Systematic notes on Asian birds. 49
A preliminary review of the Aegithalidae, Remizidae and Paridae1
S. Eck † & J. Martens

Eck, S. & J. Martens. Systematic notes on Asian birds. 49. A preliminary review of the Aegithalidae, Remizidae and Paridae.
Zool. Med. Leiden 80-5 (1), 21.xii.2006: 1-63, figs. 1-6, Plates I-III.— ISSN 0024-0672.
Siegfried Eck †, formerly Staatliche Sammlungen für Naturkunde Dresden, Museum für Tierkunde,
Königsbrücker Landstrasse 159, D-01109 Dresden, Germany.
Jochen Martens, Institut für Zoologie, Johannes Gutenberg-Universität, D-55099 Mainz, Germany (e-mail:martens@uni-mainz.de).
Key words: Paridae; Aegithalidae; Remizidae; species limits; systematics; taxonomy; eastern Palaearctic
region; Indo-Malayan region; morphology; bioacoustics; molecular genetics; secondary contacts; hybridisation; introgression.
Proposed recent taxonomic changes in Paridae, Aegithalidae, and Remizidae are reviewed within thegeographic scope of this series and their reliability is discussed in terms of the Biological Species
Concept with respect to secondary contacts, hybridization, introgression, bioacoustics, and molecular
genetics. Certain previously unpublished data are added to support the taxonomic decisions.

Introduction
This review benefits from the excellent treatment of these three passerine families in
the monograph by Harrap (1996) 2, preceded by detailed work by Vaurie (1957a, b) on the
Palearctic species set upon which Snow (1967) based his treatment for Peters’s Checklistof the Birds of the World. Only after the publication of Harrap (op. cit.) has molecular genetics begun to furnish information on parid systematics and taxonomy. This has not
only given insights into population structure and population genetics of various taxa,
but also helped understanding of species evolution and species limits. Its influence onparid taxonomy is now quite pronounced. Papers like those of Gill et al. (1989, 2005),
Kvist et al. (1996, 2001, 2003, 2006), Salzburger et al. (2002a, b), Päckert et al. (2005), Martens et al. (2006), to mention just a few, are milestones in this respect.
Research focused on contact zones in the Russian Far East and in the Himalayas(Kvist et al., 2003, Nazarenko et al., 1999, Päckert et al., 2005, Martens & Eck, 1995, Martens et al., 2006) has also provided detailed information suggestive of times of evolutionary divergence and possible refuge areas in the Himalayas and in China. These
have turned out, in detail, to be considerably more complicated than was formerly believed. The main reason is that evolutionarily deeply split population units (haplotype
clusters in terms of molecular genetics) often are not mirrored by external characters.
Only molecular genetics makes them “visible”.

1 Results of the Himalayan Expeditions of J. Martens, No. 254. For No. 253 see: Senckenbergiana biologica,

86 (1): 11-36, 2006.
2 Sometimes referred to as Harrap & Quinn, but the title page clarifies the authorship.

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Eck & Martens. Review of the Aegithalidae etc. Zool. Med. Leiden 80 (2006)

Acoustic work, so far as it is relevant to taxonomy, has also made strides forward
and now allows consideration of nearly all the species of our area (sonagram collectionsof voices of Afghan, Himalayan, South Siberian and Chinese parids in Thielcke, 1968,
Martens, 1975, Martens & Nazarenko, 1993, Martens et al., 1995, Martens & Eck, 1995,
Thönen, 1996, Ivankina et al., 1997, Päckert et al., 2005). Over these years one of us (Eck,
1976, 1977, 1979, 1980a, b, 1982, 1988, 1992, 1996, 1998 and 2006) also contributed substantial fresh information on morphology that is relevant to the relationships and taxonomy of Asian species of the genus Parus.
These more modern approaches have not yet been applied to the study of the Aegithalidae and Remizidae. Although these groups are represented by only a few species
in our area, taxonomic problems abound and in many cases these are still open toquestion.
When studying the detailed text of Harrap (1996), and the excellent plates by Quinn,
it becomes evident that present taxonomic listings are relatively superficial and thesewill certainly change when more is known about contact zones, acoustics and molecular genetics.
Methodology
This contribution critically reviews systematic and taxonomic papers on Asian titmice and their allies. Consequently, it is purely descriptive in nature. Recommendations for taxonomic changes for nearly all cases proposed here have been publishedelsewhere, but they are critically discussed and partly revised. Personal experience withparid (sensu lato) taxonomy based on morphology, acoustics and molecular genetics has
greatly influenced our views, and extended field experience mainly in the Himalayas,
China and other parts of Palaearctic and Oriental Asia has informed our decisions. This
paper was also much influenced by a detailed paper prepared by one of us, which reviews the present taxonomic situation in Palaearctic tits; published posthumously elsewhere (Eck, 2006).
The species accounts herein are often subdivided. Some or all of the following small
sections are used:•
Taxa included: here we list the subspecies that we consider valid and which occur
regularly within the area covered by this series (see Dickinson & Dekker, 2000: 5).
We have added the type localities in square brackets, or occasionally a range statement. Sometimes we have found it appropriate to list a valid subspecies that is extralimital and in such cases the term ‘EL’ is added. Although our subspecific recognition is reflected in the ‘sister’ paper on the types of these families (Dickinson et al.,2006), and we use the spellings, dates and authorships used there, the additional
geographic information given here should assist in understanding contact zones.
This arrangement also allows us to minimise the number of citations we have toinsert within each taxonomic account. Where Dickinson et al. (2006) list all relevant
synonyms, we list only those we discuss.
•
Taxonomy: generally, this section deals with the validity of subspecies and with species limits, but we also highlight the taxonomic questions which seem to us to require further research. Museum names are replaced by acronyms; these are listed at
the end of this paper. In a few cases we provide a general introduction and go on todeal with one subspecies group at a time.

Eck & Martens. Review of the Aegithalidae etc. Zool. Med. Leiden 80 (2006)
•
•
•

3

General characteristics: this section contains notes on morphological generalities andpeculiarities.
Distribution: here we comment upon matters such as range gaps, and contact zones,
and areas needing further research.
Colour images: understanding this paper will, we think, be assisted by consulting
published images, especially those we list here. Usually these are the work of
Quinn in Harrap (1996) who depicted all the species and many of the subspecies in
our area.
Aegithalidae
Aegithalos caudatus (Linnaeus, 1758)
Taxa included (7, extralimital subspecies listed for convenience: 1): Ae. c. kamtschaticus Domaniewski,
1933 [Kamchatka] EL; Ae. c. sibiricus (Seebohm, 1890) [C Siberia]; Ae. c. japonicus Prázak, 1897 [NJapan]; Ae. c. vinaceus (Verreaux, 1870) 3 [Ourato, ‘Kansu’ = Inner Mongolia]; Ae. c. glaucogularis(Moore, 1855) 4 [China = Shanghai 5]; Ae. c. magnus (Clark, 1907) [Kaiki District, Seoul, Korea]; Ae. c.
trivirgatus (Temminck & Schlegel, 1848) [Japan; restricted to Honshu by Kuroda (1923c). Range:
Honshu, Sado, Oki (Japan); Quelpart (Korea)]; Ae. c. kiusiuensis Kuroda, 1923c [N Kyushu].

Taxonomy: the Long-tailed Tit varies a great deal geographically, but the degree ofvariation is not constant. In some regions this variation is sharp and well differentiatedsubspecies have been described, but other names relate to slight differences and may becomplicated by an exceptionally high degree of individual variation (Vaurie, 1959).
Based on external morphology there are several subspecies groups; three according
to Vaurie (1959), or four in Harrap (1996). These appear to provide evidence of historic
separation followed by recent, probably post-glacial, secondary contact with interbreeding. All three of the subspecies groups of Vaurie (1959) occur in our area (caudatus, europaeus Hermann, 1804, and alpinus Hablizl, 1783); but Harrap (1996) separated a fourthgroup (glaucogularis, which Vaurie had included in his alpinus group). And according tohim, this group in which he included just glaucogularis and vinaceus occurs in the area
covered by this series, whereas his restricted alpinus does not.
Snow (1967) treated the caudatus ‘group’ as one wide-ranging form stretching acrossthe northern Palaearctic taiga zone and extending eastwards to northern Japan. Thisapproach was accepted by Harrap (1996) and Stepanyan (1990), but Stepanyan (2003)
recognised kamtschaticus, a view supported by Red’kin (in litt.), and yet, like Snow,
Stepanyan kept sibiricus Seebohm, 1890, in the synonymy of caudatus 6. Vaurie (1959)
listed japonicus, although seeing it as weakly differentiated, and Morioka, in Orn. Soc.
Japan (2000: 314), chose to follow Vaurie rather than Snow. Given our lack of fresh in-

3 Not 1871; see Dickinson et al. (2006).
4 Moore’s name has priority over Gould’s, which was used by Snow (1967: 55), see Dickinson et al.

(2006).
5 Gould (1855) redescribed this and restricted the type locality.
6 Perhaps because the name was proposed in an unorthodox and almost conditional way.

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Eck & Martens. Review of the Aegithalidae etc. Zool. Med. Leiden 80 (2006)

formation we follow Stepanyan in recognising kamtschaticus and follow Morioka (op.
cit. supra) in recognizing japonicus. This allows us to avoid the merger of populations
that may well be found to differ markedly in their molecular genetics. We have not beenable to examine specimens from NE China to see whether these best match sibiricus orjaponicus.
Apart from japonicus, which is part of the caudatus group 7, Korea and the Japaneseislands are occupied by magnus, trivirgatus and kiusiuensis which are all considered by
Vaurie, and by Harrap, to belong to the europaeus group, but from which they are widely separated. Intervening populations between the two halves make up the caudatus
group. The wisdom of seeing these eastern forms as belonging to the europaeus groupneeds revisiting when molecular information becomes available as the characters thathave suggested this relationship may instead result from morphological convergence
between quite different haplotype groupings.
Because we think variation in Siberia requires further study and we find the split in
the europaeus group questionable we consider that a detailed zoogeographic analysis
drawing upon molecular information is urgently needed.
Colour images: Quinn in Harrap (1996), pl. 33, figs. 103k-l: glaucogularis (as well as
figs. of various extralimital W Palaearctic subspecies).
Aegithalos leucogenys (Moore, 1854 8)
Taxon included (1): Ae. leucogenys (Moore, 1854) [Woods above Balu Chughur’ (= Chighu Serai), near
Gusalek, Kunar valley, NE of Jalalabad]. – Monotypic.

Distribution: the White-cheeked Tit is a restricted-range endemic of dry central Asia,
from Afghanistan and western Pakistan south to Baluchistan, and along the western
Himalayas to about 74°E. Records east of 74°E, e.g., in Baltistan and Garhwal seem
doubtful (Roberts, 1992: 351). No taxonomic problems are currently known.
Colour images: Quinn in Harrap (1996), pl. 35, figs. 104a-c.
Aegithalos concinnus (Gould, 1855)
Taxa included (6): Ae. c. iredalei (Baker, 1920a) [Himalayas = Simla 9]; Ae. c. manipurensis (Hume, 1888)[eastern hills, Manipur]; Ae. c. talifuensis (Rippon, 1903) [E Tali, NW Yunnan]; Ae. c. concinnus(Gould, 1855) [? Chu Shan, Zhejiang]; Ae. c. pulchellus (Rippon, 1900) [southern Shan States, Myanmar]; Ae. c. annamensis (Robinson & Kloss, 1919) [Langbian Mts., Vietnam].

Taxonomy: the Black-throated Tit, as currently recognised, brings together highly
distinct subspecies which were arranged in three subspecies groups (monotypic iredalei
and annamensis, and polytypic concinnus) by Harrap (1996). The subspecies are remark-7 But, as indicated, has often been treated as a synonym of nominate caudatus.
8 In full: Moore in Horsfield & Moore; Moore alone should be credited, contra Snow (1967).
9 Baker (1920a) was renaming Parus erythrocephalus Vigors, 1831, a name which was preoccupied. That

had as type locality “Himalayas”; Baker restricted this to Simla (but there has been considerable dispute
about the origin of the specimens that were described by Vigors and painted for Gould’s ‘Century’).

