ArticlePDF Available

A new species of Phylloscopus warbler from Hainan Island, China

Authors:

Abstract and Figures

Phylloscopus hainanus is a distinctive new species in the family Sylviidae. The species is endemic to Hainan Island, China, where it is restricted to tropical forest in mountains above 600 m. The most distinctive plumage features are the rather deep yellow underparts and the mainly white outer rectrices.
Content may be subject to copyright.
1993 3
A
new species of
Phylloscopus
warbler from Hainan Island, China
URBAN OLSSON', PER ALSTROMZ
&
PETER
R.
COLSTON3
University
of
Goteborg. Department
of
Zoology.
Box
25059,
S-400
3
1
Goteborg, Sweden
Kungsgatan
3,
S-462 33
Vanersborg, Sweden
Bird Group, Zoological Department, The Natural History Museum, Tring. Herts.
HP23 6AP,
UK
Phylloscopus hainanus
is a distinctive new species in the family Sylviidae. The species is
endemic to Hainan Island, China, where it
is
restricted to tropical forest in mountains
above
600
m. The most distinctive plumage features
are
the rather deep yellow underparts
and the mainly white outer rectrices.
In 1986
U.O.
examined four skins of what were labelled as
the Hainan Island subspecies of the Sulphur-breasted War-
bler
P.
ricketti goodsoni
in the Institute of Zoology, Academia
Sinica, Kunming. China (IZASK) (collected at Jian Feng Ling,
Ledong County, west Hainan [18"40'N. 108"45'E], and Diao
Lou, Lingshui County, south Hainan [18"47'N, 110"05'E]).
He was immediately struck by their distinctness from all
other species in the genus and considered that this taxon
might be a distinct species.
In mid-March 1988 P.A. and
U.O.
went to Hainan Island
in order to investigate the taxonomic status of the form.
However, despite thorough surveys
of
apparently suitable
habitat in the vicinity of Qiongzhong in central Hainan. they
were unable to find the warbler. Later the same year, P.A.
and
U.O.
found another specimen of the same form in the
Institute of Zoology, Academia Sinica. Beijing. China (col-
lected at Daio Lou. south Hainan). Colour photographs of
this specimen taken by P.R.C. are lodged in the British Mu-
seum (Natural History), Tring,
U.K.
In April 1988, Stig Jen-
sen (pers. comm.) found the bird to be common at Jian
Feng Ling Forest Reserve, Hainan, and obtained a tape re-
cording of its song. P.A. and
U.O.
noted that the song differed
significantly from that of nominate
P.
ricketti
of the mainland.
In late May 1989, P.A.. P.R.C. and
U.O.
went to Jian Feng
Ling Forest Reserve to study the species. Unfortunately,
breeding was over, and the birds were moving around singly
or in small flocks, singing only sporadically. Nevertheless,
the vocal and morphological distinctness of the form was
confirmed. In May 1990. P.A. examined the specimens in
the Institute of Zoology, Academia Sinica. Kunming, where
he confirmed the observations by
U.O.
In 1990 one of the two specimens from Hainan Island in
the collections of the American Museum of Natural History,
New York, which were considered to be
P.
ricketti goodsoni.
was loaned to the British Museum (Natural History), Tring.
There it was examined by P.R.C., who also took photographs
of it, which he sent to P.A. and
U.O.
To our surprise, it was
not the same form as we had been pursuing on Hainan
Island, but Blyth's Crowned Willow Warbler
P.
reguloides.
Later the same year, P.A. examined the holotype and again
the paratype (the specimen which had previously been sent
to P.R.C.)
of
P.
ricketti goodsoni
in the American Museum of
Natural History. He concluded that on the basis of plumage,
measurements and wing formula they could both be referred
to
P.
reguloides.
Consequently,
P.
ricketti
does not occur on
Hainan Island and
P.
ricketti goodsoni
is a synonym to
P.
reguloides
(Alstrom, Colston
&
Olsson. unpubl.).
The specimens in Beijing and Kunming thus belong to an
undescribed species, for which we propose the name
Phylloscopus
hainanus,
sp.
nov.,
Hainan Leaf Warbler
HOLOTY
PE
Institute of Zoology, Academia Sinica. Beijing no. 52154,
male collected at Diao Lou, Lingshui County, south Hainan,
Guangdong Province, China (18"47'N, 110Q5'E). on 20 April
1962. Collector unknown.
DIAGNOSIS
Medium-sized leaf warbler, with distinctive yellow super-
cilium, pale median crown-stripe and dark eye-stripe. Plum-
~
t
Plate
1.
