Drivers of fine-scale avian functional diversity with changing land use: an assessment of the effects of eco-estate housing development and management

  • Jarryd Alexander
  • David A. Ehlers Smith
  • Yvette C. Ehlers Smith
  • Colleen T. DownsEmail author
Research Article



The effects of changing land use, and especially urbanisation, on species and functional diversity are of global concern. Eco-estates are a form of urban housing development that is suggested to partially negate the effects of landscape development.


We assessed avian functional diversity at four eco-estates (previously sugarcane plantations) and one sugarcane plantation site along the north coast of KwaZulu-Natal (KZN), South Africa. We determined whether the differing development, management types, and the effects of land-cover configuration and composition at the respective study sites influenced avian functional diversity.


Fixed-radius point-count surveys were conducted within varying levels of land cover (natural, semi-natural, golf course and urban) at each eco-estate, and at a control sugarcane plantation site (natural, sugarcane-natural, and sugarcane land cover). The influence of differing land-cover covariates on all three components of functional diversity (functional richness, evenness, and divergence) were determined through a series of general linear models.


Sections of the respective study sites with increased natural land cover, mainly in the form of indigenous forest and coastal thicket/dense bush, showed improved avian functional diversity in comparison to those with increased building and road density. Negative effects of increased building density may be offset through an interconnection with or incorporation within natural land cover. Each component of functional diversity was influenced dynamically by seasonal changes and depending on land-cover type.


Eco-estates improved avian functional diversity if natural habitats are emphasised and connected. We suggest future development and management for eco-estates and agricultural land in coastal KZN follow ecological land-use complementation, to improve local ecosystem functioning.


Land use Functional richness Functional evenness Functional divergence Ecological land-use complementation Urbanisation Agriculture Land transformation 



The authors wish to thank the management of the eco-estates and sugarcane site for granting permission and access onto their sites. This research was made possible through funding from the eThekwini Municipality: Durban Action Research Partnership (D’RAP), the National Research Foundation (ZA), and the University of KwaZulu-Natal.

Supplementary material

10980_2019_786_MOESM1_ESM.docx (179 kb)
Supplementary material 1 (DOCX 179 kb)


