Free Standard AU & NZ Shipping For All Book Orders Over $80!
Register      Login
Wildlife Research Wildlife Research Society
Ecology, management and conservation in natural and modified habitats
RESEARCH ARTICLE

The potential utility of rodents and other small mammals as indicators of ecosystem ‘integrity’ of South African grasslands

Nico Avenant
+ Author Affiliations
- Author Affiliations

National Museum and Centre for Environmental Management, University of the Free State, PO Box 266, Bloemfontein, 9301, South Africa. Email: navenant@nasmus.co.za

Wildlife Research 38(7) 626-639 https://doi.org/10.1071/WR10223
Submitted: 6 December 2010  Accepted: 1 September 2011   Published: 30 November 2011

Abstract

Context: The expansive grassland biome is one of the most extensively transformed in South Africa, yet no strategy for monitoring its integrity is in place. A grassland health program, incorporating different ecosystem levels, was recently initiated. The suitability of three taxonomic groups as indicators has been tested so far: vegetation (by calculating an ecological index value, El), insects (using the South African grassland scoring system, SAGraSS) and small mammals (this study). All of these methods aim to be rapid and easy to perform. Whereas SAGraSS still needs further refinement, several factors already indicate the importance of including small mammal community parameters in integrity assessments.

Aims: This contribution reports on more than 12 years of results from various studies on small mammals in the Free State Grasslands, with the aim of exploring the utility of small mammal survey for assessment of ecosystem integrity.

Methods: The hypothesis was based on the outcomes of several short-term studies conducted in the grassland biome. Combining all previous results, this paper re-evaluates the parameters of trap success, species richness, diversity, evenness and individual species as bio-indicators.

Key results: By combining data from many sites and years, the effect of seasonal and inter-annual variations in habitat and population parameters was diminished, and a more general picture of small mammal community structure revealed. New insights were gleaned into the status of several indicator species. By providing a summary of small mammal community parameter scores and indices, the study establishes a benchmark for future small mammal assessments and monitoring. To be effective, small mammal surveys in the grassland biome of southern Africa should be carried out in autumn and early winter.

Conclusions: This study suggests that small mammal species richness and diversity decline with habitat degradation; that the generalist rodent Mastomys coucha dominates community numbers at low ecological integrity; that the number of specialist species increases towards ecological climax; and that specific species act as indicators during the successional process.

Implications: This study should benefit the monitoring, conservation and management of grassland ecosystems, make environmental impact assessments more effective, and identify new topics for ecological research.

Additional keywords: community structure, diversity, ecological integrity, species richness.


References

Abramsky, Z. (1988). The role of habitat and productivity in structuring desert rodent communities. Oikos 52, 107–114.
The role of habitat and productivity in structuring desert rodent communities.Crossref | GoogleScholarGoogle Scholar |

Abramsky, Z., and Rosenzweig, M. L. (1984). Tilman’s predicted productivity-diversity relationships shown by desert rodent. Nature 309, 150–151.
Tilman’s predicted productivity-diversity relationships shown by desert rodent.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DyaL2c7ptFGhsQ%3D%3D&md5=52aee2e37e7951e86578466409bc1d04CAS |

Avenant, M. F. (2010). Challenges in using fish communities for assessing the ecological integrity of non-perennial rivers. Water S.A. 36, 397–405.

Avenant, N. L. (1996). Identification and distribution of two Mastomys spp. in Lesotho and part of South Africa. Navorsinge van die Nasionale Museum Bloemfontein 12, 49–58.

Avenant, N. L. (1997). Mammals recorded in the QwaQwa National Park (1994–1995). Koedoe 40, 31–40.

Avenant, N. L. (1998). Mammals. EIA, Maguga Dam, Swaziland. (Afridev Consultants: Darling, South Africa.)

Avenant, N. L. (2000a). Small mammal community characteristics as indicators of ecological disturbance in the Willem Pretorius Nature Reserve, Free State, South Africa. South African Journal of Wildlife Research 30, 26–33.

Avenant, N. L. (2000b). Terrestrial small-mammal diversity in Korannaberg Conservancy, Free State, South Africa. Navorsinge van die Nasionale Museum Bloemfontein 16, 69–82.

Avenant, N. L. (2002). Mammals. In ‘Biological Resource Monitoring’. pp. 81–91. (Ed. C. Mokuku.) (NULS-Consuls: Maseru, Lesotho.)

