Community Ecology

, Volume 16, Issue 1, pp 106–114 | Cite as

Role of the Mediterranean Sea in differentiating European and North African woodland bird assemblages

  • R. A. CorreiaEmail author
  • A. M. A. Franco
  • J. M. Palmeirim


The Mediterranean Sea has separated the sclerophyllous forests of southern Europe and northern Africa for millions of years, but its role structuring forest bird assemblages remains unclear. To address this issue we sampled bird assemblages in cork oak woodlands located north and south of the Strait of Gibraltar and compared abundance, diversity, and species and guild assemblage structure between regions. Abundance and diversity patterns were remarkably similar, but dissimilarity analyses of species and guild composition revealed differences in bird assemblage structure between regions. Differences are partly attributable to the effect of the Mediterranean as barrier to the dispersal of forest birds; a few species were unable to colonize North Africa, and many that colonized it remained in sufficient isolation to evolve into distinct taxa. In addition to this divergence of biogeographic genesis, assemblages also differ because in North African cork oak woodlands forest and insectivorous specialist species are less abundant. This dissimilarity could be due to the effect of different exploitation levels present in each region. Managed cork oak woodlands are widespread in the Western Mediterranean, and are valuable because they conciliate economic exploitation with high biodiversity. In North Africa these woodlands are of greater conservation concern because they harbour endemic bird species that give its assemblage a distinct character, cover a smaller area, and are currently under greater pressure from overexploitation. These results highlight the importance to implement management practices that increase resilience and maintain biodiversity value throughout the range of cork oak woodlands.


Geographic barrier Cork oak Quercus suber Biodiversity conservation Management Guild 


Gill and Donsker (2015) 



Generalized Linar Model


Log-likelihood Ratio


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.

Supplementary material

42974_2015_16010106_MOESM1_ESM.pdf (106 kb)
Supplementary material, approximately 109 KB. (1 kb)
Supplementary material, approximately 109 KB.


