Biofuels and invasive species from an African perspective – a review

A large number of proposed biofuel crops share the same traits as known invasive plant species, many of which are already present in Africa and include species such as Prosopis glandulosa Torrey (Mimosaceae), P. juliflora (Sw.) DC, Leucaena leucocephala (Lam.) de Wit (Mimosaceae), Azadirachta indica A. Juss. (Meliaceae), and others. In this paper, we mainly assess the impacts of invasive Prosopis species in Africa, particularly in Kenya and South Africa. Introduced Prosopis species have invaded over 4 million hectares in Africa, threatening crop and pasture production, reducing underground water reserves, and displacing native plant and animal species. This has major implications for millions of people who depend on natural resources for their survival. It is therefore suggested that known invasive or potentially invasive plant species not be introduced to countries or regions for biofuel production. If (after a stringent cost–benefit analysis) the introduction of a potentially invasive species is deemed critical for economic development and the benefits clearly outweigh the potential costs, countries should endeavour to abide by the Principles and Criteria for Sustainable Biofuel Production developed by the Roundtable on Sustainable Biofuels.     

Disentangling facilitation and seed dispersal from environmental heterogeneity as mechanisms generating associations between savanna plants

Question: How can we disentangle facilitation and seed dispersal from environmental heterogeneity as mechanisms causing spatial associations of plant species? Location: Semi‐arid savanna in the Kimberley Thorn Bushveld, South Africa. Methods: We developed a two‐step protocol for the statistical differentiation of association‐promoting mechanisms in plants based on the Acacia erioloba–Grewia flava association. Individuals of the savanna shrub G. flava and the tree A. erioloba were mapped on four study plots. Disentangling the mechanism causing the association of G. flava and A. erioloba involved tests of three spatial and one non‐spatial null model. The spatial null models include homogeneous and heterogeneous Poisson processes for spatial randomness based on the bivariate spatial point patterns of the four plots. With the non‐spatial analysis, we determined the relationship between the canopy diameter of A. erioloba trees and presence or absence of G. flava shrubs in the tree understorey to find whether shrub presence requires a minimum tree canopy diameter. Results: We first showed a significant positive spatial association of the two species. Thereafter, the non‐spatial analysis supported an exclusion of environmental heterogeneity as the sole cause of this positive association. We found a minimum tree size under which no G. flava shrubs occurred. Conclusions: Our two‐step analysis showed that it is unlikely that heterogeneous environmental conditions caused the spatial association of A. erioloba and G. flava. Instead, this association may have been caused by seed dispersal and/or facilitation (e.g. caused by hydraulic lift and/or nitrogen fixation by the host tree).     

Spatial and temporal litterfall heterogeneity generated by woody species in the Central Monte desert

In arid and semiarid environments, the presence of woody species generates a series of environmental gradients that increase spatial heterogeneity and modify the pattern of distribution of the other species. We postulate that the temporal and spatial variability in litter input generated by woody species is a relevant factor in the generation of edaphic heterogeneity by redistribution of nutrients and the physical effects of litter. The objective of this study was to determine the temporal and spatial variability in the amount of litter input under the canopy of dominant woody plants (Prosopis flexuosa and Larrea divaricata) and in exposed areas at the Ñacuñán Reserve, in the central zone of the Monte desert. Litterfall was collected during 2 years from 30-cm-diameter litter traps distributed at three microsites: under P. flexuosa canopy, under L. divaricata canopy, and in exposed areas. Microhabitats beneath Prosopis showed the highest litter input per m² (between 320 and 527 g/m²), and, consequently, more than 50% of it fell to the soil beneath the canopy of P. flexuosa. Only 10% fell on exposed areas, which exhibited an annual input rate per m² of a lower order of magnitude than the sites under Prosopis. Litterfall presented a peak in summer as a consequence of convective storms, and a second one in autumn due to phenological shedding. Our results suggest that woody species have a central importance in the dynamics of nutrients in arid lands by both the increase of total productivity and litterfall, and the spatial and temporal regulation of litter input.     

