Journal für Ornithologie

, Volume 124, Issue 4, pp 393–413

Einnischung und interspezifische Territorialität überwinternder Steinschmätzer(Oenanthe isabellina, O. oenanthe, O. pleschanka) in Kenia

  • Bernd Leisler
  • Georg Heine
  • Karl-Heinz Siebenrock
Article

Zusammenfassung

Ähnliche Habitatansprüche, fehlende nahrungsökologische Trennung und Nahrungsarmut der Habitate sind vermutlich die Ursache für interspezifische Territorialität sympatrisch vorkommender Steinschmätzer(Oenanthe)-Arten. Bei zwischenartlichen Auseinandersetzungen im Brutgebiet ergibt sich die Rangordnung Isabellschmätzer — Steinschmätzer — Nonnenschmätzer. Im Winterquartier in Kenia besetzten sowohl ♂ wie ♀ von Isabell-, Stein- und Nonnenschmätzer getrennte Reviere und verteidigten sie gegen gleich- wie andersgeschlechtliche Artgenossen (einzelne Individuen mindestens 29 Tage lang). In Ausnahmefällen verteidigten kurzfristig „Pseudopaare“ aller 3 Arten Reviere. Beim Nonnenschmätzer erfolgten innerartliche Angriffe nicht bevorzugt gegen ein Geschlecht. Sowohl bei den interspezifischen Auseinandersetzungen zwischen Mitgliedern der Gilde, als auch bei den Konflikten zwischen den 3 überwinterndenOenanthe-Arten griffen Isabellschmätzer signifikant häufiger an als sie selbst angegriffen wurden. Umgekehrt war es beim Nonnenschmätzer. Beim Steinschmätzer bestand kein signifikanter Unterschied. Bis auf denO. pileata waren die afrikanischen Arten bei zwischenartlichen Konflikten stets dominant (Tab. 2 und 3). Das Dominanzverhältnis bei zwischenartlichen Auseinandersetzungen von Mitgliedern der Gilde wurde etwa gleich stark von Körpergröße und Status einer Art (ob Afrikaner oder Paläarkt) bestimmt (Tab. 4, Abb. 1). Die Verteilung der Angriffe von Isabell- und Steinschmätzer auf die jeweils beiden anderenOenanthe-Arten unterscheidet sich im Brutgebiet und Winterquartier (Tab. 6). Angriffe von Stein- gegen Isabellschmätzer in Afrika widersprechenIvanitzkys (1980 b) Ansicht, die von ihm im Brutgebiet festgestellte fehlende Aggressivität von Stein- gegenüber Isabellschmätzern sei angeboren. Nonnenschmätzer greifen weder im Brutnoch im Überwinterungsgebiet die anderen beidenOenanthe-Arten an (Tab. 6), waren aber gegen Nahrungskonkurrenten um mehr bewegliche und fliegende Beute aggressiv. In Afrika attackierten sie Braunkehlchen, Schafstelzen und Rauchschwalben. Nonnenschmätzer nutzten den Lebensraum etwas anders als die übrigen Schmätzer. Sie bewegten sich bei der Nahrungssuche vermutlich am schnellsten fort (Abb. 2), verbrachten mehr Zeit als die übrigen auf Warten und gingen nur kurz auf den Boden, um schnell Beute aufzunehmen (Abb. 3). Dadurch war es ihnen möglich, den aggressiveren, mehr bodenlebenden Arten auszuweichen. Diese Strategie, Kämpfe zu vermeiden, können als Anpassungen an den artgemäßen Lebensraum gedeutet werden. Weiter scheint das lockerere Reviersystem der spät brütenden Art die flexible Struktur gefördert zu haben. Nach Literaturangaben und eigenen Beobachtungen überlappen sich ökologisch einerseits Ruß-, Erd-, Isabell- und Steinschmätzer, andererseits Rüppell- und Nonnenschmätzer stark. Arten mit ausgeprägterem Bodenleben, zeichnen sich durch abnehmende Flügel- und zunehmende Beinbetonung, sowie durch einen kräftigeren Schnabel aus. Nonnenschmätzer zeigen als Anpassungen an Warten- und Flugjagd geringes Körpergewicht, langen Schwanz, lange Vibrissen und flachen Schnabel (Tab. 5, Abb. 4).

