Biological Invasions

, Volume 14, Issue 2, pp 295–305 | Cite as

Fitness consequences of anthropogenic hybridization in wild red-legged partridge (Alectoris rufa, Phasianidae) populations

  • F. Casas
  • F. Mougeot
  • I. Sánchez-Barbudo
  • J. A. Dávila
  • J. Viñuela
Original Paper


Hybridization is a widespread phenomenon, which plays crucial roles in the speciation of living beings. However, unnatural mixing of historically isolated taxa due to human-related activities has increased in recent decades, favouring levels of hybridization and introgression that can have important implications for conservation. The wild red-legged partridge (Alectoris rufa, Phasianidae) populations have recently declined and the releases of farm-reared partridges have become a widespread management strategy. The native range of the red-legged is limited to the south-west of Europe (from Italy to Portugal). This species does not breed in sympatry with the chukar partridge (A. chukar), whose range is Eurasian (from Turkey to China). However, red-legged partridges have often been hybridized with chukar partridges to increase the productivity of farmed birds, and game releases may have spread hybrid birds into the wild. In this study, we investigated the fitness (survival and breeding) differences between hybrid and “pure” red-legged partridges in a wild population located in central Spain. Incubation probability was similar in hybrids and “pure” partridges. Hybrid females laid larger clutches than “pure” ones, but hatching success did not differ between hybrid and “pure” partridges. Hybrid birds had lower survival rate than “pure” ones, mainly because of higher predation rates. Our results show that, despite lower survival, hybrid partridges breed in natural populations, so this could increase extinction risk of wild pure partridge populations, through releases of farmed hybrid birds. The consequences of continued releases could be of vital importance for the long term conservation of wild red-legged partridges.


Hybridization Genetic conservation Red-legged partridge Survival Breeding Alectoris rufa 



We thank all game managers and hunting societies for allowing studying partridges on their hunting estates. We are very grateful to private individuals and to the museum collection of the Estación Biológica de Doñana (EBD) that kindly provided A. rufa samples. We also thank the contribution of A. chukar samples to Alexis and Thanatos (Archipelagos Aigaiou), Ramón Fernández (Quiñón S.A.), Nai-Fa Liu (Lanzhou University), Filippo Barbanera (Università di Pisa), Giannis Kouvelos and anonymous hunters. Two anonymous referees made useful comments on previous drafts. F. Casas was supported by a pre-doctoral grant of the Junta de Comunidades de Castilla la Mancha (JCCM). This work also received financial support from the research projects “Bases científicas preliminares para un plan de conservación de la perdiz roja en Castilla-La Mancha”, and “Caracterización y diagnóstico genético de las poblaciones de perdices de Castilla-La Mancha” of the Consejería de Agricultura y Medio Ambiente de la JCCM (Junta de Comunidades de Castilla-La Mancha), and CYCIT projects MCYT-REN200307851/GLO and CGL2004-02568/BOS. IS-B was supported by a FEDENCA-Royal Spanish Federation of Hunting contract. FM was supported by Grants from the Ministerio de Educación y Ciencia, Spain (CGL 2006-11823) and from the JCCM, Spain (PAI06-0112 and PAII1C09-0227-0104).

Supplementary material

10530_2011_62_MOESM1_ESM.doc (1.3 mb)
Supplementary material 1 (DOC 1302 kb)
10530_2011_62_MOESM2_ESM.doc (304 kb)
Supplementary material 2 (DOC 304 kb)
10530_2011_62_MOESM3_ESM.doc (406 kb)
Supplementary material 3 (DOC 406 kb)
10530_2011_62_MOESM4_ESM.doc (60 kb)
Supplementary material 4 (DOC 61 kb)


