Ecol Appl. 2015 Jul;25(5):1303-18. doi: 10.1890/14-1462.1.
Dynamic models are pivotal for projecting forest dynamics in a changing climate, from the local to the global scale. They encapsulate the processes of tree population dynamics with varying resolution. Yet, almost invariably, tree mortality is modeled based on simple, theoretical assumptions that lack a physiological and/or empirical basis. Although this has been widely criticized and a growing number of empirically derived alternatives are available, they have not been tested systematically in models of forest dynamics. We implemented an inventory-based and a tree-ring-based mortality routine in the forest gap model ForClim v3.0. We combined these routines with a stochastic and a deterministic approach for the determination of tree status (alive vs. dead). We tested the four new model versions for two Norway spruce forests in the Swiss Alps, one of which was managed (inventory time series spanning 72 years) and the other was unmanaged (41 years). Furthermore, we ran long-term simulations (-400 years) into the future under three climate scenarios to test model behavior under changing environmental conditions. The tests against inventory data showed an excellent match of simulated basal area and stem numbers at the managed site and a fair agreement at the unmanaged site for three of the four empirical mortality models, thus rendering the choice of one particular model difficult. However, long-term simulations under current climate revealed very different behavior of the mortality models in terms of simulated changes of basal area and stem numbers, both in timing and magnitude, thus indicating high sensitivity of simulated forest dynamics to assumptions on tree mortality. Our results underpin the potential of using empirical mortality routines in forest gap models. However, further tests are needed that span other climatic conditions and mixed forests. Short-term simulations to benchmark model behavior against empirical data are insufficient; long-term tests are needed that include both nonequilibrium and equilibrium conditions. Thus, there is the potential to greatly improve the robustness of future projections of forest dynamics via more reliable tree mortality submodels.
动态模型对于从局部到全球尺度预测气候变化下的森林动态至关重要。它们以不同的分辨率来概括树木种群动态的过程。然而,几乎无一例外地,树木死亡率的建模都是基于缺乏生理和/或经验基础的简单理论假设。尽管这已经受到了广泛的批评,并且有越来越多的经验衍生替代方案可供选择,但它们尚未在森林动态模型中得到系统的测试。我们在森林空隙模型 ForClim v3.0 中实现了基于清单和树木年轮的死亡率例程。我们将这些例程与用于确定树木状态(存活或死亡)的随机和确定性方法相结合。我们针对瑞士阿尔卑斯山的两片挪威云杉林测试了这四个新模型版本,其中一个是经过管理的(包含 72 年的清单时间序列),另一个是未管理的(持续了 41 年)。此外,我们在三个气候情景下进行了长期模拟(-400 年),以测试模型在不断变化的环境条件下的行为。与清单数据的对比测试表明,在管理站点上,模拟的基面积和茎数与实际数据非常吻合,在未管理站点上,四个经验死亡率模型中有三个模型的拟合效果也很好,因此很难选择一个特定的模型。然而,在当前气候下的长期模拟显示,在模拟基面积和茎数的变化方面,死亡率模型的行为存在很大差异,无论是在时间还是幅度上,这表明模拟森林动态对树木死亡率假设非常敏感。我们的结果支持在森林空隙模型中使用经验死亡率例程的潜力。然而,还需要进一步的测试,涵盖其他气候条件和混合林。短期模拟来根据经验数据对模型行为进行基准测试是不够的;需要进行长期测试,包括非平衡和平衡条件。因此,通过更可靠的树木死亡率子模型,可以极大地提高森林动态未来预测的稳健性。