Termite mounds as islands: woody plant assemblages relative to termitarium size and soil properties

Questions: We investigated whether soils of small mounds resembled large mound or matrix soils, whether changes in plant composition reflected changes in soils, and the sequence in which plants colonize and disappear from mounds of increasing size. Location: Miombo woodland in northwest Zimbabwe. Methods: Macrotermitinae termitaria vary in size and soil nutrient concentrations, harbouring distinct woody plant assemblages, making them foci for plant and animal diversity, and also influencing primary, secondary and tertiary productivity. In spite of the importance of termitaria to heterogeneity and diversity, no studies have investigated changes in plant species assemblages as mound surface area increases to the point where mound vegetation is distinct from that of the matrix. We compared woody plant assemblages on 43 matrix plots with 95 Macrotermes termitaria across a range of surface areas, using ANOSIM, cluster analysis and MDS ordination. We compared soil nutrients, pH and clay, from ten large and ten small termitaria, and ten matrix sites. We also assessed how relative representation of large mound or matrix indicator species changed with mound area. Results: Change was apparent even at mound sizes of >10 m2, where both soils and plant assemblages on mounds were significantly different to those of the matrix. Plant assemblages fell into two main groups at 20% similarity; the first comprised of matrix plots, mounds 30 m2. At 40% similarity, four groups emerged: matrix, mounds 30 m2. Woody plant composition changed gradually as mound area increased. On termitaria 30 m2 in surface area, only mound indicator species were found. Conclusions: Through termite activities in concentrating nutrients and clay, termitaria provide habitat for species usually excluded from the matrix. The process of mound building and the nature of the plants that establish on them seem to establish a positive feedback for establishment of other non-woodland matrix species.

[1]  Estimating woody biomass in sub-saharan Africa; Evaluation de la biomasse ligneuse en afrique subsaharienne , 1994 .

[2]  L. Tieszen,et al.  Stable isotope analysis of termite food habits in East African grasslands , 1983, Oecologia.

[3]  P. Legendre,et al.  SPECIES ASSEMBLAGES AND INDICATOR SPECIES:THE NEED FOR A FLEXIBLE ASYMMETRICAL APPROACH , 1997 .

[4]  A. Korsaeth,et al.  Modelling the competition for nitrogen between plants and microflora as a function of soil heterogeneity , 2001 .

[5]  R. Paine,et al.  Disturbance, patch formation, and community structure. , 1974, Proceedings of the National Academy of Sciences of the United States of America.

[6]  K. R. Clarke,et al.  Non‐parametric multivariate analyses of changes in community structure , 1993 .

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

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

[9]  J. Proctor,et al.  Tropical Ecology and Physical Edaphology. , 1986 .

[10]  J. M. Moore,et al.  Heuweltjies (earth mounds) in the Clanwilliam district, Cape Province, South Africa: 4000-year-old termite nests , 1991, Oecologia.

[11]  A. Groengroeft,et al.  Fungus culturing, nutrient mining and geophagy: a geochemical investigation of Macrotermes and Trinervitermes mounds in southern Africa , 2009 .

[12]  Robert J. Scholes,et al.  The influence of soil fertility on the ecology of southern African dry savannas , 1990 .

[13]  J. Watson THE USE OF MOUNDS OF THE TERMITE MACROTERMES FALCIGER (GERSTÄCKER) AS A SOIL AMENDMENT , 1977 .

[14]  C. Canham,et al.  Forest Gaps and Isolated Savanna Trees , 1994 .

[15]  J. Midgley,et al.  Stable isotope and 14C study of biogenic calcrete in a termite mound, Western Cape, South Africa, and its palaeoenvironmental significance , 2009, Quaternary Research.

[16]  J. Leonard,et al.  The Vegetation of Africa , 1984 .

[17]  U. Bloesch Thicket clumps: A characteristic feature of the Kagera savanna landscape, East Africa , 2008 .

[18]  B. Huntley Southern African Savannas , 1982 .

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

[20]  T. Young,et al.  Long-Term Glades in Acacia Bushland and Their Edge Effects in Laikipia, Kenya , 1995 .

[21]  P. Hesse A CHEMICAL AND PHYSICAL STUDY OF THE SOILS OF TERMITE MOUNDS IN EAST AFRICA , 1955 .

[22]  S. V. Wieren,et al.  Grazing ungulates select for grasses growing beneath trees in African savannas , 2011 .

[23]  K. R. Clarke,et al.  Change in marine communities : an approach to statistical analysis and interpretation , 2001 .

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

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

[26]  D. DeAngelis,et al.  Competition and Coexistence: The Effects of Resource Transport and Supply Rates , 1994, The American Naturalist.

[27]  X. Le Roux,et al.  Influence of large termitaria on soil characteristics, soil water regime, and tree leaf shedding pattern in a West African savanna , 1999, Plant and Soil.

[28]  J. Lawton,et al.  Organisms as ecosystem engineers , 1994 .

[29]  M. Westoby,et al.  Nutrient costs of vertebrate- and ant-dispersed fruits , 1993 .

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

[31]  P. Fleming,et al.  Miombo woodland termite mounds: resource islands for small vertebrates? , 2003 .

[32]  Nathan J B Kraft,et al.  Functional Traits and Niche-Based Tree Community Assembly in an Amazonian Forest , 2008, Science.

[33]  M. Kellman Soil Enrichment by Neotropical Savanna Trees , 1979 .

[34]  F. van Langevelde,et al.  Soil clay content and fire frequency affect clustering in trees in South African savannas , 2008, Journal of Tropical Ecology.

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

[36]  E. Chidumayo,et al.  Changes in miombo woodland structure under different land tenure and use systems in central Zambia , 2002 .

[37]  G. Cumming,et al.  Large termitaria act as refugia for tall trees, deadwood and cavity-using birds in a miombo woodland , 2011, Landscape Ecology.

[38]  A. Milewski The Occurrence of Seeds and Fruits Taken by Ants Versus Birds in Mediterranean Australia and Southern Africa, in Relation to the Availability of Soil Potassium , 1982 .

[39]  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 .

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

[41]  K. R. Clarke,et al.  Statistical Design And Analysis For A Biological Effects Study , 1988 .

[42]  S. Milton,et al.  Large trees, fertile islands, and birds in arid savanna , 1999 .

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

[44]  R. Macarthur,et al.  The Theory of Island Biogeography , 1969 .

[45]  Andrew C. Millington,et al.  Estimating Woody Biomass in Sub-Saharan Africa , 1994 .

[46]  D. Wardle,et al.  Spatial soil ecology , 2002 .