  1. Aebischer NJ, Potts GR (1994) Red–legged partridge. In: Tucker GM, Heath MF (eds) Birds in Europe. Their conservation status. Birdlife conservation serie no. 3. Birdlife International, Cambridge, pp 214–215Google Scholar
  2. Allendorf FW, Leary RF, Spruell P, Wenburg JK (2001) The problems with hybrids: setting conservation guidelines. Trends Ecol Evol 16:613–622CrossRefGoogle Scholar
  3. Arnold ML (1997) Natural hybridization and evolution. Oxford University Press, UKGoogle Scholar
  4. Avise JC (2004) Molecular markers, natural history and evolution. Sinauer Associates, Inc. Publishers, SunderlandGoogle Scholar
  5. Baratti M, Ammannati M, Magnelli C, Dessi-Fulgheri F (2004) Introgression of chukar genes into a reintroduced red-legged partridge (Alectoris rufa) population in central Italy. Anim Genet 36:29–35CrossRefGoogle Scholar
  6. Barbanera F, Negro JJ, Di Giuseppe G, Bertoncini F, Cappelli F, Dini F (2005) Analysis of the genetic structure of red-legged partridge (Alectoris rufa, Galliformes) populations by means of mitochondrial DNA and RAPD markers: a study from central Italy. Biol Conserv 122:275–287CrossRefGoogle Scholar
  7. Barbanera F, Guerrini M, Hadjigerou P, Panayides P, Sokos P, Wilkinson P, Khan AA, Khan BY, Cappelli F, Dini F (2007) Genetic insight into Mediterranean chukar (Alectoris chukar, Galliformes) populations inferred from mitochondrial DNA and RAPD markers. Genetica 131:287–298PubMedCrossRefGoogle Scholar
  8. Barbanera F, Guerrini M, Khan AA, Panayides P, Hadjigerou P, Sokos C, Gombobaatar S, Samadi S, Khan BY, Tofanelli S, Paoli G, Dini F (2009a) Human-mediated introgression of exotic chukar (Alectoris chukar, Galliformes) genes from East Asia into native Mediterranean partridges. Biol Invasions 11:333–348CrossRefGoogle Scholar
  9. Barbanera F, Marchi C, Guerrini M, Panayides P, Sokos C, Hadjigerou P (2009b) Genetic structure of Mediterranean chukar (Alectoris chukar, Galliformes) populations: conservation and management implications. Naturwissenschaften 96:1203–1212PubMedCrossRefGoogle Scholar
  10. Barbanera F, Pergams ORW, Guerrini M, Forcina G, Panayides P, Dini F (2010) Genetic consequences of intensive management in game birds. Biol Conserv 143:1259–1268CrossRefGoogle Scholar
  11. Barton NH (2001) The role of hybridization in evolution. Mol Ecol 10:551–568PubMedCrossRefGoogle Scholar
  12. Bernabeu RL (2000) Evaluación económica de la caza en Castilla-La Mancha. PhD thesis, Universidad de Castilla-La Mancha, SpainGoogle Scholar
  13. Bernard-Laurent A (1984) Hybridation naturelle entre Perdrix bartavelle (Alectoris graeca saxatilis) et Perdrix rouge (Alectoris rufa) dans les Alpes Maritimes. Gibier Faune Sauvage 2:79–96Google Scholar
  14. Birdlife International (2004) Birds in the European Union: a status assessment. Birdlife International, WageningenGoogle Scholar
  15. Blanco-Aguiar JA (2007) Variación espacial en la biología de la perdiz roja (Alectoris rufa): una aproximación multidisciplinar. PhD thesis, Universidad Complutense de Madrid, SpainGoogle Scholar
  16. Blanco-Aguiar JA, Virgós E, Villafuerte R (2004) Perdiz Roja (Alectoris rufa). In: Madroño A, González C, Atienza JC (eds) Libro Rojo de las Aves de España. Dirección General para la Biodiversidad-SEO/BirdLife, Madrid, pp 182–185Google Scholar
  17. Blanco-Aguiar JA, González-Jara P, Ferrero ME, Sánchez-Barbudo I, Virgós E, Villafuerte R, Dávila JA (2008) Assessment of game restocking contributions to anthropogenic hybridization: the case of the Iberian red-legged partridge. Anim Conserv 11:535–545CrossRefGoogle Scholar
  18. Boecklen WJ, Howard DJ (1997) Genetic analysis of hybrid zones: numbers of markers and power of resolution. Ecology 78:2611–2616CrossRefGoogle Scholar
  19. Casas F, Viñuela J (2010) Agricultural practices or game management: which is the key to improve red-legged partridge nesting success in agricultural landscapes? Environ Conserv 37:177–186CrossRefGoogle Scholar
