Hybridization and recurrent evolution of left–right reversal in the land snail genus Schileykula (Orculidae, Pulmonata),Journal of Zoological Systematics and Evolutionary Research

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Hybridization and recurrent evolution of left–right reversal in the land snail genus Schileykula (Orculidae, Pulmonata)
Journal of Zoological Systematics and Evolutionary Research ( IF 2.0 ) Pub Date : 2019-12-10 , DOI: 10.1111/jzs.12353
Josef Harl _{1,

2} , Elisabeth Haring _{2,

3} , Barna Páll-Gergely ₄

Affiliation

Abstract The land snail genus Schileykula Gittenberger, 1983 is distributed in arid limestone areas from western Turkey to north‐western Iran. It comprises eight species, which display high variation in shell size and morphology. The cylindrical shells are 5–12 mm in height and the last shell whorls bear several inner lamellae and plicae. Two taxa differ in their chirality having sinistral shells, while all the others are dextrals such as the vast majority of orculids. The aim of this study was to establish a molecular genetic phylogeny of Schileykula and to test whether it conforms to the current morphology‐based classification. Furthermore, we were interested in the phylogenetic position of the two sinistral forms in order to assess whether one or two reversals happened in the evolution of the genus. Nine out of ten species, including all four subspecies of Schileykula trapezensis and three of six subspecies of Schileykula scyphus, were investigated. A section of the mitochondrial cytochrome c oxidase subunit I gene was analyzed in 54 specimens of Schileykula and from a subsample, partial sequences of the mitochondrial genes for the 12S rRNA and the 16S rRNA, and a section of the nuclear H4/H3 histone gene cluster were obtained. The phylogenetic trees based on the mitochondrial sequences feature high support values for most nodes, and the species appear well differentiated from each other. The two chiral forms evolved independently and are not sister lineages. However, some groupings disagree with the present morphology‐based classification and taxonomical conclusions are drawn. Schileykula trapezensis is polyphyletic in the molecular genetic trees; therefore, three of its subspecies are elevated to species level: Schileykula acampsis Hausdorf, 1996 comb. nov., Schileykula neuberti Hausdorf, 1996 comb. nov., and Schileykula contraria Neubert, 1993 comb. nov. Furthermore, Schileykula sigma is grouped within S. scyphus in the mitochondrial and nuclear trees and consequently treated as a subspecies of the latter (Schileykula scyphus sigma Hausdorf, 1996 comb. nov.). Schileykula nordsiecki, whose shell morphology is indistinguishable from that of the neighboring Schileykula scyphus lycaonica, but who differs in its genital anatomy, was confirmed to represent a distinct lineage. The phylogenies produced by the mitochondrial and nuclear data sets are to some extent conflicting. The patterns differ concerning the grouping of some specimens, suggesting at least two independent hybridization events involving S. contraria, S. scyphus and S. trapezensis. The results exemplify the importance of integrating both mitochondrial and nuclear sequence data in order to complement morphology‐based taxonomy, and they provide further evidence for hybridization across distantly related lineages in land snails.

中文翻译：

陆生蜗牛属Schileykula（Orculidae，Pulmonata）左右反转的杂交和循环进化

摘要陆地蜗牛属 Schileykula Gittenberger, 1983 分布于从土耳其西部到伊朗西北部的干旱石灰岩地区。它由八个物种组成，它们的贝壳大小和形态存在很大差异。圆柱形壳高 5-12 毫米，最后的壳螺纹具有数个内层片和皱襞。两个类群的手性不同，它们具有左旋贝壳，而所有其他类群都是右旋贝壳，例如绝大多数 Orculids。本研究的目的是建立 Schileykula 的分子遗传系统发育并测试它是否符合当前基于形态的分类。此外，我们对两种左旋形式的系统发育位置感兴趣，以便评估该属的进化中是否发生了一两次逆转。对十分之九的物种进行了调查，包括 Schileykula trapezensis 的所有四个亚种和 Schileykula scyphus 的六个亚种中的三个。对 54 个 Schileykula 样本中的线粒体细胞色素 c 氧化酶亚基 I 基因的一部分进行了分析，并从子样本中分析了 12S rRNA 和 16S rRNA 线粒体基因的部分序列，以及核 H4/H3 组蛋白基因簇的一部分获得。基于线粒体序列的系统发育树对大多数节点具有很高的支持值，并且物种之间似乎有很好的分化。这两种手性形式独立进化，不是姐妹谱系。然而，一些分组不同意目前基于形态的分类并得出分类学结论。 Schileykula trapezensis 在分子遗传树上是多系的；因此，其三个亚种被提升至物种水平：Schileykula acampsis Hausdorf，1996comb。十一月，Schileykula neuberti Hausdorf，1996 年梳子。 11 月，Schileykula contraria Neubert，1993 年梳。十一月此外，Schileykula sigma 在线粒体和核树中属于 S. scyphus，因此被视为后者的亚种（Schileykula scyphus sigma Hausdorf，1996comb.nov.）。 Schileykula Nordsiecki 的外壳形态与邻近的 Schileykula scyphus lycaonica 没有区别，但其生殖器解剖结构不同，被证实代表了一个独特的谱系。线粒体和核数据集产生的系统发育在某种程度上是相互矛盾的。一些标本的分组模式有所不同，表明至少有两个独立的杂交事件涉及 S. contraria、S. scyphus 和 S. trapezensis。这些结果证明了整合线粒体和核序列数据以补充基于形态的分类学的重要性，并且它们为陆地蜗牛中远缘相关谱系的杂交提供了进一步的证据。

更新日期：2019-12-10

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