  1. Adams CE, Lindsey KJ, Ash SJ (2006) Urban wildlife management. CRC Press, USAGoogle Scholar
  2. Akinnifesi FK, Sileshi GW, Ajayi OC, Akinnifesi AI, de Moura EG, Linhares JFP, Rodrigues I (2009) Biodiversity of the urban homegardens of São Luís city. Urban Ecosys, Northeastern Brazil. Google Scholar
  3. Allan E, Manning P, Alt F, Binkenstein J, Blaser S, Blüthgen N, Böhm S, Grassein F, Hölzel N, Klaus VH, Kleinebecker T, Morris KE, Oelmann Y, Prati D, Renner SC, Rillig MC, Schaefer M, Schloter M, Schmitt B, Schöning I, Schrumpf M, Solly M, Sorkau E, Steckel J, Steffen-Dewenter I, Stempfhuber B, Tschapka M, Weiner CN, Weisser WW, Werner M, Westphal C, Wilcke W, Fischer M (2015) Land use intensification alters ecosystem multifunctionality via loss of biodiversity and changes to functional composition. Ecol Lett 18:834–843PubMedPubMedCentralGoogle Scholar
  4. Bates D, Mächler M, Bolker B, Walker S (2015) Fitting linear mixed-effects models using lme4. J Stat Softw 67:1406–5823Google Scholar
  5. Blair RB (1996) Land use and avian species diversity along an urban gradient. Ecol Appl 6:506–519Google Scholar
  6. Blewett CM, Marzluff JM (2005) Effects of urban sprawl on snags and the abundance and productivity of cavity-nesting birds. Condor 107:678–693Google Scholar
  7. Burnham KP, Anderson DR (2002) Model selection and multimodel inference: a practical information-theoretic approach. Springer, New YorkGoogle Scholar
  8. Carrara E, Arroyo-Rodríguez V, Vega-Rivera JH, Schondube JE, de Freitas SM, Fahrig L (2015) Impact of landscape composition and configuration on forest specialist and generalist bird species in the fragmented Lacandona rainforest, Mexico. Biol Conserv 184:117–126Google Scholar
  9. Clergeau P, Jokimaki J, Savard JPL (2001) Are urban bird communities influenced by the bird diversity of adjacent landscapes? J Appl Ecol 38:1122–1134Google Scholar
  10. Colding J (2007) Ecological land-use complementation for building resilience in urban ecosystems. Landsc Urban Plan 81:45–55Google Scholar
  11. Colding J, Folke C (2009) The role of golf courses in biodiversity conservation and ecosystem management. Ecosystems 12:191–201Google Scholar
  12. Cornelis J, Hermy M (2004) Biodiversity relationships in urban and suburban parks in Flanders. Landsc Urban Plan 69:385–401Google Scholar
  13. Davis RA, Gole C, Roberts JD (2013) Impacts of urbanisation on the native avifauna of Perth, Western Australia. Urban Ecosyst 16:427–452Google Scholar
  14. Davis RA, Wilcox JA (2013) Adapting to suburbia: bird ecology on an urban-bushland interface in Perth, Western Australia. Pac Conserv Biol 19:110–120Google Scholar
  15. DeGraaf RM, Wentworth JM (1986) Avian guild structure and habitat associations in suburban bird communities. J Urban Ecol 9:339–412Google Scholar
  16. Dunning JB, Danielson BJ, Pulliam HR (1992) Ecological processes that affect populations in complex landscapes. Oikos 65:169–175Google Scholar
  17. Ehlers Smith DA, Ehlers Smith YC, Downs CT (2017a) Indian Ocean coastal thicket is of high conservation value for preserving taxonomic and functional diversity of forest-dependent bird communities in a landscape of restricted for availability. For Ecol Manage 390:157–165Google Scholar
  18. Ehlers Smith DA, Ehlers Smith YC, Ramesh T, Downs CT (2017b) Camera-trap data elucidate habitat requirements and conservation threats to an endangered forest specialist, the spotted ground thrush (Zoothera guttata). For Ecol Manage 400:523–530Google Scholar
  19. Ehlers Smith YC, Ehlers Smith DA, Ramesh T, Downs CT (2017c) Forest habitats in a mixed urban-agricultural mosaic landscape: patterns of mammal occupancy. Landscape Ecol 33:59–70Google Scholar
  20. Ehlers Smith YC, Ehlers Smith DA, Seymour CL, Thébault E, van Veen FJF (2015) Response of avian diversity to habitat modification can be predicted from life-history traits and ecological attributes. Landscape Ecol 30:1239–1255Google Scholar
