Termite‐induced heterogeneity in African savanna vegetation: mechanisms and patterns

Objectives: To (1) assess the strength of evidence for the role of termites in vegetation heterogeneity in African savannas, and (2) identify the mechanisms by which termites induce such heterogeneity. Location: African savannas. Methods: We conducted a review of the literature, a meta‐analysis and qualitative systems analysis to identify mechanisms to explain the observed patterns. Results: The review provided evidence for termite‐induced heterogeneity in floristic composition and vegetation patterning in savannas across Africa. Termites induced vegetation heterogeneity directly or indirectly through their nest‐building and foraging activities, associated nutrient cycling and their interaction with mammalian herbivores and fire. The literature reviewed indicated that termite mounds essentially act as islands of fertility, which are responsible for ecosystem‐level spatial heterogeneity in savannas. This was supported by the meta‐analysis, which demonstrated that mounds of Ancistrotermes, Macrotermes, Odontotermes (family Macrotermitinae), Cubitermes (family Termitinae) and Trinervitermes (Nasutitermitinae) are significantly enriched in clay (75%), carbon (16%), total nitrogen (42%), calcium (232%), potassium (306%) and magnesium (154%) compared to the surrounding savanna soil. Conclusions: Termite activity is one of the major factors that induce vegetation patterning in African savannas. The implications of this are discussed and research questions for future studies and modelling efforts are indicated.     

Root foraging strategies and soil patchiness in a humid savanna

In Lamto (Côte d'Ivoire), the savanna is a patchy environment as far as soil is concerned: tree clumps and termite mounds lead to higher nutrient contents than in the surrounding savanna. Mature Borassus aethiopum (Mart.) specimens are tall palm trees dominating the community, with aerial parts located out of these nutrient-rich patches.Palm root densities were compared under tree clumps and in the surrounding savanna, and were also sampled along transects between palm trees and nutrient-rich patches (two clumps and one mound). Palm root densities were far higher (up to 10 times) in the nitrogen-rich soil of both clumps and termite mounds than in the surrounding savanna. Evidence is given that palm trees are able to extend their root system as far as 20 m towards these nutrient-rich patches where they proliferate. These results point out a particular root foraging strategy, which is one of the first known for a woody perennial. They also provide new insights for understanding nitrogen cycling and savannas high rate of primary production.     

Origin of mound-field landscapes: a multi-proxy approach combining contemporary vegetation, carbon stable isotopes and phytoliths

Background and aims

Seasonally flooded South American savannas harbor different kinds of mound-field landscapes of largely unknown origin. A recent study used soil carbon-isotope depth profiles and other proxies to infer vegetation history in murundu landscapes in Brazil. Results suggested that differential erosion, not building-up processes (e.g., termite mounds), produced mounds. We tested this approach to inferring mound origin in a mound-field landscape in French Guiana.

Methods

We examined carbon-isotope depth profiles of soil organic matter, phytolith profiles and contemporary vegetation composition in mounds and inter-mounds.

Results

Relative abundance of C3 and C4 plants across habitats was very different from that in murundu landscapes; C3 plants were better represented in inter-mounds than on mounds. Habitat differences in C3/C4 distribution were subtler than in murundu landscapes, limiting inference of vegetation history based on carbon isotopes. Still, carbon-isotope and phytolith depth profiles gave similar pictures of vegetation history, both favoring a building-up hypothesis, corroborating other evidence that these mounds are vestiges of ancient agricultural raised fields.

Conclusions

Carbon-isotope depth profiles are unlikely to be adequate for deciphering origin of mound-field landscapes from vegetation history in seasonally flooded savannas. Including data on current vegetation and phytoliths makes inferences more robust.     

Structure and composition of savanna vegetation in northern Bolivia: A preliminary report

The vegetation of seasonally flooded savannas iin the tropical lowland of the Departamento La Paz, northern Bolivia, was studied. A first collection of vascular plants in this area includes about 600 species, at least three of which are new to the Bolivian flora and four are new to science. The eight major vegetation types distinguished within the savannas are dry bunchgrass savanna (“altura”), seasonally flooded bunch-grass community (“sartenejal”), palm swamp (“palmar”), two types of river floodplains (”bajíos”), a tall shrub community (“chaparral”), the vegetation of termite mounds, and gallery forest. The study area is among the least known in Bolivia. There is a strong need for further studies of these savannas.     