Avenant, N. L. (2003a). The use of small-mammal community characteristics as an indicator of ecological disturbance in the Korannaberg Conservancy. In ‘Rats, Mice and People: Rodent Biology and Management’. (Eds G. R. Singleton, L. A. Hinds, C. J. Krebs and D. M. Spratt.) pp. 95–98. (Australian Centre for International Agricultural Research: Canberra.)

Avenant, N. L. (2003b). Mammals. In ‘Faunal Rescue Program, Mohale’. (Ed. T. Moeti.) pp. 65–73. (National University of Lesotho: Roma, Lesotho.)

Avenant, N. L. (2004). Mammal Report. Submitted to UNDP, Lesotho, as part of the ‘Conserving Mountain Biodiversity in Southern Lesotho’ program.

Avenant, N. L. (2005). Barn owl pellets: a useful tool for monitoring small mammal communities? Belgian Journal of Zoology 135, 39–43.

Avenant, N. L., and Cavallini, P. (2007). Correlating rodent community structure with ecological integrity, Tussen-die-Riviere Nature Reserve, Free State Province, South Africa. Integrative Zoology 2, 212–219.
Correlating rodent community structure with ecological integrity, Tussen-die-Riviere Nature Reserve, Free State Province, South Africa.Crossref | GoogleScholarGoogle Scholar |

Avenant, N. L., and Kuyler, P. (2002). Small mammal diversity in the Maguga area, Swaziland. South African Journal of Wildlife Research 32, 101–108.

Avenant, N. L., and Watson, J. P. (2002). Mammals recorded in the Sandveld Nature Reserve, Free State province, South Africa. Navorsinge van die Nasionale Museum Bloemfontein 18, 1–12.

Avenant, N. L., Watson, J. P., and Schulze, E. (2008). Correlating small mammal community characteristics and ecosystem integrity in the Caledon Nature Reserve, South Africa. Mammalia 72, 186–191.
Correlating small mammal community characteristics and ecosystem integrity in the Caledon Nature Reserve, South Africa.Crossref | GoogleScholarGoogle Scholar |

Baker, S. C. (2006). A comparison of litter beetle assemblages (Coleoptera) in mature and recently clearfelled Eucalyptus obliqua forest. Australian Journal of Ecology 45, 130–136.

Beccaloni, G. W., and Gaston, K. J. (1995). Predicting species richness of neotropical forest butterflies – Ithomiinae (Lepidoptera: Nymphalidae) as indicators. Biological Conservation 71, 77–86.
Predicting species richness of neotropical forest butterflies – Ithomiinae (Lepidoptera: Nymphalidae) as indicators.Crossref | GoogleScholarGoogle Scholar |

Birney, E. C., Grant, W. C., and Baird, D. D. (1976). Importance of vegetative cover to cycles of Microtus populations. Ecology 57, 1043–1051.
Importance of vegetative cover to cycles of Microtus populations.Crossref | GoogleScholarGoogle Scholar |

Bronner, G., Rautenbach, I. L., and Meester, J. (1988). Environmental influence on reproduction in the Natal multimammate mouse Mastomys natalensis (A. Smith, 1834). South African Journal of Wildlife Research 18, 142–148.

Bultman, T., Uetz, G. W., and Brady, A. R. (1982). A comparison of cursorial spider communities along a successional gradient. The Journal of Arachnology 10, 23–33.

Cardinale, B. J., Nelson, K., and Palmer, M. A. (2000). Linking species diversity to the functioning of ecosystems: on the importance of environmental context. Oikos 91, 175–183.
Linking species diversity to the functioning of ecosystems: on the importance of environmental context.Crossref | GoogleScholarGoogle Scholar |

Carey, A. B., and Wilson, S. M. (2001). Induced spatial heterogeneity in forest canopies: responses of small mammals. The Journal of Wildlife Management 65, 1014–1027.
Induced spatial heterogeneity in forest canopies: responses of small mammals.Crossref | GoogleScholarGoogle Scholar |

Caro, T. M. (2001). Species richness and abundance of small mammals inside and outside an African national park. Biological Conservation 98, 251–257.
Species richness and abundance of small mammals inside and outside an African national park.Crossref | GoogleScholarGoogle Scholar |

Chapin, F. S., Zavaleta, E. S., Eviner, V. T., Naylor, R. L., Vitousek, P. M., Reynolds, H. L., Hooper, D. U., Lavorel, S., Sala, O. E., Hobbie, S. E., Mack, M. C., and Diaz, S. (2000). Consequences of changing biodiversity. Nature 405, 234–242.
Consequences of changing biodiversity.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD3cXjsFyjsL8%3D&md5=ecb217c98ec24ae15e56950036e75c79CAS |

Chutter, F. M. (1988). Research on the rapid biological assessment of water quality impacts in streams and rivers. Report No. 422/1/98. (Water Research Commission: Pretoria, South Africa.)

