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Andrew B Davies - One of the best experts on this subject based on the ideXlab platform.
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Termite Mounds create heterogeneity in invertebrate communities across a savanna rainfall gradient
Biodiversity and Conservation, 2020Co-Authors: Monica Leitner, Catherine L. Parr, Andrew B Davies, Mark P Robertson, Brigitte Janse Van RensburgAbstract:Termite Mounds create nutrient hotspots that serve as key resource areas for savanna vegetation and mammalian herbivores. However, despite the key ecological roles performed by Termite Mounds, few studies have investigated their influence on invertebrate communities, and none have examined such effects across environmental gradients. We hypothesised that Termite Mounds would support greater numbers of invertebrates than the surrounding savanna matrix and that assemblages would differ in composition due to the enhanced nutritional quality of vegetation on Mounds. We also predicted that the differences between on-mound and off-mound invertebrate diversity would be more pronounced in areas where the difference in nutritional value between Mounds and the savanna matrix vegetation was most prominent. We tested these hypotheses in Kruger National Park, South Africa, by sampling ground- and grass-dwelling invertebrate herbivores, omnivores and detritivores on and at various distances away from Termite Mounds at three savanna sites of varying vegetation quality across a rainfall gradient. All invertebrate groups sampled responded to Termite Mounds, but mound influence varied across trophic groups (Orthoptera showed the clearest patterns), diversity measures (changes in abundance rather than species richness) and with mean annual rainfall (strongest effects at the highest rainfall site). Orthoptera were more abundant on Mounds, particularly at the wettest site, and there was a positive relationship between mound size and Orthoptera species richness. Ant assemblage composition on Mounds differed from that off Mounds and beetle abundance was greater on Mounds, possibly as a result of concentrated mammalian herbivore activity and faecal deposition on Mounds. Our results suggest that Termite Mounds are not only important nutrient and foraging hotspots for vertebrates, but that they also introduce heterogeneity in invertebrate communities, especially in nutrient-poor savannas.
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Are Termite Mounds Always Grazing Hotspots? Grazing Variability with Mound Size, Season and Geology in an African Savanna
Ecosystems, 2018Co-Authors: Justice Muvengwi, Ed T.f. Witkowski, Francesca Parrini, Andrew B DaviesAbstract:Foraging site selection by large herbivores is influenced by multiple factors varying across landscapes and spatial scales. Termite Mounds harbour highly nutritious plants compared with the savanna matrix, making them preferred foraging patches in many savannas. However, it is unknown whether Termite Mounds equally influence herbivore grazing intensity across geological substrates and mound sizes. These knowledge gaps hamper our ability to draw general trans-ecosystem conclusions about the effect of Termite Mounds for savanna herbivores. We measured grazing intensity on Mounds of three different size classes (small, medium and large) across two geologies with differing soil nutrition (granite and basalt) in Gonarezhou National Park, Zimbabwe. We recorded measurements across three seasons (hot wet, cool dry and hot dry), and at multiple distances from Mounds. Grazing intensity on Mounds was higher on nutrient-poor granite than nutrient-rich basalt, and Termite Mounds of all sizes had a significant effect on grazing on granite during the cool dry season. Grazing was highest on large Mounds on both geologies throughout the year. Large Mounds also had the largest spatial influence on grazing in the cool dry season, up to 8 m beyond the mound edge on granite and 2 m on basalt. When scaled up to the landscape level, Mounds influenced about 15% of the granite landscape, but only about 0.5% of the basalt landscape. Our results show that the positive effects of Mounds on grazing intensity were pronounced on nutrient-poor soils but negligible on nutrient-rich soils, and that the magnitude of these effects varied across seasons and with mound size.
