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

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

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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).

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.

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Leisler, B., Heine, G. & Siebenrock, KH. Einnischung und interspezifische Territorialität überwinternder Steinschmätzer(Oenanthe isabellina, O. oenanthe, O. pleschanka) in Kenia. J Ornithol 124, 393–413 (1983). https://doi.org/10.1007/BF01640360

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