Premise of research. Floral synorganization is a structural feature of many speciose angiosperm taxa and is considered a morphological innovation paving the way for evolutionary diversification. Staphisagria is sister to the remaining Delphinieae, the only lineage of Ranunculaceae characterized by zygomorphic flowers. We aim at providing a description of floral organogenesis and morphogenesis in both Staphisagria species, presenting the disparity of Delphinieae hyperorgans in a phylogenetic framework, and proposing a scenario of likely developmental pathways underlying the different types of hyperorgans in Delphinieae. Methodology. We carried out morphological, anatomical, and developmental studies on flowers of S. macrosperma and S. picta. Pivotal results. Synorganization is complex in Staphisagria and Delphinieae as a whole and involves flower dorsoventralization, the nesting of spurs, the postgenital fusion of petals, and the formation of a shared cavity. From a choripetalous ancestor, late and partial postgenital fusion among dorsal petals evolved once or twice in the tribe. Conclusions. The Delphinieae flower includes nested spurs and nested floral parlors. These key innovations, unique in angiosperms, probably led to the diversification of this species-rich tribe in the Northern Hemisphere. The lengths of the inner (nectariferous) spurs and the nested floral parlors determine the range of pollinators able to collect nectar. These traits could be used to revise the circumscription of taxonomic groups within the tribe and should be taken into account when examining the possible coevolution between Delphinieae flowers and their pollinators. Integrating this new knowledge about the hyperorgan will be essential for future research in taxonomy, evo-devo, and pollination ecology in Delphinieae.

中文翻译：

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[斜体]金黄色葡萄球菌[/斜体]（毛茛科）的花器官发生和形态发生：对飞燕草科协同花结构进化的影响

研究的前提。花卉协同是许多物种被子植物分类群的结构特征，被认为是一种形态创新，为进化多样化铺平了道路。Staphisagria 是剩余的 Delphinieae 的姐妹，后者是毛茛科唯一以左右花为特征的谱系。我们的目标是描述两种金黄色葡萄属植物的花器官发生和形态发生，在系统发育框架中展示翠雀亚科高器官的差异，并提出了翠雀科不同类型高器官的可能发育途径的情景。方法。我们对 S. macrosperma 和 S. picta 的花进行了形态学、解剖学和发育研究。关键结果。整体而言，Staphisagria 和 Delphinieae 中的同构是复杂的，涉及花背腹化、刺的嵌套、花瓣的生殖后融合以及共享腔的形成。从一个脉络膜祖先，背花瓣之间的晚期和部分后生殖融合在部落中进化了一两次。结论。Delphinieae 花包括嵌套的马刺和嵌套的花房。这些在被子植物中独一无二的关键创新可能导致北半球这个物种丰富的部落的多样化。内部（花蜜）刺的长度和嵌套的花房决定了能够收集花蜜的传粉媒介的范围。这些特征可用于修改部落内分类群的范围，并且在检查翠雀科花与其授粉者之间可能的共同进化时应予以考虑。整合关于超器官的这些新知识对于未来在 Delphinieae 中进行分类学、进化进化和授粉生态学研究至关重要。