Modeling Method Enhances Temporal Resolution of Deep⁃Time Food Web Stability Evolution: A Case Study on Permian⁃Triassic Ecological Record from the Meishan Section
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摘要: 约2.52亿年前发生的二叠纪‒三叠纪（P-Tr）之交生物大灭绝是地质历史上规模最大的生态危机事件，被认为是当代生态危机参照案例.不过，整合多个门类、系统刻画生物群落食物网结构整体演变过程的研究仍然匮乏.特别是，化石记录的不完备性及其在地层记录的低时间分辨率成为人们“以古鉴今”的卡点.本研究基于浙江煤山P-Tr界线金钉子剖面丰富的化石物种出现记录，利用PyRate贝叶斯模拟方法对所有物种在剖面上真实首现与末现时间进行了重新估算.结果显示，物种多样性的下降分为3个阶段，第1和第3阶段迅速下降，第2阶段缓慢下降.在此基础上，探讨以不同时间分辨率划分古群落的方案，从50 ka每个群落，不断提高时间分辨率，到1 ka每群落，分析各种划分方案下相邻群落物种组成的相似度.随着时间分辨率的提高，煤山剖面相邻群落的物种组成越来越像，对于1~4 ka每群落的方案，5%以上相邻群落在物种组成上完全一样.因此选取5 ka每群落的时间分辨率，构建了连续的古群落序列，量化评估了群落组成和模拟食物网稳定性/抗灾能力在P-Tr之交的演变过程.结果表明，生态结构以及群落稳定性在第1和第2阶段的变化不大，在第3阶段骤然变化.古群落在第1和第2阶段尚具备较强的抗灾能力，可能因为这两个阶段消失的主要为冗余物种，群落的生态结构和功能维持了完整，随着第3阶段多种功能群的消失，生态结构才完全崩溃重组.本文探讨了煤山古群落划分的不同时间分辨率方案，重建了5 ka分辨率下的古生态结构演变过程，为开展高分辨率深时生态系统演化及古今对比研究提供了新思路.Abstract: The Permian-Triassic (P-Tr) mass extinction, which occurred approximately 252 million years ago, represents the largest ecological crisis in Earth's geological history and is often regarded as a reference case for modern ecological crises. However, studies that integrate multiple taxa and systematically characterize the overall evolutionary trajectory of food web structures in communities remain scarce. In particular, the incompleteness of fossil records and their low temporal resolution within stratigraphic records pose significant challenges to using deep-time crises as analogs for modern ecological crises. This study focuses on the fossil-rich Meishan Section in Zhejiang Province, the global stratotype section and point (GSSP) for the P-Tr boundary. Using Bayesian modeling with the PyRate method, we recalculated the true first and last appearances of all fossil species within this section. The results reveal a three-phase decline in species diversity: rapid drops in the first and third phases, with a gradual decline in the second phase. Building on these results, we explored schemes for delineating paleocommunities at varying temporal resolutions, ranging from 50 ka per community down to 1 ka per community. We then analyzed species composition similarity between adjacent communities under each scheme. As the temporal resolution increases, adjacent communities exhibit increasingly similar species compositions. Notably, for resolutions of 1-4 ka years per community, over 5% of adjacent communities show identical species compositions. Adopting a temporal resolution of 5 ka per community, we constructed a continuous sequence of paleocommunities and quantitatively assessed the evolutionary dynamics of community composition, simulated food web stability, and resistance across the P-Tr boundary. The results indicate minimal changes in ecological structure and community stability during the first and second phases, followed by abrupt shifts in the third phase. Communities in the first and second phases retained relatively high resistance, likely due to the loss of redundant species that did not compromise the integrity of ecological structure and function. In contrast, the third phase witnessed the collapse and reorganization of ecological structures as multiple functional groups disappeared. This study provides a detailed examination of paleocommunity delineation at varying temporal resolutions and reconstructs the evolutionary trajectory of paleoecological structures at a resolution of 5 ka. These findings offer a novel approach for investigating high-resolution deep-time ecosystem evolution and facilitate comparative studies between ancient and modern ecological systems.
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Key words:
- mass extinction /
- ecological structure /
- paleocommunities /
- food webs /
- stability /
- ecosystems /
- stratigraphy
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图 1 煤山剖面原始化石出现数据(a)；煤山剖面不同地层的物种丰度分布(b)
图a中纵轴为物种编号，每个点代表某一物种在特定时间的出现记录；图b中，纵轴为地层编号，横轴为物种丰度
Fig. 1. Original fossil occurrence data of the Meishan section (a); species richness distribution of different strata in the Meishan section (b)
图 2 PyRate模拟估算的煤山物种延限(a)及对应物种多样性变化轨迹(b)
红色和紫色虚线标记251.94 Ma和251.88 Ma时间点
Fig. 2. The species durations estimated by PyRate simulations for the Meishan section (a) and the corresponding trajectory of species diversity changes (b)
图 3 不同时间分辨率的群落划分方案，及其对应的“物种组成相同的相邻群落”比例
Fig. 3. Community division schemes at different temporal resolutions and the corresponding proportion of adjacent communities with identical species compositions
图 4 煤山剖面252.00~251.75 Ma古群落序列划分
每个古群落时间延限为5 ka
Fig. 4. Paleocommunity sequence division from 252.00 to 251.75 Ma in the Meishan section
图 5 煤山剖面古群落物种组成演变分析
其中，C1至C12代表了252.00~251.94 Ma间的古群落，C13至C20代表了251.94~251.90 Ma间的古群落，C21至C31代表了251.90~251.85 Ma间的古群落，C32至C50代表了251.85~251.75 Ma间的古群落；每个点（古群落）的颜色值反映了其与周围点（古群落）的功能群组成差异，值越高说明与周围古群落功能群组成越相似，反之亦然
Fig. 5. Paleocommunity composition evolution in the Meishan section
图 6 煤山剖面古食物网的网络营养等级、最大食物链长度、连接密度和连通度的变化
Fig. 6. Changes in network trophic position, path max, link density, and connectance of Meishan food webs
图 7 煤山剖面古食物网模块数量与模块度的变化
Fig. 7. Variations of the number of modules and modularity of the Meishan food webs
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