  21. Ehlers Smith DA, Si X, Ehlers Smith YC, Downs CT (2018a) Seasonal variation in avian diversity and tolerance of migratory forest specialists to the patch-isolation gradient across a forest system. Biodivers Conserv. Google Scholar
  22. Ehlers Smith DA, Si X, Ehlers Smith YC, Kalle R, Ramesh T, Downs CT (2018b) Patterns of avian diversity across a decreasing patch-size gradient in a critically endangered sub-tropical forest system. J Biogeogr 45:2118–2132Google Scholar
  23. ESRI (2011) ArcGIS Desktop 10.4. Environmental systems research institute. RedlandsGoogle Scholar
  24. Fahrig L (2003) Effects of habitat fragmentation on biodiversity. Ann Rev Ecol Evol Syst 34:487–515Google Scholar
  25. Fernández-Juricic E, Jokimaki J (2001) A habitat island approach to conserving birds in urban landscapes: case studies from southern and northern Europe. Biodivers Conserv 10:2023–2043Google Scholar
  26. Ferraz G, Nichols JD, Hines JE, Stouffer PC, Bierregaard RO, Lovejoy TE (2007) A large-scale deforestation experiment: effects of patch area and isolation on Amazon birds. Science 315:238–241PubMedGoogle Scholar
  27. Foresman TW, Pickett SA, Zipperer WC (1997) Methods for spatial and temporal land use and land-cover assessment for urban ecosystems and application in the greater Baltimore-Chesapeake region. J Urban Ecol 1:201–216Google Scholar
  28. Garson J, Aggarwal A, Sarkar S (2002) Birds as surrogates for biodiversity: an analysis of a data set from southern Québec. J Biosci 27:347–360PubMedGoogle Scholar
  29. Gelman A, Ys S, Yajima M, Hill J, Pittay MG, Kerman J, Zheng T, Dorie V (2009) Arm: data analysis using regression and multi-level/hierarchical models. R packageGoogle Scholar
  30. Geoterraimage (2017) KZN Province land-cover mapping (from SPOT5 Satellite imagery circa 2013). Prepared for Ezemvelo KZN Wildlife (Biodiversity Research). South AfricaGoogle Scholar
  31. Goble BJ, van der Elst R (2012) Trends in coastal development and land-cover change: the case of KwaZulu-Natal, South Africa. Western Indian Ocean J Mar Sci 11:193–204Google Scholar
  32. Gower JC (1966) Some distance properties of latent root and vector methods used in multivariate analysis. Biometrika 53:325–338Google Scholar
  33. Grey-Ross R, Downs CT, Kirkman K (2009) Using housing estates as conservation tools: a case study in KwaZulu-Natal, South Africa. Appl Geogr 29:371–376Google Scholar
  34. Groom MJ (2006) Threats to biodiversity. In: Groom MJ, Meffe GK, Carroll CR (eds) Principles of conservation biology. Sinauer Associates, Sunderland, pp 63–110Google Scholar
  35. Hockey PAR, Dean WRJ, Ryan PG (2005) Roberts birds of southern Africa, 7th edn. John Voelcker Bird Book Fund, South AfricaGoogle Scholar
  36. Hooke RL, Martin-Duque JF, Pedraza J (2012) Land transformation by humans, a review. Geol Soc Am 22:4–10Google Scholar
  37. Ke A, Sibiya MD, Reynolds C, McCleery RA, Monadjem A, Fletcher RJ (2018) Landscape heterogeneity shapes taxonomic diversity of non-breeding birds across fragmented savanna landscapes. Biodivers Conserv 27:2681–2698Google Scholar
  38. Kulkarni M, Dighe S, Sawant A, Oswal P, Sahasrabuddhe K, Patwardhan A (2001) Institutions: biodiversity hotspots in urban areas. In: Ganeshaiah KN, Uma Shnker R, Bawa KS (eds) Tropical ecosystems: structure. Diversity and Human Welfare, Oxford/IBH, India, pp 693–695Google Scholar
  39. Laliberté E, Legendre P (2010) A distance-based framework for measuring functional diversity from multiple traits. Ecology 91:299–305PubMedGoogle Scholar
  40. Laliberté E, Legendre P, Shipley B (2014) FD: measuring functional diversity from multiple traits, and other tools for functional ecology. R package version 1.0-12Google Scholar
  41. Landis DA, Gardiner MM, van der Werf W, Swinton SM (2008) Increasing corn for biofuel production reduces biocontrol services in agricultural landscapes. Proc Nat Acad Sci USA 105(51):20552–20557PubMedGoogle Scholar
