An elevational cline in leaf variegation: Testing anti-herbivory and abiotic heterogeneity hypotheses in maintaining a polymorphism.

ABSTRACT

PREMISE While some studies have found leaf variegation to reduce photosynthetic capacity, others showed that it can increase photosynthesis. Thus, what maintains variegation remains an open question. Two primary hypotheses-the anti-herbivory and abiotic heterogeneity hypotheses-have been posited, yet little empirical research explicitly investigates the maintenance of naturally occurring variegation. METHODS: We used field surveys, image analysis, and climatic associations to explore the anti-herbivory and abiotic heterogeneity hypotheses in 21 populations of Hexastylis heterophylla and H. shuttleworthii, both polymorphic for leaf variegation. We measured the frequency of variegated individuals, variegation intensity, and herbivory for each morph, assessed abiotic correlates with variegation, and measured photosynthetic efficiency. RESULTS: We found a strong elevational cline in leaf variegation strongly linked with abiotic heterogeneity; variegation was more common in lower-elevation populations characterized by higher temperatures, UV-B exposure, seasonal light change, and drier, more basic soils. Variegated and nonvariegated individuals experienced similar levels of herbivory. Morphs had similar photosynthetic quantum yields. However, nonvariegated leaves experienced more nonphotochemical quenching, an indication of photoinhibition, and had higher surface temperatures under high light. CONCLUSIONS: Our results suggest that variegation may serve as an adaptation to high temperatures and light conditions and can reduce photoinhibition in certain environmental contexts. Thus, abiotic factors can maintain variegation in wild populations and shape geographic clines in variegation.