The evolutionary success of Asteraceae, the largest family of flowering plants, has been attributed to the unique inflorescence architecture of the family, which superficially resembles an individual flower. Here, we show that Asteraceae inflorescences (flower heads, or capitula) resemble solitary flowers not only morphologically but also at the molecular level. By conducting functional analyses for orthologs of the flower meristem identity genes LEAFY (LFY) and UNUSUAL FLORAL ORGANS (UFO) in Gerbera hybrida, we show that GhUFO is the master regulator of flower meristem identity, while GhLFY has evolved a novel, homeotic function during the evolution of head-like inflorescences. Resembling LFY expression in a single flower meristem, uniform expression of GhLFY in the inflorescence meristem defines the capitulum as a determinate structure that can assume floral fate upon ectopic GhUFO expression. We also show that GhLFY uniquely regulates the ontogeny of outer, expanded ray flowers but not inner, compact disc flowers, indicating that the distinction of different flower types in Asteraceae is connected with their independent evolutionary origins from separate branching systems.In flowering plants, inflorescences are the branched structures that bear flowers. Their architecture in terms of number and arrangement of flowers shows enormous variation in nature and plays a central role in angiosperm reproductive adaptation and success. Most of our knowledge of the molecular regulation of inflorescence architecture is based on studies of three major inflorescence types: racemes in Arabidopsis (Arabidopsis thaliana) or snapdragon (Antirrhinum majus), cymes in Solanaceae species such as petunia (Petunia hybrida) or tomato (Solanum lycopersicum), and panicles in grasses (Prusinkiewicz et al., 2007;Park et al., 2014;Teo et al., 2014). In the model plant Arabidopsis, endogenous and exogenous flowering-inducing signals convert the vegetative shoot meristem into an inflorescence meristem (IM) that initiates determinate flower meristems (FMs) on its flanks. The inflorescence forms a simple, indeterminate (monopodial) raceme that elongates and never forms a terminal flower due to maintenance of the stem cells in the central zone of the meristem. In Solanaceae, the cymous IM always terminates in a flower but forms new axillary IMs that continue growth, leading to a zig-zag-like sympodial branching pattern. Panicles in grasses show more complex lateral branching, and both apical and lateral meristems may form flowers. Using mathematical modeling, Prusinkiewicz et al. (2007) showed that a single developmental model (the so-called transient model) can generate the distinct inflorescence types (racemes, cymes, and panicles) found in nature.In all basic inflorescence types, flower meristem identity (FMI) is controlled by homologs of at least three functionally conserved proteins, LEAFY (LFY), UN-USUAL FLORAL ORGANS (UFO), and SEPALLATA3 (SEP3), that diverge in their spatiotemporal expression domains, leading to differences in IM patte...