Martynov A V
Zh Obshch Biol. 2013 May-Jun;74(3):201-40.
Evolutionary history of any metazoan group is a history of the entire ontogenetic cycles instead of separate stages and genes only. Ontogeny in the most objective way links two key components of the biological systematics: historically-independent characters attribution and phylogeny itself. A general theory encompassing "static" traditional taxonomy and dynamic evolutionary process, based on the ontogenetic transformation of the organisms' shape is suggested here to term as ontogenetic systematics. As an important practical implication of the ontogenetic systematics, a new model of the bilaterian metazoans evolution is suggested. The new model considers asexual clonal reproduction as a central feature of the ancestral ontogenetic cycles of basal Bilateria. The new scenario resolves several notable contradictions, e.g. morphological, ontogenetic and molecular similarities of Pogonophora, Vestimentifera, Phoronida simultaneously to protostomian Spiralia (Lophotrochozoa) and Deuterostomia. The suggested model implies individuation (possibly multiple) of ancestral semicolonial sedentary group as a major factor of the basal Bilateria diversification. In the late Ediacaran and early Cambrian thus existed ancestral bilaterian group that shared characters of both Spiralia and Deuterostomia and possessed polyp-shape body and cephalic secretory shield (like in modern Pterobranchia and Vestimentifera), that later on reduced in various lines. This ancestral taxon in rank of supraphylum is suggested to term as Carmaphora (shield-bearers). Presence of the enigmatic sedentary fossil of the genus Cloudina with vestimentiferan-like tubes and evident clonal reproduction already in the late Ediacaran, and most recent found of an unquestionable pterobranch already in the early Cambrian support the new model of Bilateria evolution.
任何后生动物类群的进化史都是整个个体发育周期的历史,而不仅仅是各个阶段和基因的历史。个体发育以最客观的方式将生物系统学的两个关键组成部分联系起来:历史上独立的特征归属和系统发育本身。这里提出一种基于生物体形态个体发育转变的、涵盖“静态”传统分类学和动态进化过程的一般理论,称为个体发育系统学。作为个体发育系统学的一个重要实际意义,提出了一种两侧对称后生动物进化的新模型。新模型将无性克隆繁殖视为基础两侧对称动物祖先个体发育周期的核心特征。新的设想解决了几个显著的矛盾,例如须腕动物门、铠甲动物门、帚虫动物门与原口动物螺旋类(触手冠动物)和后口动物同时存在的形态学、个体发育和分子相似性。所提出的模型意味着祖先半群体固着类群的个体化(可能是多次个体化)是基础两侧对称动物多样化的主要因素。因此,在埃迪卡拉纪晚期和寒武纪早期存在一个祖先两侧对称动物类群,它兼具螺旋类和后口动物的特征,具有息肉状身体和头部分泌盾(类似于现代的鳃曳虫类和铠甲动物),后来在不同的谱系中逐渐退化。这个处于超门等级的祖先分类单元被建议称为“带甲动物”(盾携带者)。在埃迪卡拉纪晚期就已出现具有类似铠甲动物管且明显有克隆繁殖的神秘固着化石Cloudina属,以及最近在寒武纪早期发现的无疑是鳃曳虫类的化石,都支持了两侧对称动物进化的新模型。