Patterns of interspecific interactions among wintering Wheatears(Oenanthe isabellina, O. oenanthe, O. pleschanka) and resident Chats in Kenya

Summary

Wheatears have similar habitat requirements, overlapping diets, and generally live under conditions where the food supply is not stable. Where different species occur side by side spatial separation is brought about by interspecific territoriality (Panov 1974,Cornwallis 1975). Interspecific relationships in the breeding areas are largely hierarchical, e.g. where Isabelline, Common and Pied Wheatear occur togetherisabellina is the dominant andpleschanka the subordinate species (Ivanitzky 1980). We investigated the following questions on the wintering grounds of the three species in Kenya: 1) Does the interspecific dominance hierarchy remain the same as in Central Asia? 2) How do the African counterparts behave when invaded by the palearctic migrants? 3) How does the Pied Wheatear manage to coexist without fighting? Our study sites were mainly burned grass areas at Lake Nakuru where a guild of small Turdid Chats could be found which consisted of the three palearcticOenanthe species, the Rock Thrush and Whinchat and the African species, the Anteater Chat, Capped and Mourning Wheatear. Fiscal Shrikes and Yellow Wagtails were also included (Tab. 1). In all three wintering Wheatears ♂ as well as ♀ occupied separate feeding territoires and defended them intra- as well as interspecifically (single birds up to at least 29 days). As an exception we observed “pseudopairs” in all three species defending territories. In the Pied Wheatear intraspecific interactions were random between the two sexes (Tab. 2). In interspecific encounters the Isabelline Wheatear attacked more often than it was attacked. No difference between attacks and retreats could be found in the Common Wheatear, whereas the Pied Wheatear was more often the recipient than the donor of attacks (Tab. 3). There was a tendency for resident species to be dominant in interspecific clashes (Tab. 3, Fig. 1). The dominance in interspecific encounters between members of the guild was equally influenced by the body weight and status (migrant or resident) of a species (Tab. 4). The proportion of attacks of Isabelline Wheatear and Common Wheatear against the two otherOeananthe-species differed from that found on the breeding grounds (Tab. 6). Attacks of Common against Isabelline Wheatears contradictIvanitzky's (1980) interpretation that the lack of aggression he observed in the breeding areas is an innate character of Common Wheatears. Pied Wheatears are subordinate in the breeding areas as well as on the wintering grounds, although they do attack unrelated species which are possible food competitors. In Kenya they were observed to chase away Whinchats, Yellow Wagtails and Swallows. This behaviour documents that interspecific attacks in Wheatears cannot be interpreted as misdirected intraspecific aggression. Pied Wheatears differed slightly in the habitat utilization from the other congeners. They employed more flighted locomotion and a perch and pounce foraging pattern (Fig. 2, 3). These characters promote the species' fugitive strategy to avoid skirmishes with ground living, more aggressive congeners. Also the looser territorial structure of the late breeding Pied Wheatear may have favoured the evolution of a fugitive strategy. Anteater Chat, Capped, Isabelline and Common Wheatears seem to overlap most on the wintering grounds as do Mourning and Pied Wheatears. The following morphological adaptations could be found (Tab. 5, Fig. 4). The Whinchat which clings on herbaceous vegetation has the largest foot. Turdid Chats with pronounced pedal locomotion, i.e. all the African species investigated and the Isabelline Wheatear have long tarsi, short round wings and a strong beak. Adaptations of the Pied Wheatear to its use of flimsy perches and its more aerial feeding are low weight and a flat bill, long rictal bristles and a long tail.

Preview

Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.

