陕南川北地区下寒武统下部微体化石研究
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摘要
早期骨骼化石（小壳化石）是微体化石的一个重要内容。在寒武纪之初，早期骨骼化石（小壳动物群）大量出现，拉开了寒武纪生命大爆发的序幕，这些早期骨骼化石不仅前承伊迪卡拉生物群，后启澄江动物群，而且是整个生命演化史上后生动物第一次广泛出现骨骼化，在生物进化史上是一次大的飞跃，它代表了生物矿化作用质与量的突变，因此，对早期骨骼化石（小壳化石）进行深入研究，对了解寒武纪生命大爆发和生命演化具有重要意义。我国的陕南、川北地区处于扬子地台北缘，下寒武统分布广泛，富含早期骨骼化石、微古植物化石及动物胚胎化石，它们是“寒武纪生物大爆发”前夕生命多细胞化和生物多样性的见证。
此次研究通过大量的野外实地踏勘、剖面测制、样品采集和室内微体化石处理、扫描电镜照相、岩石薄片的磨制及同步辐射X射线相位衬度显微层位成像技术等一系列方法，处理出大量的早期骨骼化石、胚胎化石标本及薄片标本，并从古生物的角度对各类早期骨骼化石进行了讨论，取得了以下进展：
1、宽川铺生物群中卵裂期胚胎化石的发现及意义
本文通过醋酸浸泡获得了多枚精美的卵裂期、囊胚期及原肠胚期的三维化石标本，同时应用岩石切片技术获得了珍贵的卵裂期胚胎化石的切片标本，他们共同见证了从单个卵细胞（Olivooides）经卵裂到囊胚、原肠胚的完整胚胎发育序列。新化石材料的发现，不仅填补了宽川铺生物群中卵裂期胚胎化石的空白，完善了从单个卵细胞（Olivooides）经卵裂到囊胚、原肠胚，再到孵化的锥形幼体（Punctatus）的完整胚胎发育序列，而且通过与贵州瓮安生物群中卵裂期细胞胚胎化石及一些现生的低等动物卵裂期胚胎的对比，对揭示Olivooides胚胎的卵裂过程和特点，探讨其亲缘关系，以及后生动物的早期演化提供了新的化石证据。
2、宽川铺生物群中可能的原肠作用模式探讨
铺生物群中含有的大量原肠胚期的胚胎化石为我们研究早期生物的原肠作用模式提供了可能。本次研究主要通过对大量的原肠胚期的胚胎化石进行细致的观察，利用岩石切片及Micro-CT等技术对化石内部结构进行分析，发现了宽川铺生物群中的两种原肠作用模式，即内陷原肠作用模式和分层原肠作用模式。另外发现了两种腔囊胚形态，而囊胚的形态与原肠作用息息相关。因此目前的化石材料支持了Wolpert等人的观点，认为不同的原肠作用模式取决于胚胎本身卵黄分布的均匀程度，而对动物的发育和演化上可能并不具有重要的意义。
3、Punctatus属新种的发现及完整发育序列的完善
获得了数枚保存完整的Punctatus标本，并建立了一新种Punctatus costalis sp.nov，丰富了Punctatus属的类容。另外，在南江地区发现了保存完好的Punctatus emeiensis标本，进一步扩大了Punctatus的地理分布范围。同时通过获得的卵裂期、囊胚期、原肠胚期直至孵化幼体、成年个体的化石标本，完善了从卵细胞Olivooides到成年个体Punctatus整个发育序列，这一研究将为早寒武世早期后生动物起源、分类、辐射演化及个体发育提供不可多得的实证材料。
4、六方锥石属新种的发现
此次研究获得了大量的原始锥石类化石，其数量、种类丰富，不仅有大量的骨状锥石，亦含有大量的六方锥石，并建立描述了一个新种——宁强六方锥石（Hexangulaconularianingqiangensis sp.nov.）。新种的发现不仅丰富了六方锥石的内容，而且对地层学的研究有着重要的意义。
5、分叉状生物化石的发现及其生物属性的探讨
发现了大量的分叉状管状化石，其中包括二歧分叉和三分叉管状化石的类型，同时发现这些管状化石具有断裂生殖和出芽生殖的特征。本文通过对这些分叉状管状化石的观察和描述，对其生物属性进行了讨论。
Earliest skeletal fossils (Small shelly fossils) is an important part in microfossils. Thesmall shelly fossils thrived in the early Cambrian, marking the beginning of “Cambrianexplosion”. The small shelly fauna, lived the period between Ediacaran fauna and Chengjiangfauna, probably originated from Ediacaran and giving rise to Chengjiang fauna. Additionally,these small shelly faunas are representative of the first skeletal organisms which indicates asudden change on biomineralization. As a result, the study on earliest skeletal fossils is of animportant significance on investigating Cambrian explosion and life’s evolution. The LowerCambrian system in Southern Shanxi and Northern Sichuan, which contain rich earliestskeletal fossils, micropaleobotanical fossils and embryo fossils which witness multicellularityand biodiversity on the eve of Cambrian life’s explosion are widely distributed at the northedge of Yangtzi platform.
