接第一篇：

3 讨论

本文报道了一个线虫草属新物种，命名为羽束梗孢状线虫草，其与虫根线虫草在子座形态上均与Paraisaria 属物种相似，均在子座顶端产生球形可孕部，且子囊壳均为埋生(表3)，容易误将羽束梗孢类群错误当成虫根线虫草(Kobayasi 1937；Pérez-Villamares et al. 2017)。从形态特征来看，Paraisaria 属物种子座颜色较浅，而羽束梗孢状线虫草与虫根线虫草通常子座偏深褐色，在球形可孕部上的子囊壳顶端通常形成小突起，而Paraisaria 相对平整。本物种与虫根线虫草相比，寄主类型、子座颜色和质地等均有显著区别。从5 基因系统发育树看，O. paraisarioidea、O. entomorrhiza 与Tilachlidiopsis nigra 位于线虫草属基部，形成一个独立的分支(basal clade)。另外，ITS 序列的系统发育结果也显示O. paraisarioidea和O. entomorrhiza 与O.ravenelii clade 有紧密的亲缘关系。我们的研究较好地补充了线虫草属基部分支类群的物种多样性。该类群可能代表了线虫草属早期演化过程中一类形态与寄主特征多样、系统位置关键类型，对深入理解线虫草属的起源与早期辐射具有重要意义。从地理分布看，O. entomorrhiza 分布于日本和欧洲，而本文报道的O. paraisarioidea 目前仅见分布于中国西南生物多样性热点地区——横断山区，可能横断山区复杂的地形地貌和多样的生态环境为线虫草属基部类群的分化和O. paraisarioidea 的物种形成提供了生态位基础，显示该区域在物种形成与适应中的重要作用。

分子系统发育已成为当下物种识别鉴定及系统发育重要的研究手段(James et al. 2006；Spatafora et al. 2006；Sung et al. 2007a；Yang & Rannala 2012)，因此序列的有效性和完整性尤为关键。近年来，多基因系统发育逐渐取代单基因分析，成为线虫草属分类与系统发育研究的主流方法(Chen et al. 2013；Sanjuan et al. 2015；代永东 2018；Araújo & Hughes 2019；邹维秋 2022；Tang et al. 2023；杨涛 2023；Dai et al. 2024；周云 2024；Xu et al. 2025)。与单基因相比，多基因系统发育具备一定的容错能力，即允许部分基因缺失而不显著影响结果。本研究对多系统发育分析所用5 个基因的序列情况进行统计发现，不同基因存在不同程度的缺失(有效数据量：nrSSU 337、nrLSU 388、tef-1α 426、rpb1 401、rpb2 257)，其中rpb2 缺失最为严重，达到46%(202/439)。本研究在开展O. paraisarioidea 的分子测序过程中，成功扩增了ITS、nrSSU、nrLSU、tef-1α 和rpb1 基因，但rpb2 基因可能由于引物特异性不足或与目标物种序列匹配度较低等原因，经过多标本多次扩增，仍未获得目标序列。在系统树构建过程中，我们将O. paraisarioidea的rpb2 基因标注为“missing data”，所得结果在主要分支上的支持度依然较高，且ML 和BI 两种分析方法所得拓扑结构一致，能够有效解析O. paraisarioidea 的系统发育位置，并为其分类地位提供可靠的分子证据。但是需要注意的是，数据缺失不可避免地会对系统发育分析结果产生影响，应尽量减少此类情况的发生。后续应该设计更具有针对性的特异性引物，以补充相关数据。

本研究在构建线虫草属系统发育关系时，将寄主类型映射至线虫草属系统发育树上(图1A)，发现O. paraisarioidea 所属的线虫草属基部分支(Basel clade)及其近缘的O. ravenelii clade 的已知寄主均为鞘翅目昆虫[虽O. variabilis 寄主为双翅目食木虻幼虫，但与鞘翅目幼虫习性相似(Hodge et al. 1998)] ， 而本研究报道的O. paraisarioidea 却以鳞翅目为寄主。这一“特例”打破了该分支对寄主类型的传统一致性，为Araújo & Hughes (2019)提出的“线虫草属以鞘翅目为祖先寄主”的假说提供了新的视角，提示这一观点或需进一步探讨。这一发现暗示线虫草属在早期演化过程中可能存在更为复杂或多次发生的寄主转换事件，或在特定生态环境下形成了对非典型寄主的适应能力。尤其是在中国横断山区这一生态多样性极高的区域，宿主可获得性和环境选择压力可能促成了物种对鳞翅目等其他昆虫类群的利用，进而推动其生态位扩展和系统发育分化。

Paraisaria 是Samson & Brady (1983)根据Ophiocordyceps gracilis 的无性阶段特征即白色松散的束状孢梗束，轮生分枝的分生孢子梗，梗上具瓶状、以顶端增生方式繁殖的瓶梗，形成易聚集成孢子头状的单个透明的分生孢子而建立。Sung et al. (2007a)基于多基因系统发育研究将该属并入Ophiocordyceps 属中。随后，Mongkolsamrit et al. (2019)认为Paraisaria 在形态学上具有明显区别，且系统发育上形成一个相对独立且支持率较高的单系类群，因而将其从Ophiocordyceps 中恢复为独立属。然而，尽管Paraisaria 的形态特征具有较强的识别性，其属级地位在系统发育上仍缺乏充分支持，从本研究的结果来看，该类群位于O. ravenelii clade 分支内，且嵌套于Ophiocordyceps 属的系统发育框架之中(非单独分支)。因此目前尚不足以支持其作为一个独立属。当前Ophiocordyceps 属内界限与谱系关系仍存在争议，亟须依托更全面深入的数据、形态特征及生态信息进行综合解析，以推动该属系统分类的进一步厘清和优化。

作者贡献

常衬心：标本采集、实验设计与实施、数据分析、论文撰写；张忠霞：实验、数据分析；邓丽萍：数据分析、论文撰写、图片制作；梁建东：实验设计、数据分析、论文修改；俞琦：数据分析，论文审核与修改；代永东：论文构思、实验设计、数据分析、论文审核与修改。

利益冲突声明

该研究不存在任何潜在利益冲突的商业或财务关系。

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