Spatial variability and abiotic determinants of termite mounds throughout a savanna catchment

Termite mounds contribute to the spatial heterogeneity of ecological processes in many savannas, but the underlying patterns and determinants of mound distributions remain poorly understood. Using the Carnegie Airborne Observatory (CAO), we mapped the distribution of termite mounds across a rainfall gradient within a river catchment (～ 27 000 ha) of the Kruger National Park, South Africa. We assessed how different factors were associated with the distribution and height of termite mounds at three spatial scales: the entire catchment, among three broad vegetation types, and on individual hillslope crests. Abiotic factors such as the underlying geology and mean annual precipitation shaped mound densities at broad scales, while local hillslope morphology strongly influenced mound distribution at finer scales, emphasising the importance of spatial scale when assessing mound densities. Fire return period had no apparent association with mound densities or height. Mound density averaged 0.46 mounds ha^?1, and exhibited a clustered pattern throughout the landscape, occurring at relatively high densities (up to 2 mounds ha^?1) on crests, which are nutrient‐poor elements of the landscape. Mounds exhibited significant over‐dispersion (even spacing) at scales below 60 m so that evenly spaced aggregations of termite mounds are embedded within a landscape of varying mound densities. The tallest mounds were found in dry savanna (500 mm yr^?1) and were positively correlated with mound density, suggesting that dry granitic savannas are ideal habitat for mound‐building termites. Mound activity status also varied significantly across the rainfall gradient, with a higher proportion of active (live) mounds in the drier sites. The differential spacing of mounds across landscapes provides essential nutrient hotspots in crest locations, potentially sustaining species that would otherwise not persist. The contribution to biodiversity and ecosystem functioning that mounds provide is not uniform throughout landscapes, but varies considerably with spatial scale and context.     

Nutrient dynamics and plant assemblages of Macrotermes falciger mounds in a savanna ecosystem

Termites through mound construction and foraging activities contribute significantly to carbon and nutrient fluxes in nutrient-poor savannas. Despite this recognition, studies on the influence of termite mounds on carbon and nitrogen dynamics in sub-tropical savannas are limited. In this regard, we examined soil nutrient concentrations, organic carbon and nitrogen mineralization in incubation experiments in mounds of Macrotermes falciger and surrounding soils of sub-tropical savanna, northeast Zimbabwe. We also addressed whether termite mounds altered the plant community and if effects were similar across functional groups i.e. grasses, forbs or woody plants. Mound soils had significantly higher silt and clay content, pH and concentrations of calcium (Ca), magnesium (Mg), potassium (K), organic carbon (C), ammonium (NH₄⁺) and nitrate (NO₃⁻) than surrounding soils, with marginal differences in phosphorus (P) and sodium (Na) between mounds and matrix soils. Nutrient enrichment increased by a factor ranging from 1.5 for C, 4.9 for Mg up to 10.3 for Ca. Although C mineralization, nitrification and nitrification fraction were similar between mounds and matrix soils, nitrogen mineralization was elevated on mounds relative to surrounding matrix soils. As a result, termite mounds supported unique plant communities rich and abundant in woody species but less diverse in grasses and forbs than the surrounding savanna matrix in response to mound-induced shifts in soil parameters specifically increased clay content, drainage and water availability, nutrient status and base cation (mainly Ca, Mg and Na) concentration. In conclusion, by altering soil properties such as texture, moisture content and nutrient status, termite mounds can alter the structure and composition of sub-tropical savanna plant communities, and these results are consistent with findings in other savanna systems suggesting that increase in soil clay content, nutrient status and associated changes in the plant community assemblage may be a general property of mound building termites.     