In this study, we sampled abundant earliest skeleton fossils and embryo fossils andanalyzed these specimens by scan electronic microscope (SEM), slicing and X-ray phasecontrast micro-imaging and so on. After that, a number of earliest skeleton fossils and embryofossils were obtained. At last, we make the following progress:
1. Embryo fossils of cleavage stage in Kuanchuanpu fauna
Several exceptionally-preserved three-dimensional embryo fossils, respectivelycorresponding to cleavage, blastula and gastrula stage, were obtained through acetic acidtreatments and slicing, and a complete developmental sequence being from olivooides toblastula and gastrula then toward the juvenile stage of Punctatus, is proposed. The new fossilsfill the gap on embryo fossil of cleavage stage in Kuanchuanpu fauna and make a perfect onintegrated embryo development. In comparison with fossil embryos of Wen’an Biota inGuizhou Province and their modern equivalents, the cleavage course of embryo is revealed. Inthis paper, the developmental data facilitate a thorough integration of palaeontology intoevolutionary developmental biology on understanding the biological evolution.
2. Probable gastrulation patterns in Kuanchuanpu fauna
The rich embryo fossils in gastrulation stage in Kuanchuanpu fauna provide probabilityfor researching gastrulation patterns of early life. Based on abundant embryo fossils ingastrulation stage and their microstructure observed from rock sections and Micro-CTtechnology, we found two types of gastrulation in Kuanchuanpu fauna, namely,invagination of a coeloblastula to form a coelogastrula and delamination of a coeloblastula toform a coelogastrula, suggesting that characters of blastula are influenced by gastrulation. It also supports the viewpoint of Wolpert et al.(1992) that the gastrulation pattern depend onuniformity degree of yolk distribution in embryo, but has no significant relationship withbiological development and evolution.
3. The discovery of a new species in Punctatus and foundation of integrated developmentalsequence
Here, we obtained several intact specimens of Punctatus. They were identified as a newspecies, named Punctatus costalis sp. nov. In addition, the specimens assigned to Punctatusemeiensis were found in Nanjiang region, which expands geographical distribution ofPunctatus. The fossils being from cleavage, blastula, gastrula stage to larva, were obtained,which improves developmental sequence from Olivooides to Punctatus. This investigationprovides significant proof for genesis, systematics, evolution and ontogeny of metazoan.
4. The discovery of a new species in Hexangulaconularia
Abundant protoconulariids fossils in both number and category were obtained. Theyinclude Carinachites and Hexangulaconularia. It is worth indicating that we established anew species assigned to Hexangulaconularia ningqiangensis sp.nov., which superinduce aspecies into Hexangulaconularia. Contemporarily, it is significant to stratigraphy.
5. The discovery of Bifurcate fossils and their biological property
We found plenty of bifurcate including dichotomous and trichotomous column-shapedtube fossils characterized by fragmentation and budding. Finally, their biological property wasdiscussed based on detailed observation and description of the fossils.
此次研究通过大量的野外实地踏勘、剖面测制、样品采集和室内微体化石处理、扫描电镜照相、岩石薄片的磨制及同步辐射X射线相位衬度显微层位成像技术等一系列方法，处理出大量的早期骨骼化石、胚胎化石标本及薄片标本，并从古生物的角度对各类早期骨骼化石进行了讨论，取得了以下进展：
1、宽川铺生物群中卵裂期胚胎化石的发现及意义
本文通过醋酸浸泡获得了多枚精美的卵裂期、囊胚期及原肠胚期的三维化石标本，同时应用岩石切片技术获得了珍贵的卵裂期胚胎化石的切片标本，他们共同见证了从单个卵细胞（Olivooides）经卵裂到囊胚、原肠胚的完整胚胎发育序列。新化石材料的发现，不仅填补了宽川铺生物群中卵裂期胚胎化石的空白，完善了从单个卵细胞（Olivooides）经卵裂到囊胚、原肠胚，再到孵化的锥形幼体（Punctatus）的完整胚胎发育序列，而且通过与贵州瓮安生物群中卵裂期细胞胚胎化石及一些现生的低等动物卵裂期胚胎的对比，对揭示Olivooides胚胎的卵裂过程和特点，探讨其亲缘关系，以及后生动物的早期演化提供了新的化石证据。
2、宽川铺生物群中可能的原肠作用模式探讨
铺生物群中含有的大量原肠胚期的胚胎化石为我们研究早期生物的原肠作用模式提供了可能。本次研究主要通过对大量的原肠胚期的胚胎化石进行细致的观察，利用岩石切片及Micro-CT等技术对化石内部结构进行分析，发现了宽川铺生物群中的两种原肠作用模式，即内陷原肠作用模式和分层原肠作用模式。另外发现了两种腔囊胚形态，而囊胚的形态与原肠作用息息相关。因此目前的化石材料支持了Wolpert等人的观点，认为不同的原肠作用模式取决于胚胎本身卵黄分布的均匀程度，而对动物的发育和演化上可能并不具有重要的意义。
3、Punctatus属新种的发现及完整发育序列的完善
获得了数枚保存完整的Punctatus标本，并建立了一新种Punctatus costalis sp.nov，丰富了Punctatus属的类容。另外，在南江地区发现了保存完好的Punctatus emeiensis标本，进一步扩大了Punctatus的地理分布范围。同时通过获得的卵裂期、囊胚期、原肠胚期直至孵化幼体、成年个体的化石标本，完善了从卵细胞Olivooides到成年个体Punctatus整个发育序列，这一研究将为早寒武世早期后生动物起源、分类、辐射演化及个体发育提供不可多得的实证材料。
4、六方锥石属新种的发现
此次研究获得了大量的原始锥石类化石，其数量、种类丰富，不仅有大量的骨状锥石，亦含有大量的六方锥石，并建立描述了一个新种——宁强六方锥石（Hexangulaconularianingqiangensis sp.nov.）。新种的发现不仅丰富了六方锥石的内容，而且对地层学的研究有着重要的意义。
