黔南晚石炭世珊瑚礁生态系统研究
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摘要
中国南方晚石炭世海域广大,普遍发育了一套台地相碳酸盐岩,黔南是其中的一个代表性地区。该区碳酸盐台地浅水环境非常适宜生物生存,特别是底栖生物大量繁盛,构成了不同类型的造礁生物群落,生物造岩作用十分强烈,生物滩、丘、礁广泛出现,是开展中国南方晚石炭世生物地层和生物礁研究的理想地区。该地区大部分礁体为台地边缘礁,最具代表性的是扁坪村大型组合式珊瑚礁。
该礁体规模巨大,结构独特,根据产礁岩系特征可划分为上下叠置的3个单元(U1、U2、U3)。其中U1为一点礁层,包括一个蓝细菌泥丘、一个Ivanovia cf. manchurica点礁和一个叶状藻点礁,通过上覆的生物碎屑层与礁主体部分划分开。U2和U3具有相同的岩系特征和类似的生物组成,它们都是由Fomitchevella骨架岩和泥粒岩组成,区别在于U3的生物丰度明显高于U2。
研究区内造礁生物种类丰富,生态类型多样,成礁方式各异。在扁坪村大型组合式珊瑚礁中主要有造架生物Fomitchevella、造架—障积生物叶状藻,造架—覆盖生物Ivanovia cf. manchurica以及粘结生物蓝细菌。其中Fomitchevella为礁体的主要建造者,它能够通过出芽生殖方式形成笙状或丛状群体,从而建造了不同类型的礁体。群体中间部位的个体较为粗大,边缘部位的则细小,个体的生存能力较强。Ivanovia cf. manchurica群体呈板状,生长方式多样,群体的再生能力很强,参与了U1中的点礁建造。叶状藻是研究区最为重要的造礁生物之一,发育了多种形态,建造了不同类型的礁体,其中包括U1中的叶状藻点礁。另一种群体珊瑚Antheria数量较少,分布较为局限,它是群落演化中的一个过渡角色。蓝细菌是具有粘结功能的自养生物,它是U1中灰泥丘的主要建造者。附礁生物以蜒、腕足类为主,海百合、Tubiphytes、腹足类较为常见,含苔藓虫和少量单体珊瑚。
该礁体内划分出了6个主要的造礁生物群落,它们具有类似的生物多样性,但群落的生物丰度、空间分层性、关键种的功能作用等各有差异。根据生物的生活习性、功能结构和取食位置,将它们归类为不同的生物群团。自蓝细菌群落向Fomitchevella群落发展,群落的空间分层性越来越明显,对环境的适应能力和对内部生物的调控能力也越来越强。蓝细菌、Tubiphytes、蜓、非蜒有孔虫、腕足类、腹足类生活于群落的下层,海百合、Ivanovia cf. manchurica、Antheria在中层空间取食,Fomitchevella则有着较高的捕食位置。群落的生物组分有所不同,但群落的维持机制基本一致。群落对营养盐的吸收与现代珊瑚礁群落类似,分别适用于一种或多种吸收模型。整个生态系统的能量最终来源于太阳光,藻类、微生物、蓝细菌、Tubiphytes和水体中的溶解有机物构成了初级生产力,能量通过生物间的捕食关系层层传递和损失,最终由分解者释放到水体中,一部分参与循环,绝大多数随水流散失。
借助于对现代珊瑚礁的研究成果,对扁坪村大型组合式珊瑚礁中的生态关系类型做了详细的分析和研究。腕足类生活于叶状藻的藻丛内部,不仅获取了更多的食物,还可以在藻丛的保护环境中躲避天敌的捕食,它们不存在竞争关系,能够和谐共处,也能单独生存,因此它们是一种特殊的共栖关系。Fomitchevella群体中相邻个体为争夺更多的生长空间、更广的食物来源和更大的采光面积而出现激烈的种间空间竞争,部分个体弯曲、畸形、倾斜,甚至出向侧向的生长现象,它们由此失去了更多的生存资源,最终被直立、粗壮、高大的个体所替代。“未名造礁生物”生长和扩张速度快,攀附缠绕能力强,它们包绕在Ivanovia cf. manchurica的板状群体上,双方在接触带上出现组织加厚,形成主动包绕型的接触关系,“未名造礁生物”剥夺了Ivanovia cf. manchurica的生存资源致其死亡。Fomitchevella的个体占据基底的能力较差,Ivanovia cf. manchurica群体在侧向扩张时尽量避免与其骨架直接接触,而Fomitchevella的群体通过侧芽的方式占据了Ivanovia cf. manchurica的上方空间,从而对其形成遮蔽,二者形成了避让遮蔽型的空间竞争,最终Fomitchevella成为优势者。叶状藻对生存于藻丛中的零星Fomitchevella个体形成数量压倒型的空间竞争,竞争结果因环境条件而异。Ivanovia cf. manchurica发育在叶状藻建造的基底上,而后又被叶状藻覆盖,出现了交互式的群落替代；Ivanovia cf. manchurica群落中出现了Fomitchevella的先驱个体,后来二者的角色发生了交换,最后Fomitchevella完全成为主导者,形成了渐变式的群落替代。Fomitchevella-1群落向Fomitchevella-2群落转变的过程中,生物组成和比例出现了一定的调整,使群落的结构更加合理,对环境的适应能力更强,因此实质上是一种群落的更新。
礁体的发育与群落的演化关系密切,整个发育过程可划分为四个阶段,每个阶段都包含了群落形成、发展、衰退等状态的改变。前两个阶段建造了点礁层,后两个阶段建造了礁主体。水体的营养程度和沉积速率控制着礁体的生长速度,在一定限度内它们成正相关关系,当这些环境条件出现突变时礁体往往会停止生长。
Carbonate rocks of plateau facies well developed in vast sea area of southern China in Late-Carboniferous, and the south of Guizhou Province was a representative region. Shallow water environment on the carbonate platform was quite favourable for organisms, especially the benthic organisms, and as a result different types of reef-building communities formed. Lithogeneous process of these organisms is intensive with organic banks, bioherms and reefs built widely, and thus it is an ideal region to study Late-Carboniferous biostratum and reef of South China. Most of these reefs developed on the platform margin, and the large-scale reef complex is the mostly representative one.
