寒武纪腕足动物起源:假说、问题与展望

腕足动物与软体动物和环节动物等十多个动物门类一起构成后生动物(metazoa)中分异度最大的谱系分支——冠轮动物(Lophotrochozoa)。在冠轮动物分支概念提出20周年之际,分子系统学和基因组系统学的大量研究基本证实了腕足动物、帚虫动物和外肛动物组成的触手冠动物(Lophophorata)为单系起源(monophyly),分类上属于原口动物亚界(Protostomia),从而彻底否定了上百年传统动物学教科书上腕足动物属于后口动物亚界(Deuterostomia)或原口动物亚界与后口动物亚界之间的过渡类群的传统观点。腕足动物作为寒武纪演化动物群的重要组成部分和寒武纪大爆发期间的主要动物门类代表,因其形态发育和壳体矿化等多项特点以及海量的化石记录,历经数十余年单系、多系和并系起源的争论之后,其单系起源得到了分子生物学、基因组系统学和形态学(包括化石和现生类群)研究越来越多的强力支持。长期以来,寒武纪腕足动物门曾存在帚虫状祖先起源假说、蛤氏虫(Halkieria)起源假说和托莫特壳起源假说。目前,蛤氏虫起源假说遭到了不同国际学者的质疑和否定。澄江化石库(寒武系第三阶)最新发现的化石新种——精美玉玕囊形贝(Yuganotheca elegans),因具有砂质胶结的椎管状身体,具有封闭的腕足动物状的纤毛环取食器官,具有腕足动物状的边缘刚毛、脉管循环系统、U形的消化系统和蠕虫状的肉茎附着结构等,被誉为腕足动物的"始祖鸟化石",成为联系帚虫动物门、腕足动物门和椎管状托莫特壳的化石类群。然而,因该化石缺乏成对的矿化(磷酸钙或碳酸钙)壳体,对现今广泛使用的腕足动物高级分类方案提出了新的思考和挑战。     

云南永善肖滩剖面早寒武世Lapworthella (托莫特壳类)骨片化石

描述了采自云南永善肖滩剖面下寒武统石岩头组的托莫特壳类(tommotiids)骨片化石Lnpworthella rete Yue,1987,其骨片锥状或塔状,横向环脊明显,环脊尖棱状;骨片外表面具多边形网状纹饰,但无齿、瘤或纵脊等装饰;骨片可分为两种类型,即弯曲型和近直立型,前者数量明显多于后者。简要地讨论了托莫特壳类化石的分类及生物亲缘关系,认为目前为止,其仍为亲缘关系不明的化石类别。尚难断定所有的托莫特壳类骨片化石皆属于同一生物门类,因为难以证明将它们归人同类生物所依据的骨片性状是共同衍征(synapomorphy)还是由于趋同演化(convergent evolution)的结果;在缺乏躯干化石记录的情况下,离散骨片的生物亲缘关系研究较为困难。     

再论织金钉类(Zhijinnids)的生物亲缘

寒武纪最早期层位中获得的离散骨片化石Halkieria和Microdictyon,已在较高层位的特异埋藏化石群中找到了它们的完整生物躯体,它为研究离散骨片化石的自然分类和生物亲缘提供了直接证据。本文通过寒武纪最早期钉状化石的形态、对称性、壳质成分、矿化特征的对比,提出了离散骨片化石织金钉(Zhijinites)和拟织金钉(Parazhijinites)与澄江动物群中的怪诞虫(Hallucigcnia)和爪网虫(Onychodictyon)等体表上的钉状物有近亲关系,它们可能同属于多足缓步类(Tardipolypoda)。     

华南兰多维列世几种磷灰质微体化石*

磷灰质硬体存在于现代和地质历史中,早古生代的一些磷灰质生物硬体可以作为化石保存下来。在志留纪常见的磷灰质微体化石或磷灰质化石碎片中,有牙形刺、无颌类的鳞片、锥石、无铰纲腕足类、环节动物多毛纲的虫管,节肢动物(鲎类、剑尾目、光甲目等),似软舌螺,一些分类地位不明的小壳化石和其他未知类别经过磷酸岩矿化作用而形成的磷灰质化石。早古生代的一些零星的、分散的磷灰质骨片,早为古生物学家所瞩目。这些磷灰质微体化石,由于只是残片,不能窥其全貌,所以在鉴定、研究时都有一定的困难,但它对探讨多种生物的起源和演化历史却是至关重要的。尽管多数磷灰质骨片产地零星、数量亦少。     