(above)
Phylloscopus hainanus.
a typical individual. Important characters are the deep yellow underparts and rather faint lateral
crown stripes. Jian Feng Ling, west Hainan,
24
April
1992.
Photo: Urban
Olsson;
(below)
Phylloscopus hainanus.
another typical individual.
The lateral crown stripes are only slightly darker than the mantle, and the median crown stripe is relatively indistinct pale yellow. The outer
tail-feathers are exposed in an unnatural way in this photograph in order to show their distinctive pattern. Jian Feng Ling, west Hainan,
24
ADril
1992.
Photo: Urban Olsson.
4
U.
OLSSON.
P.
ALSTROM
&
P.
R.
COLSTON
IBIS
13
5
age green above and bright yellow below. Pale tips to greater
and median coverts form two wing-bars on each wing. The
two outer pairs of rectrices are entirely white on the inner
webs (Plate
1).
There are distinct emarginations on the outer
webs of primaries 4-8 (numbered descendantly).
DESCRIPTION
OF
HOLOTYPE
Lateral crown-stripes vivid green, marginally darker than
the mantle, turning somewhat darker toward the rear. Me-
dian crown-stripe pale yellow, extending to the bill. Super-
cilium prominent, extending to base of bill, rather deep,
warm yellow throughout its length. Eye-stripe well defined,
approximately same colour as lateral crown-stripes. Rest of
ear-coverts, throat and remainder of underparts a rather
deep, warm yellow, like supercilium. Mantle and scapulars
vivid green, rump slightly paler and more yellowish. Greater
coverts sepia on inner webs and vivid green (like upperparts)
on outer webs, with pale yellowish tips to the
six
to seven
outer coverts, forming
a
distinct wing-bar. Pale yellowish
tips to median coverts also form a wing-bar, although dis-
tinctly narrower than the one on the greater coverts. Outer
edges of tertials same colour as upperparts, as are outer
edges
of
remiges: edges to outer primaries slightly paler.
Outermost two pairs of rectrices (R5,
R6)
are entirely white
on the inner webs.
R4
shows a distinct narrow white edge
on
the inner web. Viewed from below, the inner web
of
R4
appears clearly paler than the outer web. The undenving-
coverts are yellow with some whitish intermixed.
The upper mandible is dark and the lower all pale. The
tarsus, toes and claws are medium dark, the soles paler.
Measurements (mm)-Wing 56.0 (maximum length;
Svensson 1984). tail 37.0, bill-length (to skull) 12.6, bill-
width 2.5 (measured at distal end of nostrils).
Wing formula (primaries [PI numbered descendant1y)-
Wing-point P7
=
P6, P8
-1.
P9 -8.5, P5
-1.0.
P4 -3.0.
P3 -5.0, P2
-8.0,
P1 -9.0,
P10
-25.5; P10
11.0
>
primary
coverts: P9 falls between P1 and P2: P4-P8 are emarginated
on the outer web.
PARATYPES
There are four specimens of
P.
hainanus
in the Institute of
Zoology, Academia Sinica, Kunming, China:
(1)
male, probably first-year, collected by Professor Yang
Lan at Jian Feng Ling, Ledong County, west Hainan (18"40",
108"45'E). at an altitude of 820 m on
30
July 1979;
(2) adult female, collected by Professor Yang Lan at the
same place on 4 August 1979;
(3) adult male, collected by Professor Yang Lan at Diao
Lou. Lingshui County, south Hainan (18"47'N, 11W5'E). at
an altitude
of
640 m on 26 August 1979 and
(4)
adult male collected by Professor Yang Lan at the same
dace
on
27 Aumst 1979.
PARATYPIC VARIATION
There is little variation in plumage among the five known
specimens. Measurements (in mm) of the four paratypes are
(1)
wing 55.0, tail
c.
40 (difficult to measure), bill-length
12.4, bill-width 2.4; (2) wing 49.5. tail 35.5,
bill-length 12.2. bill-width 2.4: (3) wing 52.5, tail 40.5, bill-
length 12.2, bill-width 2.4;
(4)
wing 52.5, tail 41.0. bill-
length 12.6, bill-width 2.4.
