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Vögel Amazoniens: Ökologie, Brutbiologie und Artenreichtum

Birds of Amazonia: Ecology, breeding biology, and species richness

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Zusammenfassung

Die beiden ökologischen Hauptgruppen der Waldvögel Amazoniens bestehen aus Arten, welche die terra firme Wälder besiedeln, und Bewohnern der Vegetationszonen entlang von Flußläufen. In beiden Gruppen gibt es sowohl weit verbreitete Arten als auch solche mit lokaler Verbreitung. Die Areale der letzteren sind nicht regellos verteilt, sondern treten in bestimmten Regionen gehäuft auf (Endemismus-Gebiete). Die ökologische Bedeutung breiter Flüsse als Ausbreitungsschranken ist erheblich für Vogelarten, die das Innere der Wälder besiedeln, aber nur gering für viele flugaktive Waldvögel des Baumkronenbereichs sowie für Arten der Ufervegetation der Flußläufe.

Als Erklärung für den geographischen Ausschluß zahlreicher parapatrischer Vogelarten entlang von Kontaktzonen kommt nur ökologischer Konkurrenzausschluß infrage, jedoch liegen diesbezüglich Geländestudien noch nicht vor. Einige weiträumig parapatrische Arten sind in gewissen Gebieten sympatrisch und besetzen hier unterschiedliche Habitate (terra firme Wald bzw. várzea-Wald).

Im Innern der Wälder halten syntope Vogelarten unterschiedlich schmale vertikale Nahrungszonen ein, insbesondere nahe dem Waldboden und im Baumkronenbereich. Insektenfresser müssen ihre mehr oder weniger gleichmäßig und dünn verteilte Beute mit großem Zeitaufwand suchen und sind möglicherweise aus diesem Grunde zumeist monogam und territorial. Die meisten fruchtfressenden Vögel können ihren täglichen Nahrungsbedarf mit geringem Zeitaufwand decken. Das war wahrscheinlich eine wichtige Voraussetzung für die Entstehung von Arena-Balzsystemen. Durch Samenausbreitung haben Vögel eine große Bedeutung für die Erhaltung der räumlichen Heterogenität und taxonomischen Diversität der feuchten Tropenwälder.

Die Hauptbrutzeit der Vögel entlang dem Rio Amazonas und im südlichen Amazonien liegt im südlichen Frühjahr (September–November). Ein weniger markanter Anstieg der Brutaktivität wurde bei Manaus in Zentralamazonien außerdem im Januar—Februar und bei Bélem an der Amazonas-Mündung im Mai—Juni festgestellt. Im nördlichen Amazonien übernimmt diese untergeordnete Brutsaison zunehmend die Rolle der Hauptbrutzeit wie im nördlichen Südamerika (Venezuela, Trinidad, Guyana).

Die gegenwärtige Kenntnis der Phänologie und der ökologischen Hintergründe von periodischen Wanderungen gewisser amazonischer Waldvögel ist noch sehr lückenhaft. Derartige Wanderungen deuten an, daß die Ernährungsbasis in den jeweiligen Brutgebieten temporär unsicher werden kann. Auch Vogelarten, die entlang von Flußläufen Uferbereiche und Inseln besiedeln, oder im Flachwasser fischen, weichen während der Zeiten, wenn die Flüsse Hochwasser führen, durch periodische Wanderungen in andere Flußabschnitte aus. Nur wenige Zugvögel der nördlichen Breiten überwintern in Amazonien, und zwar ausschließlich in Sekundärvegetation und entlang von Waldrändern.

Die Vogelfauna der offenen Vegetationsformationen (campina, caatinga) ist relativ artenarm. Auch Entenvögel sind in Amazonien nur mit wenigen Arten vertreten. Dagegen sind lokale Waldvogel-Gemeinschaften 5-6mal artenreicher als solche der Gemäßigten Zonen. Jedoch sind die meisten Arten unterschiedlich selten. Auf einer Probefläche in Oberamazonien waren Arten mittlerer Häufigkeit nur mit 3 Paaren/km2 vertreten. Über 100 Arten dieser Artengemeinschaft hatten geringere Populationsdichten. Die Mindestgröße von Waldreservaten, um ca. 100 Paare von Vogelarten der genannten mittleren Häufigkeit zu schützen, müßte 30 km2 betragen. Für die selteneren Arten sind wesentlich größere Flächen zu fordern. Greifvögel und seminomadische Fruchtfresser mit besonders niedrigen Populationsdichten benötigen Waldreservate von mehreren tausend km2 Größe.

