Hummingbird-Syndrome Traits, Breeding System, and Pollinator Effectiveness in Two Syntopic Penstemon Species

Red-flowered, tubular Penstemon barbatus and Penstemon pinifolius, which have probably adapted independently to hummingbird pollinators, coflower on recently burned high-elevation slopes in the Chiricahua Mountains in Arizona. Hummingbirds visit both species but visit P. barbatus at a higher rate. We compared several traits and asked whether the species differed in specialization for hummingbird pollination. The corollas of P. pinifolius were 30% narrower than those of P. barbatus, but they were similar in tube length. Although a narrower corolla appeared to decrease access by large insects, small halictid bees were the only common insect visitors, and they gathered nectar from both species. Species differences in nectar production rate and concentration were minor, being smaller than within-species differences between dry and rainy periods. Taller inflorescences and a wider corolla tube may contribute to greater hummingbird use of P. barbatus. Penstemon pinifolius was fully self-compatible and weakly autogamous, whereas P. barbatus was strongly self-incompatible and had a longer flower life. Exclusion of hummingbirds reduced seed set fourfold in both species, supporting the inference from morphology and rewards that hummingbirds are the primary pollinators. In the absence of hummingbirds, halictids increased the seed set of P. barbatus relative to exclusion of all visitors, but they did not increase the seed set of P. pinifolius. Despite sharing the basic set of hummingbird-syndrome traits, the two species differed in degree of investment in individual flowers and in adaptations for outcrossing.