Isolation and characterization of 10 microsatellite loci in the magnetic termite,Amitermes meridionalis (Isoptera: Termitidae)

The elongated mounds of the ‘magnetic termite’, Amitermes meridionalis are a prominent feature of the Northern Territory in Australia. They are restricted to habitat patches of seasonally flooded plains which are largely isolated from each other. To investigate the population structure of A. meridionalis, we developed 10 polymorphic microsatellite loci. We tested the variability of the markers on at least 20 individuals from two populations. We found three to 12 alleles per locus with a level of heterozygosity at each locus ranging from 0.05 to 0.74.     

Multi-scale pattern analysis of a mound-building termite species

Termite mounds are a widespread feature in most African savannas. These structures exhibit high nutrient contents and often host a special vegetation composition. In this study, we analysed mound distribution patterns of a fungus-growing termite species, Macrotermes michaelseni, an important ecosystem engineer in the savannas of Namibia. Inhabited mounds taller than 0.7 m were regularly distributed. We view this pattern as a result of intraspecific competition. The heights of mounds taller than 0.7 m were correlated positively with their distance, such that mounds closer together, i.e. up to inter-mound distances of approximately 50 m, tended to be smaller than average. This indicates that intraspecific competition for foraging areas controls mound distribution pattern and colony size. Differences between mound heights increased on the spatial scale up to inter-mound distances of 80 m. We assume that the foundation of new colonies is only possible in unoccupied patches. In such patches, young colonies are able to occur close together as they have a relatively low foraging demand and therefore a low spatial demand. In contrast, their critical distance to taller colonies with higher foraging demands is rather large, which leads to the observed increasing difference of mound heights with increasing distances between them.     

Mutualistic acacia ants exhibit reduced aggression and more frequent off‐tree movements near termite mounds

In many ant–plant mutualisms, ants establish colonies in hollow thorns, leaf pouches, or other specialized structures on their host plants, which they then defend from herbivores. Resource heterogeneity could affect the maintenance of these mutualisms if it leads to one or both partners altering their investment in the interaction. Such a phenomenon may be especially pertinent to the Acacia–ant mutualism found in East African savannas, where termite mounds have a profound effect on the spatial structuring of resources used by both plants and ants. Here, we examined whether the proximity to termite mounds of Acacia drepanolobium trees is associated with variation in the behavior of one of their ant associates, Crematogaster nigriceps. We found that ant colonies near termite mounds had decreased aggressive responses to simulated herbivory as well as increased off‐tree movement. We hypothesize that these changes are the result of resident ant colonies near termite mounds shifting investment from defense of their host plant to foraging for nearby resources.     

Studies on a population of large termite mounds in Uganda

• 1 Two study-sites near Kampala were mapped and all of the termite mounds within them were measured. Changes were recorded in the three succeeding years. A method was developed for estimating mound volumes from height and diameter.
• 2 Macrotermes bellicosus mounds reached a volume of about 2.4 m³ in 3 years after which they grew more slowly to a maximum from 4 to 6 m³. Growth in volume during the first 3 years was approximately linear and its rate did not vary seasonally, probably because growth is related to temperature which was nearly constant throughout the year.
• 3 M.bellicosus mounds had an average length of life of about 10 years, although colonies survived on average only 4 years. The corresponding figures for Pseudacanthotermes spp. were 5 and 2 1/2 years. Some mounds were recolonized after the death of the original colony.
• 5 Fresh earth was brought up at a minimum rate of about 10 m³ ha^-1 yr^-1 but more was lost by erosion and destruction and the populations of mounds were decreasing.
• 4 The number of ventilation shafts in M.bellicosus mounds increased with the volume of the mound. Many mounds, both dead and alive, supported vegetation.
• 6 Local opinion is that Pseudacanthotermes are harmless but M.bellicosus is regarded as a pest and attempts are made to destroy its mounds.