Literatur

  1. Amadon, D. (1943): Bird weights as an aid in taxonomy. Wilson Bull. 55, 164–177.Google Scholar
  2. Barlow, J. C. (1980): Patterns of ecological interactions among migrant and resident Vireos on the wintering grounds. In:Keast &Morton (ed.): Migrant Birds in the Neotropics 79–107.Google Scholar
  3. Bennett, S. E. (1980): Interspecific competition and the niche of the American Redstart(Setophaga ruticilla) in wintering and breeding communities. In:Keast &Morton (ed.): Migrant Birds in the Neotropics 319–335.Google Scholar
  4. Berck, K.-H. (1961): Bemerkungen zur Biologie des Steinschmätzers. Vogelwelt 82: 89–112.Google Scholar
  5. Borrett, R. P., &H. D. Jackson (1970): The European WheatearOenanthe oenanthe (L.) in southern Africa. Bull. B.O.C. 90: 124–129.Google Scholar
  6. Brittton, P. L. (ed.) (1980): Birds of East Africa. Nairobi, EANHS.Google Scholar
  7. Brown, L. H., &P. L. Britton (1980): The breeding seasons of East African birds. EANHS, Nairobi.Google Scholar
  8. Catchpole, C. K. (1978): Interspecific territorialism and competition inAcrocephalus warblers as revealed by playback experiments in areas of sympatry and allopatry. Anim. Behav. 26: 1072–1080.Google Scholar
  9. Cody, M. L. (1968): On the methods of resource division in grassland bird communities. Amer. Nat. 102: 107–147.Google Scholar
  10. Ders. (1974): Competition and the structure of bird communities. Princeton Univ. Press, Princeton, New Jersey.Google Scholar
  11. Cornwallis, L. (1975): The comparative ecology of eleven species of wheatear (GenusOenanthe) in S. W. Iran. Doctoral Ph. thesis, Univ. Oxford.Google Scholar
  12. Crook, J. H. (1961): The fodies (Ploceinae) of the Seychelles Islands. Ibis 103 a: 517–548.Google Scholar
  13. Curry-Lindahl, K. (1981): Bird migration in Africa. 2 Vol. Academic Press London, New York.Google Scholar
  14. Delius, J. D. (1963): Das Verhalten der Feldlerche. Z. Tierpsychol. 20: 297–348.Google Scholar
  15. Des Granges, J. L., &P. R. Grant (1980): Migrant Hummingbirds' accomodation into tropical communities. In:Keast &Morton (ed.): Migrant Birds in the Neotropics, 395–409.Google Scholar
  16. Dittami, J. (1981): Observations on wintering wheatears in burned grass areas at Lake Nakuru, Kenya. Vogelwarte 31: 177–178.Google Scholar
  17. Eggebrecht, E. (1943): Beitrag zur Brutbiologie des Nonnensteinschmätzers(Oenanthe pleschanka pleschanka (Lepechin)). Orn. Mber. 51: 127–135.Google Scholar
  18. Fitzpatrick, J. W. (1980): Wintering of North American Tyrant flycatchers in the Neotropics. In:Keast &Morton (ed.): Migrant Birds in the Neotropics 67–78.Google Scholar
  19. Gochfeld, M. (1979): Interspecific territoriality in Red-breasted meadowlarks and a method for estimating the mutuality of their participation. Behav. Ecol. Sociobiol. 5: 159–170.Google Scholar
  20. Haas, V. (1982): Social organisation and role of helpers in Anteater chats(Myrmecocichla aethiops) and Mourning wheatears(Oenanthe lugens schalowi). Abstracts XVIII Congr. Intern. Orn. 204, Moskau, Nauka.Google Scholar
  21. Hartley, P. H. T. (1949): The biology of the Mourning chat in winter quarters. Ibis 91: 393–413.Google Scholar
  22. Hespenheide, H. A. (1980): Bird community structure in two Panama forests: Residents, migrants, and seasonality during the nonbreeding season. In:Keast &Morton (ed.): Migrant Birds in the Neotropics 227–237.Google Scholar
  23. Hutto, R. L. (1980): Winter habitat distribution of migratory land birds in Western Mexico, with special reference to small foliagegleaning insectivores. In:Keast &Morton (ed.): Migrant Birds in the Neotropics 181–203.Google Scholar
  24. Ivanitzky, V. V. (1980 a): (Interspecific relations of sympatric species of wheatears (Oenanthe, Turdinae, Passeriformes). 1. Ecological prerequesites of competition and spatial structure of a mixed population). Zool. Sh. 59: 587–597 (russisch).Google Scholar
  25. Ders. (1980 b): (Interspecific relations of sympatric species of wheatears (Oenanthe, Turdidae, Aves). 2. Behavioural aspects of coexistence in closely related species). Zool. Sh. 59: 739–749 (russisch).Google Scholar