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Geology drives the spatial patterning and structure of Termite Mounds in an African savanna
Ecosphere, 2018Co-Authors: Justice Muvengwi, Andrew B Davies, Francesca Parrini, Ed T.f. WitkowskiAbstract:Termite Mounds perform important roles in savanna ecosystems, generating heterogeneity and influencing ecosystem processes across multiple trophic levels. However, the influence the environment and neighboring Termite colonies have on mound spatial patterning and structure is poorly understood, despite the profound implications such dynamics can have on ecosystems. To better understand these drivers, we mapped the spatial distribution and size of active and inactive Macrotermes Mounds in eight 1-km2 plots on contrasting geologies, nutrient-rich granite and nutrient-poor basalt, in a semi-arid Zimbabwean savanna. Although mound density was not significantly different between basalt (5.5 Mounds/ha) and granite (6.1 Mounds/ha), Termite mound structural attributes and spatial distribution patterns varied greatly between geologies. Mound size distributions differed between the geologies and Mounds were 2.6 times taller and 3.9 times wider and had 15 times greater lateral surface area on granite. Subsequently, 6% of the total landscape was covered by Mounds on granite compared with only 0.4% on basalt. On granite, large Mounds exhibited significant over-dispersion at scales below 30 m, signifying density-dependent thinning. Furthermore, small Mounds were clustered around large Mounds, likely a result of the budding of new colonies comprising fully fledged castes less vulnerable to competition. In contrast, random patterning was evident on comparably homogenous basalt. Our results demonstrate the powerful influence geological substrate has on mound spatial patterning and structure, suggesting that the importance of Termite Mounds for ecosystem functioning is more pronounced on nutrient-poor granitic substrates than basalts because of the pronounced over-dispersion, which maximizes mound production per unit area, and much larger mound sizes here.
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Termite Mounds vary in their importance as sources of vegetation heterogeneity across savanna landscapes
Journal of Vegetation Science, 2017Co-Authors: Justice Muvengwi, Andrew B Davies, Ed T.f. Witkowski, Francesca ParriniAbstract:Questions Termite Mounds are well known to host a suite of unique plants compared with the surrounding savanna matrix. However, most studies testing the significance of Mounds for ecosystem heterogeneity have been conducted at single sites. Mound effects on savanna heterogeneity across varying landscapes are less well understood, and how effects might vary across geological types and Mounds of different sizes is as yet unknown. Location Gonarezhou National Park, Zimbabwe. Methods We studied effects of Termite Mounds on vegetation spatial heterogeneity across two geologies (granite and basalt), including effects of mound size and the spatial extent of Termite influence. Herbaceous vegetation was sampled on Mounds and savanna matrix plots, and along distance transects away from Mounds. Soil nutrients on Mounds and in the matrix were also compared between geologies. Results Soil nutrients were more concentrated in large Mounds compared with the matrix on granite, but not on basalt, with Mounds therefore acting as nutrient hot-spots on nutrient-poor granite only. Large and medium sized Mounds hosted compositionally different grass species to the matrix on granite, but not on basalt. Large Mounds on granite also had significantly lower grass and forb species richness compared with the matrix. However, small Mounds on granite, and all mound size categories on basalt, did not have an effect on grass and forb species richness or assemblage composition, an observation attributed to a lack of difference in soil nutrients between the Mounds and matrix. Conclusion Our study shows that the significance of Termite Mounds to ecosystem spatial heterogeneity is strongly influenced by geology and mound size, with mound effects on herbaceous plant species heterogeneity more pronounced in dystrophic geologies and around large Mounds. Future studies should take greater cognisance of landscape context and mound size when seeking to understand the contribution of Termite Mounds to ecosystem structure and function. This article is protected by copyright. All rights reserved.
