Not only trees: Grasses determine African tropical biome distributions via water limitation and fire

Aim

Although much tropical ecology generally focuses on trees, grasses are fundamental for characterizing the extensive tropical grassy biomes (TGBs) and, together with the tree functional types, for determining the contrasting functional patterns of TGBs and tropical forests (TFs). To study the factors that determine African biome distribution and the transitions between them, we performed the first continental analysis to include grass and tree functional types.

Location

Sub‐Saharan Africa.

Time period

2000–2010.

Major taxa studied

Savanna and forest trees and C₄ grasses.

Methods

We combined remote‐sensing data with a land cover map, using tree functional types to identify TGBs and TFs. We analysed the relationships of grass and tree cover with fire interval, rainfall annual average and seasonality.

Results

In TGBs experiencing < 630 mm annual rainfall, grass growth was water limited. Grass cover and fire recurrence were strongly and directly related over the entire subcontinent. Some TGBs and TFs with annual rainfall > 1,200 mm had the same rainfall seasonality but displayed strongly different fire regimes.

Main conclusions

Water limitation to grass growth was fundamental in the driest TGBs, acting alongside the well‐known limitation to tree growth. Marked differences in fire regimes across all biomes indicated that fire was especially relevant for maintaining mesic and humid TGBs. At high rainfall, our results support the hypothesis of TGBs and TFs being alternative stable states maintained by a vegetation–fire feedback for similar climatic conditions.     

Demographic patterns of a widespread long‐lived tree are associated with rainfall and disturbances along rainfall gradients in SE Australia

Predicting species distributions with changing climate has often relied on climatic variables, but increasingly there is recognition that disturbance regimes should also be included in distribution models. We examined how changes in rainfall and disturbances along climatic gradients determined demographic patterns in a widespread and long‐lived tree species, Callitris glaucophylla in SE Australia. We examined recruitment since 1950 in relation to annual (200–600 mm) and seasonal (summer, uniform, winter) rainfall gradients, edaphic factors (topography), and disturbance regimes (vertebrate grazing [tenure and species], fire). A switch from recruitment success to failure occurred at 405 mm mean annual rainfall, coincident with a change in grazing regime. Recruitment was lowest on farms with rabbits below 405 mm rainfall (mean = 0–0.89 cohorts) and highest on less‐disturbed tenures with no rabbits above 405 mm rainfall (mean = 3.25 cohorts). Moderate levels of recruitment occurred where farms had no rabbits or less disturbed tenures had rabbits above and below 405 mm rainfall (mean = 1.71–1.77 cohorts). These results show that low annual rainfall and high levels of introduced grazing has led to aging, contracting populations, while higher annual rainfall with low levels of grazing has led to younger, expanding populations. This study demonstrates how demographic patterns vary with rainfall and spatial variations in disturbances, which are linked in complex ways to climatic gradients. Predicting changes in tree distribution with climate change requires knowledge of how rainfall and key disturbances (tenure, vertebrate grazing) will shift along climatic gradients.     

Nutritional values and indigenous preferences for Shea Fruits(vitellaria paradoxa C.F. Gaertn. F.) in African Agroforestry Parklands

Samples of dried shea fruit pulp were collected from tree populations in Mali, Burkina Faso, northern Cameroon, and Uganda. A variety of analytical methods was used to measure total soluble solids (TSS), crude protein, and mineral contents. The results demonstrate that shea fruits are a rich source of sugars, protein, calcium, and potassium during the “hungry season”, when food stores run low and the energy-intensive work of preparing land for planting must be done. A companion survey of indigenous shea tree and fruit classification was carried out in study area villages. Indigenous savanna inhabitants, especially men, emphasize the importance of fruit pulp taste, while women emphasize the butter content of the nuts. Shea fruits have greater importance to the inhabitants of the drier savannas such as the Sahel, where shea fruits have been shown to have higher nutritional values. While there is currently much international interest in developing the potential of shea butter production in Africa, the role of the fruit pulp in the local diet needs to be taken into consideration in development programs.     