  26. Ders. (1981): (Some aspects of the organization of intrapopulation interactions in passeriformes). J. allg. Biol. 42: 708–720 (russisch).Google Scholar
  27. Johnson, T. B. (1980): Resident and North American migrant bird interactions in the Santa Marta highlands, Northern Columbia. In:Keast &Morton (ed.): Migrant Birds in the Neotropics 239–247.Google Scholar
  28. Kalinoski, R. (1975): Intra- and interspecific aggression in House finches and House sparrows. Condor 77: 375–384.Google Scholar
  29. Keast, A. (1980 a): Spatial relationships between migratory Parulid warblers and their ecological counterparts in the Neotropics. In:Keast &Morton (ed.): Migrant Birds in the Neotropics 109–130.Google Scholar
  30. Ders. (1980 b): Migratory Parulidae: What can species co-occurrence in the North reveal about ecological plasticity and wintering patterns? In:Keast &Morton (ed.): Migrant Birds in the Neotropics 309–317.Google Scholar
  31. Ders. &E. S. Morton (ed.) (1980): Migrant Birds in the Neotropics. Ecology, behavior, distribution and conservation. Smithsonian Inst. Press, Washington.Google Scholar
  32. Kipp, F. A. (1958): Zur Geschichte des Vogelzuges auf Grundlage der Flügelanpassungen. Vogelwarte 19: 233–242.Google Scholar
  33. Lack, D. L. (1971): Ecological isolation in birds. Oxford.Google Scholar
  34. Leisler, B. (1980 a): Morphological aspects of ecological specialization in bird genera. Ökol. Vögel 2: 199–220.Google Scholar
  35. Ders. (1980 b): Ökomorphologische Freiland- und Laboratoriumsuntersuchungen. Acta XVII Congr. Intern. Orn.: 202–208.Google Scholar
  36. Ders. &E. Thaler (1982): Differences in morphology and foraging behaviour in the goldcrestRegulus regulus and firecrestR. ignicapillus. Ann. Zool. Fenn. 19: 277–284.Google Scholar
  37. Maynard Smith, J. (1974): The theory of games and the evolution of animal conflicts. J. theor. Biol. 47: 209–211.Google Scholar
  38. Meinertzhagen, R. (1954): Birds of Arabia. Edinburgh, London.Google Scholar
  39. Moreau, R. E. (1972): The Palaearctic-African bird migration systems. Academic Press, London, New York.Google Scholar
  40. Ders. &W. M. Moreau (1928): Some notes on the habits of Palaearctic migrants while in Egypt. Ibis 12: 233–252.Google Scholar
  41. Morel, G., &F. Roux (1966): Les Migrateurs Paléarctiques au Sénégal II. Passereaux et synthèse générale. Terre et la Vie 2: 143–176.Google Scholar
  42. Morse, D. H. (1974): Niche breadth as a function of social dominance. Amer. Nat. 108: 818–830.Google Scholar
  43. Morton, E. S. (1980 a): Adaptations to seasonal changes by migrant land birds in the Panama Canal zone. In:Keast &Morton (ed.): Migrant Birds in the Neotropics 437–453.Google Scholar
  44. Ders. (1980 b): The importance of migrant birds to the advancement of evolutionary theory. In:Keast &Morton (ed.): Migrant Birds in the Neotropics 555–557.Google Scholar
  45. Murray, B. G., Jr. (1971): The ecological consequence of interspecific territorial behavior in birds. Ecology 52: 414–423.Google Scholar
  46. Myers, J. P. (1980): The pampas shore bird community: Interactions between breeding and nonbreeding members. In:Keast &Morton (ed.): Migrant Birds in the Neotropics 37–49.Google Scholar
  47. Orejuela, J. E., R. J. Raitt &H. Alvarez (1980): Differential use by North American migrants of three types of Colombian forests. In:Keast &Morton (ed.): Migrant Birds in the Neotropics 253–264.Google Scholar
  48. Panov, E. N. (1974): Die Steinschmätzer der nördlichen Paläarktis. Neue Brehm Bücherei, Ziemsen; Wittenberg.Google Scholar
  49. Ders. (1978): (Kommunikationsmechanismen bei Vögeln) Moskau, Nauka (russisch).Google Scholar
  50. Ders. &V. V. Ivanitzky (1975): (Interspecific spatial relationships in a mixed population ofOenanthe finschii andpleschanka on the Mangyshlak peninsula). Zool. Sh. 54: 1357–1370 (russisch).Google Scholar
  51. Pitelka, F. A. (1951): Ecological overlap and interspecific strife in breeding populations of Anna and Allen humming birds. Ecology 32: 641–661.Google Scholar