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Termite Mounds alter the spatial distribution of African savanna tree species
Journal of Biogeography, 2015Co-Authors: Andrew B Davies, Claire A. Baldeck, Gregory P. AsnerAbstract:Aim Termite Mounds form small islands of enhanced water and soil nutrient availability on otherwise dry and nutrient-poor hill crests, which can have important impacts on the plant community. However, the way in which Termite Mounds alter the spatial distribution of particular tree species across broad savanna landscapes is poorly understood. We aimed to understand the nature and extent of the relationship between Termite Mounds and key woody savanna species at landscape scales through the use of airborne remote sensing. Location Kruger National Park, South Africa. Methods We mapped 9894 Termite Mounds and 666,679 savanna trees from 15 species across two landscapes with contrasting rainfall regimes using airborne imaging spectroscopy and LiDAR data. We then examined changes in tree species densities and community composition with respect to distance from Termite Mounds. Results In both landscapes, Termite Mounds reduced overall tree densities over distances up to 10 m from mound centres. However, the effect of Termite Mounds on tree density differed among species, with some species, typically associated with lowland and riparian habitats, showing increased density near Termite Mounds. Indeed, changes in overall tree community composition revealed that Termite Mounds harbour tree communities similar to lowland communities, with this similarity decreasing with increased distance from the nearest mound. Termite effects were more pronounced in the savanna landscape receiving higher annual rainfall, whereas a greater percentage of the landscape was affected in the drier landscape due to higher mound densities. Main conclusions Termite Mounds mediate the spatial distribution of tree species in savanna landscapes, increasing the abundance of tree species typically associated with lowland habitats. This contributes to the spatial heterogeneity of savanna vegetation within landscapes and the maintenance of savanna biodiversity.
Gregory P. Asner - One of the best experts on this subject based on the ideXlab platform.
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Termite Mounds alter the spatial distribution of African savanna tree species
Journal of Biogeography, 2015Co-Authors: Andrew B Davies, Claire A. Baldeck, Gregory P. AsnerAbstract:Aim Termite Mounds form small islands of enhanced water and soil nutrient availability on otherwise dry and nutrient-poor hill crests, which can have important impacts on the plant community. However, the way in which Termite Mounds alter the spatial distribution of particular tree species across broad savanna landscapes is poorly understood. We aimed to understand the nature and extent of the relationship between Termite Mounds and key woody savanna species at landscape scales through the use of airborne remote sensing. Location Kruger National Park, South Africa. Methods We mapped 9894 Termite Mounds and 666,679 savanna trees from 15 species across two landscapes with contrasting rainfall regimes using airborne imaging spectroscopy and LiDAR data. We then examined changes in tree species densities and community composition with respect to distance from Termite Mounds. Results In both landscapes, Termite Mounds reduced overall tree densities over distances up to 10 m from mound centres. However, the effect of Termite Mounds on tree density differed among species, with some species, typically associated with lowland and riparian habitats, showing increased density near Termite Mounds. Indeed, changes in overall tree community composition revealed that Termite Mounds harbour tree communities similar to lowland communities, with this similarity decreasing with increased distance from the nearest mound. Termite effects were more pronounced in the savanna landscape receiving higher annual rainfall, whereas a greater percentage of the landscape was affected in the drier landscape due to higher mound densities. Main conclusions Termite Mounds mediate the spatial distribution of tree species in savanna landscapes, increasing the abundance of tree species typically associated with lowland habitats. This contributes to the spatial heterogeneity of savanna vegetation within landscapes and the maintenance of savanna biodiversity.
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Termite Mounds differ in their importance for herbivores across savanna types, seasons and spatial scales
Oikos, 2015Co-Authors: Andrew B Davies, Mark P Robertson, Gregory P. Asner, Shaun R. Levick, Berndt J Van Rensburg, Catherine L. ParrAbstract:Herbivores do not forage uniformly across landscapes, but select for patches of higher nutrition and lower predation risk. Macrotermes Mounds contain higher concentrations of soil nutrients and support grasses of higher nutritional value than the surrounding savanna matrix, attracting mammalian grazers that preferentially forage on Termite mound vegetation. However, little is known about the spatial extent of such Termite influence on grazing patterns and how it might differ in time and space. We measured grazing intensity in three African savanna types differing in rainfall and foliar nutrients and predicted that the functional importance of Mounds for grazing herbivores would increase as the difference in foliar nutrient levels between mound and savanna matrix grasses increases and the Mounds become more attractive. We expected this to occur in nutrient-poor areas and during the dry season when savanna matrix grass nutrient levels are lower. Tuft use and grass N and P content were measured along transects away from Termite Mounds, enabling calculation of the spatial extent of Termite influence on mammalian grazing. Using Termite mound densities estimated from airborne light detection and ranging (LiDAR), we further upscaled field-based results to determine the percentage of the landscape influenced by Termite activity. Grasses in close proximity to Termite Mounds were preferentially grazed at all sites and in both seasons, but the strength of mound influence varied between savanna types and seasons. In the wet season, Mounds had a relatively larger effect on grazers at the landscape scale in the nutrient-poor, wetter savanna, whereas in the dry season the pattern was reversed with more of the landscape influenced at the nutrient-rich, driest site. Our results reveal that Termite Mounds enhance the value of savanna landscapes for herbivores, but that their functional importance varies across savanna types and seasons.
