Termite mounds alter the spatial distribution of African savanna tree species

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.

[1]  S. Moe,et al.  Termitaria as browsing hotspots for African megaherbivores in miombo woodland , 2004, Journal of Tropical Ecology.

[2]  P. Jouquet,et al.  Association between subterranean termites and grasses in a West African savanna: spatial pattern analysis shows a significant role for Odontotermes n. pauperans , 2004 .

[3]  Gregory Asner,et al.  Improving Remote Species Identification through Efficient Training Data Collection , 2014, Remote. Sens..

[4]  S. Moe,et al.  Mound building termites contribute to savanna vegetation heterogeneity , 2009, Plant Ecology.

[5]  J. Féret,et al.  Landscape-scale variation in plant community composition of an African savanna from airborne species mapping. , 2014, Ecological applications : a publication of the Ecological Society of America.

[6]  Han Olff,et al.  Functional traits of trees on and off termite mounds: Understanding the origin of biotically-driven heterogeneity in savannas , 2013 .

[7]  G. Asner,et al.  Variable effects of termite mounds on African savanna grass communities across a rainfall gradient , 2014 .

[8]  Gregory Asner,et al.  Mapping Savanna Tree Species at Ecosystem Scales Using Support Vector Machine Classification and BRDF Correction on Airborne Hyperspectral and LiDAR Data , 2012, Remote. Sens..

[9]  M. Lepage,et al.  Impact of Macrotermes termitaria as a source of heterogeneity on tree diversity and structure in a Sudanian savannah under controlled grazing and annual prescribed fire (Burkina Faso) , 2008 .

[10]  Kelly K. Caylor,et al.  Termite mounds can increase the robustness of dryland ecosystems to climatic change , 2015, Science.

[11]  Mulualem Tigabu,et al.  Macrotermes mounds as sites for tree regeneration in a Sudanian woodland (Burkina Faso) , 2008, Plant Ecology.

[12]  T. Eid,et al.  Termites facilitate and ungulates limit savanna tree regeneration , 2012, Oecologia.

[13]  R. Holdo,et al.  Termite Mounds as Nutrient-Rich Food Patches for Elephants1 , 2004 .

[14]  Escaping the flames: large termitaria as refugia from fire in miombo woodland , 2013, Landscape Ecology.

[15]  Gudeta W. Sileshi,et al.  Termite-induced heterogeneity in African savanna vegetation: mechanisms and patterns , 2010 .

[16]  R. Scholes,et al.  Tree-grass interactions in Savannas , 1997 .

[17]  J. Cromsigt,et al.  Restoration of a megaherbivore: landscape‐level impacts of white rhinoceros in Kruger National Park, South Africa , 2014 .

[18]  Karen Hahn,et al.  The influence of termite-induced heterogeneity on savanna vegetation along a climatic gradient in West Africa , 2012, Journal of Tropical Ecology.

[19]  J. F. Hope-Simpson,et al.  Tropical Soil-Vegetation Catenas and Mosaics: A Study in the South-Western Part of the Anglo-Egyptian Sudan , 1948 .

[20]  Kurt Hornik,et al.  Misc Functions of the Department of Statistics (e1071), TU Wien , 2014 .

[21]  D. Doak,et al.  Termites create spatial structure and govern ecosystem function by affecting N2 fixation in an East African savanna. , 2010, Ecology.

[22]  Roberta E. Martin,et al.  Large-scale impacts of herbivores on the structural diversity of African savannas , 2009, Proceedings of the National Academy of Sciences.

[23]  F. Woodward,et al.  The global distribution of ecosystems in a world without fire. , 2004, The New phytologist.

[24]  J. Muvengwi,et al.  Termite mounds may not be foraging hotspots for mega-herbivores in a nutrient-rich matrix , 2013, Journal of Tropical Ecology.

[25]  G. Asner,et al.  The spatial extent of termite influences on herbivore browsing in an African savanna , 2010 .

[26]  R. Naiman,et al.  Large Animals and System-Level Characteristics in River Corridors Implications for river management , 1997 .

[27]  Kena Fox-Dobbs,et al.  Termites, vertebrate herbivores, and the fruiting success of Acacia drepanolobium. , 2010, Ecology.

[28]  S. Moe,et al.  Large herbivores maintain termite-caused differences in herbaceous species diversity patterns. , 2012, Ecology.

[29]  Christian Hartmann,et al.  Influence of termites on ecosystem functioning. Ecosystem services provided by termites , 2011 .

[30]  J. Ellis,et al.  Landscape and climatic control of woody vegetation in a dry tropical ecosystem: Turkana District, Kenya , 1993 .

[31]  N. Grimm,et al.  Nutrient Vectors and Riparian Processing: A Review with Special Reference to African Semiarid Savanna Ecosystems , 2007, Ecosystems.

[32]  H. Prins,et al.  HERBIVORE POPULATION CRASHES AND WOODLAND STRUCTURE IN EAST-AFRICA , 1993 .

[33]  Graeme S. Cumming,et al.  Do the large termite mounds of Macrotermes concentrate micronutrients in addition to macronutrients in nutrient-poor African savannas? , 2014 .

[34]  O. Chadwick,et al.  Impact of rainfall and topography on the distribution of clays and major cations in granitic catenas , 2011 .

[35]  Effects of Erosion from Mounds of Different Termite Genera on Distinct Functional Grassland Types in an African Savannah , 2011, Ecosystems.

[36]  R. Scholes,et al.  An African Savanna: The key determinants: water, nutrients, fire and herbivory , 1993 .

[37]  Ty Kennedy-Bowdoin,et al.  Regional insight into savanna hydrogeomorphology from termite mounds. , 2010, Nature communications.

[38]  Kelly K. Caylor,et al.  Determinants of woody cover in African savannas , 2005, Nature.

[39]  Gregory P. Asner,et al.  Spatial variability and abiotic determinants of termite mounds throughout a savanna catchment , 2014 .

[40]  D. Doak,et al.  Abstracts, Reviews, and Meetings , 2011, Ecological Restoration.

[41]  W. Ellery,et al.  The mound-building termite Macrotermes michaelseni as an ecosystem engineer , 1998, Journal of Tropical Ecology.

[42]  S. Moe,et al.  Termitaria are focal feeding sites for large ungulates in Lake Mburo National Park, Uganda , 2005 .

[43]  Roberta E. Martin,et al.  Carnegie Airborne Observatory: in-flight fusion of hyperspectral imaging and waveform light detection and ranging for three-dimensional studies of ecosystems , 2007 .

[44]  Graeme S. Cumming,et al.  Termite mounds as islands: woody plant assemblages relative to termitarium size and soil properties , 2013 .

[45]  Graeme S. Cumming,et al.  Termite Mounds Increase Functional Diversity of Woody Plants in African Savannas , 2014, Ecosystems.

[46]  R. Scholes,et al.  An African Savanna: Community and landscape pattern and change , 1993 .

[47]  N. Owen‐Smith,et al.  Ecology of grazing lawns in Africa , 2015, Biological reviews of the Cambridge Philosophical Society.