Epidemiology, Parasites and Vectors, Agricultural Research Council-Onderstepoort Veterinary Research, Onderstepoort 0110, South Africa; Department of Life and Consumer Sciences, University of South Africa, South Africa; Department of Veterinary Tropical Diseases, University of Pretoria, Pretoria 0110, South Africa.
Department of Parasitology, School of Chemistry and Molecular Biosciences, The University of Queensland, Brisbane, Queensland 4072, Australia.
Ticks Tick Borne Dis. 2021 Jul;12(4):101688. doi: 10.1016/j.ttbdis.2021.101688. Epub 2021 Feb 20.
Argasid systematics remains controversial with widespread adherence to the Hoogstraal (1985) classification scheme, even though it does not reflect evolutionary relationships and results in paraphyly for the main genera of soft ticks (Argasidae), namely Argas and Ornithodoros. The alternative classification scheme, proposed by Klompen and Oliver (1993), has problems of its own: most notably paraphyly of the subgenus Pavlovskyella and the controversial grouping together of the subgenera Alectorobius, Antricola, Carios, Chiropterargas, Nothoaspis, Parantricola, Reticulinasus and Subparmatus into the genus Carios. Recent phylogenetic analyses of 18S/28S rRNA sequences and mitochondrial genomes agree with the scheme of Klompen and Oliver (1993), with regard to the paraphyly of Pavlovskyella, placement of Alveonasus, Ogadenus, Proknekalia and Secretargas in the Argasinae and placement of Carios and Chiropterargas in the Ornithodorinae (Mans et al., 2019). The Carios clade and its constituent subgenera remain controversial, since the phylogenetic position of its type species Carios (Carios) vespertilionis Latreille, 1796 (formerly Argas vespertilionis) has not been determined with confidence. The current study aimed to resolve Carios sensu lato Klompen and Oliver, 1993, and Carios sensu stricto Hoogstraal, 1985, by determining and analysing phylogenetic nuclear and mitochondrial markers for C. (C.) vespertilionis. Both the nuclear and mitochondrial markers support placement of Carios s.s. within the subfamily Ornithodorinae, but to the exclusion of the clade that includes the 6 other subgenera that are part of Carios s.l. Klompen and Oliver (1993), namely Alectorobius, Antricola, Nothoaspis, Parantricola, Reticulinasus and Subparmatus. These 6 subgenera form a monophyletic clade that might be placed as new subgenera within the genus Alectorobius, or elevated to genera. Given the substantial differences in biology among these subgenera, we propose that these 6 subgenera be elevated to genera. Thus, we propose to modify the classification scheme of Mans et al. (2019) so that the subfamily Argasinae now has six genera, Alveonasus, Argas (subgenera Argas and Persicargas), Navis, Ogadenus, Proknekalia and Secretargas, and the subfamily Ornithodorinae has nine genera, Alectorobius, Antricola (subgenera Antricola and Parantricola), Carios, Chiropterargas, Nothoaspis, Ornithodoros (subgenera Microargas, Ornamentum, Ornithodoros, Pavlovskyella and Theriodoros), Otobius, Reticulinasus and Subparmatus (genera indicated in bold).
蜱系统学仍然存在争议,尽管广泛采用 Hoogstraal(1985)分类方案,但它不能反映进化关系,导致软蜱(蜱科)的主要属 Argas 和 Ornithodoros 出现并系现象。Klompen 和 Oliver(1993)提出的替代分类方案也存在自身问题:最明显的是 Pavlovskyella 的并系现象,以及 Alectorobius、Antricola、Carios、Chiropterargas、Nothoaspis、Parantricola、Reticulinasus 和 Subparmatus 亚属被争议性地归入 Carios 属。最近对 18S/28S rRNA 序列和线粒体基因组的系统发育分析与 Klompen 和 Oliver(1993)的方案一致,涉及 Pavlovskyella 的并系现象、Alveonasus、Ogadenus、Proknekalia 和 Secretargas 在 Argasinae 中的位置以及 Carios 和 Chiropterargas 在 Ornithodorinae 中的位置(Mans 等人,2019 年)。Carios 进化枝及其组成的亚属仍然存在争议,因为其模式种 Carios(Carios)vespertilionis Latreille, 1796(以前称为 Argas vespertilionis)的系统发育位置尚未确定。本研究旨在通过确定和分析 C.(C.)vespertilionis 的核和线粒体系统发育标记物来解决 Klompen 和 Oliver(1993)的 Carios 广义概念和 Carios 狭义概念 Hoogstraal(1985)。核和线粒体标记均支持 Carios s.s. 归入 Ornithodorinae 亚科,但排除了包括 6 个其他亚属的进化枝,这些亚属是 Carios s.l. 的一部分(Klompen 和 Oliver,1993),即 Alectorobius、Antricola、Nothoaspis、Parantricola、Reticulinasus 和 Subparmatus。这 6 个亚属形成一个单系进化枝,可能被归入 Alectorobius 属内的新亚属,或提升为属。鉴于这些亚属之间在生物学上存在显著差异,我们建议将这 6 个亚属提升为属。因此,我们建议修改 Mans 等人(2019 年)的分类方案,以便 Argasinae 亚科现在有 6 个属,Alveonasus、Argas(Argas 和 Persicargas 亚属)、Navis、Ogadenus、Proknekalia 和 Secretargas,而 Ornithodorinae 亚科有 9 个属,Alectorobius、Antricola(Antricola 和 Parantricola 亚属)、Carios、Chiropterargas、Nothoaspis、Ornithodoros(Microargas、Ornamentum、Ornithodoros、Pavlovskyella 和 Theriodoros 亚属)、Otobius、Reticulinasus 和 Subparmatus(斜体显示的属)。