ABSTRACT The taxonomy of the dinoflagellate cyst-based genus Votadinium has been in need of revision. This is accomplished here by integrating morphology, large subunit (LSU) rDNA sequences, and cyst wall composition analyses through micro-Fourier transform infrared spectroscopy. Three new species, Votadinium pontifossatum, V. rhomboideum and V. reidii, are described as cyst stages of Protoperidinium paraoblongum, P. quadrioblongum, and P. steidingerae, respectively. A previously undescribed type of ornamentation is reported for V. pontifossatum. A fourth new species is assigned to V. concavum. Furthermore, Lejeunecysta psilodora Benedek (= Lejeunecysta psuchra Matsuoka), is re-described as V. psilodora and the holotype re-illustrated, as is done for the holotype of V. spinosum Reid. New illustrations are provided for V. elongatum and V. nanhaiense. LSU rDNA sequences of V. concavum reveal its placement in the Oceanica section of the genus Protoperidinium, supporting earlier evidence that the Oceanica section and the cyst-based genus Votadinium form a monophyletic clade. Thus, Votadinium could be considered the cyst equivalent of the motile-defined Oceanica section. The cyst walls of Votadinium spinosum, V. calvum and V. pontifossatum are built from carbohydrate-based macromolecules with N-containing functional groups, in agreement with the heterotrophic nature of their motile stage. Comparison with related genera Trinovantedinium and Quinquecuspis reveals that the species and genera can be separated based on cyst wall composition, supporting a phylogeny based on molecular data and/or ecological differences. Furthermore, Trinovantedinium applanatum spectra unexpectedly show an additional absorption at 1510cm–1, indicating an aromatic group that is commonly reported for sporopollenin and not seen in dinoflagellate cysts so far. This study stabilizes the taxonomy of the cyst genus Votadinium, clarifies its relationship to the Oceanica section of Protoperidinium, and further supports the application of whole cyst wall chemistry in helping address problems in dinoflagellate cyst taxonomy and ecology.

中文翻译：

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甲藻属 Votadinium Reid (Dinophyceae, Peridiniales, Protoperidiniaceae) 的分类学修订、系统发育和囊壁组成

摘要 基于甲藻囊肿的 Votadinium 属的分类需要修订。这是通过微傅立叶变换红外光谱整合形态学、大亚基 (LSU) rDNA 序列和囊壁成分分析来实现的。三个新物种 Votadinium pontifossatum、V. rhomboideum 和 V. reidii，分别被描述为 Protoperidinium paraoblongum、P. quadrioblongum 和 P. steidingerae 的囊肿阶段。V. pontifossatum 报道了一种以前未描述的装饰类型。第四个新物种被分配给 V. concavum。此外，Lejeunecysta psilodora Benedek (= Lejeunecysta psuchra Matsuoka) 被重新描述为 V. psilodora，并重新说明了正模标本，就像 V. spinosum Reid 的正模标本所做的那样。为 V. elongatum 和 V. 提供了新插图。南海恩。V. concavum 的 LSU rDNA 序列显示其位于 Protoperidinium 属的 Oceanica 部分，支持早期的证据，即 Oceanica 部分和基于囊肿的 Votadinium 属形成单系进化枝。因此，Votadinium 可以被认为是运动定义的 Oceanica 部分的囊肿等价物。Votadinium spinosum、V. calvum 和 V. pontifossatum 的胞囊壁是由具有含氮官能团的基于碳水化合物的大分子构建的，这与其运动阶段的异养性质一致。与相关属 Trinovantedinium 和 Quinquecuspis 的比较表明，物种和属可以根据胞囊壁组成进行分离，支持基于分子数据和/或生态差异的系统发育。此外，Trinovantedinium applanatum 光谱出人意料地在 1510cm-1 处显示出额外的吸收，表明一个芳香族基团通常用于孢粉质，目前在甲藻孢囊中未见。这项研究稳定了包囊属 Votadinium 的分类，阐明了它与 Protoperidinium 的 Oceanica 部分的关系，并进一步支持整个包囊壁化学在帮助解决甲藻包囊分类学和生态学问题中的应用。