新疆寒武纪早期单壳类软体动物化石分类学与生物地层学

作者系统描述了新疆阿克苏-乌什地区下寒武统玉尔吐斯组中上部单壳类软体动物化石10属10种,其中包括3个新种、3个老种和4个未定种,它们是Shabaktiella multiformis sp.nov.,Ilsanella acuta sp.nov.,Xianfengella yuertusiensis sp.nov.,Parcaconus xinjiangensis,Protostenotheca xinjiangensis,Aldanella attlebo-rensis,Obtusoconus sp.,Yochelcionella sp.,Bemella sp.和Mellopegma sp..上述10个属种基本上代表了玉尔吐斯组单壳类软体动物的面貌特征.文中介绍了本研究区10个属和属级以上分类系统在国际学术界的意见分歧后,重申了以壳的对称性和有无肌痕确定纲级界线的最佳分类原则,提出了应用壳型划分目级分类单元和口缘形状、壳顶与口缘的相对位置划分科级分类单元的可行性和有效性,但是不能接受把所有单板状软体动物化石通常被归入单板纲的做法,因为寒武纪早期一些单板状软体动物化石不一定符合单板纲的真实含义.文中还回顾了近二十年来建立的始单板纲(Eomonoplacophora)(Missarzhevsky,1989),太阳女神螺纲(Helcionelloida),背壳肌纲(Tergomya)(Peel,1991a,b)和似腹足纲(Paragastropoda)(Linsley & Kier,1984)等纲级的新概念.还有一些早期软体动物专家将许多单板状(包括帽状和螺旋状)化石归入单板纲(Runnegar et al.,1976,1985),而另有一些中青年软体动物专家继承传统理念,将它们归入了腹足纲(Parkhaev in.Alexander et al.,2001).钱逸和本格森将早期单壳类软体动物化石分成了五大形态类别,未确定科级以上分类单元名称(Qian & Bengtson,1989).总之,目前纲级分类单元界线不清,目级以下分类单元十分混乱,在没有全面清理早期单壳类软体动物百余个属和属级以上分类单元之前,是难以统一本文研究的早期单壳类软体动物10个属和属级以上分类系统.我们认为在确定属级以上分类系统及其与之密切相关的演化谱系关系时,不是利用理论前提,而是在不间断的剖面上对单壳类软体动物化石进行逐层采集并对其构造要素的详细研究,才能使得出的结论和提出的新分类系统发生错误的几率最小.文中还归纳和总结了本研究区玉尔吐斯组中上部单壳类软体动物群面貌的六大特点,其中二点最重要:1)该动物群与梅树村期微软体动物群的面貌有明显的差别,它缺失了梅树村期微软体动物群中最原始、最特征、最丰富的类群,如笠帽状的马哈螺类化石Maikhanella,Purella等和螺旋状的始旋螺类化石Archaeospira等.但是该动物群面貌却与邻区的哈萨克斯坦Atdabanian期的微软体动物和我国峡区西蒿坪动物群面貌接近,出现了许多相同或相似的属种,如Shabaktiella,Mellopegma,Aldanella,Yochelcionella等.2)该动物群共生有全球分布的相当于筇竹寺期的标准化石,如原牙形类Gapparodus,Amphigeisina,Hagionella等,有口唇并有背腹分异和复杂口盖的软舌螺类Microconus,Pachytheca等,具有肉茎孔和铰合面的有铰腕足类和像Obolus那样的原始无铰腕足类,有各种形态类型的金臂虫Dabashanella,Liangshanella等,还有可疑的三叶虫颊刺等.上述信息足以说明玉尔吐斯组中上部地层时代应属筇竹寺期而非梅树村期,从而解决了长期以来有关玉尔吐斯组时代归属的争议. 玉尔吐斯组单壳类软体动物群的研究,再次证实了寒武纪生命大爆发期间,早期单壳类软体动物与小壳动物群一起经历了三次大发展和三次大绝灭事件.这在早期单壳类软体动物化石分类学、生物地层学和早期演化史上均有较高的科学价值.     