Cole, F. R., and Wilson, D. E. (1996). Mammalian diversity and natural history. In ‘Measuring and Monitoring Biological Diversity: Standard Methods for Mammals’. (Eds D. E. Wilson, F. R. Cole, J. D. Nichols, R. Rudran and M. S. Foster.) pp. 9–40. (Smithsonian Institution: Washington, DC.)

DEAT (2005). South Africa’s National Biodiversity Strategy and Action Plan. (Department of Environmental Affairs and Tourism: Pretoria, South Africa.)

DEAT (2006). South African Environment Outlook. A Report on the State of the Environment. (Department of Environmental Affairs and Tourism: Pretoria, South Africa.)

Ecke, F., Löfgren, O., and Sörlin, D. (2002). Population dynamics of small mammals in relation to forest age and structural habitat factors in northern Sweden. Journal of Applied Ecology 39, 781–792.
Population dynamics of small mammals in relation to forest age and structural habitat factors in northern Sweden.Crossref | GoogleScholarGoogle Scholar |

Ferreira, S. M., and Avenant, N. L. (2003). Influences of trap-spacing on descriptors of hypothetical small mammal communities in Free State grasslands. Navorsinge van die Nasionale Museum Bloemfontein 19, 21–30.

Ferreira, S. M., and Van Aarde, R. J. (1997). The chronosequence of rehabilitating stands of coastal dune forest: do small mammals confirm it? South African Journal of Science 93, 211–214.

Ferreira, S. M., and Van Aarde, R. J. (1999). Habitat associations and competition in MastomysSaccostomysAethomys assemblages on coastal dune forests. African Journal of Ecology 37, 121–136.
Habitat associations and competition in MastomysSaccostomysAethomys assemblages on coastal dune forests.Crossref | GoogleScholarGoogle Scholar |

Ferreira, S. M., and Van Aarde, R. J. (2000). Maintaining diversity through intermediate disturbances: evidence from rodents colonizing rehabilitating coastal dunes. African Journal of Ecology 38, 286–294.
Maintaining diversity through intermediate disturbances: evidence from rodents colonizing rehabilitating coastal dunes.Crossref | GoogleScholarGoogle Scholar |

Fonseca, C. R., and Ganade, G. (2001). Species functional redundancy, random extinctions and the stability of ecosystems. Journal of Ecology 89, 118–125.
Species functional redundancy, random extinctions and the stability of ecosystems.Crossref | GoogleScholarGoogle Scholar |

Fox, B. J. (1982). Fire and mammalian secondary succession in an Australian coastal heath. Ecology 63, 1332–1341.
Fire and mammalian secondary succession in an Australian coastal heath.Crossref | GoogleScholarGoogle Scholar |

Fox, B. J. (1990). Changes in the structure of mammal communities over successional time scales. Oikos 59, 321–329.
Changes in the structure of mammal communities over successional time scales.Crossref | GoogleScholarGoogle Scholar |

Fox, B. J., and Fox, M. D. (1984). Small mammal recolonization of open forest following sand mining. Australian Journal of Ecology 9, 241–252.
Small mammal recolonization of open forest following sand mining.Crossref | GoogleScholarGoogle Scholar |

Glennon, M. J., and Porter, W. F. (2007). Impacts of land-use management on small mammals in the Adirondack Park, New York. Northeastern Naturalist 14, 323–342.
Impacts of land-use management on small mammals in the Adirondack Park, New York.Crossref | GoogleScholarGoogle Scholar |

Griffiths, D. (1999). On investigating local-regional species richness relationships. Journal of Animal Ecology 68, 1051–1055.
On investigating local-regional species richness relationships.Crossref | GoogleScholarGoogle Scholar |

Grime, J. P. (1998). Benefits of plant diversity to ecosystems: immediate, filter and founder effects. Journal of Ecology 86, 902–910.
Benefits of plant diversity to ecosystems: immediate, filter and founder effects.Crossref | GoogleScholarGoogle Scholar |

Hastwell, G. T., and Huston, M. A. (2001). On disturbance and diversity: a reply to Mackey and Currie. Oikos 92, 367–371.
On disturbance and diversity: a reply to Mackey and Currie.Crossref | GoogleScholarGoogle Scholar |

Hoffmann, A., and Zeller, U. (2005). Influence of variations in land use intensity on species diversity and abundance of small mammals in the Nama Karoo, Namibia. Belgian Journal of Zoology 135, 91–96.

