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Taxonomy and phylogeny of reed warblers (genusAcrocephalus) based on mtDNA sequences and morphology

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Abstract

The mitochondrial cytochrome b gene of the majority ofAcrocephalus species (76 individuals) was amplified by PCR and sequenced directly. Nucleotide sequences (1068 base pairs) were used to reconstruct phylogenetic relationships within the genusAcrocephalus as well as betweenAcrocephalus and other sylviid warblers, particularlyHippolais. Acrocephalus andHippolais share ancestry and cluster in a monophyletic clade.Hippolais appears to represent a polyphyletic assemblage sinceH. icterina figures as the sister taxon toAcrocephalus, whereas “Hippolais”pallida andcaligata cluster withinAcrocephalus. The followingAcrocephalus clades could be recognized: (1) Large reed warblers form a clade consisting of a monophyletic Palearctic-Australasian subgroup (arundinaceus, stentoreus brunnescens, orientalis, australis, andvaughani) and a monophyletic Afrotropical subgroup (brevipennis, rufescens, gracilirostris, sechellensis, andnewtoni).A. griseldis holds an isolated position at the base of the large reed warbler clade. Within the small reed warblers, two probably monophyletic clades are apparent: (2) the striped species (withbistrigiceps, melanopogon, paludicola, andschoenobaenus), and (3) the small plain-coloured complex (consisting ofdumetorum, palustris, scirpaceus, s. fuscus, baeticatus, andavicenniae plus the neighbouringagricola-complex withagricola, tangorum, andconcinens). The relationship between these groups cannot be resolved. The molecular data clarify the status of some taxa, the systematic position of which has been controversial. A morphometric analysis (PCA) of 20 external characters confirmed the basic complexes, and unveiled adaptations of general importance among clades. At species level, we found less congruence between molecular and morphological data, which can be interpreted as a consequence of specializing adaptations and convergence. The major complexes established by molecular and morphometric analyses are further supported by distributional, acoustical, and oological affinities. A sound phylogenetic framework of the genus makes it now possible to examine the distribution of ecological and behavioural characters and to differentiate informative or convergent characters.Acrocephalus may be split into four previously recognized genera with the following names:Acrocephalus for the large,Calamodus for the striped,Notiocichla for the small plain, andIduna for the brownish “Hippolais” species.

Zusammenfassung

Die artenreiche GattungAcrocephalus diente in den letzten 20 Jahren als Modellgruppe für verschiedenste ökologische, verhaltens- und evolutionsbiologische Fragestellungen, wobei sich häufig eine vergleichende Bearbeitung als besonders lohnend erwies. Da alle diese Studien auf der traditionellen Systematik basieren, war es nötig, die Phylogenie der Gattung zu rekonstruieren, um: (1) einen verläßlichen Stammbaum verfügbar zu haben, (2) phylogenetische Korrekturen bei vergleichenden Ansätzen durchführen und (3) konvergente Merkmalsentwicklungen erkennen zu können.