陕西省陆生缓步动物系统分类组成研究

缓步动物是一类微小的水生无脊椎多细胞动物,在系统分类上独列一门,与节肢动物有着较近的亲缘关系。目前,缓步动物门分为3纲、5目、13科、110多个属,超过1 250种。本文简要介绍了陕西省缓步动物门的系统分类研究进展,初步分析了陕西省缓步动物的分类组成。在陕西省已描述发表的缓步动物种类107种,隶属于2纲、3目、4科、17属。此外,展望了陕西省缓步动物区系分类的研究方法、研究方向和研究意义。     

论Yangtzechiton Yu与Paracarinachites Qian et Jiang的区别

从窄长的形状和背侧具棘刺来看,Yangtzechiton与Paracarinchites似乎有些类似,但不同点有:(1)Yangtzechiton的壳体是由头板、中间板和尾板组成,而Paracarinachites是一个弯曲的窄长骨片;(2)前者的壳板是由盖层和连接层组成,后者的整个骨片是为多层平行的薄片层组成;(3)Yangtzechiton的头板长宽近等,背侧后缘有宽深的缺四,中间板矩形,有一椭圆形孔,可能是空心棘断损后遗留下来的,后一亮板为前一壳板覆盖约1/2,尾板的后亮顶锥形;Paracarinachites前端弯曲部分有6—8个瘤粒突起,骨片后部具10—14根左右交错排列的、末端向后、向左(右)倾斜的棘刺;和(4)前者头板腹侧有一肾形凹沟和“W”形弯曲的条纹,中间板腹侧徽凹见有叠覆缝;后者腹侧分为前端弯曲部分和后部。前端弯曲部有一近圆形的盘状体,近中部有一个孔,后部空凹。因此,它们是不同门类的代表,Yangtzechiton属于软体动物门多板纲,Paracarinachites是分类位置未定的骨片。     

华南志留纪特列奇期喙壳类新材料*

喙壳纲(Rostroconchia)是Pojeta,Runnegar,Morris和Newell于1972年正式建立的软体动物门的一个纲。根据已有的材料,它们只生存于古生代的海洋中,从早寒武世出现,至晚二叠世绝灭,至今尚未有关于三叠纪喙壳类的确切报道。对现今被归于这个纲的化石的描记,至少已经有150年以上的历史。但是由于对这类化石的特征认识不足,长期以来其分类位置没有被确定。许多研究者往往根据其一般形态,将个体小、时代较老的原始类型归于节肢动物,而将其较进化的类型作为双壳类化石进行研究,或者作为分类位置未定但具有特殊形态特征的软体动物来处理。     

陆生软体动物的分类系统

自Müller(1774)正式建立属,经过186年,Zilch(1959-1960)共归纳陆生软体动物有1091个属,其中,约有58.4%的属建立于1870～1929年间,以后逐渐减少。现估计全世界陆生软体动物有35000余种,主要包括腹足纲的前鳃亚纲和肺螺亚纲两大亚纲的种类,以肺螺亚纲柄眼目的种类最多,已知有30000余种。近年来,对于陆生软体动物的系统分类,有学者陆续发表了一些文章。目前有关腹足纲的分类和系统发育的研究,主要集中于石磺科Onchidiidae、复套蛞蝓科Veronicellidae、拉氏蛞蝓科Rathouisiidae三科,研究的重点在于如何确定它们的分类地位,它们是属于后鳃亚纲还是肺螺亚纲。关于琥珀螺科Succineidae的分类问题也有诸多研究。应用分子生物学和细胞生物学的方法对某些科属进行系统分类也是当今的热点。陆生软体动物的分类特征有:1.贝壳的形态与结构;2.颚片与齿舌;3.触角的形态及其位置、构造;4.生殖系统;5.排泄系统;6.神经系统;7.其他器官;8.细胞分类学和分子分类学。本文列出了现今常用的陆生贝类分类系统。     