Johnson, K. H. (2000). Trophic-dynamic considerations in relating species diversity to ecosystem resilience. Biological Reviews of the Cambridge Philosophical Society 75, 347–376.
Trophic-dynamic considerations in relating species diversity to ecosystem resilience.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DC%2BD3cvosFemtw%3D%3D&md5=173ec951c4aa2f290a4531f51a258036CAS |

Jones, D. T., and Eggleton, P. (2000). Sampling termite assemblages in tropical forests: testing a rapid biodiversity assessment protocol. Journal of Applied Ecology 37, 191–203.
Sampling termite assemblages in tropical forests: testing a rapid biodiversity assessment protocol.Crossref | GoogleScholarGoogle Scholar |

Joubert, D. F., and Ryan, P. G. (1999). Differences in mammal and bird assemblages between commercial and communal rangelands in the Succulent Karoo, South Africa. Journal of Arid Environments 43, 287–299.
Differences in mammal and bird assemblages between commercial and communal rangelands in the Succulent Karoo, South Africa.Crossref | GoogleScholarGoogle Scholar |

Kaiser, W. (2005). The characteristics of insect and small mammal communities as a reflection of the ecological value of grasslands. Masters Thesis, University of the Free State, Bloemfontein, South Africa.

Kaiser, W., Avenant, N. L., and Haddad, C. R. (2009). Assessing the ecological integrity of a grassland ecosystem: refining the SAGraSS method. African Journal of Ecology 47, 308–317.
Assessing the ecological integrity of a grassland ecosystem: refining the SAGraSS method.Crossref | GoogleScholarGoogle Scholar |

Karr, J. R., Fausch, K. D., Angermeier, P. L., Yant, P. R., and Schlosser, I. J. (1986). Assessing biological integrity in running waters: a method and its rationale. Illinois Natural History Survey Special Publication 5.

Kern, N. G. (1981). The influence of fire on populations of small mammals of the Kruger National Park. Koedoe 24, 125–157.

Kirkland, G. L. (1990). Patterns of initial small mammal community change after clearcutting of temperate North American forests. Oikos 59, 313–320.
Patterns of initial small mammal community change after clearcutting of temperate North American forests.Crossref | GoogleScholarGoogle Scholar |

Kleynhans, C. J. (1999). The development of a fish index to assess the biological integrity of South African Rivers. Water S.A. 25, 265–278.

Klinger, R. (2006). The interaction of disturbances and small mammal community dynamics in a lowland forest in Belize. Journal of Animal Ecology 75, 1227–1238.
The interaction of disturbances and small mammal community dynamics in a lowland forest in Belize.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DC%2BD28ngtFalug%3D%3D&md5=d9afdf4eaa592c7aab4ccd8326be91fbCAS |

Kuyler, P. (2000). Veld condition assessment and small mammal community structure in the management of Soetdoring Nature Reserve, Free State, South Africa. Masters Thesis, University of the Free State, Bloemfontein, South Africa.

Leirs, H., Verhagen, R., Verheyen, W., Mwanjabe, P., and Mbise, T. (1996). Forecasting rodent outbreaks in Africa: an ecological basis for Mastomys control in Tanzania. Journal of Applied Ecology 33, 937–943.
Forecasting rodent outbreaks in Africa: an ecological basis for Mastomys control in Tanzania.Crossref | GoogleScholarGoogle Scholar |

Letnic, M., Dickman, C. R., Tischler, M. K., Tamayo, B., and Beh, C. L. (2004). The responses of small mammals and lizards to post-fire succession and rainfall in arid Australia. Journal of Arid Environments 59, 85–114.
The responses of small mammals and lizards to post-fire succession and rainfall in arid Australia.Crossref | GoogleScholarGoogle Scholar |

Linn, I. J. (1991). Influence of 6-methoxybenzoxazolinone and green vegetation on reproduction of the multimammate rat Mastomys coucha. South African Journal of Wildlife Research 21, 33–37.

Loreau, M. (2000). Biodiversity and ecosystem functioning: recent theoretical advances. Oikos 91, 3–17.
Biodiversity and ecosystem functioning: recent theoretical advances.Crossref | GoogleScholarGoogle Scholar |

Low, A. B., and Rebelo, A. G. (1996). ‘Vegetation of South Africa, Lesotho and Swaziland.’ (Department of Environmental Affairs and Tourism: Pretoria, South Africa.)