Eck & Martens. Review of the Aegithalidae etc. Zool. Med. Leiden 80 (2006)

5able in that they differ not only in coloration of the crown (chestnut brown vs. greyishdrab), but they also differ in pattern (i.e the presence or absence of supercilium andchestnut breast band).
Not surprisingly we have begun to find that these differences are paralleled by significant differences in the cytochrome-b gene. Three specimens have been sequenced.
One from Rasuwa Distr. (Nepal, ssp. iredalei) differed from one from Omei Shan (Sichuan/China, spp. talifuensis) by 5.1% sequence divergence and from one from Natmataung (Mt. Victoria, Myanmar; ssp. manipurensis) by 5.3%. The Omei and Natmatung
specimens differ by 6% (uncorrected, 663 bp). In the tree topology, the Nepal and theOmei specimens come out as ‘sisters’ and when these are compared with the Natmataung specimen the between-group distance is 5.6%. These are unexpected and remarkably strong differences at a level which, in songbirds, usually indicates a clear species
difference (J.M., M. Päckert and D.T. Tietze, unpublished). All these three samples arefrom reddish-crowned subspecies, but talifuensis from West China also has a strong reddish breast band. There are two grey-headed subspecies (pulchellus, which has the reddish breast band, and annamensis, which does not), and by this character these differ
sharply from the reddish-crowned ones.
Among these birds, crown colour seems particularly important as it may represent a
plesiomorphic character state. Just-fledged juveniles of red-headed ssp. talifuensis/concinnus have the crown colour just like the grey-headed forms; but this is quite ephemeral as
it is soon moulted, becoming chestnut apparently within a few weeks after fledging.
Thus most field guides show juveniles with the forehead and crown reddish.
It seems possible that Ae. concinnus represents a hitherto unresolved species swarm.
More data on contact zones of brownish- and greyish-crowned populations are needed,
as well as molecular and vocal information.
Distribution: the Himalayas east to large parts of C and S China, Taiwan, Myanmar
and parts of Thailand. Of the grey-headed subspecies, annamensis, in Vietnam and Laosis highly disjunct; pulchellus in northwest Myanmar seems to be in contact with thenorth-eastern Indian rufous-headed manipurensis (map in Harrap, 1996), but whether
they intergrade is unknown.
Colour images: herein Pl. I, Figs. a-b: talifuensis/concinnus; Quinn in Harrap (1996), pl.
34, figs. 105a-b iredalei (reddish-headed), 105c pulchellus (grey-headed), 105d talifuensis(reddish-headed), 105e-f annamensis (grey-headed).
Aegithalos niveogularis (Gould, 1855)
Taxa included (4): A: the niveogularis ‘group’: Ae. n. niveogularis (Moore, 1855) 10 [Northern India = N
Punjab 11]; B: the bonvaloti ‘group’ 12: Ae. n. bonvaloti (Oustalet, 1892 13) [Tatsienlu = Kangding Xian,
Sichuan, China]; Ae. n. obscuratus (Mayr, 1940) [Chengou Forks, 30 miles W of Wenchwan, Sungpan

10 Moore has priority over Gould, the author given by Snow (1967: 58), see Dickinson et al. (2006).
11 Vaurie (1957b: 20) restricted the type locality of niveogularis Gould to N. Punjab. However, Moore

published first and Gould’s name is a primary homonym and permanently invalid. See Dickinson et al.
(2006) for the parallel restriction of the type locality of niveogularis Moore.
12 This grouping is for convenient discussion and based solely on geography.
13 For use of this date in place of 1891 see Dickinson et al. (2006).

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Eck & Martens. Review of the Aegithalidae etc. Zool. Med. Leiden 80 (2006)
district, northern Sichuan; given as Maowen by Cheng, 1987]; Ae. n. sharpei (Rippon, 1904) [Mt Victoria, S Chin Hills, Myanmar].

Taxonomy: although only a few taxa are concerned, species limits within the SinoHimalayan long-tailed tits are not reliably worked out yet. The taxa niveogularis (W
Himalayas), iouschistos (Blyth, 1845) (E Himalayas east to about the Tsangpo bend), andobscuratus (adjacent SW China) occupy a more or less coherent, but not quite continuous range, whereas sharpei (Chin Hills, Myanmar) is apparently entirely disjunct. Vaurie (1959) and Snow (1967) treated all these forms as conspecific, which reflects theiroverall similarity and largely continuous distribution along the Himalayan chain extending into south-west China.
However, western niveogularis is remarkably different from the adjacent iouschistos
in coloration and slightly different in pattern. Furthermore, niveogularis is a form adapted to drier habitat, and its range ends in north-west Dhaulagiri in north-west Nepal.
This is west of the start of the range of iouschistos. It is absent in the inner valley of the
Kali Gandaki gorges (east of Dhaulagiri) and anywhere further east (Inskipp & Inskipp,1991; Martens & Eck, 1995). By contrast, iouschistos inhabits mixed broad-leafed/coniferous forests within the influence of the monsoon. Syntopic occurrences of niveogularis and iouschistos are not yet known, nor are hybrids (Harrap, 1996). Thus it seems
justified to treat niveogularis as a separate biological species; however overall relationships in this complex are unresolved. For instance, bonvaloti and sharpei are relatively
long-tailed (as is leucogenys, which differs strongly in bill shape) while iouschistos andniveogularis are relatively short-tailed] (C.S. Roselaar pers. comm.).
Martens & Eck (1995: 314-315) united western niveogularis with bonvaloti, obscuratus
and sharpei and treated them as conspecific. This arrangement is supported by external
morphology, for niveogularis, bonvaloti/obscuratus and sharpei are very similar in patternand coloration (cf. Vaurie 1957a). But this does not seem to fit with general zoogeography, and the limited ecological evidence is inconclusive. The habitat requirements of niveogularis seem to be dry forest and obscuratus lives well in dry forest too, e.g., in Yunnan(Jizu Shan; J.M.), however bonvaloti lives in wetter forest, while no clear habitat preference is recorded for sharpei of Mt. Victoria although one imagines this will not be dry
forest. Wunderlich (1991) attached obscuratus to iouschistos but this is surely erroneous.
Harrap (1996) treated niveogularis as a monotypic species and united the easternand south-eastern taxa with iouschistos. Dickinson (2003) felt unable to unite iouschistosand ‘bonvaloti’ (made up of the three eastern and south-eastern representatives: bonvaloti, obscuratus, sharpei) and listed three species. We prefer to maintain the arrangement
offered by Martens & Eck (1995), however we look forward to a molecular analysis,
which will no doubt clarify the picture.
Colour images: Quinn in Harrap (1996), pl. 34, figs. 107a-b.
Aegithalos iouschistos (Blyth, 1845 14)
Taxon included (1): Ae. iouschistos (Blyth, 1845) [Nepal].

14 The 1844 volume, and that date was used by Snow (1967: 58); however this paper appeared in 1845;

see Dickinson & Pittie (2006).

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7

Taxonomy: see above under Ae. niveogularis. – Monotypic.
Colour images: Quinn in Harrap (1996), pl. 34, figs. 107a-b.
Aegithalos fuliginosus (J. Verreaux, 1869 15)
Taxon included (1): Ae. fuliginosus (J. Verreaux, 1869) [E Tibet = W Sichuan]. – Monotypic.

Distribution: the Sooty Tit is a small-range Chinese species found in parts of Sichuan, Gansu, Shaanxi, and Hubei. It was quite common on the southern flanks of
Taibai Shan, Shaanxi in June 1997 (J.M.). – No taxonomic problems known.
Colour images: Quinn in Harrap (1996), pl. 35, figs. 108a-c.
Psaltria exilis Temminck, 1836
Taxon included (1): Psaltria exilis Temminck, 1836 [Java]. – Monotypic.

Colour images: Quinn in Harrap (1996), pl. 36, figs. 110a-b.
Remizidae
The Penduline Tit was most often treated as one wide-ranging species, R. pendulinus(as in Hartert 1905a, Vaurie 1959, Snow, 1967) until recently when Harrap (1996) accepted four species, R. pendulinus sensu stricto, R. macronyx (Severtsov, 1873), R. coronatus(Severtsov, 1873), and R. consobrinus (Swinhoe, 1870). Dickinson (2003: 530) retained thetreatment of Snow (1967) but added some comments, in a footnote in which he mistakenly suggested that the last three subspecies listed would constitute the species coronatus
if three species were recognised. In fact the name consobrinus Swinhoe, 1870, has priority
over coronatus Severtsov, 1873. In addition to differences in coloration and pattern, thegroups differ in size as well (Vaurie, 1950) and in relative bill length and structure of leg
and foot (C.S. Roselaar pers. comm.) and that two quite different nest types occur.
That three species need to be recognised flows from the evidence of Kazakhstan(Gavrilov, 1972; see maps pp. 254, 259) where it is apparent that R. r. macronyx and R. r.
coronatus are widely sympatric and that they differ not just structurally and morphologically but in habitat preference. If the appropriate recognised taxa are grouped withpendulinus, macronyx or consobrinus it is then apparent that macronyx meets with pendulinus and interbreeds in a narrow contact zone at the mouth of the river Ural and in “thesouthwest corner of the Caspian [basin]” (Harrap, 1996: 212). Whether and how any
subspecies of consobrinus actually interacts is less clear. It was suggested by Hartert(1905) that R. r. jaxarticus may interbreed with stoliczkae (Hume, 1874), but this seems tohave been based on museum specimens without field studies to determine the geography of any contact zone and how the birds interact. Thus stoliczkae is a desert form
which may or may not meet and intergrade with consobrinus. It is said to intergrade

15 Snow (1967: 58) dated this 1870, implying a delay in publication. The volume title page is dated 1869

and no evidence of delay is apparent.

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with coronatus in the Zaysan basin (Harrap. 1996: 215), and Harrap wondered whether
stoliczkae as well as consobrinus might be involved in the increase of records in eastern
China and Japan. We are unaware of any report that stoliczkae and consobrinus meet andinteract on their breeding grounds.
In sum, the evidence would allow one to retain the macronyx group as a part of thespecies pendulinus and does not clearly resolve whether coronatus and consobrinus should
be considered separate species. We prefer to treat these two as one species until moreevidence is available. To add to the morphological and biological information alreadyavailable, acoustic and molecular data are needed for sound decisions.
Remiz consobrinus (Swinhoe, 1870b)
Taxa included (3): A: the coronatus group: R. c. coronatus (Severtsov, 1873) [’Nau, on Syrdar’ya below
Khodzent’ = Leninabad, Tajikhistan]; R. c. stoliczkae (Hume, 1874) [no locality = Bora, Chinese
Turkestan 16 = Bora, SE of Yecheng, Xinjiang, China]. B: the consobrinus ‘group’: R. c. consobrinus(Swinhoe, 1870b) [Sha-She, Hubei].

Distribution: the breeding range of eastern consobrinus is disjunct from the range of
the western coronatus group.
Colour images: Quinn in Harrap (1996), pl. 11, figs. 35a-d: coronatus, 36a-d: consobrinus.
Cephalopyrus flammiceps (Burton, 1836)
Taxa included (2): C. f. flammiceps (Burton, 1836) [Himalayas = Mussoorie 17]; C. f. olivaceus (Rothschild, 1923) [Tengyueh, N Yunnan].

Taxonomy: no problems need discussion.
Colour images: Quinn in Harrap (1996), pl. 12, figs. 43a-e: flammiceps, f: olivaceus.
Paridae
For many years ornithologists have followed traditional species limits in Parus but
in recent years several of these limits have been questioned or changed, and further
modifications are likely. Acoustics and present molecular genetics are influencing thisprogress, both in splitting and in lumping taxa. It now appears that marked morphological differences, mostly in coloration but some in overall pattern, do not reliably
predict species limits in Parus. Taxa most affected by these investigations occur in fivecomplexes: first, Parus major Linnaeus, 1758, sensu lato; second, P. cyanus Pallas, 1770,
incl. flavipectus Severtsov, 1873; third, P. ater Linnaeus, 1758, incl. melanolophus Vigors,
1831; fourth, P. montanus Conrad, 1827; and fifth, P. palustris Linnaeus, 1758, incl. hypermelaenus (Berezowski & Bianchi, 1891). Each is discussed in some detail in the species
accounts that follow.

16 Designated by Hartert (1905a: 391).
17 The type locality was restricted by Whistler (1924).

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9This work on parids has offered additional insights into how well distance values of
the cytochrome-b gene of the mitochondrial genome may indicate species limits even
without additional biological data derived from morphology or full information on
behaviour at contact zones and on hybridization events. Cytochrome-b differences ofapparently “good” and long-established species, such as P. caeruleus Linnaeus, 1758,
and P. cyanus, unexpectedly display very low cytochrome-b distances and by this seemto suggest only subspecies level distances. Rather higher distance values of above 3%
are present between populations of P. ater that have traditionally been simply characterized as subspecies groups which apparently intergrade along suture zones, e.g., in China and in the western Palaearctic. These results should initiate fresh, general discussions of how reliably to apply mitochondrial distance values in avian taxonomy. Where
relevant below, such discussions are begun herein.
The subspecies in Parus accepted here differ from those recognized in Peters’s
Check-list (Snow, 1967) in two particulars. First, we include the post 1967 names ofwhich we are aware: Parus semilarvatus snowi Parkes, 1971, Parus ater martensi Eck, 1998
and Parus ater eckodedicatus Martens, Tietze & Sun, 2006. Second, species limit concepts
have changed since Snow (1967) and even since Harrap (1996), although this remains
the most useful and comprehensive treatment of the family. There are significant differences between treatments in terms of the recognition or synonymization of some names
and some views seem to have been based on limited access to specimens, or to be subjective. Such problems can be seen both in the grey tits (Poecile) and also in the great tit(P. major sensu lato) where conflicting views no doubt remain despite our attempts toelucidate Asian problems.
Taxonomy above the species level
In the 19th century attempts were already made to split the large genus Parus into
genera or subgenera. The subgenus last erected was Poeciloides Bianchi, 1902, for Parus
superciliosus. For Parus sensu lato 12 genus-level names are now in use. The species assigned to each were documented by Hellmayr (1903). Portenko (1954) and Wolters(1980) used these to label their genera. These subunits, until recently mostly used in thesense of subgenera 18, are still in use without much variation as to the species incorporated, although some authors lost track of the type species of their genera (or subgenera).
In general, the circumscription of these units based on morphological characters
was pretty good, although not based on a clear understanding of monophyly. Naturally
the advent of molecular genetic phylogeny has inspired a re-examination of generic
splits in songbirds and as Parus is a large genus it has received considerable attention.
Gill et al. (2005: 139) stated: “The 51 species of titmice and chickadees constitute a welldefined, monophyletic assemblage of sylvioid passerine birds on the basis of both morphological and molecular characters ...”. They recommended (op. cit. p. 140) splitting
Parus into six genera on grounds of cytochrome-b topology. “Each of the six recommended genera is a monophyletic group that displays distinctive behavioural and

