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[鳅科(鲤亚科、鳅亚科)单殖吸虫(扁形动物门,单殖吸虫纲)的动物区系。2. 协同进化]

[Fauna of monogeneans (Monogenea, Platyhelminthes) of gudgeons (Gobioninae, Cyprininae). 2. Coevolution].

作者信息

Gerasev P I

出版信息

Parazitologiia. 2009 Jan-Feb;43(1):12-31.

Abstract

Coevolution of parasites and in particular monogeneans and fishes, was characterized from two related angles--parallel coevolution and host switches, lead to the acquisition of new hosts from taxonomically distant but ecologically similar groups. Modern views on the evolution and taxonomy of gudgeons are discussed. This group is very heterogeneous by morphology and biology, tales unclear taxonomic position in the family Cyprynidae, and can be divided into two tribes or four groups of genera. Analysis of coevolution of dactylogirids, ancyrocephalids, gyrodactylids, and diplozoids with their hosts was carried out. Dactylogyrus spp. and gudgeons show coevolutional connections; in Ancyrocephalus s. l. spp. coevolution and host switches take place; most of Gyrodactylus spp. parasitize fishes of the genus Gobio and are the colonists of gudgeons; and none of the diplozoids parasitize exceptionally gudgeons. From 130 species of 30 genera of the World fauna of gudgeons 31 species 13 genera proved to be infected with monogeneans. We divided the dactylogyrids from gudgeons (49 species) into 13 evolution levels and 30 morphological groups, which were found equal in number with taxonomic groups of hosts. Ancyrocephalids were divided into two evolution levels, which can not be reduced to one initial type, and eight monophyletic groups. Fourteen of 18 species of these monogeneans occurred on the host from the genera Hemibarbus and Squalidus. Different directions of evolution connected with forming of different types of copulatory organ, ventral additional bar, and type of attachment on gills were revealed for Dactylogyrus spp. from different groups of genera of gudgeons (Yang et al., 2006). Monogeneans have lower evolutionary rates, as compared with their hosts, that are expressed in the taxonomic ranks (groups of species in monogeneans and genera or groups of genera in their hosts). Number of species in Dactylogyrus and Ancyrocephalus s. l. usually the same as the number of their host species. However this proportion increases in favor of the parasites owing to the explosion-like speciation of monogeneans parasitizing Hemibarbus and Saurogobio. Distribution of morphological groups of monogeneans on gudgeons in general confirms their grouping into four groups of genera on the base of molecular data. At the same time, in some cases parasitological data agree with the gudgeons' classification based on morphological characters to a greater extent (Banarescu, 1992; Naseca, 1998). The genera Sarcocheilichthys and Gnathopogon take the crucial position in the Sarcocheilichthys group of genera according to the analysis of occurrence of monophyletic groups of Dactylogyrus spp. In the Pseudogobio group of genera, the largest number of the Dactylogyrus spp. groups was found on the genera Saurogobio and Abbottina. The genus Hemibarbus, being the host of some original groups of Dactylogyrus spp. and Ancyrocephalus spp., takes the most isolated position among gudgeons by morphological, molecular, and parasitological data. Relationships of Hemibarbus and Squalidus (both from the Hemibarbus group of genera) are need in further investigations. The Sarcocheilichthys group of genera, which contains the genera Coreius and Paracanthobrama with unclear and ambiguous positions, is situated apart from the Hemibarbus group of genera. High-specialized groups of Dactylogyrus spp. are predominating on the former. The Pseudogobio group of genera is parasitized by numerous primitive groups of Dactylogyrus spp. Groups of Dactylogyrus spp. from the Gobio group of genera are derived undoubtedly from the monogeneans parasitizing the Pseudogobio group of genera, and the Gobio group of genera either derived from the Pseudogobio group of genera or must be included into the latter. Thus, we have one phylum of gudgeons with separate groups of monogeneans (Hemibarbus group of genera) and, contrariwise, other phylum of fish hosts with ancient and specialized monogeneans (Sarcocheilichthys group of genera), which ends in the gudgeons with primitive monogeneans (Pseudogobio group of genera) and evolutionary young monogeneans (Gobio group of genera). The pattern of distribution of the morphological groups of monogeneans by gudgeons is a mirror reflection of modern contradictory views on the taxonomy and evolution of these fishes and may contribute to a more profound understanding of the place of gudgeons in Cyprinidae, their taxonomy and evolution.