The reef complex has a large size and a special architectural structure, and 3 superimposed units, U1, U2 and U3 could be recognized according to features of reef rocks. U1 is a patch reef layer involving a mud mound built by cyanobacteria, an Ivanovia cf. manchurica patch reef and a phylloid algae patch reef, and it is separated from mainbody of the reef by an overlying bioclast layer. U2 has an identical lithology and a similar biological composition with U3, that is, both of them are composed of Fomitchevella framestone and packstone, the difference is that abundance of organisms in U3 is much higher than that in U2.
In the study region, the reef-building organisms are various in genera and ecotype, and their reef-building modes are diversefied. In the large-scale coral reef complex in Bianping Village, these reef-building organisms mainly include:1) Fomitchevella, frame builder; 2) phylloid algae, frame-builder and baffler; 3) Ivanovia cf. manchurica, frame-builder and covering organism; and 4) cyanobacteria, binding organism. Fomitchevella is the dominant constructor of the reef complex who grows into a phaceloid or fasciculate colony through budding and then builds reefs of diverse types. The skeletons of Fomitchevella individuals are large in the centre of the colony while small on the edge, and the individuals are viable. Ivanovia cf. manchurica develops a tabular colony and various growth forms. Their colonies get a strong power of regeneration and make a great contribution to the formation of U1. Phylloid algae is one of the most important builders in the study region which has diverse morphologies and builds different types of reefs, including the phylloid algae patch reef in U1. The other genera of colonial coral-Antheria has a low abundance and a limited distribution, and it plays a transitional role in community evolution. Cyanobacteria is a sort of autotroph with binding function, and it is the main builder of mud mound in U1. Fusulinids and brachiopods are the most common dwellers, followed by crinoids, Tubiphytes and bivalves, while bryozoans and solitary corals are few.
Six reef-building communities are recognised in the reef complex, which are similar in biological diversity while different in abundance of benthos, vertical spacial zonation, impacts of the key species and so on. The organisms are divided into various groups according to their life habits, fuctions and feeding positions. The vertical zonation becomes more and more notable along with evilution of these communities. Their adaptability to the evironment and control on organisms also get an obvious development. Cyanobacteria, Tubiphytes, fusulinids, foram, brachiopods and gastropods live in the lower part of the communities, while rinoids, Ivanovia cf. manchurica and Antheria in the middle part and Fomitchevella in the upper part. The biological components of these communities are different but the maintaining mechanisms are generally accordant.