Lynch, C. D. (1994). The mammals of Lesotho. Navorsinge van die Nasionale Museum Bloemfontein 10, 177–241.

Magurran, A. E. (2004). ‘Measuring Biological Diversity.’ (Blackwell: Oxford.)

Majer, J. D. (1983). Ants: bio-indicators of minesite rehabilitation, land-use, and land conservation. Environmental Management 7, 375–383.
Ants: bio-indicators of minesite rehabilitation, land-use, and land conservation.Crossref | GoogleScholarGoogle Scholar |

McCann, K. S. (2000). The diversity-stability debate. Nature 405, 228–233.
The diversity-stability debate.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD3cXjsFyjs7k%3D&md5=d4a809f0462a245e26b913f4198ddc5bCAS |

McGeoch, M. A., Van Rensburg, B. J., and Botes, A. (2002). The verification and application of bioindicators: a case study of dung beetles in a savanna ecosystem. Journal of Applied Ecology 39, 661–672.
The verification and application of bioindicators: a case study of dung beetles in a savanna ecosystem.Crossref | GoogleScholarGoogle Scholar |

McGeoch, M. A. (1998). The selection, testing and application of terrestrial insects as bioindicators. Biological Reviews of the Cambridge Philosophical Society 73, 181–201.
The selection, testing and application of terrestrial insects as bioindicators.Crossref | GoogleScholarGoogle Scholar |

Meester, J. A. J., Lloyd, C. N. V., and Rowe-Rowe, D. T. (1979). A note on the ecological role of Praomys natalensis. South African Journal of Science 75, 183–184.

Mendelsohn, J. M. (1982). Notes on small mammals on the Springbok Flats, Transvaal. South African Journal of Zoology 17, 197–201.

Mikola, J., and Setälä, H. (1998). Relating species diversity to ecosystem functioning: mechanistic backgrounds and experimental approach with a decomposer food web. Oikos 83, 180–194.
Relating species diversity to ecosystem functioning: mechanistic backgrounds and experimental approach with a decomposer food web.Crossref | GoogleScholarGoogle Scholar |

Monadjem, A. (1997). Stomach contents of 19 species of small mammals from Swaziland. South African Journal of Zoology 32, 23–26.

Nel, J., Avenant, N., and Purves, M. (1996). Mammals. Final Report Contract No. 1008: Baseline Biology Survey and Reserve Development, Phase 1B. (Afridev Consultants: Darling, South Africa.)

New, T. R. (1999). Untangling the web: spiders and the challenges of invertebrate conservation. Journal of Insect Conservation 3, 251–256.
Untangling the web: spiders and the challenges of invertebrate conservation.Crossref | GoogleScholarGoogle Scholar |

Norrdahl, K., and Korpimäki, E. (1995). Effects of predator removal on vertebrate prey populations: birds of prey and small mammals. Oecologia 103, 241–248.
Effects of predator removal on vertebrate prey populations: birds of prey and small mammals.Crossref | GoogleScholarGoogle Scholar |

Orgeas, J., and Andersen, A. N. (2001). Fire and biodiversity: responses of grass-layer beetles to experimental fire regimes in an Australian tropical savanna. Journal of Applied Ecology 38, 49–62.
Fire and biodiversity: responses of grass-layer beetles to experimental fire regimes in an Australian tropical savanna.Crossref | GoogleScholarGoogle Scholar |

Pearce, J., and Venier, L. (2005). Small mammals as bioindicators of sustainable boreal forest management. Forest Ecology and Management 208, 153–175.
Small mammals as bioindicators of sustainable boreal forest management.Crossref | GoogleScholarGoogle Scholar |

Pearson, D. L., and Cassola, F. (1992). World-wide species richness patterns of tiger beetles (Coleoptera: Cicindelidae): indicator taxon for biodiversity and conservation studies. Conservation Biology 6, 376–391.
World-wide species richness patterns of tiger beetles (Coleoptera: Cicindelidae): indicator taxon for biodiversity and conservation studies.Crossref | GoogleScholarGoogle Scholar |

Pearson, D. E., and Ruggiero, L. F. (2003). Transect versus grid trapping arrangements for sampling small-mammal communities. Wildlife Society Bulletin 31, 454–459.