Species-Specific Growth Responses to Climate Variations in Understory Trees of a Central African Rain Forest

Basic knowledge of the relationships between tree growth and environmental variables is crucial for understanding forest dynamics and predicting vegetation responses to climate variations. Trees growing in tropical areas with a clear seasonality in rainfall often form annual growth rings. In the understory, however, tree growth is supposed to be mainly affected by interference for access to light and other resources. In the semi-deciduous Mayombe forest of the Democratic Republic of Congo, the evergreen species Aidia ochroleuca, Corynanthe paniculata and Xylopia wilwerthii dominate the understory. We studied their wood to determine whether they form annual growth rings in response to changing climate conditions. Distinct growth rings were proved to be annual and triggered by a common external factor for the three species. Species-specific site chronologies were thus constructed from the cross-dated individual growth-ring series. Correlation analysis with climatic variables revealed that annual radial stem growth is positively related to precipitation during the rainy season but at different months. The growth was found to associate with precipitation during the early rainy season for Aidia but at the end of the rainy season for Corynanthe and Xylopia. Our results suggest that a dendrochronological approach allows the understanding of climate–growth relationships in tropical forests, not only for canopy trees but also for evergreen understory species and thus arguably for the whole tree community. Global climate change influences climatic seasonality in tropical forest areas, which is likely to result in differential responses across species with a possible effect on forest composition over time.     

Ecological adaptation of the shea butter tree (Vitellaria paradoxa C.F. Gaertn.) along climatic gradient in Bénin,West Africa

The ecological adaptation of shea butter trees was assessed based on their dendrometric and production traits in four shea butter tree parks occurring in different climatic zones of Bénin. A total of 99 rectangular plots of 50 × 30 m were established within the four parks according to a random sampling scheme. In each plot, all trees with a diameter at breast height (dbh) >10 cm were inventoried and measured for stem and crown diameters, and total height. The production of 120 productive shea butter trees was quantified. Collected data were used to compute structural parameters for each park. Moreover, stem diameter and height structures of the trees were established. Principal component analysis was performed on the dendrometric variables, and the first three components were correlated with the climatic parameters. Results revealed significant differences between parks in most of the dendrometric and production parameters of shea butter trees. For all the four parks, stem diameter and height structures present a Gaussian shape with left dissymmetry. In the Guinean zone, shea butter trees develop large crowns but produce little quantities of fruits, whereas in the Sudanian regions, the opposite trend was observed.     

Conservation value for birds of traditionally managed isolated trees in an agricultural landscape of Madagascar

Isolated trees have distinctive economic, social and cultural value for the Betsileo people living on the edge of the protected forest corridor between Ranomafana and Andringitra national parks, in South-East Madagascar. Many of these trees are Ficus spp., traditionally protected and respected. At the landscape level, they are isolated features in a heterogeneous mosaic, providing fruit, shade and aesthetic services in open cultivated areas. Within the current management system, isolated trees may also contribute significantly to the provision of ecological services by enhancing bird diversity in open areas outside the forest. We identified practices and values linked to isolated tree uses and management through ethnographic data collection. Bird presence and abundance were sampled by 338 point counts in isolated trees and open areas of the agricultural mosaic. Isolated trees were occupied by 18 out of 32 (56%) bird species in the agricultural mosaic, including 8 (25%) endemic forest species. Endemic forest birds were significantly more numerous in isolated trees than in open habitats, both in species richness and abundance (mean P value < 0.001). Overall bird species richness was significantly higher in open areas containing isolated trees, than in areas without isolated trees. Bird species richness in Ficus spp. was significantly higher than in other isolated tree species, although no differences were detected in abundance or within guilds. Community-based management of isolated trees may thus represent an opportunity for convergence between bird conservation goals outside protected areas and local management values and practices.     

Change of season-specific telomere lengths in <Emphasis Type="Italic">Ginkgo biloba</Emphasis> L.

Telomeres have lately received considerable attention in the development of tree species. Normal somatic cells have limited replicative capacity and telomeres get shorten with each round of DNA replication. For broad-leaved tree species, to determine what changes happen to their somatic cells in its annual development cycle, an exhaustive research on different ages of gingko trees telomere length changes was carried out. Analysis of changes in leaf telomere lengths in the annual development cycle of Ginkgo biloba L. showed no significant changes (P > 0.05) from April to August, but a dramatic decrease in September and October (P < 0.05). Statistical analyses showed that TRF length of males and females are equal, the p values of the three age groups comparison were all bigger than 0.05. The results showed that specific apoptotic changes occur in the annual development cycle of Ginkgo biloba L.     