Long‐term mortality patterns of the deep‐rooted Acacia erioloba: The middle class shall die!

Question: Is there a relationship between size and death in the long‐lived, deep‐rooted tree, Acacia erioloba, in a semi‐arid savanna? What is the size‐class distribution of A. erioloba mortality? Does the mortality distribution differ from total tree size distribution? Does A. erioloba mortality distribution match the mortality distributions recorded thus far in other environments? Location: Dronfield Ranch, near Kimberley, Kalahari, South Africa. Methods: A combination of aerial photographs and a satellite image covering 61 year was used to provide long‐term spatial data on mortality. We used aerial photographs of the study area from 1940, 1964, 1984, 1993 and a satellite image from 2001 to follow three plots covering 510 ha. We were able to identify and individually follow ca. 3000 individual trees from 1940 till 2001. Results: The total number of trees increased over time. No relationship between total number of trees and mean tree size was detected. There were no trends over time in total number of deaths per plot or in size distributions of dead trees. Kolmogorov‐Smirnov tests showed no differences in size class distributions for living trees through time. The size distribution of dead trees was significantly different from the size distribution of all trees present on the plots. Overall, the number of dead trees was low in small size classes, reached a peak value when canopy area was 20 ‐ 30 m², and declined in larger size‐classes. Mortality as a ratio of dead vs. total trees peaked at intermediate canopy sizes too. Conclusion: A. erioloba mortality was size‐dependent, peaking at intermediate sizes. The mortality distribution differs from all other tree mortality distributions recorded thus far. We suggest that a possible mechanism for this unusual mortality distribution is intraspecific competition for water in this semi‐arid environment.     

Classification of Prosopis juliflora invasion in the Lake Baringo basin and environmental correlations

The spread of Prosopis juliflora in the Baringo basin, Kenya, has led to severe changes in the ecosystem with negative socio‐economic impacts. The drivers that foster the invasiveness of Prosopis are not fully understood. Thus, a method to quantify the degree of infestation will support the determination of environmental preferences and the risk assessment of future Prosopis invasion. We developed a methodology for characterising and classifying degrees of Prosopis infestation in vegetation stands and propose its application in environmental correlation models. The relative cover was identified as the most suited attribute for assessing and monitoring the invasion of Prosopis. The distance of invaded stands from original plantations and environmental attributes related to water availability (ground water table, rainfall and soil water–holding capacity) have potential to predict potential or future invasion risks.     

Does Eucalyptus grandis invasion and removal affect soils and vegetation in the Eastern Cape Province,South Africa?

Many invasive alien plants drive changes in native community composition, structure and diversity. They alter soil nutrient regimes of native communities and affect native plant recovery outcomes following their removal. We assessed whether Eucalyptus grandis invasion and removal alters the soil physico‐chemical properties and native vegetation recovery in the Eastern Cape Province, South Africa. We collected samples from topsoil in E. grandis invaded sites (canopy cover > 75%), cleared sites (eight years ago) and native sites (canopy cover > 80%) and quantified soil moisture, concentrations of soil macro elements (N, C and P), pH and exchangeable cations (K, Ca, Mg, Na) as well as measured soil water repellency using the Water Droplet Penetration Time and infiltration. We conducted vegetation surveys in plots measuring 10 × 10 m. Invasion by E. grandis had varying effects on soil physico‐chemical properties, causing increase in soil pH and P, while decreasing total N and C. The removal of E. grandis also showed varying effects on soil physico‐chemical properties, but seems to have further triggered the loss of some soil nutrients (especially soil P). Soil water repellency (a measure of soil compaction) has improved in cleared sites to non‐repellent soils compared to repellent soils in invaded site. Eucalyptus grandis reduced species richness of the invaded sites. The presence of native species on cleared sites indicates a positive trajectory towards vegetation recovery. We conclude that E. grandis invasion and removal trigger varying effects on soil properties (both increases and decreases). For soil and vegetation restoration of cleared sites to be effective, active restoration techniques such as soil transfer, nutrient manipulation and native plant seeding should be considered.     