陕南寒武纪西乡化石库的马哈螺和棱管壳研究

对陕南西乡化石库发现的马哈螺类化石和棱管壳类化石做了统计和描述,补充了马哈螺类新的属征,建立新种西乡拟鳞锥Ramentoides xixiangensis sp. nov.。马哈螺和棱管壳表面装饰存在差异,并非同一物种上掉落的骨片,本文展示的马哈螺新的特征与"骨针壳"假说存在矛盾,马哈螺在形态上表现出软体动物单板类最原始的特征,而棱管壳则与halkieriids存在密切的亲缘关系。本文报道了马哈螺类化石的新形态,丰富了对马哈螺类和棱管壳类的认识,对确立马哈螺和棱管壳的亲缘关系和分类位置提供了新的材料。     

Tommotiids from the early Cambrian (Series 2, Stage 3) of Morocco and the evolution of the tannuolinid scleritome and setigerous shell structures in stem group brachiopods

An assemblage of tannuolinid sclerites is described from the Amouslek Formation (Souss Basin) of the Anti‐Atlas Mountains in Morocco. The assemblage contains two species, Tannuolina maroccana n. sp., which is represented by a small number of mitral and sellate sclerites, and Micrina sp., represented by a single mitral sclerite. Tannuolina maroccana differs from other species of the genus in the presence of both bilaterally symmetrical and strongly asymmetrical sellate sclerites. This observation suggests that the scleritome of Tannuolina was more complex than previously thought and that this tommotiid may have held a more basal position in the brachiopod stem group than previously assumed. The shell structure of both T. maroccana and Micrina sp. is well preserved and exhibits two fundamentally different sets of tubular structures, only one of which was likely to contain shell‐penetrating setae. Based on these observations, the structure of the tannuolinid shell is discussed and its implications for the evolution of tubular microstructures in stem and crown group brachiopods are analysed.     

Oikozetetes from the early Cambrian of South Australia: implications for halkieriid affinities and functional morphology

Shells of Oikozetetes and isolated halkieriid sclerites from a section of the lower Cambrian Mernmerna Formation in the Flinders Ranges, South Australia, are tentatively considered as being derived from the same scleritome. Details of shell morphology and the possible combination of biomineralized shell and sclerites suggest that Oikozetetes , if interpreted correctly, is closely related to Halkieria . A new interpretation of Oikozetetes shell morphology, in addition to the first report of paired muscle scars on the interior surface, sheds new light on the possible functional morphology of halkieriid shells and the means of attaching the shell to the body. The occurrence of Oikozetetes in South Australia extends its biostratigraphic range to the lower Cambrian and biogeographic range to East Gondwana.     

A Stem Group Brachiopod From The Lower Cambrian: Support For A Micrina (Halkieriid) Ancestry

The shell structure of the Lower Cambrian Mickwitzia , a bilaterally symmetrical bivalve hitherto doubtfully assigned to the Brachiopoda, confirms that the genus shares characters with linguliform brachiopods. The columnar lamination of its organophosphatic shell is homologous with that characterizing acrotretides. The shell, however, is also pervaded by striated apatitic tubes indistinguishable from those permeating the sclerites of the problematic organophosphatic, laminar–shelled Micrina which is close to Halkieria . No crown group brachiopods have such tubes that are presumed to have contained setae. The presence of both these features in the Mickwitzia shell suggests that the stock is a stem group brachiopod with a halkieriid ancestry.     

