Aim Climate warming reshuffles biological assemblages towards less cold‐adapted but more warm‐adapted species, a process coined thermophilization. However, the velocity at which this process is happening generally lags behind the velocity of climate change, generating a climatic debt the temporal dynamics of which remain misunderstood. Relying on high‐resolution time series of vegetation data from a long‐term monitoring network of permanent forest plots, we aim at quantifying the temporal dynamics – up to a yearly resolution – of the climatic debt in the understorey of temperate forests before identifying the key determinants that modulate it. Location France. Time period 1995–2017. Taxa studied Vascular plants. Methods We used the community temperature index (CTI) to produce a time series of understorey plant community thermophilization, which we subsequently compared to a time series of mean annual temperature changes over the same period and for the same sites. The direction and magnitude of the difference (i.e., the climatic debt) was finally analysed using linear mixed‐effect models to assess the relative contributions of abiotic and biotic determinants, including forest stand characteristics. Results We found a significant increase in CTI values over time (0.08–0.09 °C/decade), whereas the velocity of mean annual temperature changes was three times higher over the same period (0.22–0.28 °C/decade). Hence, the climatic debt increased over time and was greater in forest stands with higher basal area or older trees as well as under warmer macroclimate. By contrast, a greater frequency of anthropogenic disturbances decreased the climatic debt, while natural disturbances and herbivory had no impact. Conclusions Although often overlooked in understanding the climatic debt of forest biodiversity, changes in forest stand characteristics may modulate the climatic debt by locally modifying microclimatic conditions. Notably, the buffering effect of the upper canopy layer implies microclimate dynamics that may provide more time for understorey plant communities to locally adapt.

In this study we investigate the large and diverse Rubiaceae–Gardenieae and closely related tribes Bertiereae, Coffeeae, Cremasporeae, Octotropideae, and Pavetteae. Some of the tribes or groups have been shown to be monophyletic and strongly supported, but the phylogeny of this large complex is still far from being satisfactorily elucidated particularly for Gardenieae, both in terms of intertribal relationships as well as tribal delimitations. We reconstruct the phylogeny of the complex using an extensive sampling of 108 genera and five plastid DNA regions. Phylogenetic relationships demonstrate that Gardenieae sensu Andreasen & Bremer is polyphyletic, as Burchellia, Didymosalpinx, Monosalpinx, and Mantalania are closer to Octotropideae–Cremasporeae. In addition, Pavetteae and the investigated members of Aulacocalyceae are nested in a supported but partially unresolved Gardenieae–Pavetteae clade. Within this clade, several strongly supported groups are resolved: an Aidia group, an Alibertia group, a Gardenia group, Pavetteae including Pelagodendron, a Porterandia group, a Randia group, a Rothmannia group (including Aulacocalyx and Heinsenia), a Sherbournia group, and the two isolated genera Massularia and Schumanniophyton. The latter genus presented a high rate of genetic substitutions, which resulted in perturbations of the phylogenetic reconstruction. A revised tribal circumscription is given for Gardenieae, the Alibertia group is recognized at tribal level as an emended Cordiereae, and a new tribe, Sherbournieae, is described to accommodate the members of the Sherbournia group.