RAPD and microsatellite transferability studies in selected species of <Emphasis Type="Italic">Prosopis</Emphasis> (section Algarobia) with emphasis on <Emphasis Type="Italic">Prosopis juliflora</Emphasis> and <Emphasis Type="Italic">P. pallida</Emphasis>

The genus Prosopis (Leguminosae, Mimosoideae), comprises 44 species widely distributed in arid and semi-arid zones. Prosopis pallida (Humb. & Bonpl. ex Willd.) Kunth and P. juliflora (Sw.) DC. are the two species that are truly tropical apart from P. africana, which is native to tropical Africa (Pasiecznik et al. 2004), and they have been introduced widely beyond their native ranges. However, taxonomic confusion within the genus has hampered exploitation and better management of the species. The present study focusses primarily on evaluating the genetic relationship between Prosopis species from the section Algarobia, containing most species of economic importance, though P. tamarugo from section Strombocarpa is also included for comparison. In total, 12 Prosopis species and a putative P. pallida × P. chilensis hybrid were assessed for their genetic relationships based on RAPD markers and microsatellite transferability. The results show that P. pallida and P. juliflora are not closely related despite some morphological similarity. Evidence also agrees with previous studies which suggest that the grouping of series in section Algarobia is artificial.     

Development and characterization of microsatellite markers for Prosopis chilensis and Prosopis flexuosa and cross‐species amplification

Prosopis chilensis and Prosopis flexuosa (Fabaceae) are closely related hardwood arboreal species that are widely distributed in the arid regions of Argentina. The development of highly polymorphic markers, such as microsatellites, is desirable for genetic studies of these species. Here, we present the development and characterization of six polymorphic microsatellite markers in P. chilensis and P. flexuosa. These markers showed a polymorphism information content between 0.14 and 0.85 and the number of alleles varied from two to 13 considering both species. All markers revealed a broad cross‐species affinity when tested in seven other Prosopis species. All primers amplified in at least five species.     

Identification of Prosopis juliflora and Prosopis pallida Accessions Using Molecular Markers

There has been much taxonomic confusion over the identification of Prosopis species, especially where introduced. Prosopis juliflora is the most widespread species in the arid and semi-arid tropics, although it has been confused with other species, particularly the closely related Prosopis pallida. In this study, RAPDs markers were used for the first time to distinguish between these species. Eighteen primers were used in amplification reactions, which yielded an average of 120 bands per accession. A dendrogram showing genetic similarities among accessions was constructed using UPGMA cluster analysis and the Nei and Li similarity coefficient. The genetic similarity observed between P. juliflora and P. pallida is similar to the value in sympatric Prosopis species in North America, and reconsideration of the series rank in section Algarobia is suggested. Species-specific markers confirmed that material in Burkina Faso is P. juliflora, but suggested that material collected in Brazil, Cape Verde and Senegal is P. pallida, whereas this has previously been identified as P. juliflora.     

Variation in woody plant mortality and dieback from severe drought among soils, plant groups, and species within a northern Arizona ecotone

Vegetation change from drought-induced mortality can alter ecosystem community structure, biodiversity, and services. Although drought-induced mortality of woody plants has increased globally with recent warming, influences of soil type, tree and shrub groups, and species are poorly understood. Following the severe 2002 drought in northern Arizona, we surveyed woody plant mortality and canopy dieback of live trees and shrubs at the forest–woodland ecotone on soils derived from three soil parent materials (cinder, flow basalt, sedimentary) that differed in texture and rockiness. Our first of three major findings was that soil parent material had little effect on mortality of both trees and shrubs, yet canopy dieback of trees was influenced by parent material; dieback was highest on the cinder for pinyon pine (Pinus edulis) and one-seed juniper (Juniperus monosperma). Ponderosa pine (Pinus ponderosa) dieback was not sensitive to parent material. Second, shrubs had similar mortality, but greater canopy dieback, than trees. Third, pinyon and ponderosa pines had greater mortality than juniper, yet juniper had greater dieback, reflecting different hydraulic characteristics among these tree species. Our results show that impacts of severe drought on woody plants differed among tree species and tree and shrub groups, and such impacts were widespread over different soils in the southwestern U.S. Increasing frequency of severe drought with climate warming will likely cause similar mortality to trees and shrubs over major soil types at the forest–woodland ecotone in this region, but due to greater mortality of other tree species, tree cover will shift from a mixture of species to dominance by junipers and shrubs. Surviving junipers and shrubs will also likely have diminished leaf area due to canopy dieback.     