  1. Ajbilou, R., T. Marañón and J. Arroyo. 2006. Ecological and bio-geographical analyses of Mediterranean forests of northern Morocco. Acta Oecol. 29:104–113.CrossRefGoogle Scholar
  2. Berrahmouni, N., P. Regato, M. Ellatifi, H. Daly-Hassen, M. Bugalho, S. Bensaid, M. Diáz and J. Aronson. 2009. Ecoregional planning for biodiversity conservation. In: Aronson, J., Pereira, J. S., Pausas, J. G. (eds.), Cork Oak Woodlands on the Edge. Island Press, Washington, USA, pp. 203–216.Google Scholar
  3. Bibby, C. J., N.D. Burgess, D.A. Hill and S.H. Mustoe. 2005. Bird Census Techniques. Elsevier Academic Press, London, UK.Google Scholar
  4. Blondel, J. 1995. Biogeographie: Approche Ecologique et Evolutive. Masson, Paris, France.Google Scholar
  5. Blondel, J., J. Aronson, J.Y. Bodiou and G. Boeuf. 2010. The Mediterranean Region: Biological Diversity in Space and Time. Oxford University Press, Oxford, UK.Google Scholar
  6. Borcard, D., F. Gillet and P. Legendre. 2011. Numerical Ecology with R. Springer, New York, USA.CrossRefGoogle Scholar
  7. Brito, P.H. 2005. The influence of Pleistocene glacial refugia on tawny owl genetic diversity and phylogeography in western Europe. Mol. Ecol. 14:3077–3094.CrossRefGoogle Scholar
  8. Broderick, D., Y. Idaghdour, A. Korrida and J. Hellmich. 2003. Gene fow in great bustard populations across the Strait of Gibraltar as elucidated from excremental PCR and mtDNA sequencing. Conserv. Genet. 4:793–800.CrossRefGoogle Scholar
  9. Bugalho, M., T. Plieninger, J. Aronson, M. Ellatif and D.G. Crespo. 2009. Open woodlands: A diversity of uses (and overuses). In: Aronson, J., Pereira, J.S., Pausas, J.G. (eds.), Cork Oak Woodlands on the Edge. Island Press, Washington, USA, pp. 33–45.Google Scholar
  10. Bugalho, M.N., M.C. Caldeira, J.S. Pereira, J. Aronson and J.G. Pausas. 2011. Mediterranean cork oak savannas require human use to sustain biodiversity and ecosystem services. Front. Ecol. Environ. 9:278–286.CrossRefGoogle Scholar
  11. Carnicer, J., M. Coll, M. Ninyerola, X. Pons, G. Sánchez and J. Peñuelas. 2011. Widespread crown condition decline, food web disruption, and amplified tree mortality with increased climate change-type drought. Proc. Natl. Acad. Sci. USA 108:1475–1478.CrossRefGoogle Scholar
  12. Cherkaoui, I., S. Selmi, J. Boukhriss, R.I. Hamid and D. Mohammed. 2009. Factors affecting bird richness in a fragmented cork oak forest in Morocco. Acta Oecol. 35:197–205.CrossRefGoogle Scholar
  13. Cramp, S. and K.E.L. Simmons. 2006. BWPi: The Birds of the Western Palearctic interactive (DVD-ROM). BirdGuides Ltd., Sheffeld, UK.Google Scholar
  14. Colwell, R. 2009. EstimateS: Statistical estimation of species richness and shared species from samples. Version 8.2. User’s Guide and application published at:
  15. De Lattin, G. 1949. Beiträge zur Zoogeographie des Mittelmeer-gebietes. Verhandlungen der deutschen Zoologischen Gesell-schaft, Kiel, Suppl 13:143–151.Google Scholar
  16. Díaz, M., P. Campos and F.J. Pulido. 1997. The Spanish dehesas: a diversity in land-use and wildlife. In: Pain, D.J., Pienkowski, M.W. (eds.), Farming and Birds in Europe. Academic Press, Cambridge, UK, pp 178–209.Google Scholar
  17. Duggen, S., K. Hoernle, P. van den Bogaard, L. Rüpke and J.P. Morgan. 2003. Deep roots of the Messinian salinity crisis. Nature 422:602–606.CrossRefGoogle Scholar
  18. EBCC. 2010. Population trends of European common birds 2010. EBCC report. (accessed 31/03/2015).
  19. European Habitat Directive. url: (accessed 31/03/2015).
  20. Fishpool, L.D.C. and M.I. Evans. 2001. Important Bird Areas in Africa and Associated Islands: Priority Sites for Conservation. Birdlife International, Cambridge, UK.Google Scholar
  21. García-Mudarra, J.L., C. Ibáñez and J. Juste. 2009. The Straits of Gibraltar: barrier or bridge to Ibero-Moroccan bat diversity? Biol. J. Linn. Soc. 96:434–450.CrossRefGoogle Scholar
  22. Gill, F. and D. Donsker (eds.) 2015. IOC World Bird List (v. 5.1). DOI: 10.143/IOC.ML.5.1.Google Scholar
  23. Godinho, C. and J.E. Rabaça. 2011. Birds like it Corky: the influence of habitat features and management of ‘montados’ in breeding bird communities. Agrofor. Syst. 82:183–195.CrossRefGoogle Scholar
  24. Gotelli, N.J. and A. Chao. 2013. Measuring and estimating species richness, species diversity, and biotic similarity from sampling data. In: Levin, S.A. (ed) The Encyclopaedia of Biodiversity. Second edition, volume 5. Academic Press, Waltham, Massachusetts, USA, pp 195–211.Google Scholar
  25. Gotelli, N.J. and G.R. Graves. 1996. Null Models in Ecology. Smithsonian Institution Press, Washington, USA.Google Scholar
  26. Habel, J.C., P. Dieker and T. Schmitt. 2009. Biogeographical connections between the Maghreb and the Mediterraenan peninsulas of southern Europe. Biol. J. Linn. Soc. 98:693–703.CrossRefGoogle Scholar