Solar radiation and ENSO predict fruiting phenology patterns in a 15‐year record from Kibale National Park,Uganda

Fruiting, flowering, and leaf set patterns influence many aspects of tropical forest communities, but there are few long‐term studies examining potential drivers of these patterns, particularly in Africa. We evaluated a 15‐year dataset of tree phenology in Kibale National Park, Uganda, to identify abiotic predictors of fruit phenological patterns and discuss our findings in light of climate change. We quantified fruiting for 326 trees from 43 species and evaluated these patterns in relation to solar radiance, rainfall, and monthly temperature. We used time‐lagged variables based on seasonality in linear regression models to assess the effect of abiotic variables on the proportion of fruiting trees. Annual fruiting varied over 3.8‐fold, and inter‐annual variation in fruiting is associated with the extent of fruiting in the peak period, not variation in time of fruit set. While temperature and rainfall showed positive effects on fruiting, solar radiance in the two‐year period encompassing a given year and the previous year was the strongest predictor of fruiting. As solar irradiance was the strongest predictor of fruiting, the projected increase in rainfall associated with climate change, and coincident increase in cloud cover suggest that climate change will lead to a decrease in fruiting. ENSO in the prior 24‐month period was also significantly associated with annual ripe fruit production, and ENSO is also affected by climate change. Predicting changes in phenology demands understanding inter‐annual variation in fruit dynamics in light of potential abiotic drivers, patterns that will only emerge with long‐term data.     

Relating tree growth to rainfall in Bolivian rain forests: a test for six species using tree ring analysis

Many tropical regions show one distinct dry season. Often, this seasonality induces cambial dormancy of trees, particularly if these belong to deciduous species. This will often lead to the formation of annual rings. The aim of this study was to determine whether tree species in the Bolivian Amazon region form annual rings and to study the influence of the total amount and seasonal distribution of rainfall on diameter growth. Ring widths were measured on stem discs of a total of 154 trees belonging to six rain forest species. By correlating ring width and monthly rainfall data we proved the annual character of the tree rings for four of our study species. For two other species the annual character was proved by counting rings on trees of known age and by radiocarbon dating. The results of the climate–growth analysis show a positive relationship between tree growth and rainfall in certain periods of the year, indicating that rainfall plays a major role in tree growth. Three species showed a strong relationship with rainfall at the beginning of the rainy season, while one species is most sensitive to the rainfall at the end of the previous growing season. These results clearly demonstrate that tree ring analysis can be successfully applied in the tropics and that it is a promising method for various research disciplines.     

Annual cycles are the most common reproductive strategy in African tropical tree communities

We present the first cross‐continental comparison of the flowering and fruiting phenology of tropical forests across Africa. Flowering events of 5446 trees from 196 species across 12 sites and fruiting events of 4595 trees from 191 species across 11 sites were monitored over periods of 6 to 29 years and analyzed to describe phenology at the continental level. To study phenology, we used Fourier analysis to identify the dominant cycles of flowering and fruiting for each individual tree and we identified the time of year African trees bloom and bear fruit and their relationship to local seasonality. Reproductive strategies were diverse, and no single regular cycle was found in >50% of individuals across all 12 sites. Additionally, we found annual flowering and fruiting cycles to be the most common. Sub‐annual cycles were the next most common for flowering, whereas supra‐annual patterns were the next most common for fruiting. We also identify variation in different subsets of species, with species exhibiting mainly annual cycles most common in West and West Central African tropical forests, while more species at sites in East Central and East African forests showed cycles ranging from sub‐annual to supra‐annual. Despite many trees showing strong seasonality, at most sites some flowering and fruiting occurred all year round. Environmental factors with annual cycles are likely to be important drivers of seasonal periodicity in trees across Africa, but proximate triggers are unlikely to be constant across the continent.     