ETYMOLOGY
We have chosen to name the species after the island where
it is endemic. By emphasizing the name of this island, we
hope to focus attention on the threats to the vulnerable
natural environment there and stimulate conservation mea-
sures in order to secure the continued survival
of
its unique
ecosystem. Hainan Island harbours one more endemic bird
species, the Hainan Hill Partridge
Arborophila ardens.
and
about 60 subspecies of birds that occur nowhere else in
China (Cheng 1987). many of them nowhere else in the
world. Some of them may deserve full specific status. Most
of them are under severe threat of habitat
loss.
VOCALIZATIONS
The song is high-pitched and consists of a number of rela-
tively short, varied phrases (Fig.
l),
very unlike the rather
stereotyped song
of
P.
ricketti,
but much resembling the song
of the White-tailed Leaf Warbler
P.
davisoni.
It could be
transcribed as
tsitsitsui-tsitsitsui
. .
.
titsu-titsui-titsui
.
.
.
titsu-
titsui-titsui
.
. .
titsu-titsui-titsui
. .
.
tist tsitsui-tsitsui-tsi
.
.
.
pitsu-pitsi-tsu
. .
.
pitsu-pitsi-tsu.
The calls could be transcribed as
pitsitsui. pitsiu
or
pitsi-
pitsu.
or
similar (Fig. 2). They are very close to those of
P.
davisoni.
DISTRIBUTION
As far as we are aware, the species has only been col-
lected
at
two localities in Hainan: Diao
Lou,
south Hai-
nan, and Jian Feng Ling, west Hainan (Fig. 3). One in-
dividual was seen
at
Bawanling on 22 April
1988
by Ben
King (pers. comm.). It seems to be absent
from
suitable
forest near Qiongzhong, south central Hainan. Much
ol
the hill
forest
in Hainan has been cleared, but the species
may occur in isolated pockets where suitable habitat still
exists. In Jian Feng Ling, it is rather common, and its
existence in this
area
does not seem to be under any
major threat
at
present because
at
least part
of
the area
is protected from logging.
At Jian Feng Ling, it occurs mainly in secondary
growth
at
the forest edges, e.g. along the road just outside the
reserve (Fig.
4).
where it appeared to be one of the mosi
NEW
SPECIES
OF
PHYLLOSCOPUS
FROM
CHINA
5
1993
7.
6-
5-
-
0
I
3
g
4-
I
E
3
3-
I
>
g
4-
t
8-
7.
6-
i
I
J,
5-
>
U
D
U
LL
2-
1'
E
3-
2-
1'
a
sl
'liliz
I
i
I
2
>
C
W
U
U
?
-
I"
Y
>
C
W
U
U
E
-
0
1
2
1
I
i
5
4
I/
:i,
1
0
7
1
2
2i
1
C
0
1
2
Seconds
Figure
1.
(a-c) Sonagrams illustrating different song strophes
of
Phylloscopus hainanus. Recordings by Per Alstrom. Hainan Island,
China, April 1992: sonagrams made by Richard Rank British Li-
brary of Wildlife sounds. Band width 369 Hz.
I
i
common bird species
on
a
visit by
P.A.
and
U.O.
in late
April
1992.
Very few individuals were observed inside
the mature tropical forest.
It is not known whether
or
not the species is migratory.
We are not aware of any records between September and
March, but this may be due to there being few visits by
ornithologists
at
this time.
1992; sonagrams made
by
Richard Ranft. British Library of Wildlife
sounds. Band width
369
Hz.
BREEDING
In late April
1992
at Jian Feng Ling, P.A. and
U.O.
observed
several pairs with fledged young and found one nest with
nearly full-grown young. The nest was situated on the face
of
a steep bank by
a
road, some
170
cm above the road. It
was dome-shaped, with a side entrance, and was built of
slender panicles,
in
combination with broad grass leaves and
a few thin roots and degraded remains of leaves. It was lined
with dense pappus fibres, thin bast from tree-ferns and some
pale brown and yellow feathers.
In late May
1989,
the birds did not seem to be territorial
but moved around in feeding Rocks with other species.
RELATIONSHIPS AND RECOGNITION
Phylloscopus
hainanus
is
identifiable as belonging to the genus
Phylloscopus.
as opposed to
Seicercus,
by the relatively nar-
row-based
bill,
the rather thin and inconspicuous, although
quite long, rictal bristles and typical
Phylloscopus
head pat-
tern.
We have examined a number of specimens of most forms
of Asiatic
Phylloscopus
species and studied several in the field
(see Appendix). On morphological grounds,
P.
hainanus
ap-
pears to be most closely related to
P.
davisoni.
Size, propor-
tions, general plumage pattern and wing formula are similar.
