新元古代末期高家山生物群的生态多样性

doi:10.13745/j.esf.sf.2020.6.2

摘要/Abstract

摘要：

新元古代末期是生命演化的关键转折期,也是以微生物占主导的生态系统向显生宙以后生动物占主导的生态系统的转变期,埃迪卡拉纪大型软躯体生物以固着、底栖、食悬浮为特色,普遍缺乏运动能力。作为这一时期特殊代表的高家山生物群,是目前新元古代唯一一个以黄铁矿化三维保存的管状和锥管状化石为主导,兼有骨骼生物、原生动物、钙化蓝细菌类及遗迹化石的多门类生物组合,是研究埃迪卡拉纪末期生命演化和生态系统演变十分重要的载体。本文通过对高家山生物群古生态学的初步研究,揭示出在前寒武纪—寒武纪之交,生态系统已显示一定的多样性。为适应平底面上(level-bottom)微生物席的发育,高家山生物群的许多生物采取了适应性的生存策略,通过黏附或插入微生物席中,营底栖固着食悬浮(如Cloudina、Conotubus)或化学共生(可能的Shaanxilithes)或平躺(如Gaojiashania和Sinotubulites)食碎屑生活。底内遗迹化石表明存在可能的表栖和半内栖、可自由运动、食碎屑的造迹生物。Conotubus中常见的“回春”或“复苏”现象,Gaojiashania和Sinotubulites的身体扭转或生活姿态调整则是对频繁风暴事件的被动适应。

关键词: 高家山生物群, 新元古代末期, 生态多样性, 生命创新事件, 微生物席底, 风暴事件沉积

Abstract:

Ediacaran fossils mark a pivotal position in the evolution of life as it transitions between the predominantly microbial ecosystems of the Precambrian and the animal ecosystems of the Phanerozoic. Ediacaran communities were dominated by sessile epibenthos and their ecological structures were relatively simple. Many modern feeding modes such as macropredation, epibiosis, and interspecific competition or mutualism were absent or poorly represented. It is generally accepted that the soft-bodied Ediacara biota is part of a “failed” evolutionary experiment and it has no clear genetic relationship with Phanerozoic organisms. However, corresponding to the evolution of typical Ediacaran soft-bodied biota, there was a great proliferation of tubular animals at the end of the late Neoproterozoic, represented by the Gaojiashan biota, where Lagerstätte fossil deposit hosts a variety of soft-bodied or lightly biomineralized tubular fossils (Shaanxilithes, Gaojiashania, Sinotubulites, Conotubus, and Cloudina), calcareous microfossils (Protolagena), and calcareous cyanobacteria.
Unlike other Ediacara fossil assemblages, the Gaojiashan biota is dominated by benthic sessile suspension feeders or detritus feeders. Of them, Cloudina is a millimetre-scale conical tube tapering adapically from an aperture to a rounded apex. The bulk of the tube is constructed by a series of successively stacked, repetitive, unevenly spaced, and funnel-shaped tube wall units. Cloudina occupied an epibenthic suspension-feeding life-mode, with the apex anchoring to microbially bound muddy substrate and the aperture extending upwards into the water column. Conotubus is a centimetre-sized conical tube consisting of a series of nested cylindrical-to-funnel-shaped tube walls (cylinders hereafter). This conotubular construction of nested cylinders is similar to that of the late Ediacaran fossil Cloudina. Integrated morphological, taphonomic and palaeoecological data suggest that it likely employed an epibenthic life-mode, with the apex anchoring to the muddy substrate and the aperture extending upwards into the water column and it was probably a suspension-feeding organism. Gaojiashania is a centimetre-size tube that consists of a series of repeating units. Each unit consists of a rigid ring and a flexible bucket-shaped tube wall. Each tube maintains a constant diameter and the rigid ring elements typically range from 1 to 2 mm in length, whereas the bucket components are more varied at 0.5-5 mm in length and are ornamented with transverse annuli. Gaojiashania, however, may have been a procumbent epibenthos, possibly with the rings anchoring to microbially bound muddy substrate. Sinotubulites are characterized by millimetre-to centimetre-size multilayer tubes open at both ends. The tube consists of two morphologically different walls: a multilayer inner wall with weak ornamentation, and a multilayer outer wall with transverse or oblique corrugations and sometimes longitudinal ridges. Surface ornamentation and polygonal shape of the cross-section suggest that Sinotubulites probably lived as an epibenthos lying on the sea floor. Shaanxilithes is a ribbon-shaped impression with constant widths (1-6 mm or more) and is characterized by a series of closely spaced transverse annulations. Shaanxilithes is tentatively interpreted as a sessile epibenthos, with one end anchoring to the sandy or muddy substrate. Therefore, the Gaojiashan biota is characterized by complex morphological types and diverse body plans of variable size and lifestyle, representing a major biological innovation in Earth’s evolution.
Flourishing microbial mats on the Precambrian level-bottom forced the Gaojiashan biota to evolve some adaptive strategies, either through fixation to (such as Cloudina, Conotubus, and possibly Shaanxilithes) or free laying on (such as Gaojiashania and Sinotubilites) the substrate by adhering to or inserting into microbial mats. To cope with the frequent disturbance by storm events, ecological adaptations were manifested in many Gaojiashan organisms. The polygonal cross-section and longitudinal ridges in Sinotubulites may stabilize the tube lying on the substrate, preventing it from rolling in strong currents. After being subjected to sediment obrution, Cloudina and Conotubus employed specialised palaeoecological strategies to rejuvenate and self-right the tubes, while Gaojiashania and Sinotibulites could partially self-right through curving, extension and constriction, indicating strong burial-resistant capabilities.
Abundant meandering traces and possible burrows in the Gaojiashan biota and in the comparable Shibanta biota imply that bilaterians were an essential part of later Ediacaran ecosystems and indicate the possible presence of epibenthic and semibenthic free-moving and detritus-feeding organisms. The complexity of predator-prey dynamics at the beginning of metazoan diversification is evidenced by borings on Cloudina tubes.