Recent growth increase in endemic Juglans boliviana from the tropical Andes

The spatial coverage of tree-ring chronologies in tropical South America is low compared to the extratropics, particularly in remote regions. Tree-ring dating from such tropical sites is limited by the generally weak temperature seasonality, complex coloration, and indistinct anatomical morphology in some tree species. As a result, there is a need to complement traditional methods of dendrochronology with innovative and independent approaches. Here, we supplement traditional tree-ring methods via the use of radiocarbon analyses to detect partial missing rings and/or false rings, and wood anatomical techniques to precisely delineate tree-ring boundaries. In so doing we present and confirm the annual periodicity of the first tree-ring width (TRW) chronology spanning from 1814 to 2017 for Juglans boliviana (‘nogal’), a tree species growing in a mid-elevation tropical moist forest in northern Bolivia. We collected 25 core samples and 4 cross-sections from living and recently harvested canopy-dominant trees, respectively. The sampled trees were growing in the Madidi National Park and had a mean age of 115 years old, with certain trees growing for over 200 years. Comparison of (residual and standard) TRW chronologies to monthly climate variables shows significant negative relationships to prior year May-August maximum temperatures (r = −0.54, p < 0.05) and positive relationships to dry season May-October precipitation (r = 0.60, p < 0.05) before the current year growing season. Additionally, the radial growth of Juglans boliviana shows a significant positive trend since 1979. Our findings describe a new and promising tree species for dendrochronology due to its longevity and highlight interdisciplinary techniques that can be used to expand the current tree-ring network in Bolivia and the greater South American tropics.     

Does soil moisture availability explain liana seedling distribution across a tropical rainfall gradient?

Liana density tends to increase with decreasing rainfall and increasing seasonality. However, the pattern of liana distribution may be due to differences in soil water retention capacity, not rainfall and seasonality per se. We tested the effect of rainfall and soil substrate with respect to the distribution of liana seedlings in six sites across a rainfall gradient from the wet Atlantic to the dry Pacific in central Panama. Soils were either limestone, with low water‐holding capacity, or laterite, with higher water‐holding capacity. We sampled liana seedlings at each site using three 1 × 100 m transect. We found that relative liana seedling density was higher on limestone soils compared to laterite soils regardless of the amount of rainfall. Furthermore, liana community composition on limestone soils was more similar to dry forest sites than to adjacent wet and moist forest sites. Liana seedling species diversity relative to trees was significantly higher in a low‐fertility dry forest site compared to a high‐fertility forest, but did not differ from the other sites. Thus, liana seedling density and community structure may be driven more by soil type and thus by soil moisture availability than strictly by mean annual rainfall and the seasonality of rainfall.     

Climate response among growth increments of fish and trees

Significant correlations were found among the annual growth increments of stream fish, trees, and climate variables in the Ozark region of the United States. The variation in annual growth increments of rock bass (Ambloplites rupestris) from the Jacks Fork River was significantly correlated over 22 years with the ring width of four tree species: white oak (Quercus alba), post oak (Quercus stellata), shortleaf pine (Pinus echinata) and eastern red cedar (Juniperus virginiana). Rock bass growth and tree growth were both significantly correlated with July rainfall and stream discharge. Variations in annual growth of smallmouth bass (Micropterus dolomieu) from four streams were significantly correlated over 29 years (1939–1968) with mean May maximum air temperature but not with tree growth. The magnitude and significance of correlations among growth increments from fish and trees imply that conditions such as topography, stream gradient, organism age, and the distribution of a population relative to its geographic range can influence the climatic response of an organism. The timing and intensity of climatic variables may produce different responses among closely related species.     

Climatic and anthropogenic factors affect Ailanthus altissima invasion in a Mediterranean region

Ailanthus altissima is an aggressive invasive tree worldwide, but the ecological factors that lead to the spread of this species in Mediterranean ecosystems are still unclear. Here we aim to identify such factors, focusing on the interaction of human activity with climatic conditions. We determined the occurrence and abundance of Ailanthus in 240 sites and studied their relationship with 20 variables representing climatic, geographic, and topographic factors, as well as land use, in the region of Campania (southern Italy). Overall, we found that temperature and rainfall in Campania are suitable for Ailanthus, with the only major constraint being the temperature at an altitude exceeding 900 m a.s.l.. We found that Ailanthus is unable to spread where the mean annual temperature is lower than 11.1 °C. By contrast, precipitation variables showed poor correlation with Ailanthus distribution, suggesting that rainfall in the selected study sites is suitable to sustain the growth of this tree. About land use variables, roads were the primary landscape feature along which this species spread and invaded new areas. Roads probably combine high propagule pressure and favorable growing conditions in terms of available resources i.e., light, water, and mineral nutrients, that allow Ailanthus to establish and spread along roadside edges in different ecosystems. In conclusion, we found that climate and human-associated variables are correlated with the current occurrence of Ailanthus, with the temperature being more influential at high elevation sites and road distance playing a prominent role in low elevation areas.