Climatic signals in growth and its relation to ENSO events of two Prosopis species following a latitudinal gradient in South America

Semiarid environments throughout the world have lost a major part of their woody vegetation and biodiversity due to the effects of wood cutting, cattle grazing and subsistence agriculture. The resulting state is typically used for cattle production, but the productivity of these systems is often very low, and erosion of the unprotected soil is a common problem. Such dry‐land degradation is of great international concern, not only because the resulting state is hardly productive but also because it paves the way to desertification. The natural distribution of the genus Prosopis includes arid and semiarid zones of the Americas, Africa and Asia, but the majority of the Prosopis species are, however, native to the Americas. In order to assess a likely gradient in the response of tree species to precipitation, temperature and their connection to El Niño southern oscillation (ENSO) events, two Prosopis species were chosen along a latitudinal gradient in Latin America, from northern Peru to central Chile: Prosopis pallida from a semi‐arid land in northern and southern Peru and P. chilensis from a semiarid land in central Chile. Growth rings of each species were crossdated at each sampling site using classical dendrochronological techniques. Chronologies were related with instrumental climatic records in each site, as well as with SOI and N34 series. Cross‐correlation, spectral and wavelet analysis techniques were used to assess the relation of growth with precipitation and temperature. Despite the long distance among sites, the two Prosopis species presented similar responses. Thus, the two species' growth is positively correlated to precipitation, while with temperature it is not. In northern Peru, precipitation and growth of P. pallida present a similar cyclic pattern, with a period of around 3 years. On the other hand, P. pallida in southern Peru, and P. chilensis also present this cyclic pattern, but also another one with lower frequency, coinciding with the pattern of precipitation. Both cycles are within the range of the ENSO band.     

Impact of Parthenium hysterophorus L. invasion on plant species composition and soil properties of grassland communities in Nepal

Parthenium hysterophorus (Asteraceae) is a noxious plant that is considered one of the most invasive species in the world. We studied changes in the composition of plant species and soil properties related to the invasion of P. hysterophorus in three grassland communities of central Nepal. We collected vegetation and soil data along transects that were established in densely invaded to non-invaded areas within homogenous grassland stands. We found significant differences between invaded, transitional and non-invaded plots in species composition and soil properties. There were fewer species in non-invaded than transitional and invaded plots. By P. hysterophorus invasion both native and non-native species were supported or replaced, respectively. The concentrations of soil nitrogen and organic matter were significantly higher in transitional and invaded plots than in non-invaded plots. Soil pH, phosphorus and potassium were highest in the invaded plots, lowest in the non-invaded and intermediate in the transitional plots. Due to changes in above-ground vegetation and below-ground soil nutrient contents, P. hysterophorus invasion is likely to have an overall negative effect on the functioning of the entire ecosystem. Therefore, management of noxious P. hysterophorus is necessary to prevent future problems.     