  20. Casas F, Mougeot F, Viñuela J (2009) Double nesting behaviour and differences between sexes in breeding success in wild Red-legged Partridges Alectoris rufa. Ibis 151:743–751CrossRefGoogle Scholar
  21. Cox DR (1972) Regression models and life-tables (with discussion). J R Stat Soc B 34:187–220Google Scholar
  22. Cramp S, Simmons KEL (1980) The birds of the Western Palearctic, vol. II. Oxford University Press, OxfordGoogle Scholar
  23. Crawley MJ (1993) GLIM for ecologists. Blackwell, LondonGoogle Scholar
  24. Dávila JA (2009) Marcadores genéticos para detectar introgresión en aves del género Alectoris. Patent no. 2 323 027, Oficina Española de Patentes y MarcasGoogle Scholar
  25. Del Hoyo J, Elliott A, Sargatal J (1994) Handbook of the birds of the world, vol II. Lynx Ed, BarcelonaGoogle Scholar
  26. Deregnacourt S, Guyomarch JC, Aebischer NJ (2002) Hybridization between European Quail Coturnix coturnix and Japanese Quail Coturnix japonica. Ardea 90:15–21Google Scholar
  27. Deregnacourt S, Guyomarch JC, Spanò S (2005) Behavioural evidence of hybridization (Japanese × European) in domestic quail released as game birds. App Anim Behav Sci 94:303–318CrossRefGoogle Scholar
  28. Duarte J, Vargas JM (2004) Field interbreeding of released farm-reared Red-legged partridges (Alectoris rufa) with wild ones. Game Wildl Sci 21:55–61Google Scholar
  29. Frankham R (1999) Quantitative genetics in conservation biology. Genet Res 74:237–244PubMedCrossRefGoogle Scholar
  30. Garrido JL (2002) Capturas de perdiz roja (Economía inducida por la caza de perdiz). In: FEDENCA (ed) Aportaciones a la gestión sostenible de la caza, tomo I. FEDENCA, Madrid, pp 141–147Google Scholar
  31. Gortázar C, Villafuerte R, Martín M (2000) Success of traditional restocking of red-legged partridge for hunting purposes in areas of low density of northeast Spain Aragón. Z Jagdwiss 46:23–30CrossRefGoogle Scholar
  32. Grant PR, Grant BR (1992) Hybridization of bird species. Science 256:193–197PubMedCrossRefGoogle Scholar
  33. Grant PR, Grant BR (2000) Non-random fitness variation in two populations of Darwin’s finches. Proc Royal Soc B 267:131–138CrossRefGoogle Scholar
  34. Grant PR, Grant BR, Keller LF, Markert JA, Petren K (2003) Inbreeding and interbreeding in Darwin’s finches. Evolution 57:2911–2916PubMedGoogle Scholar
  35. Grant PR, Grant R, Markert JA, Keller LF, Petren K (2004) Convergent evolution of Darwin’s finches caused by introgressive hybridization and selection. Evolution 58(7):1588–1599PubMedGoogle Scholar
  36. Green RE (1984) Double nesting of the red-legged partridge Alectoris rufa. Ibis 126:332–346CrossRefGoogle Scholar
  37. Kandpal RP, Kandpal G, Weissman SM (1994) Construction of libraries enriched for sequence repeats and jumping clones, and hybridization selection for region-specific markers. Proc Natl Acad Sci USA 91:88–92Google Scholar
  38. Kaplan EL, Meier P (1958) Nonparametric estimation from incomplete observations. J Am Stat Assoc 53:457–481CrossRefGoogle Scholar
  39. Keane A, Brooke ML, Mcgowan PJK (2005) Correlates of extinction risk and hunting pressure in gamebirds (Galliformes). Biol Conserv 126:216–233CrossRefGoogle Scholar
  40. Keller LF, Waller DM (2002) Inbreeding effects in wild populations. Trends Ecol Evol 17:230–241CrossRefGoogle Scholar
  41. Kingsolver JG, Huey RB (2003) Introduction: the evolution of morphology, performance, and fitness. Integr Comp Biol 43:361–366PubMedCrossRefGoogle Scholar
  42. Laikre L, Schwartz MK, Waples RS, Ryman N (2010) Compromising genetic diversity in the wild: unmonitored large-scale release of plants and animals. Trends Ecol Evol 25(9):520–529PubMedCrossRefGoogle Scholar
  43. Leif AP (1994) Survival and reproduction of wild and pen-reared ring-necked pheasant hens. J Wildl Manage 58:501–506CrossRefGoogle Scholar
  44. Mallet J (2005) Hybridization as an invasion of the genome. Trends Ecol Evol 20(5):229–237PubMedCrossRefGoogle Scholar
  45. Martínez J, Viñuela J, Villafuerte R (2002) Socioeconomic and cultural aspects of gamebird hunting. REGHAB project, European Commission, BruselasGoogle Scholar