     

Proximate weather patterns and spring green‐up phenology effect Eurasian beaver (Castor fiber) body mass and reproductive success: the implications of climate change and topography

Low spring temperatures have been found to benefit mobile herbivores by reducing the rate of spring‐flush, whereas high rainfall increases forage availability. Cold winters prove detrimental, by increasing herbivore thermoregulatory burdens. Here we examine the effects of temperature and rainfall variability on a temperate sedentary herbivore, the Eurasian beaver, Castor fiber, in terms of inter‐annual variation in mean body weight and per territory offspring production. Data pertain to 198 individuals, over 11 years, using capture‐mark‐recapture. We use plant growth (tree cores) and fAPAR (a satellite‐derived plant productivity index) to examine potential mechanisms through which weather conditions affect the availability and the seasonal phenology of beaver forage. Juvenile body weights were lighter after colder winters, whereas warmer spring temperatures were associated with lighter adult body weights, mediated by enhanced green‐up phenology rates. Counter‐intuitively, we observed a negative association between rainfall and body weight in juveniles and adults, and also with reproductive success. Alder, Alnus incana, (n = 68) growth rings (principal beaver food in the study area) exhibited a positive relationship with rainfall for trees growing at elevations >2 m above water level, but a negative relationship for trees growing <0.5 m. We deduce that temperature influences beavers at the landscape scale via effects on spring green‐up phenology and winter thermoregulation. Rainfall influences beavers at finer spatial scales through topographical interactions with plant growth, where trees near water level, prone to water logging, producing poorer forage in wetter years. Unlike most other herbivores, beavers are an obligate aquatic species that utilize a restricted ‘central‐place’ foraging range, limiting their ability to take advantage of better forage growth further from water during wetter years. With respect to anthropogenic climate change, interactions between weather variables, plant phenology and topography on forage growth are instructive, and consequently warrant examination when developing conservation management strategies for populations of medium to large herbivores.     

Land‐use change outweighs projected effects of changing rainfall on tree cover in sub‐Saharan Africa

Global change will likely affect savanna and forest structure and distributions, with implications for diversity within both biomes. Few studies have examined the impacts of both expected precipitation and land use changes on vegetation structure in the future, despite their likely severity. Here, we modeled tree cover in sub‐Saharan Africa, as a proxy for vegetation structure and land cover change, using climatic, edaphic, and anthropic data (R² = 0.97). Projected tree cover for the year 2070, simulated using scenarios that include climate and land use projections, generally decreased, both in forest and savanna, although the directionality of changes varied locally. The main driver of tree cover changes was land use change; the effects of precipitation change were minor by comparison. Interestingly, carbon emissions mitigation via increasing biofuels production resulted in decreases in tree cover, more severe than scenarios with more intense precipitation change, especially within savannas. Evaluation of tree cover change against protected area extent at the WWF Ecoregion scale suggested areas of high biodiversity and ecosystem services concern. Those forests most vulnerable to large decreases in tree cover were also highly protected, potentially buffering the effects of global change. Meanwhile, savannas, especially where they immediately bordered forests (e.g. West and Central Africa), were characterized by a dearth of protected areas, making them highly vulnerable. Savanna must become an explicit policy priority in the face of climate and land use change if conservation and livelihoods are to remain viable into the next century.     

Masting in a temperate tree: Evidence for environmental prediction?

Many plant species produce large fruit crops in some years and then produce few or no fruits in others. Synchronous, inter‐annual variation in plant reproduction is known as ‘masting’ and its adaptive significance has yet to be fully resolved. For 8 consecutive years, I quantified every fruit produced by 22 females of a New Zealand tree species (Dysoxylum spectabile), which has an unusual habit of taking a full calendar year to mature fruits after flowering. Fruit production varied strongly among years and was tightly synchronized among trees. Annual variability in fruit production declined with total reproductive output, indicating trees with lower fecundity exhibited a stronger tendency to mast. Although unrelated to temperature, annual fruit production was positively related to precipitation during annual periods of fruit development, and negatively related to fruit production in the previous year. Seedlings had higher rates of survivorship in a wet, high‐seed year than in a dry, low‐seed year, suggesting that seedlings might be drought sensitive. Therefore, D. spectabile produced large fruit crops during periods of high rainfall prior to fruit maturation, which may enhance survivorship of drought‐intolerant seeds. Results were inconsistent with several hypotheses that are widely believed to be the most likely explanations for masting. Instead, results were consistent with the environmental prediction hypothesis, suggesting that this hypothesis may be more important than previously appreciated.     