     

Termite mound emissions of CH4 and CO2 are primarily determined by seasonal changes in termite biomass and behaviour

Termites are a highly uncertain component in the global source budgets of CH₄ and CO₂. Large seasonal variations in termite mound fluxes of CH₄ and CO₂ have been reported in tropical savannas but the reason for this is largely unknown. This paper investigated the processes that govern these seasonal variations in CH₄ and CO₂ fluxes from the mounds of Microcerotermes nervosus Hill (Termitidae), a common termite species in Australian tropical savannas. Fluxes of CH₄ and CO₂ of termite mounds were 3.5-fold greater in the wet season as compared to the dry season and were a direct function of termite biomass. Termite biomass in mound samples was tenfold greater in the wet season compared to the dry season. When expressed per unit termite biomass, termite fluxes were only 1.2 (CH₄) and 1.4 (CO₂)-fold greater in the wet season as compared to the dry season and could not explain the large seasonal variations in mound fluxes of CH₄ and CO₂. Seasonal variation in both gas diffusivity through mound walls and CH₄ oxidation by mound material was negligible. These results highlight for the first time that seasonal termite population dynamics are the main driver for the observed seasonal differences in mound fluxes of CH₄ and CO₂. These findings highlight the need to combine measurements of gas fluxes from termite mounds with detailed studies of termite population dynamics to reduce the uncertainty in quantifying seasonal variations in termite mound fluxes of CH₄ and CO₂.     

Role of termites in the restoration of soils and plant richness on bowé in West Africa

Bowé (hardened ferricrete soils formed by erosion, drought or deforestation) are often associated with termite mounds, but little is known about these mounds and their role in the restoration of soils and plant biodiversity on bowé. This study examined termite mounds on bowé and their effects on soil depth and plant richness. Sixty-four sampling plots were laid out randomly on bowé sites with mounds and on adjacent bowé sites without mounds. The height and circumference of each mound were measured. Species inventories were made and soil depth measured in each plot. Linear mixed effects and generalised mixed effects models with Poisson error distribution were used to assess the variation in soil depth and plant species richness in mound and nonmound microsites. Two types of mounds (small vs. large) associated with different termite species were observed on bowé, with the small mounds being most common. Plots with either large or small mounds had deeper soils and higher plant richness than the adjacent plots without mounds. Conservation of termite mounds is important for restoring soils and plant richness on bowé, and termite mounds should be taken into consideration in biodiversity and soil management strategies for bowé.     

The distribution,abundance, and the effects of fire on mound building termites (Trinervitermes and Cubitermes spp., Isoptera: Termitidae) in northern guinea savanna West Africa

Summary Termite mound densities in typical guinea savanna, Detarium, and grassland (boval) habitats in northern guinea savanna were determined by random quadratting of 2–3 sites in each habitat (100, 10x10 m quadrats per habitat). Dominant species in guinea savanna were T. geminatus (46 mounds ha^-1) and T. oeconomus (21 mounds ha^-1), in Detarium T. geminatus (59 mounds ha^-1) and C. curtatus (45 mounds ha^-1) and in boval C. curtatus (72 mounds ha^-1) and T. geminatus (22 mounds ha^-1). Only C. curtatus densities and total densities differed significantly between sites within habitats, but all species differed significantly in abundance between habitats. The composition of each community was related to general environment but no particular environmental variable was shown to be a major determinant of termite distribution. Evidence for the limitation of termite populations was obtained from indirect evidence of competition between colonies in Detarium, and by experimental manipulation of fire regimes in the typical guinea savanna habitat. Harvester termites increased four-five fold over two years in fire-protected plots as a result of increased food supplies. Total termite densities in the fire-protected community equilibrated to the new population density (100 mounds ha^-1) after only two-three years.     

Termite mounds as browsing hotspots: an exception to the rule

In African savannas, termite mounds usually serve as browsing hotspots for mammals because of their soil fertility. Van der Plas et al., in this issue, describe that browsers avoid the unpalatable, evergreen tree species on mounds of Macrotermes natalensis in a mesic savanna, preferring mainly leguminous species with high leaf N and P concentrations in the matrix. This exception is probably a consequence of the fertile soils of the study area, and highlights the importance of environmental context for assessing ecological interactions.     