Nutrient absorbing of the communities is similar to that of extant coral reef community, and one or more absorption models may be available. The energy of the whole ecosystem is derived from sunlight ultimately. Algaes, microbes, cyanobacteria, Tubiphytes and deliquescent organic materials in the water constitute the primary productivity. Energy is transferred and lost gradually through predation between organisms, and finally released into the water by decomposers. Part of the energy recycles in the ecosystem, and most is dispersed with the waterflow.
Ecological relationships in the coral reef complex in Bianping Village are analysed and studied based on research findings of extant reefs. Brachiopods lived in the mats of phylloid algae, which enabled them to get more food and also avoid being hunted by natural enemies. They could coexist without competition or live independently, and thus they were in a special relationship of commensal. In order to get more space for growth, source of food and larger lighting area, the individuals in Fomitchevella colony always competed intensely with their neigbours, resulting in some bent, misshapen, inclined or even lateral individuals. As a result, these individuals lost more living resources and would be finally replaced by those erect, strong and long ones. The "unamed reef-building organisms" grew and expanded at a high rate and were good at en winding. They wraped on the colony of Ivanovia cf. manchurica, and both of them thickened tissues on the contact zone, and as a result an aggressive-wraping relationship was formed. The "unamed reef-building organisms" deprived Ivanovia cf. manchurica of living resources and would finally lead to its death. When laterally expanding, Ivanovia cf. manchurica would as much as possible avoid direct contact with individual skeletons of Fomitchevella, who was not good at occupying the substrate but could take up the space above Ivanovia cf. manchurica throgh laterally budding and overshaded it. Then an off-overshading spatial competition was formed and Fomitchevella would be the dominant at last. An overwhelming spatial competition was formed by phylloid algae and a few Fomitchevella individuals lived in the algae mat, but the outcomes would be various with the influences of environmental conditions. Ivanovia cf. manchurica lived on substrate built by phylloid algae and the was covered by phylloid algae, which suggested an alternate community replacement. Some outrunners of Fomitchevella initialized in Ivanovia cf. manchurica community and then overthrew the former dominantor and finally a gradual replacement occurred. Fomitchevella-1 community evolved into Fomitchevella-2 community, during which adjustments on biological composition and proportion occurred to make the community structure was more appropriate with high adaptability to the environment. Thus this was actually an update of community.
Development of the reef is closely related to community evolution. The whole formation process could be divided into four stages and each stage involved transformation of initialization, development and decline of the communities. The patch reef layers were built in the first two stages, while the main body of the reef was built in the last two stages. Growth rate of the reef was controlled by both nutrient levels and sedimentation rate, and in a certain extent they were positively correlated. The reef would stop growing when environmental conditions changed abruptly.
