• Flowers have been hypothesized to contain either modules of attraction and reproduction, functional modules (pollination‐effecting parts) or developmental modules (organ‐specific). Do pollination specialization and syndromes influence floral modularity?
• In order to test these hypotheses and answer this question, we focused on the genus Erica: we gathered 3D data from flowers of 19 species with diverse syndromes via computed tomography, and for the first time tested the above‐mentioned hypotheses via 3D geometric morphometrics. To provide an evolutionary framework for our results, we tested the evolutionary mode of floral shape, size and integration under the syndromes regime, and – for the first time – reconstructed the high‐dimensional floral shape of their most recent common ancestor.
• We demonstrate that the modularity of the 3D shape of generalist flowers depends on development and that of specialists is linked to function: modules of pollen deposition and receipt in bird syndrome, and access‐restriction to the floral reward in long‐proboscid fly syndrome. Only size and shape principal component 1 showed multiple‐optima selection, suggesting that they were co‐opted during evolution to adapt flowers to novel pollinators. Whole floral shape followed an Ornstein–Uhlenbeck (selection‐driven) evolutionary model, and differentiated relatively late.
• Flower shape modularity thus crucially depends on pollinator specialization and syndrome.

中文翻译：

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花形的模块化和进化：Erica中功能，发育和翼展的作用

• 假设花朵包含吸引力和繁殖模块，功能模块（传粉影响的部分）或发育模块（特定器官的）。授粉专业化和综合症会影响花卉模块化吗？
• 为了检验这些假设并回答这个问题，我们集中研究了埃里卡（Erica）属：我们通过计算机断层扫描从19种具有多种综合症的物种的花朵中收集了3D数据，并首次通过3D几何形态计量学检验了上述假设。为了为我们的结果提供一个进化框架，我们在综合症制度下测试了花形，大小和整合的进化模式，并首次重建了其最近祖先的高维花形。
• 我们证明，通才花的3D形状的模块性取决于开发，而专家的3D形状的模块性与功能有关：鸟类综合症中的花粉沉积和接收模块，以及长喙果蝇综合症中对花卉奖励的访问限制。只有大小和形状的主要成分1才表现出多重最佳选择，表明它们在进化过程中被选为使花朵适应新型授粉媒介。整个花形遵循Ornstein–Uhlenbeck（选择驱动）进化模型，并且分化相对较晚。
• 因此，花朵形状的模块化至关重要地取决于传粉媒介的专长和综合症。