18 Although for North America generic distinctions were accepted by the A.O.U. (1998).

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Eck & Martens. Review of the Aegithalidae etc. Zool. Med. Leiden 80 (2006)

morphological characteristics.” However, this view is not unequivocal; different ones
are possible especially when the entire genus Parus sensu lato is considered and all Asianand African species are accounted for.
The scope proposed by Gill et al. for the genus Periparus Selys-Longchamps, 1884,
embraced the tropical and subtropical species (amabilis, elegans, venustulus) as well astrue Palaearctic and often high-montane species (ater, rubidiventris, rufonuchalis). The
species of these two groups by no means “...display [shared] distinctive behaviouraland morphological characteristics…”, furthermore their ecological requirements differ
considerably, and it seems to us that it will be better to employ separate generic names
for them.
In contrast, Gill et al. admit separate generic names of two Old and New World
sister taxa, Lophophanes Kaup, 1829, and Baeolophus Cabanis, 1850. Are their species that
distinct in morphology and behaviour as to sufficiently support this opinion? Their
molecular split is not deeper than that between the two units of the broad genus Periparus. Thus decisions regarding a broad genus Periparus on one hand and narrow Lophophanes and Baeolophus on the other seem to be inconsistent.
There is one other apparent inconsistency in Gill’s genus Parus sensu stricto, for thisembraces two clades: the relatives of the great tit in Eurasia on the one hand and thespecies of sub-Saharan Africa on the other. The two are separated by a deep molecularsplit and under the conditions set by Gill et al. each of these may warrant separate generic treatment as well. Finally, the phylogeny of Gill et al. lacks one important species,
P. semilarvatus (Salvadori, 1865) from the Philippines, the generic assignment of whichis unresolved, although it may be close to Poecile Kaup, 1829.
Because of these open questions we refrain, for the time being, from adopting thegeneric recommendations of Gill et al. (2005) and use the generic name Parus sensu lato
throughout. We have also avoided subgeneric names because of the Asian taxa thatremain unsampled 19 or insufficiently sampled at levels previously thought to be subspecific 20. In the circumstances we have chosen to retain the species sequence used by
Snow (1967).
The species accounts – taxonomy at the species level and below
Parus palustris Linnaeus, 1758
Taxa included (5; extralimital forms listed for convenience: 2): P. p. altaicus Johansen, 1952 [Buchtarma river, Katon-Karagai, S Altai, E Kazakhstan] EL; P. p. brevirostris (Taczanowski, 1872) [SW of
Lake Baikal, Irkutsk Region, Russia]; P. p. crassirostris (Taczanowski, 1885) [Sidemi, S. Ussuriland,
Russia]; P. p. ernsti Yamashina, 1933 [Naihoro, S. Sakhalin, Russia] EL; P. p. hensoni Stejneger, 1892
[Hakodate, Hokkaido, Japan] 21; P. p. jeholicus Kleinschmidt & Weigold, 1922a [30 km north of Bali-

19 Table 2 in Gill et al. (2005) lacked the Asian taxa: P. venustulus, P. nuchalis and P. semilarvatus.
20 Such as P. hypermelaena and P. weigoldicus.
21 Although we treat seebohmi Stejneger, 1892, as a synonym, as did Hartert (1905: 375) and later the Orn.

Soc. Japan (1942) and Snow (1967: 74), we have not located the rationale by which Stejneger’s distinctions between hensoni and seebohmi have been explained. The types may need re-examination despite
acceptance of that synonymy by Deignan (1961: 342).

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handién, NE Jehol 22 = Chengde Shi, Hebei, China]; P. p. hellmayri (Bianchi, 1902) [Peking = Beijing,
China].

Taxonomy: historically the taxonomic treatment of the Marsh Tit has been characterised by strongly opposing views: extreme splitting and extreme lumping (for example
see Kleinschmidt, 1897, for an account). Most systematists now follow Snow (1957,
1967) and Vaurie (1959). Of the numerous names given to eastern populations Vaurie(1957a) synonymised seven in brevirostris although, as he stated, he had only a single
brevirostris at his disposal for comparison. Snow (1957: 37) was initially willing to recognize two of them (altaicus and crassirostris) thus dividing the Siberian population into
three, but later (1967) decided to follow Vaurie (op. cit.).
One of us (Eck, 1980a) in an in-depth revision of the ‘Poecile’ group, which was
based on a very substantial material, was able to offer a more detailed view. Europeanand Asian Marsh Tits are not only widely disjunct, with a range gap of more than 1,000
km, they also differ significantly in morphology. These two widely disjunct subspecies
groups are currently labelled as the palustris group (west Palaearctic and, for us, extralimital) and the brevirostris group (east Palaearctic or Asian). The Asian populations havea larger bib, smaller feet, and a clumsier bill. The differences between representatives ofthese groups seem to be insignificant in terms of mitochondrial genetic distance values(Gill et al., 2005: 127; J.M. unpublished), but more taxa need to be sequenced. The Asian
populations between the Altai Mts. and Ussuriland are known by subtle colour differences and these, although placed in synonymy, should not be ignored. The populationfrom Sakhalin (ernsti) closely resembles that of Japan (hensoni; Eck, 1980a: 144).
As concerns our area we accept brevirostris, crassirostris, hensoni, jeholicus and hellmayri, and in contrast to Cheng (1987) we recognise the distinctness of both hellmayri
and jeholicus (see below). The first two of these, crassirostris and hensoni, form one morphological group, together with ernsti and extralimital brevirostris, occupying the central to eastern Palaearctic. A second morphological group encompasses jeholicus andhellmayri from more southerly north-east China. In north-east China the range of jeholicus is a continuation of that of P. p. crassirostris and presumably these intergrade, but
this needs to be verified.
The Chinese taxon hypermelaenus is probably best excluded from P. palustris (seebelow).
General characteristics: compared with west Palaearctic populations the eastern ones
have heavier bills and smaller feet. Within the group they vary in the coloration of theback and flanks, height of bill, and proportions of flight feathers. TWI (tail/wing index– see Abbreviations and Acronyms): ‘group 1’, from altaicus to crassirostris 96.6%, sd 2.26
(n = 28); group 2. ernsti 89.7%, sd 1.5 (n = 10); hensoni 89.5%, sd 2.1 (n = 24). In hellmayri
and jeholicus, the dorsal coloration is equally brown, but the two subspecies differ inwing length. Relative tail length (TWI): hellmayri 86.2%, sd 1.15 (n = 13); jeholicus 92.5%,
sd 1.52 (n = 11).
Distribution: when examining the detailed mosaic of varying populations within
what was treated as brevirostris sensu lato by Snow (1967), the distribution is as follows:

22 Jehol Province was abolished in 1955 (Zhao & Adler, 1993: 434; for Chengteh see p. 428).

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brevirostris sensu stricto occurs in southern Siberia eastwards to the Shilka and Argun rivers including Mongolia and western parts of NE China, crassirostris inhabits Amurland
and northern Korea, and probably easternmost north-eastern China, and ernsti is restricted to Sakhalin; altaicus is reported from the Altais and is sometimes considered asynonym of brevirostris, but is quite distinct in colour and is maintained by Eck (2006).
Colour images: Quinn in Harrap (1996): pl. 15, figs. 46d-e: hellmayri; Eck, 1980a: pl. 1:
brevirostris and hellmayri.
Parus hypermelaenus (Berezowski & Bianchi, 1891)
Taxa included (1): P. h. hypermelaenus (Berezowski & Bianchi, 1891) [Landzya-kou, Shaanxi (localityof lectotype according to Stepanyan & Loskot 1998, at border between S Shaanxi and Gansu, China)];
Synonyms (provisional placement only): dejeani Oustalet 1897 [Tsékou, N Yunnan, China – Kangding
according to Cheng, 1987 23]; poecilopsis (Sharpe, 1902) [Ch’ü-tung, W Yunnan, China]; P. h. ssp. (unnamed) [Mt. Victoria, Chin Hills, W Myanmar].

Taxonomy: the taxonomic rank of the Black-bibbed Tit, a local Chinese form, has often been discussed but, following Harrap (1996), Dickinson (2003) allowed it species
rank, although essentially on hypothetical grounds. The case has still to be firmly established, but is not here refuted.
There are marked morphological differences from Marsh Tits, but a close relationship of hypermelaenus with the Marsh Tit is evident and has never been questioned.
Though records of P. palustris exist close to hypermelaenus localities (maps in Cheng
1987, Harrap, 1996), no zone of contact is known and nor are intergrades or hybrids.
Consequently, intermixing with P. palustris is open to question. The value of the cytochrome-b difference between the combined west and the east Palaearctic populationgroups of P. palustris on the one hand and hypermelaenus on the other amounts to 3.3%
(J.M., unpublished). In view of the rather slight difference between the western andeastern population groups of P. palustris, of 0.65% (Gill et al., 2005), 3.3% is a remarkably
high value and indicates prolonged independent evolution of about 1.6 m.y. when the
2% rule is applied. Without further information, however, species rank for hypermelaenus
remains equivocal. In addition, we have to state that this value is of the same order ofmagnitude as those between various cytochrome-b clusters of P. ater (q.v.), in the east as
well as in the west Palaearctic. The voice of hypermelaenus is unreported.
Geographical variation of P. hypermelaenus is only subtle (see Kleinschmidt, 1940:
29-30), and within China no subdivisions seem justified. The isolated population on Mt.
Victoria (Chin Hills, Myanmar) was first considered a distinct subspecies by Stresemann (in Stresemann & Heinrich, 1940: 179), although he did not name it (see also Eck,
1980a: 144-145 and additional references; Harrap, 1996).
General characteristics: subtle geographic differences in coloration were discussed

23 This is presumably Kangding Xian, Dardo, in Sichuan at 30º03’N 102º02’E rather than in Yunnan,

but border changes that have taken place in the meantime might explain this. However, Zhao & Adler(1993: 446) thought the name equated with Tsu-kou at 28º00’N 98º52’E. This difference of opinion
awaits resolution.

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by earlier authors, but need further substantiation due to the lack of adequate series of
specimens. The bib is even larger than in Chinese P. p. jeholicus and P. p. hellmayri, andhypermelaenus is also even shorter-tailed. Measurements in Harrap (1996: 244). TWI
79.5% (range 77-81), sd 1.6 (n = 5 from China alone, including a paratype; Eck 2006).
Distribution: recorded from the provinces of Yunnan, Sichuan, Gansu and Shaanxi inmountainous western China; details in Harrap (op. cit.). It has recently been rediscovered in the Qin Ling range (southern slopes of Taibai Shan; specimens in MTD), from
where it was known historically (Cheng, 1987). This is now the northeastern-most confirmed locality.
Colour images: Quinn in Harrap (1996), pl. 15, fig. 47; Berezowski & Bianchi, (1891),
pl. II: 2; Kleinschmidt (1938), Falco 34, pl. II; Eck, 1980a: pl. I.
Parus montanus Conrad, 1827
Introduction: in terms of species limits, systematics, taxonomy, and phylogeny, theOld World Willow Tit and its chickadee relatives in the New World are among the mostintensively studied tits. There is a considerable body of knowledge of morphology, vocalizations and molecular phylogeny now available. This has influenced the recommendations made here.
In general, as in all ‘Poecile’ tits the delimitation of subspecies is often difficult andsomewhat subjective. It has been argued that too many subspecies have been describedon minor local colour differences. This, to a point, is correct, especially in parts of thewestern Palaearctic. However, this was followed by overlumping of northern forms in
the east Palaearctic by Vaurie (1957a, 1959) and Snow (1957, 1967) sometimes on thebasis of quite limited material, followed by Stepanyan (1990, 2003). We follow here thearguments presented by one of us (Eck, 1980a) but also include new molecular results(Kvist et al., 2001, Salzburger et al. 2002b).
Hartert (1905a: 376) treated the Palaearctic Willow Tits and the Nearctic chickadees(the latter now being P. atricapillus Linnaeus, 1766, and P. carolinensis Audubon, 1834) as
one single species. Mayr (1956), following several previous authors and relying quite
strongly on vocalizations, separated Old and New World taxa as different species. Thisview was tentatively adopted by Mayr & Short (1970: 64). Eck (1980a), applying a broad
species concept, stressed similarities more than the differences between Old and New
World taxa and kept them in a single species, P. atricapillus (which although not followed here might have been his preference).
The distribution of Willow Tits spans almost the whole of the Palaearctic, from the
British Isles continuously across northern Eurasia to Japan, with two disjunct range segments, one in central Asia (Tian Shan) and another in west to north-east China. The
various populations display multiple variations in coloration and in flight feather proportions. However, mitochondrial molecular genetics (Kvist et al., 2001, Salzburger etal., 2002b) indicate that all northern populations are recent divergences. The southern
populations (Tian Shan, China) are more varied, and drawing on these results a discussion of species limits is needed. Russian authors have generally treated songarus Severtsov, 1873 (Tian Shan), as a species (Stepanyan, 1990, 2003), and the Chinese endemics –affinis Przewalski, 1876, stoetzneri Kleinschmidt, 1921a, and weigoldicus Kleinschmidt,
1921b – were generally included in it despite having disjunct ranges. Harrap (1996) and14

