Variable extrafloral nectary expression and its consequences in quaking aspen

Extrafloral nectaries (EFNs) are secretory glands most commonly linked to defensive mutualisms. Both a plant's need for defense and the strength of defense provided by mutualists will vary with plant condition and local insect community. Thus, the benefit of EFNs may vary spatially and temporally. However, little attention has been paid to natural variation in the presence and abundance of EFNs within and among individuals of the same species. Quaking aspen, Populus tremuloides Michx., bear EFNs on a subset of their leaves. Here, we describe patterns of EFN expression on shoots within ramets, among ramets, and among putative clones in interior Alaska. We also examine the relationship between EFN presence and herbivory by both the very abundant aspen leaf miner, Phyllocnistis populiella Chambers, and less common chewing herbivores. The proportion of leaves bearing EFNs varied from 33% to 87% among distinct aspen stands. Within stands, short (1-2 m height) ramets had higher EFN frequency than their taller (> 4 m) neighbors. Patterns of herbivory also differed between short and tall ramets. Compared with leaves without EFNs, those with EFNs suffered less mining damage on short ramets but slightly higher damage on tall ramets. Tall ramets suffered more chewing damage than short ramets, but this damage was unrelated to the presence of EFNs. Our results suggest that variable EFN expression may be explained by variation in the benefits of EFNs. Leaves with EFNs on short ramets benefit through reduction in leaf mining, but this benefit does not extend to tall ramets or other forms of herbivory.