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spatial variability and abiotic determinants of Termite Mounds throughout a savanna catchment
Ecography, 2014Co-Authors: Andrew B Davies, Mark P Robertson, Gregory P. Asner, Shaun R. Levick, Berndt J Van Rensburg, Catherine L. ParrAbstract: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, and exhibited a clustered pattern throughout the landscape, occurring at relatively high densities (up to 2 Mounds ha) 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) 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.
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Variable effects of Termite Mounds on African savanna grass communities across a rainfall gradient
Journal of Vegetation Science, 2014Co-Authors: Andrew B Davies, Mark P Robertson, Gregory P. Asner, Shaun R. Levick, Berndt Jansen Van Rensburg, Catherine L. ParrAbstract:Questions: Termite Mounds of the genus Macrotermes are prominent features in African savannas, forming nutrient hotspots that support greater plant diversity, which is of higher nutritional value than the surrounding savanna matrix. However, little is known about grass communities on and around Mounds or how the functional importance of Mounds varies across sites. As mean annual rainfall increases, savannas in southern Africa become increasingly dystrophic through increased denitrification (including pyrodenitrification) and the leaching of soil nutrients. The functional importance of Mounds is concomitantly expected to increase as the difference in foliar nutrient levels between Mounds and the savanna matrix increases. We tested this prediction on grass communities across a rainfall gradient to: (i) determine the degree to which grass assemblages differ between Termite Mounds and the savanna matrix; (ii) determine the spatial extent to which Mounds influence grass communities; and (iii) investigate whether these patterns differ across savanna types.
Judith Korb - One of the best experts on this subject based on the ideXlab platform.
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‘Magnetic’ Termite Mounds: is their unique shape an adaptation to facilitate gas exchange and improve food storage?
Insectes Sociaux, 2014Co-Authors: A M Schmidt, Peter Jacklyn, Judith KorbAbstract:Social insects can build impressive nest Mounds but the functional significance of their architecture is rarely studied in experiments. The ‘magnetic’ Termite Mounds of monsoonal northern Australia built by Amitermes meridionalis are notable for their elongated wedge shape and north–south axial orientation. We tested whether the shape is an adaptation to facilitate gas exchange and the preservation of food stores by two experimental manipulations of Mounds in situ covering all seasons. First, Mounds were shaded to limit drying after rain and second, mound shape was amended from wedge to (approximate) sphere. Food storage, fungal contamination, and internal CO_2 concentration were unaffected by manipulation, but showed a distinct seasonal dynamic, with storage peaking towards the onset of rains and fungal load towards the end of the rainy season. Internal CO_2 concentrations were subject to a diurnal cycle, but also showed elevation during rains. We propose that one advantage of the wedge shape is the efficient use of building effort to achieve good passive ventilation for the food storage areas.
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Thermoregulation and ventilation of Termite Mounds
Naturwissenschaften, 2003Co-Authors: Judith KorbAbstract:Some of the most sophisticated of all animal-built structures are the Mounds of African Termites of the subfamily Macrotermitinae, the fungus-growing Termites. They have long been studied as fascinating textbook examples of thermoregulation or ventilation of animal buildings. However, little research has been designed to provide critical tests of these paradigms, derived from a very small number of original papers. Here I review results from recent studies on Macrotermes bellicosus that considered the interdependence of ambient temperature, thermoregulation, ventilation and mound architecture, and that question some of the fundamental paradigms of Termite Mounds. M. bellicosus achieves thermal homeostasis within the mound, but ambient temperature has an influence too. In colonies in comparably cool habitats, mound architecture is adapted to reduce the loss of metabolically produced heat to the environment. While this has no negative consequences in small colonies, it produces a trade-off with gas exchange in large colonies, resulting in suboptimally low nest temperatures and increased CO_2 concentrations. Along with the alteration in mound architecture, the gas exchange/ventilation mechanism also changes. While Mounds in the thermally appropriate savannah have a very efficient circular ventilation during the day, the ventilation in the cooler forest is a less efficient upward movement of air, with gas exchange restricted by reduced surface exchange area. These results, together with other recent findings, question entrenched ideas such as the thermosiphon-ventilation mechanism or the assumption that Mounds function to dissipate internally produced heat. Models trying to explain the proximate mechanisms of mound building, or building elements, are discussed.