However, all subspecies of
P.
davisoni
are much paler yellow
below and show distinctly darker lateral crown-stripes (at
least posteriorly) and considerably less white on the pen-
ultimate tail-feather. The Sulphur-breasted Warbler
Phyl-
loscopus
ricketti
and the Mountain Leaf Warblers
P.
trivirgatus
trivirgatus
and
P.
t.
parvirostris
share the general plumage
patterns, including rather bright yellow colouration of the
underparts, but show much darker lateral crown-stripes and
lack white outer rectrices. The latter two further differ in
lacking distinct wing-bars and in showing much dark
on
the
lower mandible.
6
U.
OLSSON,
P.
ALSTROM
&
P.
R.
COLSTON
IBIS
135
Figure
3.
and
(2)
Diao Lou Mountain.
Hainan Island. The two localities where specimens of
Phylloscopus
hainanus
have been obtained are
(1)
Jian Feng Ling Forest Reserve
The Timor Leaf Warbler
P.
presbytesfloris,
the Mountain
Leaf Warblers
P.
trivirgatus rnalindangensis.
P.
t.
flavostriatus
and
P.
t.
mindanensis
and the Sulawesi Leaf Warbler
P.
sar-
asinorurn sarasinorurn
show roughly the same amount of white
in the tail and some yellow below. However, they all differ
from
P.
hainanus
in being less brightly coloured. especially
below, in showing less distinct
or
no wing-bars and in usually
showing much dark on the lower mandible.
Vocally,
P.
hainanus
is most reminiscent
of
P.
davisoni.
This
is a further indication of the close relationship with this
species.
MOULT
One of the males in IZASK, collected
26
August 1979 at Diao
Luo. had just finished moult
of
flight feathers (traces of feath-
er sheaths). Another, collected 27 August 1979 at the same
locality, was in the final stages of a complete moult. A female
from Jian Feng Ling, collected
4
August 1979, had just fin-
ished moult, with the exception
of
secondary
6,
which was
still growing.
A
probable first year male from Jian Feng Ling,
collected
30
July 1979, showed no sign of moult. This sug-
gests that adult birds go through a complete moult in July-
Figure
4.
Typical habitat of
Phylloscopus
hainanus.
Most singing
August.
males were in secondary forest
or
scrub at the edge of mature forest.
At
least one male was holding
a
territory
in
the
area
shown
in
this
picture. Jian Feng Ling. west Hainan,
24
April
1992.
Photo: Urban
Olsson.
We are indebted
to
Professors Yang Lan and Cheng Baolai at the
Institute of
Zoology,
Academia Sinica, Kunming, for their kind as-
sistance. Professor Yang Lan provided photograohs
of
soecimens.
1993
NEW
SPECIES
OF
PHYLLOSCOPUS
FROM
CHINA
7
In Beijing, Professors Cheng Tso-hsin. Tan Yao-kuang and Lu Tai-
chun provided invaluable assistance. as did Kiyoaki
Ozaki
and Yoshi-
mitsu Shigeta at the Yamashina Institute for Ornithology, Japan.
We are indebted to Stig Jensen, who supplied information about
distribution and lent
us
tape recordings. We are also grateful to
Richard A.
Sloss
and Mary LeCroy. at the American Museum (Nat-
ural History), New York, for sending
us
specimens and granting
access to the collections, respectively. Dr
G.
F. Mees, Leiden, Kees
Rooselaar, Amsterdam, and Dr Jon Fields& Copenhagen, were all
helpful in various aspects of our research. The sonagrams were
prepared by Richard Ranft at the British Library of Wildlife Sounds,
London. We are also grateful to Dr Alan Knox and Ben King for
comments
on
an earlier draft of this manuscript. The inclusion of a
colour plate was made possible, in part, by a grant from Kungliga
Patriotiska Sallskapet.
REFERENCES
Cheng. Tso-hsin.
1987.
A Synopsis of the Avifauna of China. Ham-
Hartert.
E.
1910.
The birds of Hainan. Novit. 2001.
17:
240.
Sibley, C.G.
&
Monroe, B.L..
Jr.
1990.
Distribution and Taxonomy
of Birds of the World. New Haven: Yale University Press.
Svensson,
L.
1984.
Identification Guide to European Passerines,
3rd ed. Stockholm: Lars Svensson.
Ticehurst,
C.B.
1938.
A
Systematic Review of the Genus
Phyllos-
copus.
London: Trustees of the British Museum.
Watson,
G.E..
Traylor. M.A., Jr.
&
Mayr.
E.
1986.