Eck & Martens. Review of the Aegithalidae etc. Zool. Med. Leiden 80 (2006)

Dickinson (2003) adopted this view. Here we treat Palaearctic Willow Tits as one species, separating only weigoldicus. This is upgraded to species rank on molecular evidence (see also Eck, 2006), while the rest of ‘songarus’ is replaced in P. montanus.
Using the evidence from morphology, acoustics and to some extent molecular genetics five subspecies groups can be distinguished (Eck 1980a), of which the montanus
group and the salicarius Brehm, 1831, group are extralimital (Europe, east to the Urals).
The other three groups – borealis Selys-Longchamps, 1843, kamtschatkensis Bonaparte,
1850, and songarus – occur in our area and we treat each separately below.
A molecular cytochrome-b study by Salzburger et al. (2002b) identified four distinct
genetic clusters within trans-Palaearctic Willow Tits, represented by three individual
taxa, Parus m. weigoldicus, P. m. songarus, P. m. affinis, and a clade containing all six north
Eurasian subspecies that were investigated: borealis, montanus, rhenanus Kleinschmidt,
1900, baicalensis Swinhoe, 1871, sachalinensis Lönnberg, 1908, and restrictus Hellmayr,
1900a. The three individual taxa weigoldicus, affinis, and songarus were reciprocallymonophyletic and separated from each other by distances between 1.9 and 5.8%. Distances between the north Eurasian subspecies (central Europe to Japan) were low, only
0.65%. These latter subspecies are closely related and still share mitochondrial haplotypes, despite marked morphological and acoustic differences. The current classification which splits the species into the songarus and montanus subspecies groups and assigns them species rank is not in accord with the molecular results. However, the ‘cluster’ comprising weigoldicus appears to be a sister to all other subspecies and is deeply
split from combined montanus and songarus by distance values of 4.6-5.8%. This unexpectedly great distance supports species rank for it and is paralleled by many well-defined Palaearctic passerine species, including some sympatric ones (see, e.g., Päckert etal., 2004). So far, cytochrome-b results do not clarify any of the current subspecific delimitations in northern Eurasia.
The territorial song of Willow Tits now plays a major role in identifying evolutionary units that have a common history. Willow tit song is simple, and falls clearly into
four main types, geographically defined ‘regiolects’ (Martens, 1996) which characterize
Willow Tit populations all over the Palaearctic. These are Lowland, Alpine, Siberian,and Sino-Japanese song types (Thönen, 1962, 1996, Martens & Nazarenko, 1993), allshown in Fig. 1 24. They do not fully mirror the molecular phylogeny and apparently
differentiated on a different time scale, but they well indicate evolutionary units, too.
Current data suggest that the mono-frequency song type (Alpine) may be ancestral being retained over a long evolutionary period in certain populations, but was altered or
camouflaged by learning processes in others. Consequently, Willow Tit territorial songcontributes important information in systematics and the interpretation of distributional histories (Thönen, 1962, 1996, Martens & Nazarenko, 1993, Martens et al., 1995).
We will now discuss geographic variation and subspecies belonging to the different
groups introduced by Eck (1980a) who called them ‘Sektoren’ or sectors – and whichwere groups adopted by Harrap (1996).

24 Fig. with a capital F refers to a figure included in this paper.

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15

Fig. 1. Parus montanus: schematic sonagrams of territorial song. – a) Alpine song type; – b) Lowland songtype; – c) Siberian song type; – d) Sino-Japanese song type. – The Siberian song type is a combination of
the Lowland and Alpine song types; within its range every male disposes of both song types. Distribution: ‘Lowland’ occurs exclusively in Western Europe (extralimital to our area); ‘Alpine’ disjunctly allover the Palaearctic from the European Alps to Hokkaido, Japan; ‘Siberian’ from Scandinavia to Sakhalin; and ‘Sino-Japanese’ in central Asia, parts of China and Honshu/Japan. For more details of song type
distribution see Quaisser & Eck (2003), Martens et al. (1995), Martens & Nazarenko (1993) and Thönen(1962, 1996). – Slightly modified from Quaisser & Eck (2003).

The baicalensis group:
Taxa included (2): P. m. baicalensis (Swinhoe, 1871) [Lake Baikal, Russia]; P. m. shulpini (Portenko,
1954) [Suchan, southern Ussuriland, Khabarovsk Region, Russia].
Unresolved (see below): ? P. m. suschkini (Hachlor [sic!], 1912; Hachlov in German spelling; Khakhlov
in Russian spelling) [river Temir-Su in the Ssaur Mts. 25, the eastern extension of the Tarbagatai Mts.].

Taxonomy: the baicalensis group is most easily characterized by its Siberian song type,Fig. 1c (herein); see also Martens & Nazarenko (1993), and recordings depicted in Cramp& Perrins (1993: 179-180) and Thönen (1996: 8-9). Treatment of subspecies reaching ourarea differs: Vaurie (1957a) and Snow (1967) accepted baicalensis from western to northeastern Siberia and north-eastern China, whereas Stepanyan (1990, 2003) favoured amuch broader taxon, his subspecies borealis included all the populations of the western
taiga sector from Scandinavia to east of Lake Baikal, but this is a minority opinion.
The measurements of suschkini, according to Hachlov (1912), suggest baicalensis but
he distinguished suschkini on grounds of darker and browner upperparts than baicalensis, larger black bib, and virtual absence of white fringes along tail-feathers. These characters seem comparable to those of two skins from the Ssaur Mts in the Kazakh part of
the Tarbagatay, the type locality of suschkini, thus close to our area. Hachlov emphasized
the presence of very large bibs in his new form (specimens from June and July, with one
from March) with the underparts or belly having a beige tinge (“isabellfarbiger Anflug”).
The March skin had an ochre-brownish touch on the feathers of the shoulder of the wing.
This name seems likely to have been applied to ‘hybrids’ between songarus and borealis,view supported by the position of the Ssaur Mts which lie between Dzhungarian Alatau
25 An adult male and a juvenile female from Temir-su river were examined by SE and suggest that thisform is intermediate between songarus and baicalensis.

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and Tien Shan (songarus) and Altai (baicalensis). Clearly, such issues need better understanding but meanwhile it seems best not to recognise suschkini.
One of us (S.E.) checked about a dozen skins from the Altai. Their wing and tailmeasurements largely cover the two quite distinct size pools of borealis and baicalensis,
respectively (on the whole, however, they are closer to borealis). The crown of thesebirds is not so deeply saturated with black as in baicalensis proper. From this area several song types have been noted (Ernst, 1991; Martens et al., 1995; Ernst & Hering, 2000),
which may reflect an area of secondary contact between carriers of different song types
and may indicate a complex historical situation which includes inter-breeding of populations of different Pleistocene origin.
The subspecies shulpini from Ussuriland (in the Russian Far East, close to the borderwith north-eastern China; type and additional skins checked by S.E. at ZISP) has theback slightly more yellow than baicalensis and is valid; it is smaller and relatively shorttailed, like extralimital borealis. One of us (J.M.) has song recordings from the Russian
Far East (lower Amur and Ussuriland; shulpini); they match the Siberian song type.
General characteristics: the two subspecies accepted differ slightly in the coloration of
the upper parts, shulpini being browner, in wing length and in relative tail length (Eck,
1982); changes are clinal. TWI: baicalensis 94.1%, sd 2.08 (n = 70); shulpini 92%, sd 1.97 (n= 10). Territorial song: Siberian type (Fig. 1c).
Distribution: baicalensis group members extend from N Europe to E Siberia and NE
China, but exclude areas close to the sea of Okhotsk, the Anadyr basin, Koryakland andKamchatka [Omolon and Penzhina basins are still baicalensis]. Type specimens of P. m.
suschkuni were from the Kazakh parts of the Saur Mts, not from the Chinese part.
Colour images: Eck, 1980a: pl. IIIa: baicalensis; Quinn in Harrap (1996), pl. 16, fig. 50e:
baicalensis.
The kamtschatkensis group:
Taxa included (2; extralimital forms listed for convenience: 3): P. m. anadyrensis Belopolski, 1932
[Markovo, Anadyr basin, Magadan Region, Russia] EL; P. m. kamtschatkensis (Bonaparte, 1850) [E
Asia = Kamchatka, Russia 26] EL; P. m. sachalinensis Lönnberg, 1908 [Sakhalin; Russia] EL (vagrantto Hokkaido); P. m. ssp. [Hokkaido, Japan]; P. m. restrictus Hellmayr, 1900a [Shimotsuke, Honshu,
Japan].

Taxonomy: Stepanyan (1990: 566) thought anadyrensis and baicalensis might intergrade, but in general the subspecies of the far eastern Palaearctic ranging from the Koryak Highlands to Kamchatka, Sakhalin Is., and Japan are linked by stepwise character
gradation (Eck, 1982: 139-140). Lobkov (1997) suggested the occurrence of a sharpboundary between baicalensis and katschatkensis just north of Tymlat and Shamanka
Rivers, c. 60°N in Kamchatka; he also suggested splitting off kamtschatkensis as specifically distinct: he listed it as ‘P. (m.) kamtschatkensis’ and considered its status ‘close to
specific independence’ (p. 371). Shikoku Willow tits have been described as abei Mishima (1961) but this is not recognized as distinct (Orn. Soc. Japan, 2000).

26 As restricted by Hartert (1905a).

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The evidence of song types in relation to distribution is intriguing. Portenko (1939a:95) described anadyrensis song as “tjou-tjou-tjou”, which is difficult to relate to the song
forms currently known in eastern Asia. Instead his description seems closest to the Lowland song type (Fig. 1b, which would be unusual for the region). Adjoining kamtschatkensis, to the south, sings the Alpine song (Fig. 1a), and sachalinensis the Siberian song (Fig.
1c), while in Hokkaido the Alpine song type is known (Thönen, 1996, Thönen & Fujimaki, 1995). Finally, the Sino-Japanese song type (Fig. 1d) is reported for restrictus from
Honshu (Thönen, 1996).
The taxonomic situation of Hokkaido Willow Tits is unresolved. Hartert (1905a)
declared that these do not belong to restrictus, but later merged them into this subspecies without further comment (Hartert & Steinbacher, 1934: 196). According to the Orn.
Soc. Japan (1922: 157) Hokkaido is inhabited by sachalinensis, but Vaurie (1959) and theOrn. Soc. Japan (1974: 283; 2000: 237) accepted restrictus. However the song of thesebirds is known to be Alpine in character (Thönen, 1996: 18). This distinct song type indicates a different history for this population, implying Hokkaido Willow Tits should
neither be merged into sachalinensis nor into restrictus. Abe & Kurosawa (1984a, b) published measurements of Willow Tits from Hokkaido, but the population should be carefully circumscribed and named.
General characteristics: from north to south the populations tend to become greyer,
smaller, and relatively more short-tailed.
Distribution: kamtschatkensis group members occupy easternmost Siberia from
Anadyrland to Kamchatka, as well as Sakhalin and Japan. Apparently, there is a gap in
distribution: they are absent in the Kuril Is., while birds on the mainland shore of the
Sea of Okhotsk are baicalensis.
Colour images: Eck (1980a): pl. IIIb: anadyrensis, restrictus; Quinn in Harrap (1996), pl.
16, fig. 50f: kamtschatkensis.
The songarus group:
Taxa included: P. m. songarus Severtsov, 1873 [Tian Shan]; P. m. affinis (Przewalski, 1876) [Ala Shanand Gansu]; P. m. stoetzneri Kleinschmidt, 1921a [30 km N of Balihandien, NE Jehol 27 = Balihan, Nei
Mongol].

Taxonomy: if songarus is compared in coloration with baicalensis (or borealis) the difference is remarkable and its treatment as a separate species by Russian authors (e.g.,
Stepanyan, 1978, 1990, 2003) might seem justified. Roselaar in Cramp & Perrins (1993:
185) and Harrap (1996) both accepted this. However, Thönen (1996: 17-18) disagreed
based on vocalizations. The song type of songarus is just a slightly modified Siberiansong (Martens et al. 1995: 375-377). The subspecies affinis uses two different forms of the
Alpine song type, differing by locality (Martens et al., 1995: 377-378), while for stoetzneri
only the Alpine song type is known (but see Martens et al., 1995: 378). For the exclusionof weigoldicus from this group see below.