Strong positive effects of termites on savanna bird abundance and diversity are amplified by large herbivore exclusion

Vast areas of the African savanna landscapes are characterized by tree‐covered Macrotermes termite mounds embedded within a relatively open savanna matrix. In concert with termites, large herbivores are important determinants of savanna woody vegetation cover. The relative cover of woody species has considerable effects on savanna function. Despite the potentially important ecological relationships between termite mounds, woody plants, large herbivores, and birds, these associations have previously received surprisingly little attention. We experimentally studied the effects of termites and large herbivores on the avian community in Lake Mburo National Park, Uganda, where woody vegetation is essentially limited to termite mounds. Our experiment comprised of four treatments in nine replicates; unfenced termite mounds, fenced mounds (excluding large mammals), unfenced adjacent savanna, and fenced savanna. We recorded species identity, abundance, and behavior of all birds observed on these plots over a two‐month period, from late dry until wet season. Birds used termite mounds almost exclusively, with only 3.5% of observations occurring in the treeless intermound savanna matrix. Mean abundance and species richness of birds doubled on fenced (large herbivores excluded) compared to unfenced mounds. Feeding behavior increased when large mammals were excluded from mounds, both in absolute number of observed individuals, and relative to other behaviors. This study documents the fundamental positive impact of Macrotermes termites on bird abundance and diversity in an African savanna. Birds play crucial functional roles in savanna ecosystems, for example, by dispersing fruits or regulating herbivorous insect populations. Thus, the role of birds in savanna dynamics depends on the distribution and abundance of termite mounds.     

Effects of Erosion from Mounds of Different Termite Genera on Distinct Functional Grassland Types in an African Savannah

A key aspect of savannah vegetation heterogeneity is mosaics formed by two functional grassland types, bunch grasslands, and grazing lawns. We investigated the role of termites, important ecosystem engineers, in creating high-nutrient patches in the form of grazing lawns. Some of the ways termites can contribute to grazing lawn development is through erosion of soil from aboveground mounds to the surrounding soil surface. This may alter the nutrient status of the surrounding soils. We hypothesize that the importance of this erosion varies with termite genera, depending on feeding strategy and mound type. To test this, we simulated erosion by applying mound soil from three termite genera (Macrotermes, Odontotermes, and Trinervitermes) in both a field experiment and a greenhouse experiment. In the greenhouse experiment, we found soils with the highest macro nutrient levels (formed by Trinervitermes) promoted the quality and biomass of both a lawn (Digitaria longiflora) and a bunch (Sporobolus pyramidalis) grass species. In the field we found that soils with the highest micro nutrient levels (formed by Macrotermes) showed the largest increase in cover of grazing lawn species. By linking the different nutrient availability of the mounds to the development of different grassland states, we conclude that the presence of termite mounds influences grassland mosaics, but that the type of mound plays a crucial role in determining the nature of the effects.     

Influence of vegetation and soil types on the wheatbelt termite,Drepanotermes tamminensis (Hill), in the Western Australian wheatbelt

A survey of the distribution and density of mounds of the harvester termite,Drepanotermes tamminensis (Hill), was carried out in the Durokoppin Nature Reserve, Western Australia in 1990. Vegetation and, to a lesser extent, soil type, appear to be important factors in determining density and distribution of termite mounds within the Reserve. A more detailed study of mounds in Wandoo (Eucalyptus capillosa) woodland and Casuarina (Allocasuarina campestris) shrubland indicated that the total number and size of mounds were significantly higher in the woodland than in the shrubland. The total wet weight biomass ofD. tamminensis was calculated as 3.74 gm⁻² (37.4 kg ha⁻¹) in the woodland and 1.69 gm⁻² (16.9 kg ha⁻¹) in the shrubland. Thus, of the two favored habitats, Wandoo woodland appears to be more optimal for this termite species than the Casuarina shrubland.     

