Species distribution models have grown in complexity by incorporating fine-scale variables, including data on microclimate, physiology and species interactions. Recent studies have acknowledged the importance of the spatial scale by including higher resolution maps and more complex climatic variables. However, models rarely consider the consequences of including data related to time. Indeed, species phenology – and potential shifts in phenology due, for example, to climate change – is potentially one of the most neglected aspects of ecological modelling. We present a literature review of relevant phenological aspects at different temporal scales and across several taxa. Such elements should be considered to define better the environmental niche and project present, future and past distribution models. We considered the available studies on plants, insects, reptiles, birds and mammals to evaluate how they dealt with the phenology of the investigated species, as well as the phenology of other resources and interacting species, to infer present, past and future projections. Here we focus on four main phenological aspects that, if not considered, may easily bias any projection, namely: 1) phenology can be accompanied by a shift in distribution within the year (e.g. migratory species); 2) activity may be restricted to a portion of the year (e.g. most ectotherms from temperate climates); 3) survival and reproduction success may depend on the synchrony with other species phenology (e.g. plants–pollinators interactions); 4) changes in climatic conditions can lead to shifts in phenology (e.g. anticipated or delayed blooms or changes in migration timing). In this review, we show how neglecting such factors may quickly lead to project a biased distribution. Finally, we provide a guide on evaluating whether the case study may be affected by such factors and what actions may improve the models.

中文翻译：

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在模拟物种生态位时包括物候学的重要性

通过纳入精细尺度变量，包括微气候、生理学和物种相互作用的数据，物种分布模型变得越来越复杂。最近的研究通过包括更高分辨率的地图和更复杂的气候变量，承认了空间尺度的重要性。然而，模型很少考虑包含与时间相关的数据的后果。事实上，物种物候学——以及气候变化等导致的物候学潜在变化——可能是生态建模中最被忽视的方面之一。我们对不同时间尺度和多个分类群的相关物候方面进行了文献综述。应考虑这些要素以更好地定义环境利基和项目现在、未来和过去的分布模型。我们考虑了现有的植物研究，昆虫、爬行动物、鸟类和哺乳动物，以评估它们如何处理被调查物种的物候，以及其他资源和相互作用物种的物候，以推断现在、过去和未来的预测。在这里，我们关注四个主要的物候学方面，如果不考虑这些方面，可能很容易使任何预测产生偏差，即：1）物候学可能伴随着一年内分布的变化（例如迁徙物种）；2) 活动可能仅限于一年中的部分时间（例如来自温带气候的大多数变温动物）；3) 生存和繁殖的成功可能取决于与其他物种物候的同步性（例如植物与传粉者的相互作用）；4) 气候条件的变化会导致物候变化（例如预期或延迟的开花或迁徙时间的变化）。在这篇评论中，我们展示了忽略这些因素如何可能很快导致预测有偏差的分布。最后，我们提供了一个指南，用于评估案例研究是否会受到这些因素的影响，以及采取哪些行动可以改进模型。