27 For Balihan see Zhao & Adler, 1993: 427.

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General characteristics: all taxa of the songarus group are yellowish brown on theback, and in the eastern part of the range the populations are more greyish. Only insongarus is the bib brownish; it is blackish in affinis and stoetzneri. All members of the
songarus group have a more or less dark brownish crown, very different from the black
cap of northern willow tits of the baicalensis group. The colour of the crown of affinis wasgiven by Hartert (1905: 368) as “Pelzrobbenbraun” (“Seal Brown“ according to Ridgway, 1886, III.1), and is shown by Ridgway (1912) in his plate XXXIX; that of stoetzneri
which is blackish orange brown = “schwärzlichorangebraun“ (Anon., 2000, MichelFarbführer, pl. 17) is quite close. It is deeply brownish in songarus as well. In affinis andsongarus the typical willow tit wing patch is hardly to be seen or virtually absent (seecolour plate VI in Eck 1980a). The tail/wing relations of songarus, affinis and stoetzneri
differ considerably, stoetzneri being the smallest form but relatively long-tailed. Stoetzneri
also differs slightly in colour between western and eastern parts of its range (S.E., basedon material in IZAS). All three forms differ in size, songarus is a large form with wing up
to 73 mm, affinis up to 68 mm, stoetzneri is smallest (up to 66 mm). For detailed measurements and tail/wing proportions see Eck (1980a: 151-153, 2006).
Distribution: with weigoldicus (q.v.) excluded these forms are highly disjunct: songarus occurs in central Asia (Kazakhstan, and in our area in Xinjiang), affinis is found only
in western China (NE Qinghai, Gansu, N Sichuan, SW Shaanxi and Ningxia); andstoetzneri in northeast China (Shanxi, Henan, Hebei, Beijing and SE Nei Mongol
Zizhiqu).
Colour images: Eck, 1980a: pl. VI: stoetzneri, affinis, songarus; Quinn in Harrap (1996),
pl. 16, figs. 51a: songarus, b: stoetzneri.
Parus weigoldicus Kleinschmidt, 1921b
Taxon included (1): P. weigoldicus Kleinschmidt, 1921b [Atentze (= Atuntze, now Tehtsin)] 28. –Monotypic.

Taxonomy: the close relationships of the taxa perceived as the songarus group (songarus, affinis, stoetzneri and weigoldicus), were never called into question until Salzburger
et al. (2002b) detected a marked cytochrome-b distance of 4.6-5.9% between weigoldicus
and all other Willow Tits (incl. songarus, affinis).
In the phylogenetic topology generated weigoldicus is sister to all other Willow Tits
including affinis and songarus. The unexpectedly high distance value indicates separate
development by weigoldicus over the last two to three million years (based on the rulesuggesting 2% distance per 1 million years of separate development (Lovette, 2004).

28 Kleinschmidt (1921a) listed “z. B. Atentze”. This was thus an example of a collecting locality and he

implies that there must have been two or more type specimens. It follows that although a specimen waslabelled to suggest that it was a holotype, as indicated by Eck & Quaisser (2004: 290), we now note that it
does not qualify (see Art. 73.1.3 of the Code, ICZN, 1999). Under the terms of Art. 74.7 this error did not
accidentally result in a lectotype, and the specimen, C 23803, from Mauntschi (Barongschiba) remains a
syntype. The other specimens listed by Kleinschmidt & Weigold (1922) are not all in Dresden, one is in
the AMNH, and consultation seems desirable before any decision is made on whether lectotypification
is appropriate, especially if the current name and coordinates of Mauntschi require clarification.

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In size weigoldicus is similar to the Tian Shan Willow Tits (P. montanus songarus),
but it differs conspicuously from parapatric affinis by larger wing length, shorter taillength, smaller TWI and less pronounced tail gradation, and in coloration (see Eck,
1980a: 151-153 and Eck, 2006 for details). The song type is Sino-Japanese (Thönen,1996: 9, Martens et al., 1995) while that of affinis is the Alpine type. More work isneeded on the Chinese Willow Tits but despite the present lack of additional biological data, especially regarding interactions with adjoining affinis, which would allow abetter characterization of P. weigoldicus, the molecular evidence compels us to accept
specific status for weigoldicus. We recommend the English vernacular name Sichuan
Willow Tit.
General characteristics: conspicuously dark, earthy brown – “erdbraun”, (Kleinschmidt,
1921a) – on the back, the flanks very dark. For measurements and tail/wing length relationships see Eck (2006). No specimens are known which might indicate interbreeding with the parapatric P. montanus affinis.
Distribution: highly restricted; Sichuan and parts of northern Yunnan, China. The
range is parapatric to that of P. m. affinis (see detailed map in Quaisser & Eck, 2003). The
existence of a contact zone with the latter is so far unknown. P. weigoldicus is apparently
a rare bird and not easily encountered.
Colour images: Eck, 1980a: pl. VI, bottom figure.
Parus superciliosus (Przewalski, 1876)
Taxon included (1): Parus superciliosus (Przewalski, 1876) [alpine regions of ‘Gansu’ 29]. – Monotypic.

General characteristics: the White-browed Tit is an unmistakable rather long-tailed tit
with a white eye-brow. No taxonomic problems known.
The cytochrome-b molecular phylogeny places P. superciliosus at the base of the Poecile group or close to it (Gill et al., 2005). Detailed measurements are given in Eck(2006).
This tit inhabits tree-less high-altitude steppes, where only bushes persist. Nevertheless like all other Parus species, it is a hole-breeder, breeding in the burrows of pika
and other rodents (Martens & Gebauer, 1993), or in fissures of rock faces. The first nest
located (south of Koko Nor, Qinghai) held young and was situated in a rodent hole in aloess cliff and was entirely composed of moss (J.M., unpublished).
Distribution: high altitudes on the Tibetan plateau, China, but only locally: in NE
and SE Tibet proper, E Qinghai and NW Sichuan.
Colour images: Pleske, pl. VIII: 3, 4; Kleinschmidt, 1911; Eck, 1980a, pl. 9, upper figure; Quinn in Harrap, 1996, pl. 18, fig. 56; Anderton in Rasmussen & Anderton, 2005, pl.
155, fig. 7.

29 Vaurie (1972: 88, 88fn), who used the name Tibet in a geographic, not political sense, wrote of Przew-

alski’s specimens labelled ‘Gansu’ “I was not sure at the time that they had been collected in Tibet, but
I believe now that these were probably taken in Tibet, chiefly in the South Tatung Range, not in Kansu
proper”. Vaurie (1972: ix) mapped the Tatung River (now Datong He) north of Hsi-ning (now Xining),
and the range, now the Daban Shan, is close to southern bank of the river. It is in fact in present day
Qinghai.

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Parus davidi (Berezowski & Bianchi, 1891)
Taxon included (1): Parus davidi (Berezowski & Bianchi, 1891) [S Gansu]. – Monotypic.

Taxonomy: one of us (S.E.) has held varied views on the conspecificity of P. davidi
Père David’s Tit with west Palaearctic P. lugubris Temminck, 1820, the Sombre Tit; these
forms seem related by morphology and geography and it has been hypothesised thatconnection was through P. l. hyrcanus. Eck (1976: 33) did not consider this, but later (Eck,
1980a), after a comparison with hyrcanus, came to accept it and used a broad species
concept (see Eck, 1996).
However, one of the cytochrome-b topologies places P. davidi and P. lugubris as paraphyletic groups at the basis of the ‘Poecile’ tree (Gill et al., 2005) and the conspecificityof P. davidi and P. lugubris is not maintained here.
Distribution: central China.
General characteristics: a small, vivid and intensely-coloured tit with a thick bill.
Wing length up to 71 mm, tail length up to 54 mm. TWI: 74.7%, sd 2.13 (n = 24).
Colour images: Berezowski & Bianchi, 1891, pl. II: 4; Eck, 1980a: pl. II; Quinn in Harrap, 1996, pl. 18, fig. 57.
Parus cinctus Boddaert, 1783
Taxon included (1, extralimital form listed for convenience: 1): P. c. cinctus (Boddaert, 1783) [Siberia],
P. c. sayanus (Sushkin, 1904) [W Sayan, Altai] EL.

General characteristics: sayanus, a southerly montane form, is similar to nominate
cinctus in the relative length of wing and tail (TWI: 95.9%, sd 1.17, n = 10), but is markedly larger, wing up to 76 mm.
Taxonomy: the breeding localities in Mongolia are close to the type localitiy of Parus(Póecila) obtectus Cabanis, 1871 (‘south of Lake Baikal’). That form is usually treated as asynonym of nominate cinctus (Snow, 1967, Stepanyan, 1990). Consequently, we list
nominate cinctus for our area. However, according to Mauersberger (1983), Nazarenko(1978) identified new material from southern Transbaikalia with sayanus and treatedthis as extending into northern Mongolia.
Distribution: early suggestions that the Siberian Tit occurred in China have been
discussed and finally rejected by Vaurie (1957a, 1959). Cheng (1976, 1987) also expressed
doubts about old records and knew of no new records. Zheng (2005) did not even list
the species for China. The species nonetheless reaches north-west Mongolia where it
breeds in an area just west and east of Hövsgöl Nuur and in N Kentei (Hentyin Nuruu)(C.S. Roselaar pers. comm.). Whether sayanus reaches our area (as a breeding bird) remains unclear. Dickinson et al. (2006) consequently list it as ‘probably extralimital’.
Colour images: Eck 1980a, pl. VIII: lapponicus (extralimital); Quinn in Harrap, 1996,
pl. 19, fig. 58a-c: lapponicus, 58d: cinctus (both extralimital).
Parus rubidiventris Blyth, 1847
Taxa included (4): P. r. rubidiventris Blyth, 1847 [Nepal]; P. r. beavani (Jerdon, 1863) [Mt. Tongloo, Sikkim, India]; P. r. whistleri Stresemann, 1931 [near Lau-hu-kou, S Tetung (= Datong) Mts., NW Gansu,

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China]; Syn.? P. r. szetschwanensis Meise, 1937; see below [Wa Shan, E Hsikang, now W Sichuan]; P.
r. saramatii Ripley, 1961 [Mt. Saramati, Naga Hills 30, Myanmar].

Taxonomy: until recently the Rufous-vented Tit posed no taxonomic problems except for the fact that the two Sino-Himalayan subspecies, P. r. rubidiventris (western
Himalayas) and P. r. beavani (eastern Himalayas, China), were sometimes considered
to represent different species (Wolters, 1980; Haffer, 1993), though on scant evidence
except for marked differences in coloration of grey-bellied and rufous-bellied populations. No hybrid population of these strongly different subspecies had been detected,
but, apparently, nobody paid special attention to this question. Allegedly, the range ofrubidiventris overlapped with that of beavani (see Martens 1975); but there had been no
recent confirmation. Even though such a zone of contact might have been expected in
the upper course of Bhote Kosi valley (somewhat northeast of Kathmandu).
Cytochrome-b data depict a more complicated evolutionary situation (Martens etal., 2006). Within the continuous Sino-Himalayan region (Kashmir to Shaanxi, China)
all P. rubidiventris populations fall into two deeply split clusters (with a distance value
between them of about 2.5%) and these do not coincide with the current subspecificdelimitation of rufous-bellied rubidiventris and grey-bellied beavani (Fig. 2).
30 Mt. Saramati lies exactly on the border with India and is judged to be a part of the Naga Hills.

Fig. 2. Distributional map of subspecies and of cytochrome-b haplotype clusters of Parus rubidiventris.
Solid bar (in Himalayas, 1) indicates the distribution limit of reddish-bellied western nominate rubidiventris and of grey-bellied eastern beavani and whistleri; isolated area: saramatii (grey-bellied). Thin bar(SW China, 2) separates cytochrome-b haplotype clusters, i.e. the Himalayan cluster (nominate rubidiventries and beavani) and the Chinese cluster (whistleri). Haplotype affiliation of saramatii unknown. –Slightly modified from Martens et al. (2006).

