The establishment of permanent animal communities on land was a defining event in the history of evolution, and one for which the ichnofauna and facies of the Tumblagooda Sandstone of Western Australia have been considered an archetypal case study. However, terrestrialization can only be understood from the rock record with conclusive sedimentological evidence for non-marine deposition, and original fieldwork on the formation shows that a marine influence was pervasive throughout all trace fossil-bearing strata. Four distinct facies associations are described, deposited in fluvial, tidal and estuarine settings. Here we explain the controversies surrounding the age and depositional environment of the Tumblagooda Sandstone, many of which have arisen due to the challenges in distinguishing marine from non-marine depositional settings in lower Palaeozoic successions. We clarify the terminological inconsistency that has hindered such determination, and demonstrate how palaeoenvironmental explanations can be expanded out from unambiguously indicative sedimentary structures. The Tumblagooda Sandstone provides a unique insight into an early Palaeozoic ichnofauna that was strongly partitioned by patchy resource distribution in a littoral setting. The influence of outcrop style and quality is accounted for to contextualize this ichnofauna, revealing six distinct low-disparity groups of trace fossil associations, each related to a different sub-environment within the high-ichnodisparity broad depositional setting. The formation is compared with contemporaneous ichnofaunas to examine its continued significance to understanding the terrestrialization process. Despite not recording permanent non-marine communities, the Tumblagooda Sandstone provides a detailed picture of the realm left behind by the first invertebrate pioneers of terrestrialization.

中文翻译：

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重新审视 Tumblagooda 砂岩：异常丰富的微量化石和地质露头为无脊椎动物陆生化之前的古生代沿海栖息地提供了一个窗口

在陆地上建立永久性动物群落是进化史上的一个决定性事件，西澳大利亚 Tumblagooda 砂岩的动物群落和相被认为是一个典型的案例研究。然而，陆地化只能从具有非海洋沉积确凿沉积学证据的岩石记录中理解，并且对地层的原始实地工作表明，海洋影响在所有微量含化石地层中普遍存在。描述了四种不同的相组合，沉积在河流、潮汐和河口环境中。在这里，我们解释了围绕 Tumblagooda 砂岩的年龄和沉积环境的争议，其中许多是由于在下古生代演替中区分海洋和非海洋沉积环境的挑战而产生的。我们澄清了阻碍这种确定的术语不一致，并展示了如何从明确指示的沉积结构中扩展古环境解释。Tumblagooda 砂岩提供了对早期古生代鱼类动物群的独特见解，该动物群在沿海环境中被不规则的资源分布强烈划分。露头样式和质量的影响被解释为这种 ichnofauna 的背景，揭示了六个不同的低差异微量化石关联组，每个组都与高 ichnodisparity 广泛沉积环境中的不同亚环境有关。将该地层与同时代的动物群进行比较，以检验其对理解陆地化过程的持续意义。尽管没有记录永久性的非海洋社区，Tumblagooda 砂岩提供了第一批陆地化无脊椎动物先驱留下的领域的详细图片。