Brachiopod-Supported Benthic Communities from Guanshan Biota Uncover Early Cambrian Ecological Complexities
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摘要: 关山生物群产自滇东地区寒武系第4阶，是一个由多门类后生动物化石组成的原位埋藏的布尔吉斯页岩型化石库，该生物群在数量上以腕足动物主导，它们大量出现并密集成层，特色鲜明.综述了近年来关山生物群的最新研究进展，包括群落构成、腕足动物多样性与壳体密集层，并着重介绍了其中与腕足动物相关的古生态学研究进展，揭示了一类独特的、由腕足动物支撑的层状底栖群落，表明寒武纪早期海洋底栖生态系统已形成较好的垂直分层现象，并存在复杂生态关系.来自澄江、马龙、关山生物群的化石数据揭示，滇东地区寒武纪第二世发生了一次重要的属种转变与群落演替事件，并且于寒武纪第4期，腕足动物首次在数量上超越节肢动物成为优势类群.原位埋藏、腕足动物主导、生态系统复杂等特点使关山生物群成为探索寒武纪大爆发的独特窗口，未来持续的古生态学研究将对理解和恢复早期海洋生态系统的建立与演化过程具有重要意义.同时，开展该时期古环境的纵向演变研究，探索影响群落演替的关键控制因素，将对寒武系第二统全球地层对比工作提供重要参考.Abstract: The Guanshan Biota is an in situ preserved Lower Cambrian (Series 2, Stage 4) Burgess shale-type Konservat-Lagerstätten from East Yunnan, South China, which is composed of multiple metazoan groups. Recent studies revealed that Guanshan Biota is dominated by brachiopods on relative abundance. Those brachiopods formed a geographically widespread phenomenon of many monospecific or paucispecific shell-bed concentrations during the Cambrian Age 4. The repaired shell damages and encrusted kleptoparasites on brachiopod shells reveal complex ecological relationships of benthic communities. Associations of sessile epibenthic brachiopods encrusted with kleptoparasitic tube-dwelling worms, infaunal paleoscolecidans, and swimming Tuzoia were discovered on the same micro-bedding plane. It represents a unique, brachiopod-supported, well-developed vertically stratified benthic community, which is composed of the different phyla that fill multiple ecological niches, exhibiting a complex trophic network and community structure with increased utilization of eco-space during the Cambrian Age 4. Furthermore, the fossil record shows an unusual ecological transition from trilobite- to brachiopod-dominated communities from the Guanshan Biota for the first time. Generally, the Chengjiang, Malong and Guanshan assemblages share similar taxa, however, the main trilobite and brachiopod assemblages are unique. This indicates an important ecological event of faunal overturn and benthic community succession during the Cambrian Epoch 2 in East Yunnan. In Situ preserved, brachiopod-dominated Guanshan Biota with complex ecosystem provides a unique view of the seafloor during the Early Cambrian. Paleoecology, especially biological interactions between other benthic taxa in the Guanshan Biota are required in the future. Temporal changes of paleoenvironment during the Cambrian Explosion and key factors that affect benthic community succession are still unclear in East Yunnan. Further studies will provide important evidence for Cambrian Series 2 global stratigraphic correlation.
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Key words:
- Guanshan Biota /
- brachiopod /
- benthic community succession /
- ecosystem complexity /
- Cambrian Age 4 /
- East Yunnan /
- ecology
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图 1 滇东地区寒武纪早期的沉积露头分布、化石产地及马龙砍斧箐剖面地层柱状图
橙色星号指示本文主要的化石采集剖面（砍斧箐、石将军、大海剖面）以及产出层位
Fig. 1. Fossil localities and stratigraphic column of the Cambrian Series 2 in East Yunnan, Yangtze
图 2 滇东地区寒武系第4阶乌龙箐组下部野外露头照片
a.马龙砍斧箐剖面乌龙箐组（WLQ）泥岩、粉砂岩与下伏红井哨组（HJS）厚层石英砂岩整合接触界线；b.武定县石将军剖面乌龙箐组与红井哨组界线；c.乌龙箐组底部砾岩；d.乌龙箐组下部泥岩、粉砂岩互层
Fig. 2. Field photographs of the lower Wulongqing Formation (Cambrian Stage 4) in East Yunnan
图 3 关山生物群中的部分代表门类
a.保存软组织的腕足动物Neobolus wulongqingensis；b.小嘴贝类腕足动物Nisusia（白色箭头）和Kutorgina（黑色箭头）；c.古虫动物前体；d.保存原始磷酸钙质壳体成分的腕足动物Eoobolus malongensis；e.未定物种；f.三叶虫Redlichia；g.等刺虫；h.奇虾类前附肢；i.保存软组织的软舌螺Linevitus Malongensis
Fig. 3. Fossils collected from the Guanshan Biota in East Yunnan
图 4 关山生物群中磷酸钙质壳和钙质壳腕足动物的属种分类(据段晓林，2021修改)
Fig. 4. Phosphatic and calcareous brachiopod taxa from the Guanshan Biota (modified from Duan, 2021)
图 5 关山生物群中的腕足动物和三叶虫单属种密集层
a.腕足动物Linnarssonia；b.三叶虫Palaeolenus；c.腕足动物Neobolus；d.腕足动物Schizopholis
Fig. 5. Brachiopod and trilobite fossil concentrations from the Guanshan Biota in East Yunnan
图 6 马龙砍斧箐、会泽大海剖面采集的腕足动物Neobolus wulongqingensis-管状寄生蠕虫-古蠕虫-吐卓虫化石组合的光学照片
a.腕足动物及其管状附着生物、吐卓虫（左下角）共同保存；b.腕足动物壳体和古蠕虫躯干相互叠压保存（白色细箭头指示古蠕虫躯干，红色粗箭头指示腕足动物）
Fig. 6. General view of the paleoscolecidans-Neobolus wulongqingensis-parasite tube worm-Tuzoia association from the Guanshan Biota at the Kanfuqing and Dahai sections
图 7 复原图显示寒武纪第4期关山生物群中腕足动物Neobolus wulongqingensis支撑的层状底栖群落，显示关山生物群化石标本中游泳的吐卓虫、内栖的古蠕虫、表栖的腕足动物及寄生的管状蠕虫共存的现象
a.游泳的吐卓虫；b.表栖的腕足动物N.wulongqingensis和壳体上寄生的管状蠕虫；c.内栖的古蠕虫；d.钟形关山迹Guanshanichnus glockerichnoides Weber，Hu，et，Steiner，2012；e.水平遗迹侧视图；f.线形迹属Gordia Emmons，1844；g.环圈棋盘山迹Qipanshanichnus gyrus Luo et Tao，1994
Fig. 7. Simplified reconstruction shows a brachiopod Neobolus wulongqingensis -supported vertical, stratified benthic community in the Guanshan Biota during Cambrian Stage 4
图 8 滇东武定、马龙地区关山生物群门类个体丰度饼状图(据Chen et al., 2021修改)
Fig. 8. Pie charts of relative abundance of the Guanshan Biota in Wuding County and Malong County respectively(modified from Chen et al., 2021)
图 9 武定石将军剖面乌龙箐组下部岩性柱状图及壳体富集层(a)和不同采样层位(A~J)化石组合相对丰度变化图(b)
红色线条之下地层内节肢动物的数量较多，之上腕足动物占据主导地位；据Chen et al.（2021）修改
Fig. 9. Stratigraphic fluctuation in the relative abundance of the community at the phylum level (a) and brachiopod genus level (b) from the Shijiangjun Section
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