Mallatt Jon, Chen Jun-yuan
School of Biological Sciences, Washington State University, Pullman, Washington 99164-4236, USA.
J Morphol. 2003 Oct;258(1):1-31. doi: 10.1002/jmor.10081.
This study investigates whether the recently described Cambrian fossil Haikouella (and the very similar Yunnanozoon) throws light on the longstanding problem of the origin of craniates. In the first rigorous cladistic analysis of the relations of this animal, we took 40 anatomical characters from Haikouella and other taxa (hemichordates, tunicates, cephalochordates, conodont craniates and other craniates, plus protostomes as the outgroup) and subjected these characters to parsimony analysis. The characters included several previously unrecognized traits of Haikouella, such as upper lips resembling those of larval lampreys, the thick nature of the branchial bars, a mandibular branchial artery but no mandibular branchial bar, muscle fibers defining the myomeres, a dark fibrous sheath that defines the notochord, conclusive evidence for paired eyes, and a large hindbrain and diencephalon in the same positions as in the craniate brain. The cladistic analysis produced this tree: (protostomes, hemichordates (tunicates, (cephalochordates, (Haikouella, (conodonts + other craniates))))), with the "Haikouella + craniate" clade supported by bootstrap values that ranged from 81-96%, depending on how the analysis was structured. Thus, Haikouella is concluded to be the sister group of the craniates. Alternate hypotheses that unite Haikouella with hemichordates or cephalochordates, or consider it a basal deuterostome, received little or no support. Although it is the sister group of craniates, Haikouella is skull-less and lacks an ear, but it does have neural-crest derivatives in its branchial bars. Its craniate characters occur mostly in the head and pharynx; its widely spaced, robust branchial bars indicate it ventilated with branchiomeric muscles, not cilia. Despite its craniate mode of ventilation, Haikouella was not a predator but a suspension feeder, as shown by its cephalochordate-like endostyle, and tentacles forming a screen across the mouth. Haikouella was compared to pre-craniates predicted by recent models of craniate evolution and was found to fit these predictions closely. Specifically, it fits Northcutt and Gans' prediction that the change from ciliary to muscular ventilation preceded the change from suspension feeding to predatory feeding; it fits Butler's claim that vision was the first craniate sense to start elaborating; it is consistent with the ideas of Donoghue and others about the ancestor of conodont craniates; and, most strikingly, it resembles Mallatt's prediction of the external appearance of the ancestral craniate head. By contrast, Haikouella does not fit the widespread belief that ancestral craniates resembled hagfishes, because it has no special hagfish characters. Overall, Haikouella agrees so closely with recent predictions about pre-craniates that we conclude that the difficult problem of craniate origins is nearly solved.
本研究调查了最近描述的寒武纪化石海口虫(以及非常相似的云南虫)是否能为长期存在的有头类动物起源问题提供线索。在对该动物关系的首次严格分支系统分析中,我们从海口虫和其他类群(半索动物、被囊动物、头索动物、牙形有头类动物和其他有头类动物,外加原口动物作为外类群)选取了40个解剖学特征,并对这些特征进行了简约分析。这些特征包括一些此前未被识别的海口虫特征,如类似七鳃鳗幼体的上唇、鳃弓的厚实特征、一条下颌鳃动脉但没有下颌鳃弓、界定肌节的肌纤维、界定脊索的深色纤维鞘、成对眼睛的确凿证据,以及与有头类动物大脑相同位置的大后脑和间脑。分支系统分析得出了这样的树形图:(原口动物,半索动物(被囊动物,(头索动物,(海口虫,(牙形动物 + 其他有头类动物))))),“海口虫 + 有头类动物”分支得到的自展值在81% - 96%之间,具体数值取决于分析的结构方式。因此,得出结论海口虫是有头类动物的姐妹群。将海口虫与半索动物或头索动物归为一类,或认为它是基础后口动物的其他假说,几乎没有得到支持或完全没有得到支持。尽管海口虫是有头类动物的姐妹群,但它没有头骨且没有耳朵,不过它的鳃弓中有神经嵴衍生物。它的有头类特征大多出现在头部和咽部;其间距宽且粗壮的鳃弓表明它是通过鳃节肌进行呼吸,而非通过纤毛。尽管海口虫具有有头类动物的呼吸方式,但它不是捕食者,而是悬浮取食者,这一点由其类似头索动物的内柱以及在口部形成滤网的触手所表明。将海口虫与有头类动物进化的近期模型所预测的前有头类动物进行了比较,发现它与这些预测非常吻合。具体而言,它符合诺思卡特和甘斯的预测,即从纤毛呼吸到肌肉呼吸的转变先于从悬浮取食到捕食取食的转变;它符合巴特勒的观点,即视觉是最早开始进化的有头类动物感官;它与多诺霍等人关于牙形有头类动物祖先的观点一致;最引人注目的是,它类似于马拉特对祖先有头类动物头部外观所做的预测。相比之下,海口虫不符合普遍认为的祖先有头类动物类似盲鳗的观点,因为它没有盲鳗的特殊特征。总体而言,海口虫与近期关于前有头类动物的预测非常吻合,因此我们得出结论,有头类动物起源这一难题几乎已得到解决。