Key words: Gaojiashan biota, terminal Neoproterozoic, ecological diversification, biological innovation, microbial-mat substrate, storm event deposit

中图分类号:

华洪, 蔡耀平, 闵筱, 柴姝, 代乔坤, 崔再航. 新元古代末期高家山生物群的生态多样性[J]. 地学前缘, 2020, 27(6): 28-46.

HUA Hong, CAI Yaoping, MIN Xiao, CHAI Shu, DAI Qiaokun, CUI Zaihang. Ecological diversity in the terminal Ediacaran Gaojiashan biota[J]. Earth Science Frontiers, 2020, 27(6): 28-46.

图/表 14

图1 高家山生物群化石产地

Fig.1 Geographical location of the Gaojiashan biota

图2 高家山—牛落坑剖面灯影组高家山段下部野外剖面层位1~5,图中黑色线条表示高家山段起始层位。

Fig.2 Outcrop of the lower Gaojiashan Member in the Dengying Formation of the Gaojishan-Niuluokeng section

图3 高家山—牛落坑剖面灯影组高家山段中上部野外剖面层位6~9,图中黑色线条表示高家山段终结层位。

Fig.3 Outcrop of the middle and upper Gaojiashan Member in the Dengying Formation of the Gaojishan-Niuluokeng section

图4 高家山段事件沉积与微生物岩构造(部分图件据文献[20,23]修改) (a)—黄绿色粉砂岩中的正粒序。(b)—Gaojiashania化石保存于风暴事件层中的正粒序粉砂岩中,上部的均一块状泥质岩中未见化石。(c)—黄铁矿化Conotubus化石产出在微生物席覆盖的粉砂岩中,显示多角状MISS构造。(d)—群集的黄铁矿化Conotubus管体保存于风暴事件层中的正粒序粉砂岩中,箭头所示为粉砂岩层底面上出现的石英颗粒层。(e)—定向(顶面朝上)的手标本显示两套粉砂岩-泥岩构成的层偶。数字示意每层粉砂岩层的底面。层偶由下部发育正粒序的粉砂岩层和上部的均一块状泥岩层构成。粉砂岩层的底面与下伏岩层界线截然,顶面与上覆岩层呈过渡关系。(f)—高家山段上部发育具明暗交替纹层的微生物灰岩。

Fig.4 Event deposits and MISS structures. Partially modified from [20,23].

