Forb ecology research in dry African savannas: Knowledge, gaps, and future perspectives.

Savannas are commonly described as a vegetation type with a grass layer interspersed with a discontinuous tree or shrub layer. On the contrary, forbs, a plant life form that can include any nongraminoid herbaceous vascular plant, are poorly represented in definitions of savannas worldwide. While forbs have been acknowledged as a diverse component of the herbaceous layer in savanna ecosystems and valued for the ecosystem services and functions they provide, they have not been the primary focus in most savanna vegetation studies. We performed a systematic review of scientific literature to establish the extent to which forbs are implicitly or explicitly considered as a discrete vegetation component in savanna research. The overall aims were to summarize knowledge on forb ecology, identify knowledge gaps, and derive new perspectives for savanna research and management with a special focus on arid and semiarid ecosystems in Africa. We synthesize and discuss our findings in the context of different overarching research themes: (a) functional organization and spatial patterning, (b) land degradation and range management, (c) conservation and reserve management, (d) resource use and forage patterning, and (e) germination and recruitment. Our results revealed biases in published research with respect to study origin (country coverage in Africa), climate (more semiarid than arid systems), spatial scale (more local than landscape scale), the level at which responses or resource potential was analyzed (primarily plant functional groups rather than species), and the focus on interactions between life forms (rather seldom between forbs and grasses and/or trees). We conclude that the understanding of African savanna community responses to drivers of global environmental change requires knowledge beyond interactions between trees and grasses only and beyond the plant functional group level. Despite multifaceted evidence of our current understanding of forbs in dry savannas, there appear to be knowledge gaps, specifically in linking drivers of environmental change to forb community responses. We therefore propose that more attention be given to forbs as an additional ecologically important plant life form in the conventional tree-grass paradigm of savannas.

The impact of livestock grazing on plant diversity: an analysis across dryland ecosystems and scales in southern Africa.

A general understanding of grazing effects on plant diversity in drylands is still missing, despite an extensive theoretical background. Cross-biome syntheses are hindered by the fact that the outcomes of disturbance studies are strongly affected by the choice of diversity measures, and the spatial and temporal scales of measurements. The aim of this study is to overcome these weaknesses by applying a wide range of diversity measures to a data set derived from identical sampling in three distinct ecosystems. We analyzed three fence-line contrasts (heavier vs. lighter grazing intensity), representing different degrees of aridity (from arid to semiarid) and precipitation regimes (summer rain vs. winter rain) in southern Africa. We tested the impact of grazing intensity on multiple aspects of plant diversity (species and functional group level, richness and evenness components, alpha and beta diversity, and composition) at two spatial scales, and for both 5-yr means and interannual variability. Heavier grazing reduced total plant cover and substantially altered the species and functional composition at all sites. However, a significant decrease in species alpha diversity was detected at only one of the three sites. By contrast, alpha diversity of plant functional groups responded consistently across ecosystems and scales, with a significant decrease at heavier grazing intensity. The cover-based measures of functional group diversity responded more sensitively and more consistently than functional group richness. Beta diversity of species and functional types increased under heavier grazing, showing that at larger scales, the heterogeneity of the community composition and the functional structure were increased. Heavier grazing mostly increased interannual variability of alpha diversity, while effects on beta diversity and cover were inconsistent. Our results suggest that species diversity alone may not adequately reflect the shifts in vegetation structure that occur in response to increased grazing intensity in the dryland biomes of southern Africa. Compositional and structural changes of the vegetation are better reflected by trait-based diversity measures. In particular, measures of plant functional diversity that include evenness represent a promising tool to detect and quantify disturbance effects on ecosystems.

How best to quantify soil seed banks in arid rangelands of the Nama Karoo?

Sampling design and three sample treatments prior the application of the seedling emergence method were tested in order to find the best method for seed bank quantification in arid Nama Karoo rangelands. I analyzed species composition and seed densities by contrasting undercanopy and open-matrix samples from two soil depths and by comparing the effects of cold-, heat-, and no stratification on germination rates of species in a greenhouse setting. The soil seed bank showed minimal similarity to the standing vegetation, with only 20 plant species germinated. Spatial distribution of seeds was highly heterogeneous. Nearly 90% of germinated seeds were located in 0- to 4-cm compared to >4- to 8-cm soil depth. Undercanopy seed banks contained significantly more species and seeds than open-matrix seed banks. Neither the number nor the diversity of seeds germinated differed significantly among the three treatments. Cold stratification tended to detect more species and seeds only at >4- to 8-cm soil depth. The results highlight the importance of spatial heterogeneity in the accurate evaluation of soil seed banks in the arid Nama Karoo and the importance of considering seasonal variability in the availability of readily germinable seeds. Data also suggest that sample pretreatment in germination trials may give little return for cost and effort, which emphasizes that it is more important to choose the sampling design most likely to give a representative number of seed bank species. Further studies are needed to analyze seed bank dynamics and species-specific germination requirements to promote recruitment of plant taxa underrepresented in the seed bank.