Pine invasion impacts on plant diversity in Patagonia: invader size and invaded habitat matter

Conifers, which are widely planted as fast growing tree crops, are invading forested and treeless environments across the globe, causing important changes in biodiversity. However, how small-scale impacts on plant diversity differ according to pine size and habitat context remains unclear. We assessed the effects of different stages of pine invasion on plant communities in forest and steppe sites located in southern Chile. In each site, we sampled plant diversity under and outside the canopy of Pinus contorta individuals, using paired plots. We assessed the relative impact of pine invasion on plant species richness and cover. In both sites, richness and cover beneath pine canopy decreased with increasing pine size (i.e. height and canopy area). A significant negative impact of pines on species richness and plant cover was detected for pines over 4 m in height. The impact of pines on plant richness and cover depended on pine size (i.e. canopy area) and habitat type. Larger pines had more negative impacts than smaller pines in both sites, with a greater impact for a given pine size in the Patagonian steppe compared to the A. araucaria forest. Species composition changed between under and outside canopy plots when pines were 4 m or taller. Pine presence reduced cover of most species. The impacts of pine invasions are becoming evident in forested and treeless ecosystems of southern Chile. Our results suggest that the magnitude of pine invasion impacts could be related to how adapted the invaded community is to tree cover, with the treeless environment more impacted by the invasion.     

Prosopis L. woody growth in relation to hydrology in South America: A review

Arboreal species of the genus Prosopis L. have played an important role in the development of tree-ring research in arid and semi-arid ecoregions of South America. Given the distribution of Prosopis across a broad precipitation gradient from 0 to 2000 mm y⁻¹ and its unique role as a phreatophyte, the relationship between Prosopis species growth and water has been a recurring theme over the past century. We conducted a systematic review of the literature addressing Prosopis and water research in South America, and combined site coordinates with GIS data of mean annual precipitation (MAP), elevation, biome, and soil moisture from online databases to understand the spatial distribution of research to date. We compiled 40 publications from 1931 to 2022, including results from 11 species of Prosopis among four countries, on the relationship between Prosopis spp and precipitation, groundwater levels, soil humidity, among other hydrological parameters. The spatial distribution of research sites spans tropical-subtropical and temperate latitudes from 4° to 35°S, excluding regions where the genus is present in Patagonia and northeastern South America. Studies covered a broad range of elevations from 30 to 3500 m a.s.l. but was limited to 1–730 mm y⁻¹ MAP, excluding more humid climates where Prosopis occurs. Results obtained from 32 dendrochronological studies and eight studies relating to Prosopis and hydrology, were grouped into sub-disciplines of tree-ring formation and the hydrosystem, dendroclimatology, dendrohydrology, and dendroecology. The review highlights the unique affinities of Prosopis to arid conditions, and the use of tree rings as a proxy for historical droughts and variability in water tables. Nonetheless, there are opportunities to expand the geographical-climatological extent of Prosopis growth research to humid climates, as well as to incorporate novel techniques such as stable isotopes and vessel size chronologies to understand how this genus records hydrological change throughout South America.     

Performances of machine learning algorithms for mapping fractional cover of an invasive plant species in a dryland ecosystem

In recent years, an increasing number of distribution maps of invasive alien plant species (IAPS) have been published using different machine learning algorithms (MLAs). However, for designing spatially explicit management strategies, distribution maps should include information on the local cover/abundance of the IAPS. This study compares the performances of five MLAs: gradient boosting machine in two different implementations, random forest, support vector machine and deep learning neural network, one ensemble model and a generalized linear model; thereby identifying the best‐performing ones in mapping the fractional cover/abundance and distribution of IPAS, in this case called Prosopis juliflora (SW. DC.). Field level Prosopis cover and spatial datasets of seventeen biophysical and anthropogenic variables were collected, processed, and used to train and validate the algorithms so as to generate fractional cover maps of Prosopis in the dryland ecosystem of the Afar Region, Ethiopia. Out of the seven tested algorithms, random forest performed the best with an accuracy of 92% and sensitivity and specificity >0.89. The next best‐performing algorithms were the ensemble model and gradient boosting machine with an accuracy of 89% and 88%, respectively. The other tested algorithms achieved comparably low performances. The strong explanatory variables for Prosopis distributions in all models were NDVI, elevation, distance to villages and distance to rivers; rainfall, temperature, near‐infrared and red reflectance, whereas topographic variables, except for elevation, did not contribute much to the current distribution of Prosopis. According to the random forest model, a total of 1.173 million ha (12.33% of the study region) was found to be invaded by Prosopis to varying degrees of cover. Our findings demonstrate that MLAs can be successfully used to develop fractional cover maps of plant species, particularly IAPS so as to design targeted and spatially explicit management strategies.     