Family Sylviidae.
Old World warblers.
In
Mayr.
E.
&
Cottrell. G.W. (eds) Checklist
of Birds of the World,
Vol.
11:
3-294.
Cambridge, Mass.: Harvard
Museum of Comparative Zoology.
Williamson,
K.
1967.
Identification for Ringers
2:
the genus
Phyl-
loscopus.
BTO Field Guide
No.
8. Tring: British Trust for Orni-
thology.
burg: Science Press.
Submitted
16
January
1992;
revision received
9
June
1992
APPENDM: SPECIMENS EXAMINED
We have examined the/ollowing relevantjorms mentioned by Hartert (1
910).
Ticehurst
(1938).
Williamson
(1967).
Watson
et al.
(1986)
and Sibley
b
Monroe
(I
990)
in
the American Museum oJNatura1 History, New York, USA;
British Museum (Natural History). Tring.
UK;
Institute
of
Zoology. Academia
Sinica. Beijing. China; Institute oJZoology, Academia Sinica, Kunming, China;
Yamashina Institute
o/
Ornithology, Chiba. Japan; Rijksmuseum van Natuur-
lijke Historie, Leiden, Netherlands, and Zoological Museum, University
o/
Co-
penhagen. Denmark. We have not been able to examine every subspecies ofthe
islandforms. None
o/
the tara
below,
however, appeared to be closely related
to
Phylloscopus hainanus.
Number of Number
of
specimens field
ob-
Species studied servations
Phylloscopus hainanus
P. reguloides reguloides
P. r. kashmiriensis
P. r. assamensis
P.
r.
claudiae
P.
r.
Jokiensis
P. r. ticehursti
P. davisoni davisoni
P.
d. disturbans
P. d. ogilvie-granti
P. d. klossi
P. ricketti ricketti
P. r. goodsoni'
P. trivirgatus trivirgatus
P.
t. mindanensis
P.
t.
malindangensis
P.
t.
peterseni
P.
t.
pavostriatus
P.
t.
parvirostris
P.
t.
kinabaluensis
P.
1.
nigrorum
P. cebuensis cebuensis
P. c. luzonensis
P. olivaceus
P. poliocephalus poliocephalus
P. p. matthiae
P.
p.
henrietta
P.
p.
albigularis
P.
p.
cyclopum
P.
p.
hamlini
P.
p.
moorhousei
P.
p.
majorensis
P.
p.
bougainvillei
P.
p.
pallescens
P.
p.
becki
P.
p.
ceramensis
P.
p.
everetti
P.
p.
waterstradti
P.
p.
giulianettii
P. sarasinorum sarasinorum
P.
s.
nesophilus
P. presbytes presbytes
P.
p.
pons
P. makirensis
P. amoenus
P. ijimae
P. coronatus
5
501
lo+
60
+
20
40
8
70+
10
6
14
40+
3
60
+
1
6
2
1
23
14
35
23
12
38
9
2
7
6
3
4
14
7
24
1
20
10
15
33
54
28
45
14
12
16
5
lo+
120
1
This
form
is.
in
our
opinion. not
a
subspecies
of
P.
ricketti
but
belongs with
P. requloides
(Alstrom.
Colston
&
Olsson.
unouhl.).
... P. reguloides itself is known to be polytypic, embracing seven subspecies (Dickinson 2003) but the group's taxonomic situation is not at all stable and in recent years changes have abounded. One little-known taxon from Hainan Island, goodsoni Hartert, declared to be similar to the tropical P. ricketti by Hartert (1910) and Ticehurst (1938), was assigned to the P. reguloides group by Olsson et al. (1993), and accepted as such by Dickinson (2003). ...
... In goodsoni, the outermost (R6) bears only a narrow (1mm or less wide) white margin, similar to that of fokiensis, and the size and general coloration of these two taxa are also alike. In the meantime, Olsson et al. (1993) had described P. hainanus (which see), an apparently genuine Hainan Island endemic differing markedly in coloration, size and vocalizations from all accepted mainland representatives of P. reguloides sensu lato, goodsoni and fokiensis included. ...
... Distribution: A Chinese endemic described from two mid-winter specimens from Hainan Island (Hartert 1910). None was found there during several days of observation in spring (author's data, Olsson et al. 1993) and in the absence of further records the types of goodsoni are assumed to have been winterers outside the breeding range of the species. Olsson et al. (1993) recorded a March-dated specimen from the opposite mainland and (2005) indicated three additional localities in Guangxi, Guangdong and Hongkong, but without giving precise observation data or saying whether specimens were collected. ...
