Abstract
At least half of the 3600 species of Bromeliaceae are pollinated by hummingbirds. There is little doubt that the four to 12 evolutionary shifts towards and c. 32 shifts away from hummingbird pollination opened new evolutionary spaces for bromeliad diversification, and that hummingbird pollination has led to increased bromeliad diversification rates. However, the mechanisms leading to these increased rates remain unclear. We here propose that there are four main types of mechanisms that may increase diversification rates of hummingbird-pollinated bromeliad clades: (1) bromeliad speciation through adaptation to different hummingbird species; (2) increased allopatric speciation in hummingbird-pollinated clades due to lower pollen transfer efficiency compared with other pollinators; (3) differential speciation rates in hummingbird-pollinated clades dependent on of flowering phenology and hummingbird behaviour; and (4) higher speciation rates of bromeliads in montane environments (where hummingbird pollination predominates) due to topographic population fragmentation. To date, none of these hypotheses has been appropriately tested, partly due to a lack of data, but also because research so far has focused on documenting the pattern of increased diversification in hummingbird-pollinated clades, implicitly assuming that this pattern supports an underlying mechanism while ignoring the fact that several competing mechanisms may be considered. The aim of the present review is to increase awareness of these mechanisms and to trigger research aimed at specifically testing them. We conclude that much additional research on the roles of hummingbird behaviour and gene flow between bromeliad species is needed to elucidate their contribution to the evolution of diversity in bromeliads and other plant families.
Kessler, Michael