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ventilation of Termite Mounds new results require a new model
Behavioral Ecology, 2000Co-Authors: Judith Korb, Karl Eduard LinsenmairAbstract:In 1955, Luscher proposed a ventilation mechanism for cathedral-shaped Termite Mounds to exchange respiratory gases. This mechanism was generally accepted, although it had never been tested critically. We tested this mechanism by investigating temperatures, CO2 concentrations, and air currents in and around two types of Macrotermes bellicosus Mounds: cathedral-shaped Mounds with many ridges and thin walls located in the savanna and dome-shaped Mounds without ridges and with thick walls in the forest. These two mound shapes have two different mechanisms of ventilation, depending on the environmental tem- perature. In the savanna during the day, sun heats the air in the peripheral air channels inside the ridges of the mound above the central nest temperatures and produces a temperature gradient in the peripheral air channels with decreased temperatures at the top of the mound. This temperature gradient leads to convection currents with air rising inside the air channels of the ridges to the top of the mound, meanwhile exchanging CO2. In contrast, in the savanna during the night and generally in the forest, the temperatures inside the air channels are lower than those of the central nest, and no air currents rising upward inside the air channels were detected. The CO2 concentrations in the air channels of savanna Mounds at night and forest Mounds in general were higher than during the day in the savanna. Therefore, our data do not support Luscher's proposed mechanism. Key words: carbon dioxide, gas exchange, Ivory Coast, Macrotermes bellicosus, metabolism, Termite Mounds, ther- moregulation. (Behav Ecol 11:486-494 (2000))
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the architecture of Termite Mounds a result of a trade off between thermoregulation and gas exchange
Behavioral Ecology, 1999Co-Authors: Judith Korb, Karl Eduard LinsenmairAbstract:We examined the influence of gas exchange on the architecture of Termite Mounds. In Comoe National Park (Cote d’Ivoire), Macrotermes bellicosus builds, as an adaptation to ambient temperature conditions, differently shaped Mounds in the shrub savanna and the gallery forest. Previous studies suggested that there might be a constraint that limits the degree of thermal insulation of the interior (i.e., nest) of the Mounds in environments with relatively low ambient temperatures. This factor causes, in proximate terms, suboptimal low nest temperatures and ultimately leads to reduced reproductive success in the gallery forest. In this study, we examined whether the necessity for gas exchange might constrain mound architecture. We measured CO2 concentrations in the air channels of Mounds in different habitats and under manipulated temperature regimes. During both the dry and the rainy season we found higher CO2 concentrations in Mounds of the gallery forest than in Mounds of the savanna. Additional measurements in forest Mounds, architecturally resembling those of the savanna due to an experimental increase in ambient temperatures, revealed lower CO2 concentrations than unmanipulated Mounds in this habitat. Generally, concentrations were higher during the rainy season compared to the dry season and lower during day than during night. Summarizing these results we present a model that illustrates this trade-off between thermoregulation and gas exchange under different temperature regimes. Both factors together result in different mound architectures under different environmental temperatures and may finally limit the distribution of this species.
Grant S. Joseph - One of the best experts on this subject based on the ideXlab platform.