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The populations in the Himalayas proper, i.e., western rufous-bellied nominate rubidiventris and grey-bellied beavani, belong to the same cytochrome-b haplotype cluster
and can carry the same haplotype. Thus most of the western, Himalayan, part of therange of beavani belongs to a cytochrome-b cluster that is deeply different from the eastern cluster which relates to populations in the eastern parts of the range of beavani, i.e.
in China. Representatives of the two clusters also differ in body size.
For the genetically different Chinese cluster (eastern grey-bellied “beavani”) the oldest available synonym is whistleri and the use of this was revalidated by Martens et al.
(2006). Originally, whistleri (from Gansu) was separated from beavani on the basis of its
smaller size and lighter slaty-grey upperparts. Further south, birds from Sichuan wereseparated as szetschwanensis. Like whistleri they were considered to be smaller than beavani and they differed by being darker slaty-grey above (Meise in Stresemann et al.,1937). Both these small forms, which also differ from beavani in flight feather proportions, were unfortunately lumped with beavani by Vaurie (1957a). For the moment, we
keep szetschwanensis in the synonymy of whistleri, which is clearly a valid form. A morethorough revision of the eastern grey-bellied tit ‘cluster’ is needed. P. r. saramatii, a disjunct population from the Naga Hills, is apparently smaller and differs in colorationfrom beavani and whistleri, its cytochrome-b cluster assignment is unknown.
The cytochrome-b sequence difference between the western cluster (rufous- andgrey-bellied forms in the western central and eastern Himalayas) and the eastern cluster (in China and probably in parts of the easternmost Himalayas, but see below) is up
to 2.5%. This is the same magnitude of difference as between the three southern cytochrome-b clusters of P. ater and to the values between the well-established species P.
venustulus, P. amabilis and P. elegans (see below).
General characteristics: centre of belly rufous or rusty red in rubidiventris from thewestern and west-central Himalayas. Measurements can be found in Martens & Eck(1995: 327). Belly grey in the eastern forms beavani and whistleri; and also in saramatii,
but the nature of the grey tone differs (Ali & Ripley, 1973). Between the Himalayas andGansu/China wing length decreases. Overall coloration differs locally. P. r. beavani sensu
stricto has a TWI of 70.3%, sd 1.22 (n = 44) and a wing length of up to 74 mm (like rubidiventris); whistleri/szetschwanensis tend to have longer tails, their TWI is 71.9%, sd 1.30
(n = 31), maximum wing length 69 mm (see also Eck, 2006).
Distribution: nominate rubidiventris ranges from Kashmir (as a rare breeding bird,
Jamdar & Price, 1990) east to at least Syng Gyang monastery in the Gosainkund Lekh at
85°21’E., just north of Kathmandu (Martens & Eck, 1995; specimen ZFMK). Even further east, in Helambu, Kandak-Kosi watershed, Malemchi Khola, rufous-vented birds
were observed at about 85°31’E (Proud, 1951). Slightly further east, at appropriate altitude in the upper Bothe Kosi valley (at about 86°E, probably not in Nepal but in southern Tibet), nominate rubidiventris and P. r. beavani are thought likely to meet, but no such
locality has yet been spotted. There is no information on the area where representativesof the two ‘beavani’ clusters may meet; this may be somewhere in the eastern Himalayas
or in SW China. C.S. Roselaar (in litt.) suggests that if size is a reliable basis for telling
beavani from whistleri, the border between them is probably formed by the upper Mekong/Lancang rivers. According to Vaurie (1950), N Myanmar and NW Yunnan birds
are still large, while Sichuan and Gansu birds are small. It is not yet clear whether birds
from NE Burma should be associated with whistleri or with szetschwanensis.

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Colour images: herein Pl. II, Fig. b: rubidiventris, c: beavani; rubidiventris; Martens &Eck (1995), pl. 2 is the same image; Quinn in Harrap (1996): pl. 20, figs. 62a-b: rubidiventris, c-d: beavani.
Parus rufonuchalis Blyth, 1849
Taxon included (1): Parus rufonuchalis Blyth, 1849 [“Range beyond Simla, near the snow-line”, NW
Himalayas. Synonyms? blanchardi Meinertzhagen, 1938 [Gardez Forest, E Afghanistan]; parvirostris
Keve, 1943 [Naryn, Tian Shan]. – Monotypic.

Taxonomy: Martens (1971) discovered sympatric and even syntopic occurrences of P.
rubidiventris rubidiventris, the Rufous-vented Tit, and the Rufous-naped Tit P. rufonuchalis during the breeding season in dry trans-Himalayan coniferous forests in west Nepal,
proving the latter to be an independent species.
At different places either P. a. melanolophus or P. a. martensi, and P. rubidiventris occur in the same forest patches as P. rufonuchalis. Thus ‘Periparus’ is represented in the
Himalayas by three different-sized species rufonuchalis (largest), rubidiventris (medium-sized) and ater Linnaeus, 1758 (smallest), and these size differences allow syntopic breeding. Sympatric occurrence of the two larger species in the western Himalayas has also been reported by Biswas (1963), Jamdar & Price (1990) and Gaston et al.
(1993).
General characteristics: nuchal spot rusty; bib widely extended on lower breast. One
of us (S.E.) found a maximum wing length of 79 mm and maximum tail length of 58 mm
in P. rufonuchalis while P. r. beavani has a maximum wing length of 75 mm and a maximum tail length of 52 mm (Ali & Ripley, 1973: 183). When sexed correctly there seems
to be considerable variation of measurements within sexes so that one may doubt
whether rufonuchalis is uniform throughout its range (see measurement details in Eck,
2006). However, mitochondrial cytochrome-b haplotype evidence from Nepal and Kyrgyzstan (Martens et al., 2006) so far suggests there is no geographical variation. Should
subspecific treatment be needed synonyms are available (see above).
Distribution: from the Tian Shan to the western Himalayas in west Nepal (eastern
Dhaulagiri area), as far east as the dry Kali Gandaki river gorge in east Dhaulagiri/west
Annapurna (Martens, 1971, Inskipp & Inskipp, 1991, Martens & Eck, 1995). In China it
is restricted to SW Kokshaaltau and the E Pamir Mts in W Xinjiang (Vaurie, 1972). It also
occurs in the uppermost Sutlej valley just within SW Tibet, but this is not mapped inCheng (1987). A record (dot) in Cheng (1987: 887, map 701; under P. rubidiventris) ineastern Tian Shan needs verification as this is far east of known distribution in thisrange (C.S. Roselaar pers. comm.).
Colour images: herein Pl. II, Fig. a; Martens & Eck, 1995, pl. 2 fig. a (same image);
Quinn in Harrap, 1996, pl. 20, figs. 61a-b.
Parus ater Linnaeus, 1758
Introduction: the taxonomy of Coal Tits in the Himalayas is still in debate (Martens& Eck, 1995, Harrap, 1996, Dickinson, 2003), but a fresh evaluation can be offered basedon surprising molecular genetic data obtained recently (Martens et al., 2006).

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Eck & Martens. Review of the Aegithalidae etc. Zool. Med. Leiden 80 (2006)

Fig. 3. Distribution of the Coal Tit complex (Parus ater sensu lato) in Nepal. In the upper Myagdi valley
melanolophus and martensi meet and hybridise, martensi occupies a small area in the eastern Dhaulagiri
and Annapurna ranges; east of the Annapurna massif aemodius occurs, extending to the easternmost
Himalayas. Cinnamon-bellied morphs of melanolophus are known from the Dhorpatan valley only. Blackshading: potential breeding zone of the P. ater group, roughly between 3000 and 4000 m. – Slightly modified from Eck (1998). Bars shown: a. eastern limit of P. a. melanolophus; b. eastern limit of P. a. martensi; c.
the Buri Kandaki valley (Coal Tit population awaiting determination). Further east P. a. aemodius occurs.

Marked colour and pattern differences between western and eastern Himalayan
Coal Tit populations (and Coal Tits in general) led to a conclusion with which nearly all
recent authors agreed (Hartert 1905a, Harrap 1996, Inskipp et al., 1996, Dickinson, 2003),
namely that two small ‘Periparus’ species inhabit the Himalayas: monotypic P. melanolophus Vigors, 1831, in the west, and representatives of P. ater (martensi Eck, 1998, andaemodius Blyth, 1845 31) in the east, ranging into western China.
However the situation has steadily been revealed to be considerably more complex.
In the south-western Dhaulagiri area of west Nepal “cinnamon-bellied hybrids” werediscovered (Diesselhorst & Martens, 1972) (Fig. 3). These have a striking colour pattern,which differs from melanolophus in the dark chestnut-red centre of the belly, and differs
strongly from typical Himalayan Coal Tits as well (Diesselhorst & Martens, 1972, Martens, 1994; Martens & Eck, 1995: 330-332, pl. 3 fig. d). An alternative hypothesis wasthat these birds were a local colour morph, not hybrids (Diesselhorst & Martens, 1972,
Martens & Eck, 1995) – possibly a vanishing form (Eck, 2006). This ater population lives
at low density within the range of melanolophus-coloured birds in a single high-altitude
valley, the Dhorpatan valley at the south-western corner of the Dhaulagiri massif, andthey interbreed freely with local melanolophus (Martens, 1975). Subsequently, smallsized ‘Periparus’ were found more continuously along the Himalayan chain and P.

31 Dated 1844 in Snow (1967: 93); for reasons to change see Dickinson & Pittie (2006).

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Fig. 4. Distribution of cytochrome-b haplotype clusters of Parus ater sensu lato in the Palaearctic. Numbers indicate the cytochrome-b lineages and the bars signify the appropriate distribution limits between
them (of which details are unknown). – 1: North African cluster; – 2: Central/South European cluster;– 3: Siberian cluster (several subspecies and includes disjunct Japanese insularis and Tian Shan rufipectus); – 4: West Himalayan cluster (melanolophus); – 5: East Himalayan cluster (martensi, aemodius); – 6:
Chinese cluster (eckodedicatus). Cluster not yet determined and area without number: southeast China(kuatunensis) and Taiwan (ptilosus). Extralimital to the area under consideration are lineages 1 and 2. –Slightly modified from Martens et al. (2006).

melanolophus was reduced to a “semispecies” (Martens, 1975), and later to subspecies
rank (Eck, 1996, Martens & Eck, 1995). More recently a population of the Coal Tit complex with a highly variable colour pattern was located in the upper course of the Myagdi valley in the south-eastern Dhaulagiri massif, some 50 km from Dhorpatan; thesebirds appeared to be ‘hybrids’ between aemodius and melanolophus (Martens & Eck,
1995: 337) and this was supported by their molecular genetics (Martens et al., 2006).
About 30 km further northeast of the upper Myagdi Valley, along the upper course of
the Kali Gandaki river another quite distinct and highly restricted Coal Tit population
exists, which in colour is close to eastern Himalayan aemodius, not to melanolophus (colour plates in Martens, 1993, Martens & Eck, 1995, republished here). This was describedas P. ater martensi by Eck (1998). From the neighbouring Marsyandi valley eastwards theHimalayan chain is inhabited by P. ater aemodius; its range extends to the mountains ofwestern China.
More complete recent molecular genetic analyses paint a complicated, but different picture of the Parus ater/P. melanolophus complex (Martens et al., 2006). In the Palaearctic it is divided into (at least) six distinct cytochrome-b haplotype clusters, ofwhich four occur in our area (Fig. 4). The Sino-Himalayan area contains three of them:
one is typical melanolophus (western Himalayas), the second is in the eastern Himalayas and the third in western China while the contact area between these two is not yet
known. Fourthly, we have a variable ‘hybrid’ population (melanolophus × martensi) in a

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restricted area in western Nepal, where the western cluster (melanolophus) mixes withthe eastern Himalayan cluster as confirmed from cytochrome-b haplotype details anddiscussed above.
P. a. aemodius sensu lato is subdivided into two cytochrome-b clusters, which are restricted to the Himalayas and to western China respectively. The Chinese cluster, forwhich no name was previously available, has recently been described as P. a. eckodedicatus Martens, Tietze & Sun, 2006.
Another geographically extended cytochrome-b cluster ranges from northern andeast-central Europe and central Asia via Siberia to Japan. It includes the proven subspecies, ater Linnaeus, 1758, rufipectus Severtsov, 1873, amurensis Buturlin, 1907, and insularis Hellmayr, 1902.
This clear-cut genetic structure is somewhat inconsistent with current subspecies limits (see below). The value distances between the eastern Palaearctic clusters are remarkably high, ranging from 2.6% to 3.2%. They indicate differentiation times of roughly 1.5
m.y. (based on the 2% rule) and the value distances are even higher (up to 3.4%) when thetwo remaining clusters from Europe and North Africa are compared with them.
This arrangement of geographical “forms” makes the Coal Tit one of the most variable passerine species of the eastern Palaearctic and in the Old World in general.
Below we arrange the Asian subspecies according to their cytochrome-b cluster affiliation. Within clusters some subspecies may be morphologically similar and others
very different (as in the Siberian cluster for example); equally subspecies of different
clusters may be very similar (East Himalayan and Chinese clusters).
The Siberian cluster:Taxa included (4, extralimital forms listed for convenience: 1): P. a. rossosibiricus Johansen, 1952
[Popowo, 20 km SE of Kainsk, Baraba steppe, west Siberia] EL; P. a. rufipectus Severtsov, 1873 [C
Tian Shan]; P. a. amurensis (Buturlin, 1907) [Amur Valley] 32; P. a. insularis Hellmayr, 1902 [Surugano-kumi, Honshu, Japan]; P. a. pekinensis David, in Swinhoe 1870a [Peking = Beijing].

Taxonomy: based on coloration and proportions this cluster can be recognised in a
number of different forms. In our area, the populations in Siberia are rather uniform,
and not all subspecies have been universally accepted. The individuals belonging to theSiberian populations are uncrested. In view of unexpectedly distinct mitochondrial genetic characteristics, even among extremely similar populations (e.g., the east Himalayan cluster, see below) it seems best to accept some less pronounced subspecies inorder to signal evolutionary units within the genetic clusters. In regard to amurensis
note that Buturlin (1907) had enough specimens to give a range of measurements but
did not specify a type and the whereabouts of his type material is unknown (see Dickinson et al. 2006, this issue). He wrote “Diese Form bewohnt die Stromgebiete des Amurund Ussuri, ferner Sachalin und die südöstliche Küste des Ochotskischen Meeres” indi-

32 The type locality given by Snow (1967) appears to be a derivation of the scientific name that Buturlin

selected and not a restriction of type locality. As quoted above the range Buturlin gave included both theAmur and Ussuri river basins and Sakhalin as well as the south-east coast of the Sea of Okhotsk.