摘要

寄生虫尤其是单殖吸虫与鱼类的协同进化,从两个相关角度进行了表征——平行协同进化和宿主转换,这导致从分类学上遥远但生态相似的类群中获取新宿主。讨论了关于鳅科鱼类进化和分类学的现代观点。该类群在形态和生物学上非常多样化,在鲤科中的分类地位不明确,可分为两个族或四个属组。对指环虫科、锚首虫科、三代虫科和双身虫科与其宿主的协同进化进行了分析。指环虫属物种与鳅科鱼类表现出协同进化关系;在广义锚首虫属物种中发生了协同进化和宿主转换;大多数三代虫属物种寄生于雅罗鱼属鱼类,是鳅科鱼类的殖民者;而没有双身虫专门寄生于鳅科鱼类。在世界鳅科动物区系的30个属的130个物种中,有13个属的31个物种被证明感染了单殖吸虫。我们将鳅科鱼类的指环虫(49个物种)分为13个进化水平和30个形态学组,发现其数量与宿主的分类学组相等。锚首虫科分为两个进化水平,不能归为一个初始类型,以及八个单系类群。这些单殖吸虫的18个物种中有14个寄生于半刺厚唇鱼属和麦穗鱼属的宿主上。从鳅科不同属组的指环虫属物种中揭示了与不同类型交配器官、腹侧附加杆以及鳃上附着类型形成相关的不同进化方向(Yang等人,2006)。与宿主相比,单殖吸虫的进化速率较低,这在分类学等级中表现出来(单殖吸虫中的物种组以及其宿主中的属或属组)。指环虫属和广义锚首虫属中的物种数量通常与其宿主物种数量相同。然而,由于寄生于半刺厚唇鱼属和似刺鳊鮈属的单殖吸虫的爆发式物种形成,这个比例对寄生虫更为有利。单殖吸虫在鳅科鱼类上的形态学类群分布总体上证实了它们基于分子数据被分为四个属组。同时,在某些情况下,寄生虫学数据在更大程度上与基于形态特征的鳅科鱼类分类一致(Banarescu,1992;Naseca,1998)。根据指环虫属单系类群出现情况的分析,厚唇鱼属和颌须鮈属在厚唇鱼属组的属中占据关键位置。在似鲴属组的属中,在似刺鳊鮈属和棒花鱼属上发现的指环虫属物种组数量最多。半刺厚唇鱼属作为一些原始指环虫属物种组和锚首虫属物种的宿主,在形态、分子和寄生虫学数据方面在鳅科鱼类中处于最孤立的位置。半刺厚唇鱼属和麦穗鱼属(均来自半刺厚唇鱼属组的属)之间的关系需要进一步研究。包含铜鱼属和似刺鳊鮈属且位置不明确和模糊的厚唇鱼属组的属,与半刺厚唇鱼属组的属分开。前者上以高度特化的指环虫属物种为主。似鲴属组的属被许多原始的指环虫属物种寄生。来自雅罗鱼属组的指环虫属物种组无疑源自寄生于似鲴属组的单殖吸虫,并且雅罗鱼属组的属要么源自似鲴属组的属,要么必须归入后者。因此,我们有一个鳅科鱼类门类,其中有单独的单殖吸虫类群(半刺厚唇鱼属组的属),相反,还有其他鱼类宿主门类,其中有古老和特化的单殖吸虫(厚唇鱼属组的属),在鳅科鱼类中以原始单殖吸虫(似鲴属组的属)和进化较年轻的单殖吸虫(雅罗鱼属组的属)结束。单殖吸虫在鳅科鱼类上的形态学类群分布模式是对这些鱼类分类学和进化的现代矛盾观点的一种镜像反映,可能有助于更深入地理解鳅科鱼类在鲤科中的地位、它们的分类学和进化。

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