27The orchid flower is renowned for complexity of flower organ morphogenesis and has 28 attracted great interest from scientists. The YABBY genes encode plant-specific transcription 29 factors with important roles in vegetative and reproductive development in seed plants. 30 DROOPING LEAF/CRABS CLAW (DL/CRC) orthologs are involved in reproductive organ 31 development (especially carpels) of angiosperms. Orchid gynostemium (the fused organ of 32 the androecium and gynoecium) and ovule development are unique developmental processes. 33 Understanding the DL/CRC-like genes controlling the developmental program of the 34 gynostemium and ovule could provide accessible information for reproductive organ 35 molecular regulation in orchids. Two DL/CRC-like genes, named PeDL1 and PeDL2, were 36 cloned from Phalaenopsis equestris. The orchid DL/CRC forms a monophyletic clade with 37 two subclades including AshDL, PeDL1 and DcaDL1 in subclade I, and PeDL2 and DcaDL2 38 in subclade II. The temporal and spatial expression analysis indicated PeDL genes are 39 specifically expressed in the gynostemium and at the early stages of ovule development. Both 40 PeDLs could partially complement an Arabidopsis crc-1 mutant. Transient overexpression of 41 PeDL1 in Phalaenopsis orchids caused abnormal development of ovule and stigmatic cavity 42 of gynostemium. PeDL1, instead of PeDL2, could form a heterodimer with PeCIN8. 43 Paralogue retention and subsequent divergence of the gene sequence of PeDL1 and PeDL2 in 44 P. equestris might result in the differentiation of function and protein behaviors. These results 45 reveal the important roles of PeDLs involved in orchid gynostemium and ovule development 46 and provide new insights for further understanding the molecular mechanisms underlying 47 orchid reproductive organ development. 48 49 52 Over the last couple of decades, genes involved in angiosperm floral organ specification 68 were identified and the 'ABCDE model' was proposed, with the combined effect of the A-, B-, 69 C-, D-, and E-class MADS-box genes determining the floral organ identity. In monocot 70 orchids, the Phalaenopsis species was used as a model plant to delineate the B-, C-, D-, and 71 E-class functions of MADS-box genes participating in specialized floral organ development 72 (Pan et al., 2014; Tsai et al., 2014). The most notable feature is that P. equestris contains four 73 B-class AP3-like genes with differential expression largely responsible for differentiation of 74 the two closely-spaced tripartite perianth whorls into three sepals of the outer whorl, versus 75 the inner whorl of two lateral petals and a median labellum (Tsai et al., 2004). Later, the 76 refined 'orchid code' and 'Homeotic Orchid Tepal (HOT)' models were, respectively, 77proposed to illustrate the regulation of perianth morphogenesis in orchids 78 4 (Mondragon- Palomino & Theissen, 2011; Pan et al., 2011). The studies of floral terata from 79Phalaenopsis and Cymbidium agreed that C-and D-class MADS-box genes correlated with 80 the gynostemium and...