Shrubs are a characteristic component of savannas, where they coexist with trees and grasses. They are often part of woody encroachment phenomena, which have been observed globally, and the determinant of shrub encroachment cases, which are particularly of concern in African savannas. In response to climate change and land use change, African savannas are vulnerable to biome shifts and shrub encroachment is a process driving and explaining this risk. We contribute to furthering the understanding of shrubs biogeography and ecology by considering the number of stems of woody plants to characterise shrubs phenotype and strategy. We postulate that shrubs are multi-stemmed, compared to single-stemmed trees and integrate this assumption in aDGVM2 (adaptive Dynamic Global Vegetation Model 2). Modelling a trait representing the number of stems of a woody plant implies a trade-off between single-stemmed plants having higher height growth potential and multi-stemmed plants having higher hydraulic capacity but limited height growth. Multi-stemmed individuals, being shorter, are more likely to suffer severe damage from fires than tall single-stemmed trees managing to grow their crown out of the flame zone. We simulate potential vegetation over sub-Saharan Africa at 1° spatial resolution, with aDGVM2 and compare it to simulations without our shrub model turned on. We also test the impact of fire by including or excluding it from our simulations. To assess the accuracy and relevance of our approach, we benchmark our overall model’s performance against multiple satellite derived products of above ground biomass (AGBM), and against specific field measurements of AGBM. We further benchmark our results against vegetation cover type derived from satellite data. We demonstrate that shrubs can be modelled as multi-stemmed woody plants in African savannas based on whole-plant trait trade-off without being predefined as static functional types. Indeed, the addition of our shrub model to aDGVM2 allows for shrubs to emerge dynamically through community assembly processes without a priori categorisation. Our shrub model also improves the simulated vegetation patterns simulated by aDGVM2 in sub-Saharan Africa, particularly in savannas. The simulated pattern of stem number per woody individual broadly follows our assumptions about biogeographic patterns as it is lowest in equatorial African forests and increases in savannas and grasslands as precipitation decreases. Shrubs are more abundant in more water-stressed regions where they have a competitive advantage over trees due to their increased relative water transport potential. However, in arid and hyper-arid regions, further investigations are required. Simulated shrub prevalence is higher in more open and fire prone landscapes, where woody cover and biomass are reduced. Adding shrubs to aDGVM2, while increasing complexity allows for greater simulated diversity. As resilience and resistance of ecosystems have been shown to be influenced by diversity, such model development is necessary to improve our ability to forecast ecosystems responses to changes. However, there are challenges to fully tap this benefit. Assessing the accuracy and relevance of our approach is challenging. Data and simulations are conceptually different which limit the possibility to conclude based on comparison. Benchmarking challenge is exacerbated by the variability existing among satellite derived products and site studies observations. In areas of extremely low biomass and vegetation cover, such as deserts and semi-deserts, the accuracy of our model is more concerning as small differences in absolute values are relatively more important. Categorisation of life-forms shapes our understanding of their ecology and biogeography, thus, consensus about their definition is direly needed. To contribute to this debate, we investigate how vegetation distribution patterns arising from our shrub model inform our understanding of shrub biogeography. First, shrub distribution in trait space (considering stem number), relatively to environmental drivers, concurs with our assumptions. Second, shrub spatial distribution is consistent with our characterisation assumptions. Third, the role of simulated shrubs in an ecosystem supports realistic ecological dynamics. Our model allows for, shrubs to exhibit a specific phenotype, but also a specific life-strategy, which we characterise in terms of persistence strategy (shrubs are mainly resprouters, in contrast to trees, which can be either resprouters or reseeders) and in terms of resource acquisition (rooting strategy) and allocation (carbon investment). Adding stem count as a trait to aDGVM2 increase the range of simulated functional diversity. Our shrub model allows for aDGVM2 to simulate realistic ratio of grass to woody vegetation across sub-saharan Africa. Similarly, it simulates ratio of shrubs to trees consistent with our hypotheses. ...
Status of the Amargosa niterwort (Amaranthaceae) in California and Nevada