Bei 23Acrocephalus-Arten (in 27 Unterarten) wurde das mitochondrielle Cytochrom b-Gen mittels PCR amplifiziert und sequenziert. Anhand der Unterschiede in den Nukleotidsequenzen (1068 Basenpaare) erstellten wir einen Stammbaum der GattungAcrocephalus und untersuchten die verwandtschaftlichen Beziehungen zu anderen Sylviiden, besondersHippolais. Hippolais erweist sich als eine polyphyletische Gruppe.Hippolais (icterina und andere Arten) ist das Schwestertaxon vonAcrocephalus. “H.” pallida undcaligata gehören in die GattungAcrocephalus (subgenusIduna, Abb. 3, 4). Alle molekularen Stammbäume (Abb. 1–4) unterteilenAcrocephalus in drei monophyletische Hauptgruppen: 1. die großen Arten und die kleinenIduna Arten als wahrscheinliche Schwestergruppe, 2. die gestreiften und 3. die kleinen einfarbigen Arten. Die großen Rohrsänger umfassen eine paläarktisch-australasiatische Untergruppe (mitarundinaceus, stentoreus (brunnescens),orientalis, australis undvaughani) sowie eine afrotropische (mitbrevipennis, rufescens, gracilirostris, sechellensis undnewtoni).A. griseldis nimmt eine isolierte Stellung an der Basis der großen Rohrsänger ein und ist möglicherweise eine ursprüngliche Form. Die gestreiften Arten (mitschoenobaenus, bistrigiceps, melanopogon undpaludicola) bilden ebenso eine geschlossene Abstammungsgemeinschaft wie die kleinen einfarbigen, bei denen sich zwei Gruppen abheben (eine mitdumetorum, palustris, scirpaceus, s. fuscus, baeticatus undavicenniae und eine mitagricola, tangorum undconcinens). Die verwandtschaftlichen Beziehungen zwischen diesen drei Hauptgruppen lassen sich nicht auflösen. Morphologische Untersuchungen von 20 äußeren Merkmalen bestätigen einerseits die nach genetischen Ähnlichkeiten aufgestellten Hauptgruppen, die sich durch unterschiedliche Basisanpassungen auszeichnen (Abb. 6, 7). Auf Artniveau stimmen dagegen die aufgrund morphologischer bzw. genetischer Ähnlichkeiten gebildeten Gruppen weniger gut überein (Abb. 5). Diese Diskrepanz ist auf die spezialisierenden Anpassungen der einzelnen Arten an ihre Habitate und auf Konvergenz zurückzuführen. Die Gliederung der Großgruppen entspricht der traditionellen Systematik. Sie wird auch durch die Verbreitung der Arten und durch bioakustische und oologische Merkmale gestützt. In der Feinsystematik klären die molekularen Ergebnisse die verwandtschaftliche Stellung einiger Taxa, deren Einordnung bisher umstritten war: z. B. bildetA. tangorum mitA. concinens undA. agricola eine Gruppe;A. orientalis scheint näher mitA. stentoreus als mitA. arundinaceus verwandt;A. avicenniae scheint eine eigene Art in derscirpaceus-Gruppe zu sein. In weiteren Studien kann nun die Verteilung phänotypischer Merkmale (z. B. Zeichnungsmuster, Gesangsmerkmale, Nestbauweisen) auf den verschiedenen Niveaus des Stammbaums untersucht und historisch oder adaptiv erklärt werden. Z. B. haben sich dunkle Brauenstreifen und Schwanzstelzen verwandtschaftsunabhängig (konvergent) sowohl bei den gestreiften Rohrsängern als auch in deragricola-Gruppe entwickelt; extrem unterschiedliche Paarungssysteme entstanden bei den gestreiften Rohrsängern usw. Wollte man die GattungAcrocephalus aufteilen, sollteAcrocephalus für die großen Arten,Calamodus für die gestreiften,Notiocichla für die kleinen einfarbigen stehen undIduna für die bräunlichen “Hippolais”-Vertreter.

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Authors and Affiliations

  1. Max-Planck-Institut für Verhaltensphysiologie, Vogelwarte Radolfzell, D-78315, Radolfzell

    Bernd Leisler

  2. Institut für Pharmazeutische Biologie der Universität Heidelberg, Im Neuenheimer Feld 364, D-69120, Heidelberg

    Petra Heidrich & Michael Wink

  3. Bleichgrabenstr. 37, D-41063, Mönchengladbach

    Karl Schulze-Hagen

Authors
  1. Bernd Leisler
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  2. Petra Heidrich
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  3. Karl Schulze-Hagen
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  4. Michael Wink
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Leisler, B., Heidrich, P., Schulze-Hagen, K. et al. Taxonomy and phylogeny of reed warblers (genusAcrocephalus) based on mtDNA sequences and morphology. J Ornithol 138, 469–496 (1997). https://doi.org/10.1007/BF01651381

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