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Microclimates mitigate against hot temperatures in dryland ecosystems: Termite Mounds as an example
Ecosphere, 2016Co-Authors: Grant S. Joseph, Colleen L. Seymour, Bernard W. T. Coetzee, Mduduzi Ndlovu, A. De La Torre, R. Suttle, N. Hicks, S. Oxley, Stefan H. FoordAbstract:Termite Mounds have recently been shown to protect against drought by providing refuges for plants and foci for revegetation, but whether Mounds modulate temperature remains untested. Organisms tend to experience climate at finer scales than those captured by models predicting how distributions alter with global change, so microclimates represent important “climate refuges.” Using data we collected from African savanna sites, generalized linear mixed-effects models and linear quantile regression analysis confirm for the first time that the woody species associated with large Termite Mounds establish microclimates that are significantly cooler than surrounding savannas, a cooling effect that is even greater at warmer extremes. As air temperatures approached 40°C, a cooling effect of up to 4°C occurred, representing a doubling from that observed at 34°C. African savannas encompass 10 million km2, and much of this harbors evenly dispersed termitaria. The temperature-modulating effect of Mounds facilitates agricultural and conservation decision-making as global change begins to impact the integrity of both human well-being and ecological processes.
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Termite Mounds increase functional diversity of woody plants in african savannas
Ecosystems, 2014Co-Authors: Grant S. Joseph, Colleen L. Seymour, Graeme S Cumming, David H M Cumming, Zacheus MahlanguAbstract:Fine-scale spatial heterogeneity influences biodiversity and ecosystem productivity at many scales. In savanna systems, Macrotermes Termites, through forming spatially explicit Mounds with unique woody plant assemblages, emerge as important sources of such heterogeneity. Despite a growing consensus regarding the importance of functional diversity (FD) to ecosystem processes, no study has quantified how Termite Mounds affect woody plant FD. We address whether Termite Mounds alter the distribution of functional traits, and increase FD of woody plant communities within Africa’s largest savanna woodland, the 2.7 million km2 miombo system. Using plant traits that change according to soil resources (for example, water and nutrients), and disturbance (for example, fire and elephant herbivory), we identified response functional groups and compared relative representation of these groups between mound and matrix habitats. We also asked whether mound and matrix habitats differed in their contribution to FD within the system. Although species representing most functional groups were found in both mound and matrix habitats, relative abundance of functional groups differed between mound and matrix. Mound plant assemblages had greater response diversity to soil resources than matrix plots, but there was no difference in response diversity to disturbance. High trait values on Mounds included tree height, leaf nitrogen, phosphorus, and palatability. Species with root ectomycorrhizae dominated the matrix. In conclusion, these small patches of nutrient-enriched substrate emerge as drivers of FD in above-ground woody plant communities.
Zacheus Mahlangu - One of the best experts on this subject based on the ideXlab platform.
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Termite Mounds increase functional diversity of woody plants in african savannas
Ecosystems, 2014Co-Authors: Grant S. Joseph, Colleen L. Seymour, Graeme S Cumming, David H M Cumming, Zacheus MahlanguAbstract:Fine-scale spatial heterogeneity influences biodiversity and ecosystem productivity at many scales. In savanna systems, Macrotermes Termites, through forming spatially explicit Mounds with unique woody plant assemblages, emerge as important sources of such heterogeneity. Despite a growing consensus regarding the importance of functional diversity (FD) to ecosystem processes, no study has quantified how Termite Mounds affect woody plant FD. We address whether Termite Mounds alter the distribution of functional traits, and increase FD of woody plant communities within Africa’s largest savanna woodland, the 2.7 million km2 miombo system. Using plant traits that change according to soil resources (for example, water and nutrients), and disturbance (for example, fire and elephant herbivory), we identified response functional groups and compared relative representation of these groups between mound and matrix habitats. We also asked whether mound and matrix habitats differed in their contribution to FD within the system. Although species representing most functional groups were found in both mound and matrix habitats, relative abundance of functional groups differed between mound and matrix. Mound plant assemblages had greater response diversity to soil resources than matrix plots, but there was no difference in response diversity to disturbance. High trait values on Mounds included tree height, leaf nitrogen, phosphorus, and palatability. Species with root ectomycorrhizae dominated the matrix. In conclusion, these small patches of nutrient-enriched substrate emerge as drivers of FD in above-ground woody plant communities.