The Importance of Termites to the CH<Subscript>4</Subscript> Balance of a Tropical Savanna Woodland of Northern Australia

Termites produce methane (CH₄) as a by-product of microbial metabolism of food in their hindguts, and are one of the most uncertain components of the regional and global CH₄ exchange estimates. This study was conducted at Howard Springs near Darwin, and presents the first estimate of CH₄ emissions from termites based on replicated in situ seasonal flux measurements in Australian savannas. Using measured fluxes of CH₄ between termite mounds and the atmosphere, and between soil and the atmosphere across seasons we determined net CH₄ flux within a tropical savanna woodland of northern Australia. By accounting for both mound-building and subterranean termite colony types, and estimating the contribution from tree-dwelling colonies it was calculated that termites were a CH₄ source of +0.24 kg CH₄-C ha⁻¹ y⁻¹ and soils were a CH₄ sink of −1.14 kg CH₄-C ha⁻¹ y⁻¹. Termites offset 21% of CH₄ consumed by soil resulting in net sink strength of −0.90 kg CH₄-C ha⁻¹ y⁻¹ for these savannas. For Microcerotermes nervosus (Hill), the most abundant mound-building termite species at this site, mound basal area explained 48% of the variation in mound CH₄ flux. CH₄ emissions from termites offset 0.1% of the net biome productivity (NBP) and CH₄ consumption by soil adds 0.5% to the NBP of these tropical savannas at Howard Springs.     

Effect of landscape position on the acidification of loess‐derived soils under Pinus radiata

Abstract Interactions between landscape position and the acidifying effect of trees planted into loess‐derived grassland soils were studied in the Ventana region of Argentina. Forests of Pinus radiata planted at the end of 1940, were selected in two different positions from the landscape, plains and slopes. Samples of the soil surface mineral horizon were taken from landscape positions at four distances from the trees and compared with grassland soils. The values of the main soil chemical properties changed significantly with distance from trees, with a decrease in pH, base saturation, exchangeable Ca²⁺, Mg²⁺ and K⁺, and increase in Na⁺, Al³⁺ and particularly H⁺ closer to the trees. This pattern confirms the prominent role of vegetation in bringing about changes in soil properties. Regression models showed high levels of explanation (r² > 0.85) indicating that a high percentage of the spatial variability of soil chemical properties is systematic and predictable with distance from the trees. The pH in KCl proved an excellent tool for predicting the cationic composition of soils. Organic carbon and total nitrogen were significantly higher in the plains positions than in the slopes under the trees, whereas there was no difference under grassland. The slopes of the regression lines indicated that acidification is more intense in soils on the plains. Vegetation was the main factor influencing acidification of the studied soils. The landscape position regulates the bio‐hydrological factor and thus the speed of acidification process.     

Soil CO2 Emissions Associated with Termitaria in Tropical Savanna: Evidence for Hot-Spot Compensation

Our understanding of carbon (C) dynamics within savannas is very limited, especially how source/sink dynamics are influenced by the resident biota. Previous measurements of epigeal termite mounds (termitaria), ubiquitous in many savannas, have shown that they are considerable point sources of soil carbon dioxide (CO₂), whereas CO₂ measurements collected outside the mounds were generally assumed to be independent of termite activity. However, no measurements were conducted along gradients away from the mounds to confirm this. We quantified daytime soil CO₂ emissions (soil respiration) along gradients from the center to 20?m from the mound edge in Serengeti National Park, and measured soil temperature/moisture, macro-invertebrate abundance, and vegetation height as variables potentially influencing these emissions. Further, we quantified how far into the savanna termitaria impact CO₂ emissions. As in other studies, we found the highest soil CO₂ fluxes at the termitaria-center and considerably lower fluxes in the surrounding savanna. Macro-invertebrate abundance was associated with the differences in emissions measured, whereas the other variables were not. The analysis of spatial autocorrelation revealed significantly lower fluxes between the termitaria edge and up to 9?m from the edge compared to the values measured at the termitaria-center and between 10 and 20?m from the termitaria edge. When extrapolating the emissions across the landscape our results suggest that the lower CO₂ emissions found between the edge and 9?m fully compensate for the high fluxes measured at the termitaria center. Consequently, our findings provide evidence that termitaria might influence the savanna C source-sink dynamics differently than previously thought.