Article
Full-text available
Recent and proposed taxonomic changes to the warbler genera Phylloscopus and Seicercus are reviewed within the geographic scope of this series. Their reliability relative to secondary contacts, hybridization, introgression, bioacoustics and molecular genetics is discussed in terms of current species concepts, mainly the Biological Species Concept, and certain previously unpublished data are added in support of taxonomic decisions made. Within the area under consideration, this review records an increase in the number of Phylloscopus species from the 33 in Mayr et al. (1986) and 44 in del Hoyo et al. (2006) to 50. In Seicercus, S. burkii, a previously merged taxon swarm, has been split into six partly allopatric, partly sympatric species. This enlarging of species number emerges from due application of the results of bioacoustic and molecular genetic analyses to passerine systematics. Homogeneous external morphology largely hid the phylogenetic complexity and richness of these genera, but their species totals are not yet fixed; several populations presently accepted at species level need further substantiation.
... The genetic study by Olsson et al. (2005) was not backed by specimens; a shortfall that has also recently been criticized for other molecular systematic studies (Peterson et al., 2007). Type specimens had been investigated only in the case of P. r. goodsoni (Olsson et al., 1993). Regardless of these reservations, in the present study, we refer to taxon names as suggested by Olsson et al. (2005). ...
... By contrast, the high-pitched whistled songs of P. ricketti and P. cantator (sister clade to P. occipitalis in the cyt b ML and NJ trees) are quite unlike the songs of all members of the P. reguloides group (Päckert et al., 2004; Olsson et al., 2005), highlighting another non-molecular character uniquely shared by all taxa of the P. reguloides clade (including P. occipitalis ): a characteristic wing-flicking of males in case of excitement during the breeding season (cf. Alström & Olsson, 1993; Rasmussen & Anderton, 2005). ...
Article
The present study evaluated the degree of differentiation between closely-related species of Blyth's leaf warbler complex (Phylloscopus reguloides s.l.) and white-tailed leaf warbler complex (Phylloscopus davisoni s.l.) by molecular (cytochrome b and 16S rRNA) and bioacoustic markers (sonagraphic analysis of songs). Molecular phylogenetic results corroborate the recently suggested splittings of the two former taxa into three and two species, respectively. By contrast to previous opinion, differentiation of territorial songs parallels the genetic diversification in both groups. In the P. reguloides group, all recently established species can be distinguished by a characteristic song pattern. Regularly structured songs composed of repeated two- or three-note syllables were found in all species of the group, whereas trill songs with single notes repeated are unique to Phylloscopus claudiae. The phylogenetic relationships of Phylloscopus occipitalis are ambiguous among different molecular trees, but song pattern clearly corroborates a close relationship to the P. reguloides group. In the white-tailed leaf warbler group, songs are irregularly structured and often descending in frequency with exception of P. davisoni and Phylloscopus ogilviegranti disturbans. Song pattern corroborates the genetic affiliation of Phylloscopus hainanus and Phylloscopus xanthoschistos to the molecular white-tailed leaf warbler clade. All target species correspond to distinct acoustic clusters resulting from discriminant analysis of several spectral and structural song parameters. Linear correlation between genetic and acoustic differentiation (pairwise p-distances versus difference of discriminant functions) is significant within both groups, except for one case, in which discriminant function 1 was correlated with a spectral parameter (minimum frequency) only. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 96, 584–600.
... As the largest continental island in the northern region of the South China Sea, Hainan Island possesses a variety of habitats that facilitate species diversification. Previously, many new taxa have been documented on this island, including birds (Olsson et al., 1993), amphibians (Chou et al., 2007), and reptiles (Grismer et al., 2002). The continuous discovery of new species has broadened our insights into understanding the evolutionary processes on this island, although the interaction between Hainan Island and its nearby continental landmass has received limited attention in previous studies (but see Chen et al., 2015;Mao et al., 2010;Wang et al., 2016). ...
... Olsson et al. 1993; ...
Article
Among Palearctic warblers of the genus Phylloscopus those species that breed farther north occupy larger geographical ranges than those which breed farther south (Rapoport's rule). We suggest that much of this pattern is a consequence of the differential ability of species to occupy areas rendered inhospitable during the Pleistocene. In support of this suggestion, the midpoint of breeding range in a north-south direction has been an exceptionally labile trait through evolutionary time. Comparisons of ecological attributes of those species breeding in the Himalayas with close relatives in Siberia implies a role for habitat tracking in determining which species have been able to colonize northern areas; hypotheses based on climate and climatic variability have less support. In addition there is a likely role for geographic barriers and/or biotic interactions in preventing some taxa from spreading from small southern ranges.