Petchey, O. L. (2000). Species diversity, species extinction, and ecosystem function. American Naturalist 155, 696–702.
Species diversity, species extinction, and ecosystem function.Crossref | GoogleScholarGoogle Scholar |

Petit, S., and Usher, M. B. (1998). Biodiversity in agricultural landscapes: the ground beetle communities of woody uncultivated habitats. Biodiversity and Conservation 7, 1549–1561.
Biodiversity in agricultural landscapes: the ground beetle communities of woody uncultivated habitats.Crossref | GoogleScholarGoogle Scholar |

Pocock, M. J. O., and Jennings, N. (2008). Testing biotic indicator taxa: the sensitivity of insectivorous mammals and their prey to the intensification of lowland agriculture. Journal of Applied Ecology 45, 151–160.
Testing biotic indicator taxa: the sensitivity of insectivorous mammals and their prey to the intensification of lowland agriculture.Crossref | GoogleScholarGoogle Scholar |

Rainio, J., and Niemelä, J. (2003). Ground beetles (Coleoptera: Carabidae) as bioindicators. Biodiversity and Conservation 12, 487–506.
Ground beetles (Coleoptera: Carabidae) as bioindicators.Crossref | GoogleScholarGoogle Scholar |

Rich, T. D. (2002). Using breeding land birds in the assessment of western riparian systems. Wildlife Society Bulletin 30, 1126–1139.

Rodríguez, J. P., Pearson, D. L., and Barrera, R. R. (1998). A test for the adequacy of bioindicator taxa: are tiger beetles (Coleoptera: Cicindelidae) appropriate indicators for monitoring the degradation of tropical forests in Venezuela? Biological Conservation 83, 69–76.
A test for the adequacy of bioindicator taxa: are tiger beetles (Coleoptera: Cicindelidae) appropriate indicators for monitoring the degradation of tropical forests in Venezuela?Crossref | GoogleScholarGoogle Scholar |

Rosenzweig, M. L. (1995). ‘Species Diversity in Space and Time.’ (Cambridge University Press: Cambridge.)

Rowe-Rowe, D. T. (1995). Small-mammal recolonization of a fire-exclusion catchment after unscheduled burning. South African Journal of Wildlife Research 25, 133–137.

Rowe-Rowe, D. T., and Lowry, P. B. (1982). Influence of fire on small-mammal populations in the Natal Drakensberg. South African Journal of Wildlife Research 12, 130–139.

Rowe-Rowe, D. T., and Meester, J. (1982). Habitat preferences and abundance relations of small mammals in the Natal Drakensberg. South African Journal of Zoology 17, 202–209.

Seaman, M. T., and Louw, S. vdM. (1999). SAGraSS, a biomonitoring method for grasslands. In ‘IAIAsa’99 Conference Proceedings’. pp. 231–238. (University of the Free State: Bloemfontein, South Africa.)

Skinner, J. D., and Chimimba, C. T. (2005). ‘The Mammals of the Southern African Subregion.’ (Cambridge University Press: Cape Town.)

Tilman, D. (1982). ‘Resource Competition and Community Structure.’ (Princeton University Press: Princeton.)

Tilman, D., Wedin, D., and Knops, J. (1996). Productivity and sustainability influenced by biodiversity in grassland ecosystems. Nature 379, 718–720.
Productivity and sustainability influenced by biodiversity in grassland ecosystems.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK28Xht1OisL8%3D&md5=ac7bc6e628f0618cea04525508cc0dfdCAS |

Van Rooyen, N. (2002). Veld management in the savannas. In ‘Game Ranch Management’. (Ed. J. du P. Bothma.) pp. 571–620. (Van Schaik Publishers: Pretoria, South Africa.)

Vorster, M. (1982). The development of the ecological index method for assessing veld condition in the Karoo. Proceedings of the Grassland Society of South Africa 17, 84–89.

Wang, G., Wang, Z., Zhou, Q., and Zhong, W. (1999). Relationship between species richness of small mammals and primary productivity of arid and semi-arid grasslands in north China. Journal of Arid Environments 43, 467–475.
Relationship between species richness of small mammals and primary productivity of arid and semi-arid grasslands in north China.Crossref | GoogleScholarGoogle Scholar |

Wootton, J. T. (1998). Effects of disturbance on species diversity: a multitrophic perspective. American Naturalist 152, 803–825.
Effects of disturbance on species diversity: a multitrophic perspective.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DC%2BD1cnit12jug%3D%3D&md5=23ab6609d104f931427cc1cd8070f76fCAS |