5、分叉状生物化石的发现及其生物属性的探讨
发现了大量的分叉状管状化石，其中包括二歧分叉和三分叉管状化石的类型，同时发现这些管状化石具有断裂生殖和出芽生殖的特征。本文通过对这些分叉状管状化石的观察和描述，对其生物属性进行了讨论。
Earliest skeletal fossils (Small shelly fossils) is an important part in microfossils. Thesmall shelly fossils thrived in the early Cambrian, marking the beginning of “Cambrianexplosion”. The small shelly fauna, lived the period between Ediacaran fauna and Chengjiangfauna, probably originated from Ediacaran and giving rise to Chengjiang fauna. Additionally,these small shelly faunas are representative of the first skeletal organisms which indicates asudden change on biomineralization. As a result, the study on earliest skeletal fossils is of animportant significance on investigating Cambrian explosion and life’s evolution. The LowerCambrian system in Southern Shanxi and Northern Sichuan, which contain rich earliestskeletal fossils, micropaleobotanical fossils and embryo fossils which witness multicellularityand biodiversity on the eve of Cambrian life’s explosion are widely distributed at the northedge of Yangtzi platform.
In this study, we sampled abundant earliest skeleton fossils and embryo fossils andanalyzed these specimens by scan electronic microscope (SEM), slicing and X-ray phasecontrast micro-imaging and so on. After that, a number of earliest skeleton fossils and embryofossils were obtained. At last, we make the following progress:
1. Embryo fossils of cleavage stage in Kuanchuanpu fauna
Several exceptionally-preserved three-dimensional embryo fossils, respectivelycorresponding to cleavage, blastula and gastrula stage, were obtained through acetic acidtreatments and slicing, and a complete developmental sequence being from olivooides toblastula and gastrula then toward the juvenile stage of Punctatus, is proposed. The new fossilsfill the gap on embryo fossil of cleavage stage in Kuanchuanpu fauna and make a perfect onintegrated embryo development. In comparison with fossil embryos of Wen’an Biota inGuizhou Province and their modern equivalents, the cleavage course of embryo is revealed. Inthis paper, the developmental data facilitate a thorough integration of palaeontology intoevolutionary developmental biology on understanding the biological evolution.
2. Probable gastrulation patterns in Kuanchuanpu fauna
The rich embryo fossils in gastrulation stage in Kuanchuanpu fauna provide probabilityfor researching gastrulation patterns of early life. Based on abundant embryo fossils ingastrulation stage and their microstructure observed from rock sections and Micro-CTtechnology, we found two types of gastrulation in Kuanchuanpu fauna, namely,invagination of a coeloblastula to form a coelogastrula and delamination of a coeloblastula toform a coelogastrula, suggesting that characters of blastula are influenced by gastrulation. It also supports the viewpoint of Wolpert et al.(1992) that the gastrulation pattern depend onuniformity degree of yolk distribution in embryo, but has no significant relationship withbiological development and evolution.
3. The discovery of a new species in Punctatus and foundation of integrated developmentalsequence
Here, we obtained several intact specimens of Punctatus. They were identified as a newspecies, named Punctatus costalis sp. nov. In addition, the specimens assigned to Punctatusemeiensis were found in Nanjiang region, which expands geographical distribution ofPunctatus. The fossils being from cleavage, blastula, gastrula stage to larva, were obtained,which improves developmental sequence from Olivooides to Punctatus. This investigationprovides significant proof for genesis, systematics, evolution and ontogeny of metazoan.
4. The discovery of a new species in Hexangulaconularia
Abundant protoconulariids fossils in both number and category were obtained. Theyinclude Carinachites and Hexangulaconularia. It is worth indicating that we established anew species assigned to Hexangulaconularia ningqiangensis sp.nov., which superinduce aspecies into Hexangulaconularia. Contemporarily, it is significant to stratigraphy.
5. The discovery of Bifurcate fossils and their biological property
We found plenty of bifurcate including dichotomous and trichotomous column-shapedtube fossils characterized by fragmentation and budding. Finally, their biological property wasdiscussed based on detailed observation and description of the fossils.
引文
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