该礁体规模巨大,结构独特,根据产礁岩系特征可划分为上下叠置的3个单元(U1、U2、U3)。其中U1为一点礁层,包括一个蓝细菌泥丘、一个Ivanovia cf. manchurica点礁和一个叶状藻点礁,通过上覆的生物碎屑层与礁主体部分划分开。U2和U3具有相同的岩系特征和类似的生物组成,它们都是由Fomitchevella骨架岩和泥粒岩组成,区别在于U3的生物丰度明显高于U2。
研究区内造礁生物种类丰富,生态类型多样,成礁方式各异。在扁坪村大型组合式珊瑚礁中主要有造架生物Fomitchevella、造架—障积生物叶状藻,造架—覆盖生物Ivanovia cf. manchurica以及粘结生物蓝细菌。其中Fomitchevella为礁体的主要建造者,它能够通过出芽生殖方式形成笙状或丛状群体,从而建造了不同类型的礁体。群体中间部位的个体较为粗大,边缘部位的则细小,个体的生存能力较强。Ivanovia cf. manchurica群体呈板状,生长方式多样,群体的再生能力很强,参与了U1中的点礁建造。叶状藻是研究区最为重要的造礁生物之一,发育了多种形态,建造了不同类型的礁体,其中包括U1中的叶状藻点礁。另一种群体珊瑚Antheria数量较少,分布较为局限,它是群落演化中的一个过渡角色。蓝细菌是具有粘结功能的自养生物,它是U1中灰泥丘的主要建造者。附礁生物以蜒、腕足类为主,海百合、Tubiphytes、腹足类较为常见,含苔藓虫和少量单体珊瑚。
该礁体内划分出了6个主要的造礁生物群落,它们具有类似的生物多样性,但群落的生物丰度、空间分层性、关键种的功能作用等各有差异。根据生物的生活习性、功能结构和取食位置,将它们归类为不同的生物群团。自蓝细菌群落向Fomitchevella群落发展,群落的空间分层性越来越明显,对环境的适应能力和对内部生物的调控能力也越来越强。蓝细菌、Tubiphytes、蜓、非蜒有孔虫、腕足类、腹足类生活于群落的下层,海百合、Ivanovia cf. manchurica、Antheria在中层空间取食,Fomitchevella则有着较高的捕食位置。群落的生物组分有所不同,但群落的维持机制基本一致。群落对营养盐的吸收与现代珊瑚礁群落类似,分别适用于一种或多种吸收模型。整个生态系统的能量最终来源于太阳光,藻类、微生物、蓝细菌、Tubiphytes和水体中的溶解有机物构成了初级生产力,能量通过生物间的捕食关系层层传递和损失,最终由分解者释放到水体中,一部分参与循环,绝大多数随水流散失。
借助于对现代珊瑚礁的研究成果,对扁坪村大型组合式珊瑚礁中的生态关系类型做了详细的分析和研究。腕足类生活于叶状藻的藻丛内部,不仅获取了更多的食物,还可以在藻丛的保护环境中躲避天敌的捕食,它们不存在竞争关系,能够和谐共处,也能单独生存,因此它们是一种特殊的共栖关系。Fomitchevella群体中相邻个体为争夺更多的生长空间、更广的食物来源和更大的采光面积而出现激烈的种间空间竞争,部分个体弯曲、畸形、倾斜,甚至出向侧向的生长现象,它们由此失去了更多的生存资源,最终被直立、粗壮、高大的个体所替代。“未名造礁生物”生长和扩张速度快,攀附缠绕能力强,它们包绕在Ivanovia cf. manchurica的板状群体上,双方在接触带上出现组织加厚,形成主动包绕型的接触关系,“未名造礁生物”剥夺了Ivanovia cf. manchurica的生存资源致其死亡。Fomitchevella的个体占据基底的能力较差,Ivanovia cf. manchurica群体在侧向扩张时尽量避免与其骨架直接接触,而Fomitchevella的群体通过侧芽的方式占据了Ivanovia cf. manchurica的上方空间,从而对其形成遮蔽,二者形成了避让遮蔽型的空间竞争,最终Fomitchevella成为优势者。叶状藻对生存于藻丛中的零星Fomitchevella个体形成数量压倒型的空间竞争,竞争结果因环境条件而异。Ivanovia cf. manchurica发育在叶状藻建造的基底上,而后又被叶状藻覆盖,出现了交互式的群落替代；Ivanovia cf. manchurica群落中出现了Fomitchevella的先驱个体,后来二者的角色发生了交换,最后Fomitchevella完全成为主导者,形成了渐变式的群落替代。Fomitchevella-1群落向Fomitchevella-2群落转变的过程中,生物组成和比例出现了一定的调整,使群落的结构更加合理,对环境的适应能力更强,因此实质上是一种群落的更新。
礁体的发育与群落的演化关系密切,整个发育过程可划分为四个阶段,每个阶段都包含了群落形成、发展、衰退等状态的改变。前两个阶段建造了点礁层,后两个阶段建造了礁主体。水体的营养程度和沉积速率控制着礁体的生长速度,在一定限度内它们成正相关关系,当这些环境条件出现突变时礁体往往会停止生长。
Carbonate rocks of plateau facies well developed in vast sea area of southern China in Late-Carboniferous, and the south of Guizhou Province was a representative region. Shallow water environment on the carbonate platform was quite favourable for organisms, especially the benthic organisms, and as a result different types of reef-building communities formed. Lithogeneous process of these organisms is intensive with organic banks, bioherms and reefs built widely, and thus it is an ideal region to study Late-Carboniferous biostratum and reef of South China. Most of these reefs developed on the platform margin, and the large-scale reef complex is the mostly representative one.
The reef complex has a large size and a special architectural structure, and 3 superimposed units, U1, U2 and U3 could be recognized according to features of reef rocks. U1 is a patch reef layer involving a mud mound built by cyanobacteria, an Ivanovia cf. manchurica patch reef and a phylloid algae patch reef, and it is separated from mainbody of the reef by an overlying bioclast layer. U2 has an identical lithology and a similar biological composition with U3, that is, both of them are composed of Fomitchevella framestone and packstone, the difference is that abundance of organisms in U3 is much higher than that in U2.