表1 高家山生物群已知化石类群及其初步认识

Table 1 List of distinct taxa in the Gaojiashan biota and their possible interpretation

属种名称						生态位
管状动物化石	Cloudina	可能多毛类	漏斗套漏斗锥管状结构,具矿化壁	底栖	固着滤食	初级消费者
	Conotubus	可能多毛类	漏斗套漏斗锥管状结构,有机质壁	底栖	固着滤食	初级消费者
	Gaojiashania	后生动物	由一系列坚硬的圆环构成的空心管体,圆环间由软体组织连接	表栖	游移食腐殖质	初级消费者
	Sinotubulites	可能多毛类	管套管结构; 多层壁,表面具纵脊及横向皱褶	表栖	滤食或食腐殖质	初级消费者
	Shaanxilithes	后生动物化石	带状构造,一系列片状构造组合	底栖	固着化学共生	自养
瓶状化石	Protolagena	原生动物,可能有孔虫	瓶状,颈长短不一,颈短,口缘圆滑或具外翻的漏斗状口缘	底栖	滤食	初级消费者
瓶状化石	Sicylagena	原生动物,可能有孔虫	亚葫芦状, 具颈、口孔居中或侧位	底栖	滤食	初级消费者
蓝细菌或藻类化石	Obruchevella	钙化蓝细菌	螺旋状盘卷管丝体	底栖	固着	生产者
	Eiphyton	钙化蓝细菌	二歧式分枝管丝体	底栖	固着	生产者
	Cambricodium	钙化蓝细菌	束发状管丝体	底栖	固着	生产者
	Girvanella	钙化蓝细菌	相互缠绕管丝体	底栖	固着	生产者
遗迹化石	Bucerusichnus	动物遗迹	可能垂直潜穴	底内	食腐殖质	初级消费者
	Planolites	动物遗迹	简单表生爬迹	底内	食腐殖质	初级消费者
	Helminthopsis	可能遗迹化石	蛇曲形遗迹	底内	食腐殖质	初级消费者
可能的小壳化石先驱	Anabarites sp.	动物化石	三个管规则旋卷	底栖	食腐殖质	初级消费者
可能的小壳化石先驱	Paragloborilus sp.	动物化石	圆锥形,始部球形肿大	底栖	食腐殖质	初级消费者

表1 高家山生物群已知化石类群及其初步认识

Table 1 List of distinct taxa in the Gaojiashan biota and their possible interpretation

属种名称						生态位
管状动物化石	Cloudina	可能多毛类	漏斗套漏斗锥管状结构,具矿化壁	底栖	固着滤食	初级消费者
	Conotubus	可能多毛类	漏斗套漏斗锥管状结构,有机质壁	底栖	固着滤食	初级消费者
	Gaojiashania	后生动物	由一系列坚硬的圆环构成的空心管体,圆环间由软体组织连接	表栖	游移食腐殖质	初级消费者
	Sinotubulites	可能多毛类	管套管结构; 多层壁,表面具纵脊及横向皱褶	表栖	滤食或食腐殖质	初级消费者
	Shaanxilithes	后生动物化石	带状构造,一系列片状构造组合	底栖	固着化学共生	自养
瓶状化石	Protolagena	原生动物,可能有孔虫	瓶状,颈长短不一,颈短,口缘圆滑或具外翻的漏斗状口缘	底栖	滤食	初级消费者
瓶状化石	Sicylagena	原生动物,可能有孔虫	亚葫芦状, 具颈、口孔居中或侧位	底栖	滤食	初级消费者
蓝细菌或藻类化石	Obruchevella	钙化蓝细菌	螺旋状盘卷管丝体	底栖	固着	生产者
	Eiphyton	钙化蓝细菌	二歧式分枝管丝体	底栖	固着	生产者
	Cambricodium	钙化蓝细菌	束发状管丝体	底栖	固着	生产者
	Girvanella	钙化蓝细菌	相互缠绕管丝体	底栖	固着	生产者
遗迹化石	Bucerusichnus	动物遗迹	可能垂直潜穴	底内	食腐殖质	初级消费者
	Planolites	动物遗迹	简单表生爬迹	底内	食腐殖质	初级消费者
	Helminthopsis	可能遗迹化石	蛇曲形遗迹	底内	食腐殖质	初级消费者
可能的小壳化石先驱	Anabarites sp.	动物化石	三个管规则旋卷	底栖	食腐殖质	初级消费者
可能的小壳化石先驱	Paragloborilus sp.	动物化石	圆锥形,始部球形肿大	底栖	食腐殖质	初级消费者

图5 Cloudina形态学特征 a—薄片显示管体套合特征;b, c—扫描电镜显示Cloudina的漏斗套漏斗结构;d—Cloudina漏斗套漏斗结构及生长方向的转变,可能仅末端始终向上;e—箭头所示为Cloudina管体上的生物钻孔。

Fig.5 Growth patterns of Cloudina tubes

图6 Conotubus形态学特征(b、c据文献[12]) a—弯曲生长的Conotubus化石,可能仅开口端的管节直立伸向水体中;b, c—黄铁矿化的Conotubus化石标本,箭头所示为可能的“回春”或“复苏”现象;d—薄片显示的Conotubus漏斗套漏斗结构及可能的“回春”或“复苏”现象:可能仅末端始终向上;e—黄铁矿化的Conotubus化石标本。

Fig.6 Growth patterns of Conotubus tubes. b and c adapted from [12].