Tree-shrub interactions in a subtropical savanna parkland: competition or facilitation?

Abstract. Prosopis glandulosa var. glandulosa has played a central role in the encroachment of woody plants in southern Texas, grasslands and savannas by acting as a nurse plant for various shrubs that establish in its understory. To test for continued facilitation of established understory shrubs by Prosopis and to determine if established shrubs compete with the Prosopis nucleus, selective removal experiments were conducted and monitored over a 2–5 yr period. Short-term (1–3 days) and long-term (2 yr) growth and physiological activities (midday net photosynthesis and leaf/shoot water potential) of two common understory shrubs, Zanthoxylum fagara and Berberis trifoliolata, growing with Prosopis, were generally comparable to those of individuals occurring in clusters where Prosopis was removed. Shrubs growing with an intact Prosopis occasionally showed significantly higher leaf-[N] and pre-dawn water potentials than those in clusters lacking a live Prosopis, especially under drought conditions; however, these differences did not translate into greater midday leaf gas exchange or shoot growth. By comparison, removal of understory shrubs elicited large increases in Prosopis net photosynthesis, annual trunk growth in each of the 5 yr monitored, and seed pod production in three of the four years monitored. Seven of 26 Prosopis plants in experimental clusters with an intact understory died over a 5-yr period, compared to only two of the 26 plants in clusters with the cleared understory. Results indicate that (1) the founding overstory Prosopis plant may continue to facilitate understory shrubs following their establishment, but these beneficial effects appear to be small and transitory, and (2) the understory shrubs have a pronounced negative effect on Prosopis, such that competition between overstory and understory woody plants is strongly asymmetrical. These findings suggest that understory shrubs will likely persist despite changes in microclimate and soils (potentially) that occur after the Prosopis plant, which facilitated their ingress or establishment, has died. Soil resource depletion by shallow-rooted understory shrubs appears to be a primary factor contributing to the demise of the deeply rooted, overstory Prosopis plants, especially on upland sites with duplex soils where below-ground competition is accentuated.     

Dendroecology of Prosopis flexuosa woodlands in the Monte desert: Implications for their management

In the Monte desert of Argentina open woodlands of several species of Prosopis occur in areas with accessible underground water. The great latitudinal extent of the Monte (26–43°S) exhibits strong climatic gradients involving temperature, rainfall seasonality, and wind regime. Prosopis woodlands have been a source of subsistence for human communities for several centuries and continue to be exploited by the local inhabitants. The “mining” of this resource has led to severe desertification and consequent impoverishment of the local people. In order to suggest strategies for the better management and recuperation of these woodlands we studied the population structure and productivity of Prosopis flexuosa from multiple plots at Pipanaco (27°58′S), Telteca (32°20′S), and Ñacuñán (34°03′S). For each plot we measured the density of P. flexuosa trees, number of stems, basal diameter (DAB), height and canopy diameter of each tree. Tree ring data were used to determine the growth rates, annual wood production and biological rotation age for each area. The ecological structure of the woodlands differs between the three sites. Along this north–south transect, there is a decrease in adult tree density, mean basal diameter, mean tree height, canopy cover, productivity and total wood biomass. Consequently, the potential sustainable use of these woodlands varies. Only the northern, Pipanaco, woodlands have the potential for lumber production. In contrast, the short, multi-stem and low-productivity trees in the Telteca and Ñacuñán areas can only sustain a combination of local firewood production and activities such as extensive grazing by livestock. The present, uniform regulations for harvesting wood in these areas must be changed to acknowledge these differences in order to optimize wood production in, and conservation of, these woodlands.     