... goodsoni fokiensis, and P. reguloides reguloides-P. occipitalis (cf. Baker, 1997;Olsson et al., 1993;Rasmussen and Anderton, 2005;Ticehurst, 1938;Williamson, 1967). Moreover, being endemic to Hainan island, hainanus is geographically isolated from the others. ...
Article
Weaverbirds are a diverse passerine group with species diversity concentrated in sub-Saharan Africa. No comprehensive phylogenetic hypothesis regarding relationships of weaverbirds has been produced, however, so we developed a first extensive phylogeny for the family Ploceidae, based on a multilocus dataset of three mitochondrial loci and four nuclear markers. Analysis of these data offered strong support for monophyly of the family and revealed seven distinct clades within Ploceidae. A major feature of our results is broad polyphyly of Ploceus: Asian Ploceus species should retain the generic name, whereas African Ploceus, together with Anaplectes, should be placed in Malimbus. In light of deep divergence, we assign the Malagasy Ploceus species to their own genus, Nelicurvius. Divergence time analysis based on DNA substitution rates suggests a mid-Miocene origin of the family. This study lays a foundation for an array of future studies of character evolution, biogeography, and evolutionary history of the family.
... The genera Phylloscopus and Seicercus have undergone dramatic taxonomic changes in the past 20 years, due to studies of especially vocalizations and DNA. No fewer than six species new to science have been described in that period: P. hainanus (Olsson et al., 1993), P. emeiensis (Alström and Olsson, 1995), S. omeiensis (Martens et al., 1999), S. soror (Alström and Olsson, 1999), P. occisinensis (Martens et al., 2008) and P. calciatilis . Moreover, taxonomic revisions have led to the recognition of several new species: P. canariensis and P. ibericus (previously treated as subspecies of P. collybita; P. ibericus formerly incorrectly called P. brehmii) (Helbig et al., 1996;Salomon et al., 1997;Helbig et al., 2001;Bensch et al., 2002;Salomon et al., 2003); P. orientalis (previously subspecies of P. bonelli; Helbig et al., 1995); P. borealoides (previously synonym of P. tenellipes ;Martens, 1988); P. chloronotus (previously subspecies of P. proregulus; Martens et al., 2004); P. forresti (previously synonym of P. proregulus chloronotus; Martens et al., 2004); P. kansuensis (previously synonym of P. p. proregulus;Alström et al., 1997;Martens et al., 2004); P. yunnanensis (previously synonym of P. proregulus chloronotus; nensis in latter study]; Martens et al., 2004); P. claudiae, P. goodsoni and P. ogilviegranti (two former previously subspecies of P. reguloides, third previously subspecies of P. davisoni; Olsson et al., 2005;Päckert et al., 2009); P. examinandus and P. xanthodryas (previously subspecies of P. borealis; Saitoh et al., 2006Saitoh et al., , 2008Saitoh et al., , 2010Reeves et al., 2008;Martens, 2010;Alström et al., 2011d); S. 6 Family Phylloscopidae based on concatenated nuclear MB and mitochondrial cytochrome b and 12S sequence data, based on Olsson et al. (2004Olsson et al. ( , 2005, Alström et al. (2006Alström et al. ( , 2010Alström et al. ( , 2011d, Saitoh et al. (2010) and Johansson et al. (2007), reanalyzed by Bayesian inference in three partitions (by locus). ...
Article
Full-text available
The systematics of the avian superfamily Sylvioidea are reviewed, focusing on studies of relationships among families and within genera, more superficially on taxonomic studies at the species level. For the families Bernieridae and Phylloscopidae, new analyses based on already published sequence data are presented. Our understanding of relationships has been vastly improved in recent years due to a large number of molecular studies. However, the relationships among the different families remain largely obscured, probably mainly as a result of rapid divergence of the different primary lineages (families). Also, species level taxonomy has been much improved in recent years due to a large number of studies applying molecular markers and/or vocalizations and other life-history data. It seems likely that the number of species will continue to increase, as new groups are being studied with modern integrative methods.