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cating a substantial range. It may prove desirable to restrict the type locality and thebest means of doing so would be to designate a lectotype hence we appeal for information on any specimens that may be his types.
The disjunct central Asian rufipectus is very similar in coloration to Himalayan aemodius but has no distinct crest although the feathers on the hind crown are slightly elongated. This subspecies is proportionately extremely long-tailed (measurements andproportions in Eck, 2006).
Distribution: this subspecies group spans the northern Palaearctic from northern
Scandinavia and eastern Germany to Japan and reaches our area via Mongolia andnorth-east China (amurensis and pekinensis). Isolated populations occur in central Asia
(rufipectus) and Japan (insularis).
Colour images: herein Pl. III, Fig. b: rufipectus Tian Shan; Martens & Eck, 1995, pl. 3b:is the same image; Quinn in Harrap, 1996, pl. 21, fig. 64f: pekinensis.
Southern clusters:
General characteristics: members of these southern clusters are found in the Himalayas and the mountainous areas of western China with outliers in Fujian/Jiangxi and
Taiwan, and they have a long crest in common. They are disjunct from the northern
Euro-Siberian-Japanese area (which includes the Siberian cluster). They vary strongly
in coloration, the western Himalayan form (melanolophus) being dark slaty grey, and theeastern Himalayan and Chinese forms beige or chamois coloured on the belly. There isa cline of increasing wing length from the western Himalayas to western China (melanolophus via martensi and aemodius to eckodedicatus).
a) The west Himalayan cluster:Taxon included (1): P. a. melanolophus Vigors, 1831 [Himalayas = Simla-Almora 33]

Taxonomy: where in the eastern range of melanolophus this taxon meets P. a. martensi(of the east Himalayan cluster) a variable local ‘hybrid’ population arises in the upper
Myagdi valley of the south-western Dhaulagiri massif. This population incorporates
haplotypes of the west and the east Himalayan cytochrome-b clusters. Measurements
are given in Martens & Eck (1995) and in Eck (2006); and a detailed map of melanolophus
distribution is found in Wunderlich (1989). Löhrl (1994) crosbred Afghan P. ater melanolophus and German P. ater abietum in captivity. Offspring were quite variable and included individuals similar to the cinnamon-bellied hybrids of west Nepal, though lessintense in reddish coloration.
Colour images: herein Pl. III, Figs. c-d (d is the cinnamon-bellied morph); Martens &Eck, 1995, pl. 3 is the same plate). Quinn in Harrap, 1996, pl. 63, figs. a-c; cinnamon-bellied morph: fig. e.

33 Restricted by Ticehurst & Whistler (1924: 471) but not without some dispute.

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Eck & Martens. Review of the Aegithalidae etc. Zool. Med. Leiden 80 (2006)b) The east Himalayan cluster:Taxa included (2): P. a. martensi Eck, 1998 [west-central Nepal, Chadziou Khola valley, Thakkhola,
Mustang District, Nepal 34]; P. a. aemodius Blyth, 1845 35, sensu stricto [Nepal 36].

Taxonomy: martensi differs from both aemodius (sensu stricto, eastern Himalayas)
and eckodedicatus (China) by its bright ochre-beige vent and belly, and this character
makes it a clear-cut and easily distinguished subspecies (Eck, 1998). Measurements in
Eck (2006).
Colour images: herein Pl. III, Fig. e: martensi, f: aemodius; Martens & Eck, 1995, pl. 3cis the same plate; Quinn in Harrap, 1996, pl. 21, figs. 63d: martensi (subspecies still undescribed when Harrap, 1996, was published), 64k-l: aemodius.
c) The Chinese ‘cluster’:Taxon included (1): P. a. eckodedicatus Martens, Tietze & Sun, 2006 [Wawu Shan, Sichuan, China].

Taxonomy: in outward appearance this newly described subspecies is very similar toHimalayan aemodius, but it forms a distinct cytochrome-b haplotype cluster and differs
morphologically by having slightly smaller wing measurements and a slightly different
vent colour. Through haplotype analysis this subspecies can be confirmed from Gansu,
Sichuan, Shaanxi and northern Yunnan (Martens et al., 2006). The contact area withaemodius is not yet known, but is presumably in southwest Yunnan or in the easternmost Himalayas.
Colour images: Martens et al., 2006, fig. 6 (colour photograph p. 117).
d) Undefined cluster(s):Taxa included (2): P. a. kuatunensis la Touche, 1923c [Kuatun = Guadun, Fujian, China], P. a. ptilosus
Ogilvie-Grant, 1912 [Taiwan, China].

These are highly disjunct, restricted-range populations in Fujian, Jiangxi and Anhui, and in Taiwan, respectively, which may represent cytochrome-b clusters of theirown. Their molecular genetic arrangement is not yet known. Both have pointed crests.
Colour images: Wang et al. (1991: 209).
Parus venustulus Swinhoe, 1870b
Taxon included (1): Parus venustulus Swinhoe, 1870b [gorges of Yangtze = Chang Jiang]. Monotypic.

34 This lies west of the range of aemodius sensu stricto (see Fig. 3 herein).
35 Dated 1844 in Snow (1967: 93); for reasons to change see Dickinson & Pittie (2006).
36 Eck (1998: 130) noted that due to the wide distribution of P. a. aemodius, from EC Nepal to C China, andthe corresponding establishment of this name in the literature it is justifiable to refer the name aemodius
to the form which is widely distributed in the eastern Himalayas (combined with the erection of thesubspecies martensi this is implicitly a restriction of the type locality to eastern Nepal. – Ed.).

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Taxonomy: Parkes (1958: 96) for the first time, and no doubt correctly, combined P.
venustulus, the Yellow-bellied Tit, P. amabilis Sharpe, 1877, and P. elegans Lesson, 1831, in
a superspecies. Based on cytochrome-b molecular genetics both Gill et al. (2005) andMartens et al. (2006) found these species form a monophyletic group.
However, the cytochrome-b distance values are quite low in so far as these values
are derived from forms to which species rank is assigned: 3.3% between amabilis andelegans, 3.4% between venustulus and amabilis, and 3.8% between venustulus and elegans.
These values are of much the same order of magnitude as those between the cytochrome-b clusters of Parus ater (q.v.) 37 which are all treated as belonging to one species.
Apparently, and more puzzling, there is no clear lineage sorting as to species. Threesamples of P. venustulus belonged to a single haplotype (Sichuan, Shaanxi; Martens &al., 2006) and were sister to a sample of P. e. elegans (Luzon). One of P. e. mindanensis(Mearns, 1905) (Mindanao) appeared as sister to one P. amabilis (Palawan) and also to
the P. venustulus and P. e. elegans sisters. Material from several more Philippine islands
is needed to shed more light on these unexpected findings.
Morphologically, P. venustulus, P. elegans, and P. amabilis differ not only in size, coloration and colour pattern, but also in flight feather proportions. P. venustulus is extremely short-tailed – TWI: 54.6%, sd 1.65 (n = 12); WTI 23.7%. sd 1.42 (n = 13). The large
P. amabilis, the allospecies from Balabac, Palawan and Calauit, is relatively long-tailed –TWI: 62.2%, sd 2.1 (n = 4); WTI: 19.6%, sd 1.76. The last allospecies, the smaller polytypic P. elegans (for distribution in the Philippines see Dickinson et al., 1991, or Kennedy
et al., 2000) is similar in tail/wing proportions to P. amabilis.
General characteristics: there is marked sexual dimorphism in colour, moult, andfeather generations (Löhrl, 1988, Harrap, 1996: 311). The behaviour of captive P. venustulus was investigated by Löhrl (1987, 1988) and compared to other species. Löhrl discovered interesting similarities to Parus ater and to Sylviparus modestus Burton, 1836.
Colour images: Berezowski & Bianchi, 1891, pl. II: 3; Quinn in Harrap, 1996, pl. 18,
figs. 65a-f.
Parus elegans Lesson, 1831
Taxa included (9): Not detailed here; no taxonomic changes have occurred since Snow (1967) andDickinson et al. (1991).

Taxonomy: the Elegant Tit, endemic to the Philippines, is currently divided into ninesubspecies (Snow, 1967; Dickinson et al., 1991) which indicates the high evolutionary
potential of this insular species. The unexplained intraspecific molecular-genetic situation of elegans has been discussed above under P. venustulus. Parkes (1958) placed panayensis (Mearns, 1916) in the synonymy of ssp. elegans, but with some doubt. The resultant range of the lumped taxon although seemingly contiguous is zoogeographicallysurprising since Panay usually hosts the same form as is found on Negros (E.C. Dickinson, pers. comm.).

37 Although that between venustulus and elegans is somewhat higher.

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Eck & Martens. Review of the Aegithalidae etc. Zool. Med. Leiden 80 (2006)

Colour images: Quinn in Harrap, 1996, pl. 22, figs. 66a-d: elegans, 66e: gilliardi, 66f:
albescens, 66g-h: mindanensis.
Parus amabilis Sharpe, 1877
Taxon included (1): Parus amabilis Sharpe, 1877 [Balabac, Philippines]. – Monotypic.

Taxonomy: no special comments on the Palawan Tit, a restricted-range species endemic to Balabac, Palawan and Calauit in the Philippines (Dickinson et al., 1991). Placement on the topology of Parus based on the cytochrome-b gene was discussed by Gill etal. (2005) and Martens et al. (2006).
Colour images: Quinn in Harrap, 1996, pl. 22, figs. a-d.
Parus dichrous Blyth, 1845 38
Taxa included (5): P. d. kangrae (Whistler, 1932) [Koti State = Kangra, N Punjab, India]; P. d. dichrous
Blyth, 1845 [Nepal = northern C Nepal 39]; P. d. izzardi Biswas, 1955 [Thammu, Bhote Kosi valley,
Khumbu, E Nepal; for recognition of this 40 see Martens & Eck (1995: 321-322)]; P. d. wellsi Baker,
1917 [‘Big Bend of the Yangtze’ = Chang Jiang, NW Yunnan, China]; P. d. dichroides (Przewalski,1876) [‘Gansu’ 41, China].

Taxonomy: Vaurie (1957a, 1959) accepted four subspecies of the Grey Crested Tit,
and in his 1959 synoptic revision he described the colour differences between them asslight but well marked. Martens & Eck (1995: 321-322) pointed to a marked differencein wing length and relative tail length between dichrous (western Himalayas) and izzardi (eastern Himalayas). The easternmost known locality of dichrous (in Rasuwa Distr.;
specimen in MTD) and the westernmost of izzardi (in Ramechap Distr., specimen in
ZFMK), both in Nepal, are about 125 km apart. It remains unknown whether there is acontinuous cline connecting these two populations or whether they range closer and an
abrupt character change occurs, but we repeat here that we feel it best to recognise izzardi. Though the material is still somewhat sparse (dichrous 4么, 2乆, izzardi 9么, 4乆; allfrom Nepal, the Martens and Diesselhorst collections, see Martens & Eck, 1995; Diesselhorst, 1968, and one additional specimen in MTD), the difference between the two is
obvious: wing length of 么: dichrous 65.5-69 mm (mean 67.1); izzardi 71-75 mm (mean72.4). TWI of 么: dichrous 63.8-66.4% (mean 65.4), izzardi 66.7-72.2% (mean 68.8, sd 1.68).
P. dichrous forms a superspecies with P. cristatus of the western Palaearctic (Martens &Eck, 1995: 331). These species share reddish eyes and have a similar voice (for sonagrams of dichrous see Martens & Eck, 1995, fig. 103a-i). Accordingly, in the cytochromeb topology they fit as sisters (Gill et al., 2005).
Distribution: from the Himalayas to the Taibai Shan in the Qin Ling range, Shaanxi,

38 Dated 1844 in Snow (1967: 93); for reasons to change see Dickinson & Pittie (2006).
39 Restricted by Biswas (1955: 88).
40 Vaurie (1959) and Snow (1967: 98) made this a synonym of nominate dichrous.
41 See footnote attached to Parus superciliosus. Most probably from Tibet.

