Korovchinsky Nikolai M
A.N. Severtsov Institute of Ecology and Evolution of the Russian Academy of Sciences, Leninsky prospect 33, 119071 Moscow, Russian Federation.; Email:
Zootaxa. 2015 May 1;3955(1):1-44. doi: 10.11646/zootaxa.3955.1.1.
Two core species of the genus Bythotrephes Leydig, B. longimanus Leydig and B. cederströmii Schӧdler, are redescribed on the basis of the type material, topotypic material, and other materials from Western and Northern Europe. They were investigated with an application of detailed morphological analysis, including intra- and interpopulation variability and the original scheme of morphometric measurements. The neotype of B. longimanus was selected from the topotypic population of the species from Bodensee (Switzerland-Germany). The taxonomy of B. cederströmii and of the whole genus is complicated by the presence of a variety of similar forms with intermediate morphology supposed to be interspecific hybrids, which much confused previous researches. The evolutionary transformations of some morphological structures, such as thoracic limbs, abdomen, and caudal process, are analyzed and discussed. In particular, it is suggested that the internal side of the endopodites of the thoracic limbs of the second-fourth pairs grew in proximal direction, occupying the part of protopodite with the "gnathobasic" process, which remains to be homologous to those of the thoracic limbs of the first pair. The strongly reduced thoracic limbs of the fourth pair lost two terminal segments and have proved to be represented by three segments, two proximal of which are those of protopodite, while the third terminal segment is composed of the fused distal part of the protopodite and the first proximal endopodital segment, bearing transformed and specifically arranged setae. The specificity of the "gnathobasic" processes of Bythotrephes is that they are the derivatives of the second, not of the first, inner endite of the thoracic limb and for this reason they are not homologues to the gnathobases of other cladocerans and should be called pseudognathobases.The same position of the "gnathobasic" processes is characteristic for other Onychopoda, which stresses the uniqueness of the representatives of the order among Cladocera and the whole class Branchiopoda. The abdomen of Bythotrephes was found to be three-segmented, but the third distal segment has proved to be transformed and fused with the postabdomen, participating in the formation of a long and massive caudal process. The comparison of morphological features of Bythotrephes and Leptodora clearly shows their independent homoplasic evolutionary development and distant phylogenetic position. According to new available data, the range of B. longimanus embraces large and deep lakes of Switzerland, Southern Germany, North Italy, Austria, and Great Britain, whereas B. cederströmiis. str. is certainly known only in the south of Sweden (probably also in Finland). All other B. cederströmii-like forms, supposed to be interspecific hybrids, are much more widely distributed in Eurasia from Northern Europe and Central European Russia to the Lower Volga River and Yakutia in Eastern Siberia. It is suggested that just these hybrids invaded and conquered the North American inland waters.
基于模式标本、地模标本以及来自西欧和北欧的其他标本,对Bythotrephes Leydig属的两个核心物种——长吻Bythotrephes longimanus Leydig和谢氏Bythotrephes cederströmii Schӧdler进行了重新描述。运用详细的形态学分析方法对它们进行了研究,包括种群内和种群间的变异性以及形态测量的原始方案。长吻Bythotrephes的新模标本是从博登湖(瑞士 - 德国)该物种的地模种群中选取的。谢氏Bythotrephes以及整个属的分类因存在各种形态介于两者之间的类似形式(推测为种间杂种)而变得复杂,这使得先前的研究陷入了极大的困惑。对一些形态结构的进化转变进行了分析和讨论,如胸肢、腹部和尾突。特别指出的是,第二至四对胸肢内肢的内侧向近端生长,占据了具有“颚基”突起的原肢部分,该部分与第一对胸肢的相应部分仍为同源。第四对胸肢强烈退化,失去了两个末端节段,经证实由三个节段组成——两个近端节段是原肢的节段,而第三个末端节段由原肢的融合远端部分和内肢的第一个近端节段组成,其上着生有经过改造且排列特殊的刚毛。Bythotrephes的“颚基”突起的特殊性在于它们是胸肢第二对内叶而非第一对内叶的衍生物,因此它们与其他枝角类的颚基不同源,应称为假颚基。“颚基”突起的这种相同位置是其他猛水蚤目的特征,这突出了该目在枝角类以及整个鳃足纲中的独特性。发现Bythotrephes的腹部由三个节段组成,但第三个远端节段已发生转变并与后腹部融合,参与形成了一个长而粗壮的尾突。Bythotrephes和Leptodora形态特征的比较清楚地表明了它们独立的趋同进化发展和较远的系统发育位置。根据新获得的数据,长吻Bythotrephes的分布范围包括瑞士、德国南部、意大利北部、奥地利和英国的大型深水湖泊,而谢氏Bythotrephes指名亚种仅在瑞典南部(可能也在芬兰)被确切知晓。所有其他类似谢氏Bythotrephes的形式,推测为种间杂种,在欧亚大陆分布更为广泛,从北欧和俄罗斯中部到西伯利亚东部的伏尔加河下游和雅库特地区均有分布。有人认为正是这些杂种入侵并征服了北美内陆水域。