Benthic Ecosystem Engineer after the Cambrian Explosion: An Example from Henan Province
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摘要: 扰动生物作为生态系统工程建造者，通过对沉积底质的改造直接影响沉积物和水体之间的相互作用（营养物质的循环、迁移和存储），对底栖生态系统工程有重要的影响.遗迹化石作为底栖生物活动的载体，为研究寒武纪大爆发之后底栖生态系统工程建造者的演化规律，并评价底栖生态系统工程建造者对生态系统工程产生的影响提供了依据.对河南地区8个寒武系剖面的遗迹化石数据进行统计，应用现代生物学中生态系统工程影响值的概念，从功能群、阶层、生物灌溉3个方面对底栖生态系统工程建造者进行研究.研究结果表明，寒武系第三阶至鼓山阶底栖生态系统工程建造者功能群类型由生物扩散型扰动者往向上/下运输者转变，阶层逐渐加深，生物灌溉越来越普及，生态系统工程影响值不断增大.鼓山阶至第十阶，遗迹化石类型及结构变化不大，生态系统工程影响值趋于稳定.这表明从寒武纪生物大爆发至鼓山阶，底栖生态系统工程建造者的行为策略发生着明显的进化与创新，生态系统工程建造者不仅开拓了重要的生态空间，而且为其他底栖生物和微生物创造了新的生境，在寒武纪海洋生态系统中扮演着重要的角色.Abstract: The bioturbators, which are ecosystem engineers, act on the interaction between sediments and water (the circulation, migration and storage of nutrients) through the reworking of sediments directly. As a result, they have an significant impact on benthic ecosystems engineering. The trace fossils provide evidence for studying the evolution of ecosystem engineers and assessing the impact of engineers on ecosystem engineering after the Cambrian explosion, as they provide records for activities of benthic organism. This paper studies benthic ecosystem engineers from three aspects-functional group, tiering and biological irrigation, by applying the modern biology concept of 'ecosystem engineering impact'. Statistics on the trace fossils of eight Cambrian sections in Henan Province show that the functional group types of the ecosystem engineers have changed from biodiffusive bioturbators to upward/downward conveyors, infaunal tier has gotten deeper, biological irrigation has became more common, and the ecosystem engineering impact has been increasing from Stage 3 to Drumian. But the type and structure of trace fossils have no changes, and the ecosystem engineering impact tends to be stable from Drumian to the Stage 10.This means that the behavioral strategy of benthic ecosystem engineer sevolve and innovate obviously during the time from the ending of the Cambrian Explosion to Drumian, by expanding ecospaces and creating new habitats for other macrobiotic and microbiotic organisms. In a word, these ecosystem engineers played an important role in the Cambrian marine ecosystem.
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Key words:
- Cambrian /
- benthic organism /
- ecosystem engineer /
- functional groups /
- tiering /
- biological irrigation /
- paleontology
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图 1 河南华北型寒武系剖面地理位置及岩性特征
Fig. 1. Map showing the geographical location and lithological characteristics of the North China type Cambrian in Henan Province
图 2 河南寒武系第三阶典型遗迹化石
a. Oldhamia（Ol），由一个中心向外放射的细线组成的放射状花纹，覆盖于层面；Torrowangea（To），蛇曲形潜穴，遗迹表面具有横环节；b. Helminthoidichnites（He），层面蛇曲形光滑通道，互不相交；c. Planolites（Pl），柱形潜穴，水平或略倾斜分布，充填物无结构；d. Treptichnus（Tr），锯齿状潜穴；e. Phycodes（Ph），水平束状的柱形潜穴，表面光滑或具细环纹；f. Skolithos（Sk），垂直潜穴，层面上呈点状
Fig. 2. Typical trace fossils of the Cambrian Stage 3 in Henan Province
图 3 河南寒武系第四阶Thalassinoides潜穴
a. Thalassinoides层面可见Y形分支（Yb）和T形分支（Tb）；b. Thalassinoides水平方向呈多枝网格状互相连接；c.剖面上可见Thalassinoides垂直管与水平管相连成三维潜穴系统，具Y形和T型分支；d.剖面上可见Thalassinoides潜穴多期扰动，显著地改变了原始沉积组构