  27. Heath, M.F., C. Borggreve, N. Peet and W. Hagemeijer. 2000. European Bird Populations: Estimates and Trends. Birdlife Conservation Series No 10. Birdlife International/European Bird Census Council, Cambridge, UK.Google Scholar
  28. Hoogeveen, Y., J.E. Petersen, K. Balazs and I. Higuero. 2004. High Nature Value Farmland – Characteristics, Trends and Policy Challenges. EEA Report. No 1/2004. European Environment Agency, Copenhagen, Denmark.Google Scholar
  29. Husemann, M., T. Schmitt, F.E. Zachos, W. Ulrich and J.C. Habel. 2014. Palaearctic biogeography revisited: evidence for the existence of a North African refugium for Western Palaearctic biota. J. Biogeogr. 41:81–94.CrossRefGoogle Scholar
  30. Leal, A.I., R.A. Correia, J.P. Granadeiro and J.M. Palmeirim. 2011a. Impact of cork extraction on birds: Relevance for conservation of Mediterranean biodiversity. Biol. Conserv. 144:1655–1662.CrossRefGoogle Scholar
  31. Leal, A.I., R.C. Martins, J.M. Palmeirim and J.P. Granadeiro. 2011b. Influence of habitat fragments on bird assemblages in Cork Oak woodlands. Bird Study 58:309–320.CrossRefGoogle Scholar
  32. Leal, A.I., R.A. Correia, J.M. Palmeirim and J.P. Granadeiro. 2013. Does canopy pruning affect foliage-gleaning birds in managed cork oak woodlands? Agrofor. Syst. 87:355–363.CrossRefGoogle Scholar
  33. Lepage, D. 2013a. Checklist of the Birds of Morocco. Avibase, the world bird database. Retrieved from 9/07/2013.
  34. Lepage, D. 2013b. Checklist of the Birds of Spain. Avibase, the world bird database. Retrieved from 9/07/2013.
  35. Marañón, T., R. Ajbilou, F. Ojeda and J. Arroyo. 1999. Biodiversity of woody species in oak woodlands of southern Spain and northern Morocco. For. Ecol. Manage. 115:147–156.CrossRefGoogle Scholar
  36. Myers, N., R.A. Mittermeier, C.G. Mittermeier, G.A.B. Fonseca and J. Kent. 2000. Biodiversity hotspots for conservation priorities. Nature 403:853–858.PubMedPubMedCentralGoogle Scholar
  37. Ojeda, F., T. Marañón and J. Arroyo. 1996. Patterns of ecological, chorological and taxonomic diversity at both sides of the Strait of Gibraltar. J. Veg. Sci. 7:63–72.CrossRefGoogle Scholar
  38. Plieninger, T. and C. Wilbrand. 2001. Land use, biodiversity conservation, and rural development in the dehesas of Cuatro Lugares, Spain. Agrofor. Syst. 51:23–34.CrossRefGoogle Scholar
  39. Pons, J.M., G. Olioso, C. Cruaud and J. Fuchs. 2011. Phylogeography of the Eurasian green woodpecker (Picus viridis). J. Biogeogr. 38:311–325.CrossRefGoogle Scholar
  40. R Development Core Team. 2011. R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria.Google Scholar
  41. Rodríguez-Sánchez, F., R. Pérez-Barrales, F. Ojeda, P. Vargas and J. Arroyo. 2008. The Strait of Gibraltar as a melting pot for plant biodiversity. Quaternary. Sci. Rev. 27:2100–2117.CrossRefGoogle Scholar
  42. Rosalino, L.M., D. Ferreira, I. Leitão and M. Santos-Reis. 2011. Usage patterns of Mediterranean agro-forest habitat components by wood mice Apodemus sylvaticus. Mamm. Biol. 76:268–273.CrossRefGoogle Scholar
  43. Saetre, G.P., T. Borge and T. Moum. 2001. A new bird species? The taxonomic status of the “Atlas Flycatcher” assessed from DNA sequence analysis. Ibis 143: 494–497.CrossRefGoogle Scholar
  44. Salzburger, W., J. Martens and C. Sturmbauer. 2002. Paraphyly of the Blue Tit (Parus caeruleus) suggested from cytochrome b sequences. Mol. Phylogenet. Evol. 24:19–25.CrossRefGoogle Scholar
  45. Silva, P.M., C.A.S. Aguiar, J. Niemelä, J.P. Sousa and A.R.M. Serrano. 2009. Cork-oak woodlands as key-habitats for biodiversity conservation in Mediterranean landscapes: a case study using rove and ground beetles (Coleoptera: Staphylinidae, Carabidae). Biodivers. Conserv. 18:605–619.CrossRefGoogle Scholar
  46. Sutherland, W.J., I. Newton and R.E. Green. 2004. Bird Ecology and Conservation: A Handbook of Techniques. Techniques in Ecology and Conservation Series. Oxford University Press, Oxford, UK.CrossRefGoogle Scholar
  47. Telleria, J. L. and T. Santos. 1993. Distributional patterns of insectivorous passerines in the Iberian forests: Does abundance decrease near the border? J. Biogeogr. 20:235–240.CrossRefGoogle Scholar
  48. Telleria, J.L. 2001. Passerine bird communities of Iberian dehesas: a review. Anim. Biodivers. Conserv. 24:67–78.Google Scholar
  49. Warton, D.I., S.T. Wright and Y. Wang. 2012. Distance-based mul-tivariate analysis confound location and dispersion effects. Methods Ecol. Evol. 3:89–101.CrossRefGoogle Scholar

Copyright information

© Akadémiai Kiadó, Budapest 2015

This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (, which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made.

Authors and Affiliations

  • R. A. Correia
    • 1
    • 2
    Email author
  • A. M. A. Franco
    • 2
  • J. M. Palmeirim
    • 1
  1. 1.Universidade de LisboaLisboaPortugal
  2. 2.University of East AngliaNorwichUK

Personalised recommendations