  46. McCarthy EM (2006) Handbook of avian hybrids of the world. Oxford University Press, New YorkGoogle Scholar
  47. McPhee ME (2003) Generations in captivity increases behavioural variance: considerations for captive breeding and reintroduction programs. Biol Conserv 115:71–77CrossRefGoogle Scholar
  48. Nadal J (1992) Problemática de las poblaciones de perdiz roja, bases ecoetológicas para tener éxito con las repoblaciones. In: La perdiz roja (ed) Gestión del Hábitat. Fundación La Caixa, Barcelona, Spain, pp 87–100Google Scholar
  49. Negro JJ, Torres MJ, Godoy JA (2001) RAPD analysis for detection and eradication of hybrid partridges (Alectoris rufa × A. Graeca) in Spain. Biol Conserv 9:19–24CrossRefGoogle Scholar
  50. Olden JD, Douglas ME, Douglas MR (2005) The human dimensions of biotic homogenization. Conserv Biol 19:2036–2038CrossRefGoogle Scholar
  51. Pérez JA, Alonso ME, Gaudioso VR, Olmedo JA, Díez C, Bartolomé D (2004) Use of radiotracking techniques to study a summer repopulation with red-legged partridge (Alectoris rufa) chicks. Poultry Sci 83:882–888Google Scholar
  52. Pollock KH, Winterstein SR, Bunck CM, Curtis PD (1989) Survival analysis in telemetry studies: the staggered entry design. J Wildl Manage 53:7–15CrossRefGoogle Scholar
  53. Potts GR (1989) The impact of releasing hybrid partridges on wild red-legged populations. Game Conserv Rev 20:81–85Google Scholar
  54. Puigcerver M, Vinyoles D, Rodríguez-Teijeiro JD (2007) Does restocking with Japanese quail or hybrids affect native populations of common quail Coturnix coturnix? Biol Conserv 168:628–635CrossRefGoogle Scholar
  55. Putaala A, Hissa R (2003) Breeding dispersal and demography of wild and hand-reared grey partridges Perdix perdix in Finland. Wildl Biol 4:137–145Google Scholar
  56. Randi E (2008) Detecting hybridization between wild species and their domesticated relatives. Mol Ecol 17:285–293PubMedCrossRefGoogle Scholar
  57. Rhymer JM, Simberloff D (1996) Extinction by hybridization and introgression. Annu Rev Ecol S 27:83–109CrossRefGoogle Scholar
  58. Ryan ME, Johnson JR, Fitzpatrick BM (2009) Invasive hybrid tiger salamander genotypes impact native amphibians. Proc Natl Acad Sci USA 106:11166–11171PubMedCrossRefGoogle Scholar
  59. Sáenz de Buruaga M, Lucio A, Purroy FJ (2001) Reconocimiento de sexo y edad en especies cinegéticas. EDILESA eds, LeónGoogle Scholar
  60. Sambrook J, Fritsch EF, Maniatis T (1989) Molecular cloning: a laboratory manual, 2nd edn. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, New YorkGoogle Scholar
  61. SAS (2001) SAS/STAT User’s guide, version 8.01. SAS Insitute Inc., CaryGoogle Scholar
  62. Skutch AF (1982) Parent birds and their young. Texas University Press, AustinGoogle Scholar
  63. StatSoft Inc (2002) STATISTICA data analysis software system, version 6.
  64. Tejedor MT, Monteagudo LV, Mautner S, Hadjisterkotis E, Arruga MV (2007) Introgression of Alectoris chukar genes into a Spanish wild Alectoris rufa population. J Hered 98:179–182PubMedCrossRefGoogle Scholar
  65. Templeton AR (1986) Coadaptation and outbreeding depression. In: Soulé ME (ed) Conservation biology; the science of scarcity and diversity. Sinauer Associates Inc., Sunderland, pp 105–116Google Scholar
  66. Vargas JM, Guerrero JC, Farfán MA, Barbosa AM, Real R (2006) Land use and environmental factors affecting red-legged partridge (Alectoris rufa) hunting yields in southern Spain. Euro J Wildl Res 52:188–195CrossRefGoogle Scholar

Copyright information

© Springer Science+Business Media B.V. 2011

Authors and Affiliations

  • F. Casas
    • 1
  • F. Mougeot
    • 2
  • I. Sánchez-Barbudo
    • 1
  • J. A. Dávila
    • 1
  • J. Viñuela
    • 1
  1. 1.Instituto de Investigación en Recursos Cinegéticos (IREC, CSIC-UCLM-JCCM)Ciudad RealSpain
  2. 2.Estacion Experimental de Zonas Áridas (EEZA-CSIC)La Cañada de San Urbano, AlmeríaSpain

Personalised recommendations