41Grasses have varying inflorescence shapes; however, little is known about the genetic mechanis ms 42 specifying such shapes among tribes. We identified the grass-specific TCP transcription factor 43 COMPOSITUM 1 (COM1) expressed in inflorescence meristematic boundaries of differe nt 44 grasses. COM1 specifies branch-inhibition in Triticeae (barley) versus branch-formation in non- 45 Triticeae grasses. Analyses of cell size, cell walls and transcripts revealed barley COM1 regulates 46 cell growth, affecting cell wall properties and signaling specifically in meristematic boundaries to 47 establish identity of adjacent meristems. COM1 acts upstream of the boundary gene Liguleless1 48 and confers meristem identity independent of the COM2 pathway. Furthermore, COM1 is subject 49 to purifying natural selection, thereby contributing to specification of the spike inflorescence shape. 50 This meristem identity module has conceptual implications for both inflorescence evolution and 51 molecular breeding in Triticeae. 52 53 54 55 56 57 58 59 60 61 4 Main Text: 62 63The grass family (Poaceae), one of the largest angiosperm families, has evolved a striking diversity 64 of inflorescence morphologies bearing complex structures such as branches and specialized 65 spikelets 1 . These structural features are key for sorting the grass family into tribes 1 . Current grass 66 inflorescences are proposed to originate from a primitive ancestral shape exhibiting "a relative ly 67 small panicle-like branching system made up of primary and secondary paracladia (branches), each 68 one standing single at the nodes" 2 ( Fig. 1A). This ancestral panicle-like inflorescence is also 69 known as a compound spike [3][4][5] . Several independent or combined diversification processes 70 throughout the evolutionary history of the grass family have resulted in the broad diversity of 71 today's grass inflorescences 2,3,6 . Some tribes, e.g. Oryzeae (rice) and Andropogoneae (maize and 72 sorghum), still display ancestral and complex compound shapes, keeping true-lateral long primary 73 and secondary branches. Other grasses, such as Brachypodium distachyon, show lower 74 inflorescence complexity with branch length and number reduced to lateral, small pedicels ending 75 in only one multi-floretted spikelet ( Fig. 1A-C). Inflorescences within the tribe Triticeae, e.g. 76 barley (Hordeum vulgare L.), probably evolved from the ancestral compound spike into the typical 77 unbranched spike (Fig. 1D). The spike displays the least-complex inflorescence shape due to the 78 sessile nature of spikelets and reduction in rachis internodes 2,7 . Architectural variation is often 79 manifested through subtle modifications of transcriptional programs during critical transitio na l 80 windows of inflorescence meristem (IM) maturation 7,8 or functional divergence of key 81 transcriptional regulators and/or other genes 9,10 . Identification of key genetic determinants is 82 crucial for better understanding and explaining both the origin of grass inflores...