confer benefits to plants through reduction of herbivore damage. This species is not

commonly found in C. alliodora, however, and therefore this ant species probably has

little impact on the C. alliodora-ant relationship at the population level or across the

range of the plant.

 The results of the experimental leaf palatability trial with the specialist beetle

herbivore, Coptocycla leprosa, corroborated those from the survey of natural herbivory in

some ways but also differed on several important points. In the field, leaves from 1-yr-

old plants had higher proportional leaf damage than those from 5-yr-old plants, but in the

laboratory there was no difference between the two plant ages in leaf area consumed.

Although the insects responsible for the leaf damage in the field are unknown, this

difference could support the finding that ants limit C. leprosa damage in 5-yr-old trees

because it was so abundant at the site. Whereas in the laboratory trial there was less

herbivory on leaves from fertilized plants regardless of plant age, in the field the

fertilization effect was only marginally significant and appeared to have no effect at all

for the 5-yr-old plants. The contrast between the results of the leaf palatability trial and

the observed patterns of leaf damage in the field suggest that the beetles do not base their

foraging choices in nature solely on preference for the most palatable leaves. Rather,

they are most effective at attacking young plants that do not house large colonies of

defending ants.

 Fertilization significantly reduced the leaf area consumed in the palatability trial

and reduced herbivory of 1-yr-old, but not 5-yr-old, plants in the field survey of

herbivory. Together, these findings suggest that nutrient augmentation increases the

production of defensive chemicals in C. alliodora, and that the relative importance of