Abstract The late Olenekian to early Ladinian marine carbonates of the epicontinental European (Germanic) Basin accumulated during the several-million-year-long biotic recovery following the Permian/Triassic (P/Tr) mass extinction. In this paper, we discuss the environmental and evolutionary controls on the temporal ichnodiversity and ichnofabric changes within Anisian open-marine limestones (called Muschelkalk) deposited in the south-eastern margin of the basin, close to the Tethys Ocean, and their implications for the timing of biotic recovery. The focus is on the lithologically similar transgressive systems tracts (TST) of three consecutive 3rd-order depositional sequences, each of which start with high-energy shoal-sand packstones-grainstones overlain by deeper-platform lime mudstones with skeletal and conglomeratic tempestites. In general, these facies are characterized by a high degree of bioturbation, high ichnotaxonomical diversity (20 softground ichnotaxa, 2 firmground ichnotaxa, 3 hardground ichnotaxa, and undetermined mottles and spots in a soupground suite), and high ethological diversity (domichnia, fodinichnia, cubichnia, repichnia, pascichnia, praedichnia). Despite the high overall diversity, monoichnospecific and almost monoichnospecific assemblages of Rhizocorallium, Oravaichnium, and Thalassinoides prevail. With these assemblages dominance changes stratigraphically, from Rhizocorallium dominated assemblages in TST1, to Oravaichnium dominated in TST2, and Thalassinoides dominated in TST3. Concurrently, the width of Thalassinoides burrows increases, from 1 to 2 cm in TST1 and TST2, to 3–7 cm in the TST3. The two secular changes may represent a gradual recovery of burrowing organisms following the P/Tr biotic crisis, given that environmental conditions remained more-or-less stable through time, as evidenced by facies analysis and stable carbon and oxygen isotope data. The occurrence of large Rhizocorallium and medium-sized Thalassinoides represents the penultimate stage in recovery, whereas the widespread appearance of large Thalassinoides – comparable in size with typical pre-extinction Permian forms – marks the final recovery of trace-maker communities.

中文翻译：

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来自波兰南部上西里西亚的日耳曼盆地东南部（Wellenkalk 相）的早中三叠世（阿尼阶）踪迹化石、地层结构和基质类型：二叠纪/三叠纪大灭绝后生物恢复的意义

摘要 在二叠纪/三叠纪 (P/Tr) 大灭绝之后的数百万年的生物恢复过程中，欧列克纪晚期至拉丁纪早期海相碳酸盐岩积累起来。在本文中，我们讨论了沉积在盆地东南缘靠近特提斯洋的 Anisian 开放海相石灰岩（称为 Muschelkalk）内的时间地层多样性和地层结构变化的环境和演化控制，以及它们对生物恢复的时间。重点是三个连续三级沉积序列的岩性相似的海侵系统域 (TST)，每个序列都以高能浅滩-砂砾岩-颗粒岩开始，上面覆盖着具有骨骼和砾岩风暴岩的更深台地灰泥岩。一般来说，这些相的特点是高度的生物扰动、高度的鱼类分类多样性（20 个软地鱼类、2 个硬地鱼类、3 个硬地鱼类，以及汤地套房中未确定的斑点和斑点）和高度的生态多样性（domichnia、fodinichnia、cuvichnia、repichnia , pascichnia, praedichnia)。尽管总体多样性很高，但Rhizocorallium、Oravaichnium 和 Thalassinoides 的单鱼种和几乎单鱼种组合占优势。随着这些组合在地层上的优势发生变化，从 TST1 中的根珊瑚主导组合，到 TST2 中的 Oravaichnium 主导，TST3 中的 Thalassinoides 主导组合。同时，Thalassinoides 洞穴的宽度增加，从 TST1 和 TST2 的 1 到 2 厘米，到 TST3 的 3-7 厘米。考虑到环境条件随着时间的推移或多或少保持稳定，这两种长期变化可能代表着穴居生物在 P/Tr 生物危机之后逐渐恢复，正如相分析和稳定的碳和氧同位素数据所证明的那样。大型根珊瑚和中型海藻的出现代表了恢复的倒数第二阶段，而大型海藻的广泛出现——其大小与典型的灭绝前二叠纪形态相当——标志着示踪者群落的最终恢复。