Pleistocene climate fluctuations had profound influence on the biogeographical history of many biota. As large areas in high mountain ranges were covered by glaciers, biota were forced either to peripheral refugia (and possibly beyond to lowland refugia) or to interior refugia (nunataks). However, nunatak survival remains controversial as it relies solely on correlative genetic evidence. Here, we test hypotheses of glacial survival using two high alpine plant species (the insect-pollinated Pedicularis asplenifolia and wind-pollinated Carex fuliginosa) in the European Alps. Employing the iDDC (integrative Distributional, Demographic and Coalescent) approach, which couples species distribution modelling, spatial and temporal demographic simulation and Approximate Bayesian Computation, we explicitly test three hypotheses of glacial survival: (a) peripheral survival only, (b) nunatak survival only and (c) peripheral plus nunatak survival. In P. asplenifolia the peripheral plus nunatak survival hypothesis was supported by Bayes factors (BF> 100), whereas in C. fuliginosa the peripheral survival only hypothesis, although best supported, could not be unambiguously distinguished from the peripheral plus nunatak survival hypothesis (BF = 5.58). These results are consistent with current habitat preferences (P. asplenifolia extends to higher elevations) and the potential for genetic swamping (i.e., replacement of local genotypes via hybridization with immigrating genotypes [expected to be higher in the wind-pollinated C. fuliginosa]). Although the persistence of plants on nunataks during glacial periods has been debated and studied over decades, this is one of the first studies to explicitly test the hypothesis instead of solely using correlative evidence.

中文翻译：

---

在两个高山植物物种的外围与努纳塔克族避难所中更新世存活的显式测试。

更新世的气候波动对许多生物区系的生物地理历史产生了深远的影响。由于高山上的大片区域被冰川覆盖，因此生物群被迫要么进入外围避难所（甚至可能越过低地避难所），要么被迫流向内部避难所（努纳塔克人）。但是，努纳塔克人的生存仍然有争议，因为它仅依赖于相关的遗传证据。在这里，我们在欧洲阿尔卑斯山使用两种高寒植物物种（昆虫授粉的小孢子虫和风生的Carex fuliginosa）测试了冰川生存的假设。利用结合物种分布模型，时空人口模拟和近似贝叶斯计算的iDDC（综合分布，人口统计学和聚结）方法，我们明确检验了冰河生存的三个假设：（a）仅周边生存，（b）仅努纳塔克生存，以及（c）周边加努纳塔克生存。在P. asplenifolia中，贝叶斯因子（BF> 100）支持外围加努纳克生存假设（BF> 100），而在C. fuliginosa中，只有最好的外围生存假说与外围加Nunatak生存假说（BF）明确区分。 = 5.58）。这些结果与当前的栖息地偏好（P. asplenifolia延伸到更高的海拔）和遗传沼泽的潜力（即，通过与移民基因型杂交[替换为风传粉红色的C. fuliginosa预计会更高]来取代局部基因型）保持一致。 。尽管数十年来人们一直在辩论和研究冰河时期植物在小菜蛾上的持久性，