  52. Potapova, E. G., &E. N. Panov (1977): (Structural patterns of jaw apparatus in wheatears of the genusOenanthe, with special reference to the ways of decrease of food competition between closely related species). Zool. Sh. 56: 743–752 (russisch).Google Scholar
  53. Powell, G. V. N. (1980): Migrant participation in neotropical mixed species flocks. In:Keast &Morton (ed.): Migrant Birds in the Neotropics 477–483.Google Scholar
  54. Rabenold, K. N. (1980): The Black-throated Green Warbler in Panama: Geographic and seasonal comparison of foraging. In:Keast &Morton (ed.): Migrant Birds in the Neotropics 297–307.Google Scholar
  55. Rappole, J. H., &D. W. Warner (1980): Ecological aspects of migrant bird behavior in Veracruz, Mexico. In:Keast &Morton (ed.): Migrant Birds in the Neotropics 353–393.Google Scholar
  56. Remmert, H. (1980): Ökologie. Zweite Aufl. Springer Berlin, Heidelberg.Google Scholar
  57. Reynolds, J. F. (1974): Palaearctic birds in East Africa. Brit. Birds 67: 70–76.Google Scholar
  58. Ripley, S. D. (1961): Aggressive neglect as a factor of interspecific competition in birds. Auk 78: 366–371.Google Scholar
  59. Root, R. B. (1967): The Niche Exploitation Pattern of the Blue-Gray Gnatcatcher. Ecol. Monogr. 37: 317–350.Google Scholar
  60. Russell, S. M. (1980): Distribution and abundance of North American migrants in lowlands of Northern Columbia. In:Keast &Morton (ed.): Migrant Birds in the Neotropics 249–252.Google Scholar
  61. Sachs, L. (1972): Statistische Auswertungsmethoden. 3. Aufl. Springer Berlin, Heidelberg, New York.Google Scholar
  62. Simmons, K. E. L. (1951): Interspecific territorialism. Ibis 93: 407–413.Google Scholar
  63. Sinclair, A. R. E. (1978): Factors effecting the food supply and breeding season of resident birds and movements of Palaearctic migrants in a tropical African savannah. Ibis 120: 480–497.Google Scholar
  64. Sluys, R., &M. van den Berg (1982): On the specific status of the Cyprus Pied WheatearOenanthe cypriaca. Ornis Scand. 13: 123–128.Google Scholar
  65. Smith, K. D. (1971): Notes onOenanthe species in winter in Africa. Bird Study 18: 71–79.Google Scholar
  66. Stresemann, E. (1950): Interspecific competition in chats. Ibis 92: 148.Google Scholar
  67. Terborgh, J. W., &J. R. Faaborg (1980): Factors affecting the distribution and abundance of North American migrants in the Eastern Caribbean region. In:Keast &Morton (ed.): Migrant Birds in the Neotropics 145–155.Google Scholar
  68. Thiollay, J. M. (1979): Structure et dynamique du peuplement avien d'un mattoral aride (Bolson de Mapimi, Mexique). Terre et Vie 33: 563–589.Google Scholar
  69. Tramer, E. J., &T. R. Kemp (1980): Foraging ecology of migrant and resident warblers and vireos in the highland of Costa Rica. In:Keast &Morton (ed.): Migrant Birds in the Neotropics 285–296.Google Scholar
  70. Ursprung, J. (1980): Interspezifische Territorialität bei Rohrsängern. — Fragestellung, Methodik und erste Ergebnisse einer laufenden Untersuchung am Neusiedlersee. Biol. Forsch. Inst. Burgenland Ber. 37: 67–71.Google Scholar
  71. Vaurie, C. (1959): The Birds of the Palaearctic Fauna — Passeriformes. Witherby, London.Google Scholar
  72. Vogl. R. J. (1974): Effects of fire on grasslands. In:T. T. Kozlowski (ed.): Fire and Ecosystems. Academic Press, New York.Google Scholar
  73. Wiens, J. A. (1965): Behavioral interactions of red-winged blackbirds and common grackles on a common breeding ground. Auk 82: 356–374.Google Scholar
  74. Winkler, H. (1971): Die Bedeutung der Organisation angeborenen Verhaltens für das Verständnis der Ökologie der Wirbeltiere. Sitz.ber. Österr. Akad. Wiss., Mathem.-naturw. Kl., Abt. I, 179: 3–4.Google Scholar
  75. Wolters, H. E. (1975-82): Die Vogelarten der Erde. Parey, Hamburg, Berlin.Google Scholar

Copyright information

© Verlag der Deutschen Ornithologen-Gesellschaft 1983

Authors and Affiliations

  • Bernd Leisler
    • 1
  • Georg Heine
    • 1
  • Karl-Heinz Siebenrock
    • 1
  1. 1.Aus dem Max-Planck-Institut für Verhaltensphysiologie, Vogelwarte RadolfzellGermany

Personalised recommendations