Journal of Ornithology

, Volume 145, Issue 3, pp 206–222

Radiation and species limits in the Asian Pallas’s warbler complex (Phylloscopus proregulus s.l.)*

  • Jochen Martens
  • Dieter Thomas Tietze
  • Siegfried Eck
  • Michael Veith
Original Article


Pallas’s Warbler (Phylloscopus proregulus) of the Eastern Palaearctic (Siberia, China, Himalayas) resolves into a complex of four closely related but distinct allospecies forming a superspecies, P. [proregulus]. Assignment of species rank is based on cytochrome-b gene sequences and on vocalisations (territorial song and calls). Morphological divergence is poor, as is typical in Phylloscopus. It applies to size (nominate Siberian proregulus as opposed to the Sino-Himalayan taxa), and slight coloration differences among the latter taxa do not correspond to the cyt-b topology and are considered to be adaptive. Vocal differences relate to syntax (verse song vs endless song), frequency (broad vs narrow bandwidth, allocation of elements within the frequency band), form of elements (ascending vs descending parts, combination of these), and repertoire size. Strong premating isolating mechanisms are assigned to vocalisations. Contrary to current taxonomy, we propose the following constituent taxa of the Phylloscopus [proregulus] complex: Phylloscopus proregulus s.str. (Siberia, disjunct from the following taxa), P. kansuensis (China: Gansu, Qinghai), P. forresti (W/SW China, part of E Himalayas?), P. chloronotus with two subspecies: P. c. simlaensis (W Himalayas), and P. c. chloronotus (C and E Himalayas). Contact zones and possible hybridisation between the Sino-Himalayan taxa remain undetermined. A closely related isospecies is Phylloscopus yunnanensis (syn. P. sichuanensis), which is locally sympatric with and extremely similar morphologically to P. forresti and P. kansuensis. Cyt-b distance values range from 3.1% to 4.6% within the P. proregulus complex and from 7.5% to 8.4% between the latter and P. yunnanensis. This corresponds to differentiation times of 4.1 to 5.5 Myears (P. proregulus complex vs P. yunnanensis) and 1.7 to 3.2 Myears (within the P. proregulus complex).


Phylloscopus proregulus complex Phylloscopus yunnanensis Cytochrome-b gene Vocalisations Species limits 

Supplementary material

appendices.pdf (252 kb)
Appendix 1 +2 (PDF 253 KB)

Sound archive

Phylloscopus proregulus s. str., part of the “endless” territorial song, recorded by S. Ernst (Fig. 3 a–h); background: Parus ater.

pro-s.wav (1.3 mb)
pro-s.wav (1.2 MB)

Phylloscopus kansuensis, territorial song, 2 verses, recorded by J. Martens (Fig. 31); background: Carpodacus thura.

kan-s.wav (374 kb)
kan-s.wav (347 KB)

Phylloscopus forresti, territorial song, song form A, 2 verses, recorded by J. Martens (Fig. 3 m); background: Seicercus valentini.

chla-s.wav (654 kb)
chla-s.wav (655 KB)

Phylloscopus forresti, part of the “endless” territorial song, song form B, recorded by J. Martens (Fig. 3 n–p).

chlb-s.wav (672 kb)
chlb-s.wav (672 KB)

Phylloscopus chloronotus simlaensis, part of the “endless” territorial song, recorded by J. Martens (Fig. 4 a–g).

sim-s.wav (1.6 mb)
sim-s.wav (1.6 MB)

Phylloscopus yunnanensis, part of the “endless” territorial song, recorded by J. Martens (Fig. 4 k).

yun-s.wav (1 mb)
yun-s.wav (1.0 MB)

Phylloscopus yunnanensis, part of a “call song”, China, Shaanxi, Taibai Shan, recorded by J. Martens (similar to Fig. 4 u–v).

yun-c.wav (1.1 mb)
yun-c.wav (1.1 MB)

Phylloscopus proregulus s. str., call, recorded by J. Martens (Fig. 7 a).

pro-c.wav (43 kb)
pro-c.wav (44 KB)

Phylloscopus kansuensis, call, recorded by J. Martens (Fig. 7 f).

kan-c.wav (43 kb)
kan-c.wav (44 KB)

Phylloscopus forresti, call, recorded by J. Martens (Fig. 7 h).

chl-c.wav (41 kb)
chl-c.wav (41 KB)

Phylloscopus chloronotus simlaensis, call, recorded by J. Martens (Fig. 7 p).

sim-c.wav (42 kb)
sim-c.wav (42 KB)

Copyright information

© Dt. Ornithologen-Gesellschaft e.V.  2004

Authors and Affiliations

  • Jochen Martens
    • 1
  • Dieter Thomas Tietze
    • 1
  • Siegfried Eck
    • 2
  • Michael Veith
    • 1
  1. 1.Institut für ZoologieJohannes Gutenberg-UniversitätMainzGermany
  2. 2.Staatliche Naturhistorische Sammlungen DresdenMuseum für TierkundeDresdenGermany

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