  42. MacArthur RH, Wilson OE (1967) The theory of island biogeography. Princeton University Press, PrincetonGoogle Scholar
  43. Magle SB, Hunt VM, Vernon M, Crooks KR (2012) Urban wildlife research: past, present, and future. Biol Conserv 155:23–32Google Scholar
  44. Mason NWH, Mouillot D, Lee WG, Wilson JB (2005) Functional richness, functional evenness and functional divergence: the primary components of functional diversity. Oikos 111:112–118Google Scholar
  45. McPherson SC, Brown M, Downs CT (2016) Crowned eagle nest sites in an urban landscape: requirements of a large eagle in the Durban Metropolitan Open Space System. Landsc Urban Plan 146:43–50Google Scholar
  46. Melles S, Glenn S, Martin C (2003) Urban bird diversity and landscape complexity: species–environment associations along a multiscale habitat gradient. Conserv Ecol 7:22Google Scholar
  47. Mönkkönen M, Rajasärkkä A, Lampila P (2014) Isolation, patch size and matrix effects on bird assemblages in forest reserves. Biodiv Conserv 23:3287–3300Google Scholar
  48. Morimoto T, Katoh K, Yamaura Y, Watanabe S (2006) Can surrounding land-cover influence the avifauna in urban/suburban woodlands in Japan? Landsc Urban Plan 74:46–69Google Scholar
  49. Mouchet MA, Villéger S, Mason NWH, Mouillot D (2010) Functional diversity measures: an overview of their redundancy and their ability to discriminate community assembly rules. Funct Ecol 24:867–876Google Scholar
  50. Pardini R, de Souza SM, Braga-Neto R, Metzger JP (2005) The role of forest structure, fragment size and corridors in maintaining small mammal abundance and diversity in an Atlantic forest landscape. Biol Conserv 124:253–266Google Scholar
  51. Patterson L, Kalle R, Downs CT (2018) Factors affecting presence of Vervet monkey troops in a suburban matrix in KwaZulu-Natal, South Africa. Landsc Urban Plan 169:220–228Google Scholar
  52. Petchey WL, Gaston KJ (2002) Functional diversity (FD), species richness and community composition. Ecol Lett 5:402–411Google Scholar
  53. Pope SE, Fahrig L, Merriam HG (2000) Landscape complementation and metapopulation effects on leopard frog populations. Ecology 81:2498–2508Google Scholar
  54. Primack RB (1993) Essentials of conservation biology. Sinauer Associates, SunderlandGoogle Scholar
  55. Quin A, Aviron S, Dover J, Burel F (2004) Complementation/supplementation of resources for butterflies in agricultural landscapes. Agric Ecosyst Environ 103:473–479Google Scholar
  56. R Core Team (2013) R: a language and environment for statistical computing. R Foundation for Statistical Computing, ViennaGoogle Scholar
  57. Ramesh T, Downs CT (2013) Impact of farmland use on population density and activity patterns of serval in South Africa. J Mammal 94:1460–1470Google Scholar
  58. Ramesh T, Downs CT (2014) Land use factors determining occurrence of red-necked spurfowl (Pternistis afer) in the Drakensberg Midlands, South Africa. J Ornith 155:471–480Google Scholar
  59. Ramesh T, Downs CT (2015) Impact of land use on occupancy and abundance of terrestrial mammals in the Drakensberg Midlands, South Africa. J Nat Conserv 23:9–18Google Scholar
  60. Ricketts T, Imhoff M (2003) Biodiversity, urban areas, and agriculture: locating priority ecoregions for conservation. Conserv Ecol 8:1Google Scholar
  61. Rollinson DP, Coleman J, Downs CT (2014) Roost temperature and fidelity of Wahlberg’s epauletted fruit bat, Epomophorus wahlbergi, in an urban environment. J Afr Zool 49:173–180Google Scholar
  62. Russell C, Downs CT (2012) Effect of land use on anuran species composition in north-eastern KwaZulu-Natal, South Africa. Appl Geogr 35:247–256Google Scholar
  63. Sala OE, Chapin FS, Armesto JJ, Berlow R, Bloomfield J, Dirzo R, Huber-Sanwald E, Huenneke LF, Jackson RB, Kinzig A, Leemans R, Lodge D, Mooney HA, Oesterheld M, Poff NL, Sykes MT, Walker BH, Walker M, Wall DH (2000) Global biodiversity scenarios for the year 2100. Science 287:1770–1774PubMedGoogle Scholar