Paterimitra pyramidalis from South Australia: scleritome,shell structure and evolution of a lower Cambrian stem group brachiopod

The tommotiid Paterimitra pyramidalis Laurie, 1986, is redescribed based on well‐preserved material from the lower Cambrian Wilkawillina, Wirrapowie and Ajax limestones of the Flinders Ranges, South Australia. The material shows that the scleritome of Paterimitra pyramidalis includes three sclerite morphotypes (S1, S2 and L). Detailed shell microstructure studies show striking similarities with both the paterinid brachiopod Askepasma toddense and the tommotiid Eccentrotheca helenia, which strengthens the suggested evolutionary link between tommotiids and brachiopods. Based on the partly articulated specimens and similarities in shell microstructure and sclerite morphology with Eccentrotheca, Paterimitra pyramidalis is reconstructed as a tube‐dwelling, epifaunal, sessile, filter‐feeder with an organic pedicle‐like attachment structure. The proposed reconstruction of the scleritome comprises a basal unit composed of one S1 and one S2 sclerite, as well as an unresolved number of L sclerites lining a coniform tubular structure.     

The Early Cambrian Halkieria is a mollusc

Halkieria evangelista was originally interpreted as a stem-group brachiopod. It was later proposed that the brachiopods evolved from a halkieriid ancestor that curved ventrally, lost the foot and radula, developed a lophophore as a new feeding organ, and modified the sclerites to chaetae. Molluscan affinities have been discussed by a number of authors. We have studied some of the original material of Halkieria and found that all the characters that can be observed with certainty are compatible with characters in living molluscs. The combination of characters observed in Halkieria does not fit with any of the living classes of molluscs, so we propose placing it in a new class, Diplacophora. Living polyplacophorans have hairs secreted by a number of epithelial cells and containing nerve cells; they are clearly not homologous with annelid chaetae. Thus, there is no support for theories that regard Halkieria as belonging to the stem group of either the annelids or brachiopods.     

The tommotiid Kelanella and associated fauna from the early Cambrian of southern Montagne Noire (France): implications for camenellan phylogeny

A carbonate bed of the Pardailhan Formation, early Cambrian, southern Montagne Noire (southern France), provided microfossils such as Hyolithellus sp., Torellella cf. mutila and Torellella sp. along with numerous disarticulated pieces of composite skeletons such as valves of the brachiopod Eoobolus priscus and of the bradoriid Monceretia erisylvia, and chancelloriid sclerites (Chancelloria sp.). The assemblage also furnished a rich set of sclerites from the tommotiid Kelanella altaica. Five morphological variations of the latter have been identified. The presence of concentric ribs formed by distal inflation of selected shell laminae in Kelanella supports its assignment to the camenellans. More particularly, the presence of transverse structures within the internal cavity (septa) of Kelanella suggests a close relationship with the Lapworthellidae. However, the latter differ from Kelanella by the continuous morphological variation along their scleritome which is also composed of simple conical elements with uniform ornamentation. Several forms of Kelanella are similar to mitral and sellate sclerites of Camenella, whereas some other forms are comparable to Kennardia. The new material suggests that Kelanella occupies a transitional position between Lapworthellidae and the grouping of Tommotiidae and Kennardiidae. Such a phylogenetic position also implies that the number of sclerite morphotypes tends to decrease within the camenellan scleritome during evolution.     

How the mollusc got its scales: convergent evolution of the molluscan scleritome

Radiation of dramatically disparate forms among the phylum Mollusca remains a key question in metazoan evolution, and requires careful evaluation of homology of hard parts throughout the deep fossil record. Enigmatic early Cambrian taxa such as Halkieria and Wiwaxia (in the clade Halwaxiida) have been proposed to represent stem‐group aculiferan molluscs (Caudofoveata + Solenogastres + Polyplacophora), as complex scleritomes were considered to be unique to aculiferans among extant molluscs. The ‘scaly‐foot gastropod’ (Neomphalina: Peltospiridae) from hydrothermal vents of the Indian Ocean, however, also carries dermal sclerites and thus challenges this inferred homology. Despite superficial similarities to various mollusc sclerites, the scaly‐foot gastropod sclerites are secreted in layers covering outpockets of epithelium and are largely proteinaceous, while chiton (Polyplacophora: Chitonida) sclerites are secreted to fill an invaginated cuticular chamber and are largely calcareous. Marked differences in the underlying epithelium of the scaly‐foot gastropod sclerites and operculum suggest that the sclerites do not originate from multiplication of the operculum. This convergence in different classes highlights the ability of molluscs to adapt mineralized dermal structures, as supported by the extensive early fossil record of molluscs with scleritomes. Sclerites of halwaxiids are morphologically variable, undermining the assumed affinity of specific taxa with chitons, or the larger putative clade Aculifera. Comparisons with independently derived similar structures in living molluscs are essential for determining homology among fossils and their position with respect to the enigmatic evolution of molluscan shell forms in deep time. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 949–954.     

