential showed a concave down quadratic relationship with latitude, peaking at roughly 30°. No strong evidence that any of the other six predictors correlated with attachment potential was found.5. We demonstrate that protection from granivores is a widespread convergent function of seed mucilage and is associated with mucilage production. We highlight the need for increased mechanistic investigations into this common but poorly studied trait, particularly in relation to functional drivers of the broad patterns we found.

An undercited 1983 paper in Ecology by Patricia Fuller and Mark Hay found that seeds of a mint, when the mucilage was wetted and subsequently dried, became encrusted by the sandy desert substrate and that this coating reduced seed predation by harvester ants and rodents. We tested Fuller and Hay's defensive hypothesis in 53 plant species, finding that a mucilage-bound sand coating reduced predation by harvester ants in almost every species tested. While mucilage has many possible functions and probably did not evolve for the purpose of seed defense, in many dry areas it likely has this ecological function.

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Background and Aims Pendulous flowers (due to a flexible pedicel) are a common, convergent trait of hummingbird-pollinated flowers. However, the role of flexible pedicels remains uncertain despite several functional hypotheses. Here we present and test the ‘lever action hypothesis’: flexible pedicels allow pendulous flowers to move upwards from all sides, pushing the stigma and anthers against the underside of the feeding hummingbird regardless of which nectary is being visited. Methods To test whether this lever action increased pollination success, we wired emasculated flowers of serpentine columbine, Aquilegia eximia, to prevent levering and compared pollination success of immobilized flowers with emasculated unwired and wire controls. Key Results Seed set was significantly lower in wire-immobilized flowers than unwired control and wire control flowers. Video analysis of visits to wire-immobilized and unwired flowers demonstrated that birds contacted the stigmas and anthers of immobilized flowers less often than those of flowers with flexible pedicels. Conclusions We conclude that flexible pedicels permit the levering of reproductive structures onto a hovering bird. Hummingbirds, as uniquely large, hovering pollinators, differ from flies or bees which are too small to cause levering of flowers while hovering. Thus, flexible pedicels may be an adaptation to hummingbird pollination, in particular due to hummingbird size. We further speculate that this mechanism is effective only in radially symmetric flowers; in contrast, zygomorphic hummingbird-pollinated flowers are usually more or less horizontally oriented rather than having pendulous flowers and flexible pedicels.

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