  64. Sandström UG, Angelstam P, Mikusiński G (2006) Ecological diversity of birds in relation to the structure of urban green space. Landsc Urban Plan 77:39–53Google Scholar
  65. Savard JPL, Clergeau P, Mennechez G (2000) Biodiversity concepts and urban ecosystems. Landsc Urban Plan 48:131–142Google Scholar
  66. Schütz C, Schulze CH (2015) Functional diversity of urban bird communities: effects of landscape composition, green space area and vegetation cover. Ecol Evol 5:5230–5239PubMedPubMedCentralGoogle Scholar
  67. Sekercioğlu CH (2012) Bird functional diversity and ecosystem services in tropical forests, agroforests and agricultural areas. J Ornithol 153:153–161Google Scholar
  68. Seto KC, Güneralp B, Hutyra LR (2012) Global forecast of urban expansion to 2030 and direct impacts on biodiversity and carbon pools. Proc Nat Acad Sci USA 109:16083–16088PubMedGoogle Scholar
  69. Singh P, Downs CT (2016) Hadedas in the hood: Hadeda Ibis activity in suburban neighbourhoods of Pietermaritzburg, KwaZulu-Natal, South Africa. Urban Ecosyst 19:1283–1293Google Scholar
  70. Sorace A, Visentin M (2007) Avian diversity on golf courses and surrounding landscapes in Italy. Landsc Urban Plan 81:81–90Google Scholar
  71. Sperling C, Lortie C (2009) The importance of urban backgardens on plant and invertebrate recruitment: a field microcosm experiment. Urban Ecosyst 13:223–235Google Scholar
  72. Tanner RA, Gange AC (2005) Effects of golf courses on local biodiversity. Landsc Urban Plan 71:137–146Google Scholar
  73. Terman MR (1997) Natural links: naturalistic golf courses as wildlife habitat. Landsc Urban Plan 38:197–283Google Scholar
  74. Theobald DM, Hobbs NT, Bearly T, Zack JA, Shenk T, Riebsame WE (2000) Incorporating biological information in local land-use decision making: designing a system for conservation planning. Landscape Ecol 15:34–45Google Scholar
  75. Threlfall CG, Mata L, Mackie JA, Hahs AK, Stork NE, Williams NS, Livesley SJ, Beggs J (2017) Increasing biodiversity in urban green spaces through simple vegetation interventions. J Appl Ecol 54:1874–1883Google Scholar
  76. Trimble MJ, van Aarde RJ (2014) Amphibian and reptile communities and functional groups over a land-use gradient in a coastal tropical forest landscape of high richness and endemicity. Anim Conserv 17:441–453Google Scholar
  77. Tscharntke T, Klein AM, Kruess A, Steffan-Dewenter I, Thies C (2005) Landscape perspectives on agricultural intensification and biodiversity and ecosystem service management. Ecol Lett 8:857–874Google Scholar
  78. Tscharntke T, Sekercioğlu CH, Dietsch TV, Sodhi NS, Hoehn P, Tylianakis JM (2008) Landscape constraints on functional diversity of birds and insects in tropical agroecosystems. Ecology 89:944–951PubMedGoogle Scholar
  79. van Heezik Y, Smyth A, Mathieu R (2008) Diversity of native and exotic birds across an urban gradient in a New Zealand city. Landsc Urban Plan 87:223–232Google Scholar
  80. Villéger S, Mason NWH, Mouillot D (2008) New multidimensional functional diversity indices for a multifaceted framework in functional ecology. Ecology 89:2290–2301PubMedGoogle Scholar
  81. White CL, Main MB (2005) Waterbird use of created wetlands in golf-course landscapes. Wildl Soc Bull 33:411–421Google Scholar
  82. Widdows CD, Ramesh T, Downs CT (2015) Factors affecting the distribution of large spotted genets (Genetta tigrina) in an urban environment in South Africa. Urban Ecosyst 18:1401–1413Google Scholar
  83. Wilby RL, Perry GLW (2006) Climate change, biodiversity and the urban environment: a critical review based on London, UK. Prog Phys Geogr 30:73–98Google Scholar
  84. Wood BC, Pullins AS (2002) Persistence of species in fragmented urban landscapes: the importance of dispersal ability and habitat availability for grassland butterflies. Biodiv Conserv 11:1451–1468Google Scholar

Copyright information

© Springer Nature B.V. 2019

Authors and Affiliations

  1. 1.School of Life SciencesUniversity of KwaZulu-NatalPietermaritzburgSouth Africa

Personalised recommendations