Zu den ökologischen Mechanismen für die Erhaltung des hohen tropischen Artenreichtums gehören die Bildung engerer ökologischer Nischen der Arten im Vergleich zu höheren Breiten sowie eine größere Zahl von Predatoren, welche die Populationsdichte der verschiedenen Beute-Arten niedrig halten. Die historische Ursache der Entstehung des tropischen Artenreichtums ist eine reiche Artdifferenzierung wahrscheinlich verbunden mit einer im Vergleich zu den Faunen höherer Breiten geringeren Aussterberate tropischer Tierarten. Dadurch könnte es im Laufe der geologischen Geschichte der letzten 60 Millionen Jahre (Tertiär-Quartär) zu einer „Akkumulation” von Arten in vielen tropischen Bereichen gekommen sein.

Summary

The two main ecological groups of Amazonian forest birds consist, respectively, of species inhabiting the terra firme forest and of species inhabiting vegetation zones along the river courses (sandbar scrub, riveredge forest, flood forest). Species in both groups are either rather widespread or fairly localized in distribution. Localized species of both ecological groups cluster in several areas of endemism located in peripheral portions of Amazonia (Napo, Inambari, Imerí, Rondônia, Guiana, Belém).

Broad river courses are formidable barriers to dispersal for bird species of the forest interior but are insignificant barriers for birds of the forest canopy and for species inhabiting the vegetation zones along river courses.

Geographical exclusion of numerous parapatric members of species pairs along well defined contact zones probably is due to ecological competition, although no field studies have been carried out so far to substantiate this assumption. Some of the regionally parapatric species are sympatric in certain areas and here occupy different habitats (terra firme forest and várzea forest, respectively).

Birds of the forest interior are restricted to varyingly narrow vertical feeding zones, especially near the forest floor and in the canopy. Insect eaters use much time in the search of their more or less evenly and rather thinly spread prey. This may be the reason why they are mostly monogamous and territorial. In contrast, most fruiteating birds are able to satisfy their daily food requirements in a short time. This was probably an important precondition for the development of arena courtship systems in some families of fruiteating birds. In view of their role as effective seed dispersal agents birds are important for the maintenance of the spatial heterogeneity and taxonomic diversity of humid tropical forests.

The main breeding season of birds along the Amazon River and in southern Amazonia falls in the months of the southern spring (September–November). A less conspicuous increase of breeding activity has been recorded near Manaus in central Amazonia during January and February and near Belém at the mouth of the Amazon River during the months May–June. Going north from central Amazonia, the subordinate peak of breeding activity during the first half of the year probably becomes increasingly more conspicuous until it represents the main breeding season as is the case in northern South America (Venezuela, Trinidad, Guyana).

The current knowledge of the phenology and ecological background of periodical migrations of certain Amazonian forest birds is still very meagre. These migrations indicate that the food supply of the species involved becomes temporarily uncertain in their breeding areas. Birds which inhabit river margins and islands or which fish in shallow water retreat to other portions of the Amazon river system during periods of high water level in their home range. Only few migrant birds from the north Temperature Zone spend the winter in Amazonia where they are restricted to secondary vegetation and forest borders.

The bird fauna of the open vegetation formations (campina, caatinga) is relatively poor in species number compared to the forest avifauna. Among waterbirds ducks (Anatidae) are represented in Amazonia by less species than in extratropical areas. On the other hand, local forest bird communities in Amazonia are 5–6 times richer in species than those of north Temperate Zone forests. However, individual Amazonian species are comparatively rare. On a test plot in upper Amazonia, species of median abundance had a density of only three pairs per 100 ha. More than 100 species in this community were represented by fewer pairs per 100 ha. A forest reserve of 30 km2 would be required to protect a minimum population of 100 pairs of the species of median abundance. Larger areas are needed for the numerous rarer species. The size of forest reserves to protect raptors and seminomadic fruiteaters with particularly low population densities is estimated at several thousand km2.

Some of the ecological mechanisms responsible for the maintenance of the high tropical species richness are the formation of narrower ecological niches compared to species of the higher latitudes as well as the occurrence of a higher number of predators in the tropics which keep the population densities of the various prey species low. The historical cause of the development of the high tropical species richness was a prolific species differentiation probably combined with a relatively lower extinction rate of the species compared to the faunas of the higher latitudes. In this way species in tropical faunas may have “accumulated” during the course of the Tertiary and Quaternary periods, i. e. over the last 60 Million years.

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Haffer, J. Vögel Amazoniens: Ökologie, Brutbiologie und Artenreichtum. J Ornithol 129, 1–53 (1988). https://doi.org/10.1007/BF01641531

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