图7 Gaojiashania形态学特征 a—弯曲生长的黄铁矿化Gaojiashania,显示管体可能具一定的柔韧性;b—叠压的圆环或管体碎片;c—Gaojiashania化石,显示加厚的圆环;d—黄铁矿化保存的Gaojiashania化石,显示加厚的圆环和环间细纹;e—平行粉砂岩-泥岩层面产出的至少5个Gaojiashania化石个体组成的化石群体。化石标本均以有机碳质压膜-黏土矿物方式保存。

Fig.7 Growth patterns of Gaojiashania tubes

图8 Sinotubulites形态学特征 a,b—弯曲生长的Sinotubulites,红色虚线显示管体具一定程度的扭曲,推测为风暴事件沉积后为防止被悬浮的碎屑掩埋,管体的被动姿态调整;c—3-辐射状Sinotubulites管体,可能为增大平躺贴合面积;d—Sinotubulites管体横切面薄片,显示多层套合管体特征,内部管壁平直,外部通过部分加厚形成皱褶。

Fig.8 Growth patterns of Sinotubulites tubes

图9 Shaanxilithes形态学特征 a,b,c—不同保存状态下的Shaanxilithes管体。

Fig.9 Growth patterns of Shaanxilithes tubes

图10 瓶状化石形态学特征 a,c—虚线显示示顶底构造,近于30°与沉积底面斜交,可能反映口部朝下以一定角度斜交黏附在藻席覆盖的基底上生长的特征;b—示顶底构造,瓶口部碎屑物充填;d—瓶状化石扫描电镜照片。

Fig.10 Growth patterns of vase-shaped microfossils

图11 磷酸盐化保存的钙化蓝细菌(据文献[17,18]) a—Obruchevella显示左旋的旋卷特征;b—复杂树丛状分枝Epiphyton管体,分枝角约30°~45°;c, d—Girvanella,显示空心的束状丝体;e—Cambricodium,管状丝状体形成的束状群体;f—Subtifloria,排列成束的管状。

Fig.11 SEM images of phosphatized calcified cyanobacteria. Adapted from [17-18].

表2 根据营养阶层、运动方式及取食策略对高家山生物群相关动物生态类型划分

Table 2 Ecological categories in the Gaojiashan biota based on tiering position, motility level and feeding mechanism

生态分类		特征描述	高家山生物群动物化石代表
营养层	1.浮游游泳	生活在脱离基底的水体中
	2.底栖直立	底栖,直立于水体中生活	Conotubus,Shaanxilithes
	3.表栖	在基底表面生活,不明显向上生长	Cloudina,Sinotubulites,Gaojiashania,遗迹化石
	4.半内栖	半埋半暴露在水体中生活	遗迹化石
	5.浅表内栖	在基底-水界面下5 cm之内的沉积物中生活	遗迹化石
	6.深部内栖	在基底之下大于5 cm的沉积物内生活
活动能力	1.自由快速运动	有规律地不受阻碍的移动或游移
	2. 自由慢速运动	有规律地移动,与基底保持紧密接触	瓶状化石
	3.兼性非固着	只有在必要的时候才会移动,自由平躺
	4.兼性固着	固着生活,只有在必要的时候才移动	Sinotubulites,Gaojiashania
	5.非固着,不能移动	不具备自我推移的能力,平躺生活
	6. 固着,不能移动	不具备自我推移的能力,固着生活	Cloudina,Conotubus,Shaanxilithes
取食策略	1.食悬浮	捕捉水中的悬浮食物颗粒	Cloudina,Conotubus,瓶状化石
	2.底表食碎屑	从基底上捕捉松散的颗粒	Sinotubulites,Gaojiashania
	3.底内挖掘	在基底内取食埋在地下的食物	根据遗迹化石推测
	4.牧食	从基质上刮削或啃食食物
	5. 捕食	捕获具一定抵抗力的生物	Cloudina管体表明的钻孔
	6. 其他营养方式	包括光合自养或化学共生及寄生等多种类型	Shaanxilithes?

图12 高家山生物群生态空间占用与埃迪卡拉白海组合、纳玛组合及现代海洋动物对比(改自文献[82,112])

Fig.12 Ecospace occupancy in the Gaojiashan biota, as compared with the White Sea and the Nama assemblages and modern animals. Modified from [82,112].

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