Abstract Determining the extrinsic (physical) factors controlling speciation and diversification of species through time is of key interest in paleontology and evolutionary biology. The role of sea-level change in shaping species richness patterns of marginal marine species has received much attention, but with variable conclusions. Recent work combining genetic data and Geographical Information Systems (GIS)-based habitat modeling yielded a framework for how geomorphology of continental margins mediates genetic connectivity of populations during sea-level change. This approach may ultimately yield insights on how distinct lineages, species, and biodiversity accumulate in coastal settings. Here, we expand this GIS work globally to different geomorphic settings to model estuarine habitat in a larger geographic framework and test how tectonic setting, oceanographic setting, climate, and margin age affect habitat distribution during sea-level change. In addition, independent of estuaries we explore paleobiologic (e.g. Olsson, 1961) and neontolologic effects of sea-level change on evolution, and test the relation between overall shelf area and species richness using data of 1721 fish species. We find 82% global reduction of estuarine habitat abundance at lowstand relative to highstand, and find large habitats change in size much more than small habitats. Consistent with prior work, narrow continental margins have significantly less habitat at highstand and lowstand than wide margins, and narrow margins significantly associate with fore-arc settings, effectively linking tectonic setting to habitat abundance. Surprisingly, narrow margins host greater species richness, a finding which violates the canonical species-area relation. This finding can be explained if: 1) the physical isolation imposed by narrow margins facilitates the formation of new species over time; 2) the size-stability of small habitats, which disproportionately occur on narrow margins, accumulate and retain species extirpated in the more variable habitats on wide margins; or 3) the smaller habitats on narrow margins facilitate greater species richness through greater habitat heterogeneity. These results are generally at odds with prior interpretations, but the combination of richness data and population genetic principles offer a different perspective on these long-studied questions. Finally, we emphasize that the nuance of Pleistocene-Holocene sea level oscillations should be more explicitly considered in genetic studies.

中文翻译：

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海平面变化期间河口栖息地分布的全球物理控制：随着时间的推移遗传多样化的后果

摘要 确定随时间控制物种形成和多样化的外在（物理）因素是古生物学和进化生物学的关键兴趣所在。海平面变化在塑造边缘海洋物种物种丰富度模式中的作用受到了广泛关注，但结论不一。最近的工作结合了遗传数据和基于地理信息系统 (GIS) 的栖息地建模，为大陆边缘的地貌如何在海平面变化期间调节种群的遗传连通性提供了一个框架。这种方法最终可能会深入了解不同的谱系、物种和生物多样性如何在沿海环境中积累。这里，我们将这项 GIS 工作在全球范围内扩展到不同的地貌环境，以在更大的地理框架中模拟河口栖息地，并测试构造环境、海洋环境、气候和边缘年龄如何影响海平面变化期间的栖息地分布。此外，独立于河口，我们探索了海平面变化对进化的古生物学（例如 Olsson，1961）和新生动物学的影响，并使用 1721 种鱼类的数据测试了整个大陆架面积与物种丰富度之间的关系。我们发现低位相对于高位的河口栖息地数量在全球范围内减少了 82%，并且发现大栖息地的大小变化比小栖息地大得多。与先前的工作一致，狭窄的大陆边缘在高位和低位的栖息地明显少于宽边缘，和狭窄的边缘与前弧环境显着相关，有效地将构造环境与栖息地丰度联系起来。令人惊讶的是，狭窄的边缘拥有更大的物种丰富度，这一发现违反了典型的物种-面积关系。这一发现可以在以下情况下得到解释：1) 狭隘边缘强加的物理隔离促进了新物种的形成；2) 不成比例地出现在狭窄边缘的小型栖息地的大小稳定性，积累并保留了在广阔边缘的变化更大的栖息地中灭绝的物种；或 3) 狭窄边缘的较小栖息地通过更大的栖息地异质性促进了更大的物种丰富度。这些结果通常与先前的解释不一致，但是丰富度数据和种群遗传原理的结合为这些长期研究的问题提供了不同的视角。最后，我们强调在遗传研究中应该更明确地考虑更新世-全新世海平面振荡的细微差别。