Ontogeny,morphology and taxonomy of the soft‐bodied Cambrian ‘mollusc’ Wiwaxia

The soft‐bodied Cambrian organism Wiwaxia poses a taxonomic conundrum. Its imbricated dorsal scleritome suggests a relationship with the polychaete annelid worms, whereas its mouthparts and naked ventral surface invite comparison with the molluscan radula and foot. 476 new and existing specimens from the 505‐Myr‐old Burgess Shale cast fresh light on Wiwaxia's sclerites and scleritome. My observations illuminate the diversity within the genus and demonstrate that Wiwaxia did not undergo discrete moult stages; rather, its scleritome developed gradually, with piecewise addition and replacement of individually secreted sclerites. I recognize a digestive tract and creeping foot in Wiwaxia, solidifying its relationship with the contemporary Odontogriphus. Similarities between the scleritomes of Wiwaxia, halkieriids, Polyplacophora and Aplacophora hint that the taxa are related. A molluscan affinity is robustly established, and Wiwaxia provides a good fossil proxy for the ancestral aculiferan – and perhaps molluscan – body plan.     

Shell Structure And Inferred Growth, Functions And Affinities Of The Sclerites Of The Problematic Micrina

The stratiform laminae of Micrina sclerites originally consisted of rheomorphic successions of monolayers of micrometric–sized, apatitic tablets, presumably interleaved with chitin and glycosaminoglycans (GAGs). Paired laminae enclose slot–like chambers swelling into lobes distally that originally contained GAGs and deposits of spherulitic and prismatic apatite. The laminae are pervaded by apatitic tubes, apparently secreted by microvillous setoblasts and containing, at the surface, chitinous setae. Internal markings suggest that the triangular (sellate) sclerite supported a pair of muscles and the planospiral (mitral) sclerite, a medial muscle and gonadal sacs flanked by a pair of crescentic muscle bases. Both sclerites were secreted by a mantle with a circumferential fold. The sellate and mitral sclerites are homologized with the anterior and posterior shells of Halkieria and could have become the dorsal and ventral valves of the ancestral brachiopod by a sequence of transformations. These include: the folding of the halkieriid body axis; accelerated mixoperipheral growth of the anterior (dorsal) shell to enclose, with the posterior (ventral) shell, a mantle cavity lined with modified ciliated epithelium of the foot; reduction of sclerite–secreting epithelium to the locus of the brachiopod pedicle epithelium; and the anterior (dorsal) spread of gonadal lamellae.     

A comparative study of Lower Cambrian Halkieria and Middle Cambrian Wiwaxia

Two Cambrian lepidote metazoans known from different respective types of preservation have been compared in order to elucidate their biology and affinities. The widely distributed Lower Cambrian Halkieria is represented by isolated hollow sclerites, probably of originally calcareous composition. The Middle Cambrian Wiwaxia is known from the Burgess Shale as isolated sclerites (scales and spines) and as more or less complete individuals. Although Halkieria sclerites were mineralized and those of Wiwaxia were probably not, there are fundamental structural and morphological similarities between the two. Both bad an imbricating scaly and spiny armour consisting of hollow sclerites with a longitudinally fibrous structure. The sclerites did not grow, but were probably moulted during the course of ontogenetic growth. Halkieria and Wiwaxia are regarded as closely related. Both are referred to the Order Sachitida He 1980. The sclerite armour of Halkieria is reconstructed on the template provided by Wiwaxia. The interpretation of sachitid sclerites as protective armour is an alternative to the interpretation by Jell (1981, Alcheringa 5 )that sachitid sclerites were respiratory organs in an animal of probable annelid affinities. Sachitids are interpreted as sluggish, benthic deposit feeders that do not belong to any recognized phylum.