Using plant functional types to compare vegetation structure of alien-invaded and uninvaded Acacia nilotica savannas

Four plant functional types (PFTs) were used to compare the vegetation structure of an alien-invaded Acacia nilotica savanna with one of negligible invasions. Heights, canopy covers and species richness of three native PFTs (woody plants, grasses and herbs) and one alien PFT (woody plants) were measured in 14, 1-m² quadrats sampled in a stratified-random pattern in a 400-m² plot demarcated in each savanna. In the uninvaded plot, mean heights of native PFTs were stratified. In the invaded plot, the mean height of aliens extended into the native woody stratum with the lower range of native woody PFT heights reduced to the grass stratum. Discriminant analysis of canopy covers and species richness of the four PFTs revealed significant differences in composition between plots with the alien PFT being the most important variable correlated with these differences. Univariate analysis confirmed the dominance of alien woody plants in the invaded plot but also showed significant reductions in the canopy covers and species richness of native herbs and grasses compared to those in the uninvaded plot. These results suggest that PFTs can rapidly measure small-scale, spatial differences in the physiognomy, composition and species richness of A. nilotica savannas when invaded by alien woody plants.     

Experimental evidence for a delayed response of the above-ground vegetation and the seed bank to the invasion of an annual exotic plant in deciduous forests

Invasions by alien plants significantly affect native biodiversity and ecosystem functioning. We conducted a 5-year field experiment to investigate potential effects of the annual invasive plant Impatiens glandulifera on both the native above-ground vegetation and the soil seed bank in a deciduous forest in Switzerland. Eight years after the establishment of I. glandulifera, we set up plots in patches invaded by the alien plant, in plots from which the invasive plant had been manually removed and in plots which were not yet colonized by the invasive plant. We examined plant species richness, diversity and plant species composition in the above-ground vegetation and soil seed bank in all plots one year and five years after the initiation of the experiment. The 36 plots (3 plot types × 6 replicates × 2 sites) were equally distributed over two forest sites. Neither the native above-ground vegetation nor the soil seed bank was influenced by the presence of I. glandulifera one year after the start of the field experiment. After five years, however, plant species richness of both the above-ground vegetation and the soil seed bank was reduced by 25% and 30%, respectively, in plots invaded by the alien plant compared to plots from which I. glandulifera had been removed or uninvaded plots. Furthermore, plots invaded by the alien plant had a lower total seedling density (reduction by 60%) and an altered plant species composition in the soil seed bank compared to control plots. Our field experiment indicates that negative effects of the annual invasive plant on the native above-ground vegetation and soil seed bank of deciduous forests become visible with a delay of several years.     

Grass, rainfall and herbivores as determinants of Acacia erioloba (Meyer) recruitment in an African savanna

Acacia species in arid environments are thought to only establish in years of above-average rainfall, so should exhibit cohorted or pulsed recruitment. I studied population demography of Acacia erioloba Meyer in semi-arid savanna in the Kimberley area (mean annual precipitation = 425 ± 132 mm), South Africa, to establish whether they recruit episodically. This species was found to have a sapling bank at the sites, indicating that even though cohorted recruitment probably occurs, it is not the primary factor limiting recruitment to larger size classes. A. erioloba saplings given supplementary water and protection from herbivory showed significantly less height growth than saplings given water but not protected from herbivores, and grass within exclusion plots was taller and denser than outside of exclusion plots. The generalized linear model, although finding a significant difference between watered and protected and watered and unprotected saplings, explained only 16% of variation in growth, demonstrating the importance of factors other than competition from grass when rainfall is above-average. Average height increase across all treatments for a growing season was small (43.1 ± 30.81 mm), suggesting most investment is belowground. A. erioloba sapling below surface stem diameter correlated positively with aboveground growth, indicating larger, and presumably older, individuals grow faster. When grass was removed around saplings, growth rates were not significantly greater than for saplings surrounded by grass, over a growing season. Thus, in these study sites, pulsed events allow seedlings to establish, and then saplings slowly accumulate over many growing seasons within the grass sward, owing to slow aboveground growth combined with competition from grass in absence of grazers. Release events, e.g., heavy grazing combined with good rainfall, may allow mass release, giving the impression of cohorted or pulsed recruitment.