16Managing forests to preserve biodiversity requires a good knowledge not only of the factors 17 driving its dynamics but also of the structural elements that actually support biodiversity. Tree-18 related microhabitats (cavities, cracks, conks of fungi) are tree-borne features that are reputed 19 to support specific biodiversity for at least a part of species' life cycles. While several studies 20 have analysed the drivers of microhabitats number and occurrence at the tree scale, they 21 remain limited to a few tree species located in relatively narrow biogeographical ranges. We 22Promoting large living and dead trees of several tree species may be an interesting, and nearly 36 universal, way to favour microhabitats and enhance the substrates needed to support specific 37 biodiversity. In addition, a better understanding of microhabitat drivers and dynamics at the 38 tree scale may help to better define their role as biodiversity indicators for large-scale 39 monitoring. 40 41 42 Small natural features are structural habitat elements that have a disproportionately important 43 role for biodiversity related to their actual size [1]. Taking these features into account in 44 biodiversity conservation strategies is a crucial step in science-based decision making [2]. 45Identifying such structural features in a tri-dimensional forest environment is quite challenging 46 since their number and variety is potentially infinite. Small natural features include, for example, 47 large old trees [3] as well as tree-borne structures. While large old trees are disappearing at 48 the global scale [4], their importance for biodiversity has not yet been fully elucidated, not to 49 mention the peculiar structures they may bear (eg. cracks, cavities, epiphytes), also known as 50 'tree-related microhabitats' (hereafter 'microhabitats' [5]). Microhabitats have recently aroused 51 the interest of scientists and forest managers alike since these structures can be a substrate 52for specific forest biodiversity [6], and can ultimately serve as forest biodiversity indicators [5, 53 7, 8]. Their conservation has hence become an issue in day-to-day forest management, as 54 have large old trees and deadwood [9, 10]. However, our understanding of the drivers and 55 dynamics influencing these microhabitats, notably at the tree scale, remains incomplete [11]. 56Tree diameter and living status (living vs. dead trees) are key factors for microhabitat diversity 57 at the tree scale [12][13][14]. Larger trees are likely to bear more microhabitats than smaller ones, 58as they have experienced more damage, injuries and microhabitat-creating events (e.g. 59 woodpecker excavation, storms, snowfalls). Similarly, gradually decomposing dead trees are 60 likely to bear more microhabitats than living trees and play a role as habitat and food sources 61 for many microhabitat-creating species [15]. Nevertheless, the relationships between 62 microhabitats and tree characteristics have only been demonstrated on a limited number of 63 tree species involving at...