In the study region, the reef-building organisms are various in genera and ecotype, and their reef-building modes are diversefied. In the large-scale coral reef complex in Bianping Village, these reef-building organisms mainly include:1) Fomitchevella, frame builder; 2) phylloid algae, frame-builder and baffler; 3) Ivanovia cf. manchurica, frame-builder and covering organism; and 4) cyanobacteria, binding organism. Fomitchevella is the dominant constructor of the reef complex who grows into a phaceloid or fasciculate colony through budding and then builds reefs of diverse types. The skeletons of Fomitchevella individuals are large in the centre of the colony while small on the edge, and the individuals are viable. Ivanovia cf. manchurica develops a tabular colony and various growth forms. Their colonies get a strong power of regeneration and make a great contribution to the formation of U1. Phylloid algae is one of the most important builders in the study region which has diverse morphologies and builds different types of reefs, including the phylloid algae patch reef in U1. The other genera of colonial coral-Antheria has a low abundance and a limited distribution, and it plays a transitional role in community evolution. Cyanobacteria is a sort of autotroph with binding function, and it is the main builder of mud mound in U1. Fusulinids and brachiopods are the most common dwellers, followed by crinoids, Tubiphytes and bivalves, while bryozoans and solitary corals are few.
Six reef-building communities are recognised in the reef complex, which are similar in biological diversity while different in abundance of benthos, vertical spacial zonation, impacts of the key species and so on. The organisms are divided into various groups according to their life habits, fuctions and feeding positions. The vertical zonation becomes more and more notable along with evilution of these communities. Their adaptability to the evironment and control on organisms also get an obvious development. Cyanobacteria, Tubiphytes, fusulinids, foram, brachiopods and gastropods live in the lower part of the communities, while rinoids, Ivanovia cf. manchurica and Antheria in the middle part and Fomitchevella in the upper part. The biological components of these communities are different but the maintaining mechanisms are generally accordant.
Nutrient absorbing of the communities is similar to that of extant coral reef community, and one or more absorption models may be available. The energy of the whole ecosystem is derived from sunlight ultimately. Algaes, microbes, cyanobacteria, Tubiphytes and deliquescent organic materials in the water constitute the primary productivity. Energy is transferred and lost gradually through predation between organisms, and finally released into the water by decomposers. Part of the energy recycles in the ecosystem, and most is dispersed with the waterflow.
Ecological relationships in the coral reef complex in Bianping Village are analysed and studied based on research findings of extant reefs. Brachiopods lived in the mats of phylloid algae, which enabled them to get more food and also avoid being hunted by natural enemies. They could coexist without competition or live independently, and thus they were in a special relationship of commensal. In order to get more space for growth, source of food and larger lighting area, the individuals in Fomitchevella colony always competed intensely with their neigbours, resulting in some bent, misshapen, inclined or even lateral individuals. As a result, these individuals lost more living resources and would be finally replaced by those erect, strong and long ones. The "unamed reef-building organisms" grew and expanded at a high rate and were good at en winding. They wraped on the colony of Ivanovia cf. manchurica, and both of them thickened tissues on the contact zone, and as a result an aggressive-wraping relationship was formed. The "unamed reef-building organisms" deprived Ivanovia cf. manchurica of living resources and would finally lead to its death. When laterally expanding, Ivanovia cf. manchurica would as much as possible avoid direct contact with individual skeletons of Fomitchevella, who was not good at occupying the substrate but could take up the space above Ivanovia cf. manchurica throgh laterally budding and overshaded it. Then an off-overshading spatial competition was formed and Fomitchevella would be the dominant at last. An overwhelming spatial competition was formed by phylloid algae and a few Fomitchevella individuals lived in the algae mat, but the outcomes would be various with the influences of environmental conditions. Ivanovia cf. manchurica lived on substrate built by phylloid algae and the was covered by phylloid algae, which suggested an alternate community replacement. Some outrunners of Fomitchevella initialized in Ivanovia cf. manchurica community and then overthrew the former dominantor and finally a gradual replacement occurred. Fomitchevella-1 community evolved into Fomitchevella-2 community, during which adjustments on biological composition and proportion occurred to make the community structure was more appropriate with high adaptability to the environment. Thus this was actually an update of community.
Development of the reef is closely related to community evolution. The whole formation process could be divided into four stages and each stage involved transformation of initialization, development and decline of the communities. The patch reef layers were built in the first two stages, while the main body of the reef was built in the last two stages. Growth rate of the reef was controlled by both nutrient levels and sedimentation rate, and in a certain extent they were positively correlated. The reef would stop growing when environmental conditions changed abruptly.
引文
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