Article
Full-text available
The leaf warbler radiation (Aves: Phylloscopidae) has undergone a c. 50% increase in the number of recognised species over the last three decades, mainly as a result of analyses of vocalisations and DNA. Using a multilocus dataset for all of the species in this family, and multispecies coalescent-based as well as concatenation methods, we provide the first complete species-level phylogeny for this important group, as well as an estimate of the timing of diversification. The most recent common ancestor for the family was dated at 11.7 million years ago (mya) (95% highest posterior density 9.8-13.7 mya), and divergence times between sister species ranged from 0.5 mya (0.3-0.8 mya) to 6.1 mya (4.8-7.5 mya). Based on our results, we support synonymising Seicercus with Phylloscopus, which results in a monogeneric Phylloscopidae. We discuss the pros and cons of this treatment,and we argue againstproliferation of taxonomic names,and conclude that a large monogeneric Phylloscopidae leads to the fewest taxonomic changes compared to traditional classifications. We briefly discuss morphological evolution in the light of the phylogeny. The time calibrated phylogeny is a major improvement compared to previous studies based on a smaller number of species and loci and can provide a basis for future studies of other aspects of phylloscopid evolution.
Article
Newly described animal species are often assumed to be relatively small-bodied, probably because small animals are considered to be more easily overlooked We show that bird species first described in the period 1966-90 average smaller-bodied than birds in general, but suggest that other factors, with which body size is correlated, are more important determinants of the probability of description than is body size
Article
Cryptic species of passerine birds lack notable morphological differentiation and can best be identified by molecular and bioacoustic markers. Here we investigate seven cryptic species of the golden-spectacled warbler (Seicercus burkii complex) with respect to territorial song and cytochrome-b (cyt-b) sequences. Their phylogenetic relations to other Seicercus species and to members of the genus Phylloscopus are inferred by the same methods. Three separate lineages of Seicercus are nested within different branches of the molecular Phylloscopus tree. The S. burkii complex is a monophyletic unit comprising seven species (S. burkii s. str., S. whistleri, S. valentini, S. soror, S. omeiensis, S. tephrocephalus and S. affinis). S. xanthoschistos turned out to be a close relative of Phylloscopus davisoni within the P. reguloides group. Two isolated sister taxa, S. grammiceps and S. castaniceps, also branch together with the P. reguloides group. Within the S. burkii complex the overall haplotype and nucleotide diversity is highest in taxa from the Chinese middle and upper mountain belt (S. valentini, S. omeiensis and S. soror), indicating at least partially restricted gene flow in these species. This is explained by the fragmentation of high-altitude habitats in China while in the Himalayas the vicariant species S. whistleri inhabits a more continuous mountain belt at the same altitude. For the Chinese species from medium and high altitudes, past range expansion is indicated by significantly negative Tajima Ds. According to pairwise genetic distances, most species of the S. burkii complex have diverged 5 myr ago, the most recent split between S. burkii and S. tephrocephalus is dated 2 myr ago. Coalescence times for haplotype lineages of the different species range from 9 up to 12 myr, and between 5 and 6 myr for S. burkii and S. tephrocephalus.
  • Watson
Ticehurst (1938). Williamson (1967). Watson et al. (1986) and Sibley b
Tso-hsin. 1987. A Synopsis of the Avifauna of China
  • Cheng
Cheng. Tso-hsin. 1987. A Synopsis of the Avifauna of China.
( I 990) in the American Museum oJNatura1 History
  • Monroe
Monroe ( I 990) in the American Museum oJNatura1 History, New York, USA;
Natural History) Tring. UK; Institute of Zoology China; Institute oJZoology
  • British Museum
British Museum (Natural History). Tring. UK; Institute of Zoology. Academia Sinica. Beijing. China; Institute oJZoology, Academia Sinica, Kunming, China;
Family Sylviidae. Old World warblers
  • G E Watson
  • M A Traylor
  • Jr
  • E Mayr
Watson, G.E.. Traylor. M.A., Jr. & Mayr. E. 1986. Family Sylviidae. Old World warblers. In Mayr. E. & Cottrell. G.W. (eds) Checklist of Birds of the World, Vol. 11: 3-294. Cambridge, Mass.: Harvard Museum of Comparative Zoology.
Identification Guide to European Passerines
  • L Svensson
Svensson, L. 1984. Identification Guide to European Passerines, 3rd ed. Stockholm: Lars Svensson.
A Synopsis of the Avifauna of China. Ham-Hartert. E. 1910. The birds of Hainan
  • Cheng
  • Tso-Hsin
Cheng. Tso-hsin. 1987. A Synopsis of the Avifauna of China. Ham-Hartert. E. 1910. The birds of Hainan. Novit. 2001. 17: 240.