Fig. 3. The ichnogenus Thalassinoides of the Cambrian Stage 4 in Henan Province
图 4 河南寒武系乌溜阶典型潜穴
a. Cruziana，由三叶虫形成的二叶形爬痕; b. Scolicia，平行层面分布，潜穴两侧光滑平整，不发育横向纹饰，遗迹中间的脊条特别明显；c. Arenicolites，垂直层面的宽阔U形潜穴，无蹼状构造，受出露位置影响，其在剖面上往往呈I形或J形；d. Palaeophycus，柱形潜穴微弯曲，不发育分支，表面具纵纹；e. Diplocraterion，潜穴垂向延伸，圆柱形栖管末端呈烟筒状延伸到层面之上，两栖管之间距离3~5 cm; f. Skolithos，垂直或略倾斜管状潜穴，垂向延伸深度5~15 cm
Fig. 4. Typical trace fossils of the Cambrian Wuliuan in Henan Province
图 5 河南寒武系鼓山阶Thalassinoides isp.潜穴
图a~c为剖面照片，可见Thalassinoides isp.的分布主要呈高角度倾斜或垂直于层面发育，部分呈水平或近水平分布发育纵切面多呈I/J形，少数也可观察到扁平的U形，且有分枝；图d~e为层面照片，遗迹化石聚集呈簇状、线状或点状分布，层面上也可观察到一些潜穴呈排或列分布，具有厚衬壁是该遗迹化石的一个显著特征
Fig. 5. The Thalassinoides isp. of the Cambrian Drumian in Henan Province
图 6 河南省寒武系鲕粒灰岩中的Planolites montanus
图a, b为层面照片；图c, d为剖面照面；鲕粒灰岩中的Planolites montanus（Pl）近水平分布，潜穴内部被灰泥充填，揭示了其造迹生物生活在深阶层中
Fig. 6. The Planolites montanus in Cambrian oolitic limestone in Henan Province
图 7 寒武系遗迹化石多样性、差异性及生态系统工程影响值变化趋势
Fig. 7. The variation trend of Cambrian ichnodiversity, ichnodisparity, ecosystem engineering impact
表 1 河南寒武系第三阶遗迹化石的生态系统工程影响值
Table 1. The ecosystem engineering impact value of the Cambrian Stage 3 in Henan Province
遗迹化石 (a)功能群α (b)阶层β (c)生物灌溉γ (d)生态系统工程影响值δ Didymaulichnus 1 1 1 3 Gordia 1 1 1 3 Helminthopsis 1 2 1 4 Lumbricarina 1 2 2 5 Oldhamia 1 2 2 5 Palaeophycus 1 2~3 2 5~6 Pelecypodichnus 1 1 1 3 Phycodes 1 3 2 6 Planolites 1 2 1 4 Skolithos 2 2~3 2 6~7 Taenidium 1 2 1 4 Torrowangea 1 2 1 4 Treptichnus 3 3 2 8 注：α: 1.扩散型扰动者; 2.再造者; 3.向上/下运送者; 4.巷道式扩散者; β: 1.表层; 2.半内栖层; 3.浅阶层; 4.中阶层; 5.深阶层；γ: 1.不具有（大部分表栖层运动或觅食迹.不太可能具有生物灌溉）; 2.具有（平流主导型）; 3.具有（扩散主导型）；δ:生态系统工程影响值即a，b，c 3部分最小值和最大值相加所得到的范围. 
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表 2 河南寒武系第四阶遗迹化石的生态系统工程影响值
Table 2. The ecosystem engineering impact value of the Cambrian Stage 4 in Henan Province
遗迹化石 (a)功能群 (b)阶层 (c)生物灌溉 (d)生态系统工程影响值 Beaconites 1 1 1 3 Oniscoidichnus 1 1 1 3 Planolites 1 2 1 4 Thalassinoides 4 3-5 3 10-12
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表 3 河南寒武系乌溜阶遗迹化石的生态系统工程影响值
Table 3. The ecosystem engineering impact value of the Cambrian Wuliuan in Henan Province
遗迹化石 (a)功能群 (b)阶层 (c)生物灌溉 (d)生态系统工程影响值 Arenicolites 3 3 2 8 Beaconites 1 1 1 3 Bergaueria 2 2 1 5 Bifungites 2 2 1 5 Cruziana 1 1 1 3 Didymaulichnus 1 1 1 3 Dimorphichnus 1 1 1 3 Diplichnites 1 1 1 3 Diplocraterion 3 4 3 10 Gordia 1 1 1 3 Monocraterion 3 3 3 9 Monomorphichnus 1 1 1 3 Palaeophycus 1 3 3 7 Phycodes 1 3 2 6 Planolites 1 2 1 4 Qipanshanichnus 1 1 1 3 Rusophycus 1 1 1 3 Scolicia 1 1~2 1 3~4 Skolithos 2 3~5 2~3 7~10 Thalassinoides 4 3 3 10 Treptichnus 3 3 2 8
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表 4 河南寒武系鼓山阶至第十阶遗迹化石的生态系统工程影响值
Table 4. The ecosystem engineering impact value of the Cambrian Drumian to Stage 10 trace fossils in Henan Province
遗迹化石 (a)功能群 (b)阶层 (c)生物灌溉 (d)生态系统工程影响值 Arenicolites 3 3 2~3 8~9 Bergaueria 2 3 3 8 Cruziana 1 1 1 3 Diplichnites 1 1 1 3 Palaeophycus 1 3 3 7 Helminthopsis 1 2 1 4 Planolites 1 2~5 1~2 4~8 Skolithos 2 3~5 2~3 7~10 Treptichnus 3 3 2 8 Thalassinoides 3~4 3~4 3 9~11
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