Fruit production is influenced by tree size and size‐asymmetric crowding in a wet tropical forest

In tropical forest communities, seedling recruitment can be limited by the number of fruit produced by adults. Fruit production tends to be highly unequal among trees of the same species, which may be due to environmental factors. We observed fruit production for ~2,000 trees of 17 species across 3 years in a wet tropical forest in Costa Rica. Fruit production was modeled as a function of tree size, nutrient availability, and neighborhood crowding. Following model selection, tree size and neighborhood crowding predicted both the probability of reproduction and the number of fruit produced. Nutrient availability only predicted only the probability of reproduction. In all species, larger trees were more likely to be reproductive and produce more fruit. In addition, number of fruit was strongly negatively related to presence of larger neighboring trees in 13 species; presence of all neighboring trees had a weak‐to‐moderate negative influence on reproductive status in 16 species. Among various metrics of soil nutrient availability, only sum of base cations was positively associated with reproductive status, and for only four species. Synthesis Overall, these results suggest that direct influences on fruit production tend to be mediated through tree size and crowding from neighboring trees, rather than soil nutrients. However, we found variation in the effects of neighbors and nutrients among species; mechanistic studies of allocation to fruit production are needed to explain these differences.     

Acorn production prediction models for five common oak species of the eastern United States

Acorn production varies considerably among oak (Quercus) species, individual trees, years, and locations, which directly affects oak regeneration and populations of wildlife species that depend on acorns for food. Hard mast indices provide a relative ranking and basis for comparison of within- and between-year acorn crop size at a broad scale, but do not provide an estimate of actual acorn yield—the number of acorns that can potentially be produced on a given land area unit based on the species, number, and diameter at breast height (dbh) of oak trees present. We used 10 years of acorn production data from 475 oak trees to develop predictive models of potential average annual hard mast production by five common eastern oak species, based on tree diameter and estimated crown area. We found a weak (R² = 0.08–0.28) relationship between tree dbh and acorn production per unit crown area for most species. The relationship between tree dbh and acorn production per tree was stronger (R² = 0.33–0.57). However, this is because larger-dbh trees generally have larger crowns, not because they have a greater capacity to produce more acorns per unit crown area. Acorn production is highly variable among individual trees. We estimated that dbh of at least 60 dominant or codominant oak trees per species should be randomly sampled to obtain an adequate representation of the range of dbhs (≥12.7 cm dbh) in a given forest area, and achieve precise estimates when using these equations to predict potential acorn production. Our predictive models provide a tool for estimating potential acorn production that land managers and forest planners can apply to oak inventory data to tailor estimates of potential average annual acorn production to different forest management scenarios and multiple spatial scales. © 2011 The Wildlife Society.     

The ecology of Maesopsis eminii Engl. in tropical Africa

Maesopsis eminii is referred to as one of the most widely distributed African tree species. However, its occurrence in Africa has never been mapped and little is known as to how this species can sustain in different environments. To gain insight into Maesopsis’ ecology, we (i) made a synthesis of its functional trait data from the literature, (ii) investigated phenological patterns using data on four M. eminii trees from Yangambi, DR Congo, (iii) assessed an empirical provenance trial from Uganda on 600 Maesopsis trees and (iv) synthesized geo‐referenced point location maps of Maesopsis entailing WorldClim precipitation and temperature and FAO soils, rainfall and ecological zones for Africa. We found M. eminii to straddle the equator equidistantly in terms of latitude (10.97°N and 10.98°S) covering five forest types where twenty soil types and variable rainfall regimes support complex plant biodiversity. Maesopsis eminii was, however, largely concentrated in the tropical rainforest ecosystem which contains fertile Orthic Ferralsol soils. More than 97% of the point locations were found where annual precipitation was >1000 mm, and 82% occurred where average annual temperature was 22–28°C. Its functional traits, phenology and provenance trial findings explained its occurrence in Africa.