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in central China (Cheng, 1987) and north to NE Qinghai in the Qilian Shan (the former
Richthofen Mts.) to about 38°N in N China (Stresemann et al., 1937).
Colour images: Quinn in Harrap, 1996, pl. 20, fig. 69a: kangrae, b: wellsi, c: dichroides.
Parus major Linnaeus, 1758, sensu lato
Introduction: the Great Tit is one of the most discussed species of Old World tits andthis applies to its systematics and taxonomy. This is partly due to its vast range, which
extends from North Africa across temperate Eurasia and into large parts of tropicalSouth East Asia (Portenko & Wunderlich, 1984). This interest is also partly due to theconsiderable differences in colour, in mensural data and in morphology (e.g., tail/wingindex, bill shape) of large population groups. This variety gave rise to numerous different named forms and to different ranking of the groups that seem to be composed ofsimilar forms, rankings that even now differ markedly from author to author. Recently,
substantial bioacoustic and molecular genetic work has been extended to the Great Tit.
Fresh collecting, and field work in perceived contact zones, has helped to clarify sometaxonomic problems.
Presently, four Sektoren 42 (singular Sektor), a term used by one of us (S. E.) in various papers, are accepted. Originally, these Sektoren were distinguished by colour alone(Stegmann, 1931), but later flight feather proportions were examined (Eck, 1977, 1980b,
1988), as well as vocalizations (Gompertz, 1968, Martens, 1994, 1996, Ivankina et al.,1997) and molecular genetics (Kvist et al., 2003, Päckert et al., 2005). Representative
subspecies of all four Sektoren occur in the area we have under consideration, and two
Sektoren are nearly confined to it (Fig. 5).
General characteristics: the major Sektor (group 1): yellow belly, greenish back; theminor Temminck & Schlegel, 1848, Sektor: greyish belly, greenish back; the cinereus
Vieillot, 1818, Sektor: light grey belly, bluish-grey back; the bokharensis Lichtenstein,
1823, Sektor (= group 2 when placed within major): white belly, light grey back. Molecular phylogeny identifies the major and bokharensis groups and the minor and cinereus
Sektoren, respectively, as close relatives (Kvist et al., 2003, Päckert et al., 2005) and in thecase of the major and bokharensis groups we have merged the two into one Sektor (while
retaining them as distinct groups of subspecies).
Formerly, the Great Tit was considered to be the classic example of a ring species(Rensch, 1933; Mayr, 1942: 182; a term still used by Kerimov & Formozov, 1986), but thiswas erroneous (Päckert et al., 2005). In a ring species the ‘end’ members of an uninterrupted and continuous ring of populations meet. In this case it was considered that they
met in the middle Amur valley of eastern Siberia, and they were believed not to hybridize (Stegmann, 1928, 1929, 1931, Nazarenko, 1971). But for a long time the degree of
reproductive isolation was not known.
Nevertheless, most western ornithologists grouped all these Sektoren into one extended, highly polymorphic species, P. major sensu lato, and the Sektoren were treated

42 Sektoren are groups of subspecies and may sometimes themselves comprise two or more identifiablegroups of subspecies.

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Fig. 5. Distribution of the component species of the great tit superspecies (Parus [major]) and their zones
of secondary contact. Different shading indicates the areas of P. major sensu stricto (major and bokharensis
Sektors, P. minor and P. cinereus and of P. monticolus (line only). Zones of contact and (restricted) hybridization are: – I. middle Amur valley, E Siberia (P. major sensu stricto and P. minor); – II. SW Mongolia,
Bulugun valley (major Sektor and bokharensis Sektor of P. major sensu stricto); – III. Kyrgystan and Kazakhstan (man-made sympatry of major Sektor and bokharensis Sektor of P. major sensu stricto), – IV. transCaspian area of south Turkmenistan and north Iran (major Sektor and bokharensis Sektor of P. major sensu
stricto and P. cinereus). – V. South China and parts of Indochina (P. minor and P. cinereus, hybridization
resulting in P. minor ‘commixtus’, full extent of hybridization unknown). – Slightly modified from Päckert et al. (2005).

as subspecies groups, with varying numbers of subspecies (Delacour & Vaurie, 1950).
Vaurie (1957a) differed slightly, splitting off the bokharensis group as an independent
species, P. bokharensis, from the rest of P. major on grounds of being “very distinct morphologically and apparently ‘overlapping’ in some regions [with P. major]… during thebreeding season”. Most subsequent authors (Snow, 1967, Harrap, 1996, Dickinson, 2003)
followed this view, but see Eck (1992; 1996: 47). By contrast Russian authors, adopting
a typological view, treated all four Sektoren as independent species (e.g., Stepanyan,
1990, 2003), but rarely explained their rationale or logic, although Stepanyan (1983,
1988) was an exception.
Recent studies of vocalizations have clarified that at least two of the four Sektoren
have different voice parameters for their territorial song (Martens, 1994, Ivankina et al.,1997, Päckert et al., 2005), and in part these are so different that an alien voice is notunderstood by members of another Sektor – for example minor group voices in the ma-

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33

jor group area (JM, unpublished). But vocalizations, in this case, may not be the mostimportant pre-mating isolating mechanisms preventing hybridisation. Molecular genetic studies have demonstrated quite strong, but geographically differing, sequence
distances between the Sektoren, and without doubt long-independent evolutionary
histories of these Sektoren.
Here we treat the Great Tit as a superspecies, Parus [major], which includes threeallospecies, each of which requires species rank under the Biological Species Concept(Mayr, 1942): Parus [major] major, the Northern Great Tit, including the bokharensis
group, Parus [major] minor, the Eastern Great Tit, and Parus [major] cinereus, the Southern Great Tit. Reasons are given below. We believe that treating these three taxa as allospecies within a superspecies best represents their close relationships and allows theirgeographic replacement to be kept in mind (see Helbig et al., 2002), as do the vernacularnames suggested.
Parus [major] major Linnaeus, 1758 sensu stricto
Taxa included: A: major Sektor (2 relevant subspecies): P. m. kapustini Portenko, 1954 [Sretensk, eastern Siberia]; P. m. bargaensis Yamashina, 1939 [Lamagulusu, Lake Dalainor (now Hulun Nur), Manchuria, China 43]. B: bokharensis Sektor (2 relevant subspecies, plus 2 extralimital ones added forclarity): P. m. bokharensis Lichtenstein, 1823 [Bukhara; ? Synonym panderi Zarudny & Härms, 1913
[Transcaspia] EL; P. m. turkestanicus Zarudny & Loudon, 1905 [Dsungaria, Semiratschja and SyrDarja; implicitly restricted to the Syr-Darya area by Zarudny & Bilkewitsch, 1912 and further restricted to Tashkent by Eck, 1977: 197 44; Synonym ferghanensis Buturlin, 1912 [partem merid.-orient.
Ferghanae (Mts. Alai) “hiemalis” 45]; Synonym meinertzhageni Koelz, 1939 [Balkh, Afghanistan]; P.
m. iliensis Zarudny & Bilkewitsch, 1912 [Djarkent (Semiratschja) = Zharkent, NE Kazakhstan] EL?;P. m. dzungaricus Zarudny & Bilkewitsch, 1912 [Orchu river, Dzungaria 46] 47.

Taxonomy and distribution: P. major sensu stricto covers most of temperate continental
Eurasia and extends into eastern Siberia and northern China (Cheng, 1987). Nominate
major is replaced by the lighter-coloured kapustini just east of Lake Baikal, and thatmeets P. [major] minor in the middle Amur valley, on both the Russian and Chinese sidesof the Amur river.
Yellow-bellied and green-backed subspecies of the major Sektor that should berecognized within Chinese borders are not yet fully clarified. Cheng (1987) indicated

43 Type in YIO, No. 00128; Yamashina No. 19037, adult male, 22.IV.1935; type locality given by Cheng(1987) as Lamuquer Miao, near Hulun Lake, Nei Mongol Aut. Reg.
44 Because Zarudny & Loudon (1905) listed three areas and Zarudny & Bilkevitch (1912) named new
taxa from two of those the effect was to restrict the type locality of turkestanicus to Syr Darya. The restriction by Laubmann (1913) to the Orchu River was thus invalid and Vaurie (1950: 32) followed by
Snow (1967: 110-111) were mistaken in following this, missing the fact that von Jordans (1923: 20-23) had
already pointed out Laubmann’s error.
45 Snow (1967: 110) gave the type locality as “Kurshab, Alay valley, Kyrgyzstan; Ferghana (winter)” andSudilovskaya (1959) reported that the type is from the ‘Gul’cha area, Ferghana basin’.
46 Dzungaria is in the northern part of Xinjiang and is indicated on modern maps by the Junggar Pendi(basin).
47 For which recent authors, as explained in the earlier footnote, wrongly used the name turkestanicus.

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kapustini for north-western Xinjiang Province and a disjunct presence in Nei Mongol.
The first of these populations probably refers to birds representative of Siberian major
which were released by hundreds between 1960 and 1965 in various localities in Kazakhstan, mostly close to Almaty (Dolgushin et al., 1972). These did well, enlarged theirarea considerably, e.g., into Kyrgyzstan (lake Issyk Kul basin) and also into SE Kazakhstan, in the Charyn valley close to the border of Xinjiang (Martens 1996), and theycertainly have reached north-west China. The Nei Mongol population is indeed kapustini extending from the middle Amur into northern China, whose biological characters
are now pretty well known (Päckert et al., 2005).
The type of bargaensis, held by the YIO and said to originate from Barga steppe near
Hailar in Nei Mongol, has been kindly photographed by Hiraoka Takashi and, contrary
to its placement by Cheng (1987: 879) in the synonymy of artatus Thayer & Bangs, 1909,
which is part of the minor Sektor, belongs to the yellowish-greenish major Sektor.
The type localities of kapustini (Sretensk) and bargaensis (Dalai Nor, Barga steppe)are possibly only 350 km apart and Cheng (1987) may be correct that former Lamagusulu Monastery is now Lamuquer Miao, though Lamagusulu is some 100 km north of
Lamuquer. Also, Barga steppe, from which the name bargaensis is derived, now Barag,
is between Hailar and Hulun Nur (C.S. Roselaar pers. comm.). Consequently, kapustiniand bargaensis may be synonyms, bargaensis being the older name. But see comments
on bargaensis by Snow in Vaurie (1957: 42). The type of bargaensis seems unlikely tohave been directly compared with topotypes of kapustini and until then we prefer torecognise both.
In the middle Amur valley, the yellow-bellied major Sektor meets the westernmost
grey-bellied birds of the minor Sektor. The two differ in wing/tail proportions and size
(Eck 1980b, 1988). The territorial songs of the two differ considerably in their elements,
frequency range, and lower and upper frequency, and these differences are believed toact as premating isolating mechanisms, but they apparently do not hold in P. major(Päckert et al., 2005). Mitochondrial genome studies (control region [CR], Kvist et al.,
2003; cytochrome-b, Päckert et al., 2005) showed that the major and minor aggregates are
separated by a considerable genetic distance. In the faster evolving CR gene, this distance amounts to 5.9% (Tamura-Nei); using cytochrome-b the distance is 2.3% (uncorrected). This difference, at least in the case of cytochrome-b, is situated at a level which,in passerine birds, is sometimes used to define species limits, but at the lower end of therange for that indicator (Helbig et al., 1996). It seems important to us to add additional
criteria to the complex of arguments.
Hybrids between the major and minor Sektors were reported by Portenko (1954), but
not found by Nazarenko (1971). Later Eck (1980b) wrote of them again based on material in ZISP collected by I. A. Neufeldt, and Nazarenko et al. (1999) confirmed their occurrence. A detailed study by Päckert et al. (2005) discovered hybrids in a limited area ofcontact and overlap. But hybrids, determined in a collection of newly gathered skins bycolour, flight feather proportions, and cytochrome-b cluster assignment, were quite rare:
only seven of 52 specimens from the area of contact. Kvist et al. (2006) and Kvist &Rytkönen (2006), largely based on the same material as Päckert et al. (2005), found ahigher percentage of hybrids using nuclear microsatellite analyses rather than mitochondrial markers and morphological characters, indicating that hybridisation eventsare under-estimated by a cytochrome-b-analysis alone. A further population study in theEck & Martens. Review of the Aegithalidae etc. Zool. Med. Leiden 80 (2006)

35

Amur contact zone by Fedorov et al. (2006) largely confirmed the results of Päckert et al.
(2005), Kvist at al. (2006) and Kvist & Rytkönen (2006). Thus there is, apparently, a zone ofhybridization in existence but a large-scale swamping effect does not currently occur.
The two groups are also ecologically different; major Sektor tits are common in open
habitat in villages, and minor birds on the outskirts of settlements and in nearby forests.
Major seems to be a more aggressive species; its occurrence in the Amur valley is recent
following human cultivation, as was first implied by Stegmann (1931), and it is still
actively enlarging its range to the east (with single records in Khabarovsk close to the
Pacific coast in 2000, see Päckert et al., 2005).
In the area of contact, major birds are also able to enlarge their vocal repertoire bylearning and adding minor song elements. However, minor aggregate birds apparently
do not learn major Sektor song elements. Consequently, their repertoire remains smaller
and it seems difficult for them to find mates among the major birds and among hybrids
within the zone of contact (Nazarenko et al., 1999). All these individual factors point toisolating mechanisms between major and minor, which prevent a swamping effect in thearea of contact even though hybrids occur in considerable numbers.
Surprisingly, the bokharensis group is closest to the major group with cytochrome-b
distances between them of only 0.8% (Päckert et al., 2005), a value typical of subspecies oreven smaller population-level differentiation. In addition, territorial songs of the two Sektoren are remarkably similar with respect both to frequency range and form of elements(compare Fig. 6a-b with c-d). In a zone of contact in the Bulugun valley deserts of extreme
south-west Mongolia, all specimens collected, about ten, turned out to be hybrids (Eck &Piechocki, 1977; Eck unpublished). In a new zone of secondary contact, caused by introduction, with resultant overlap of major and bokharensis birds in Kazakhstan, all specimens checked were considered to be hybrids (Formozov et al., 1993), yet in other areas(e.g., Charyn valley, SE Kazakhstan) they presently breed side by side apparently without
large-scale mixing (Martens, 1996), and hybridization is not discernible from external
morphology. Only one bokharensis from the Charyn valley carried a major haplotype (Kvist
et al., 2003, Päckert et al., 2005). Thus, formation of mixed pairs between these morphologically very different forms is quite easily explicable and should not be seen as a remarkable occurrence as stated by Chalikova (2001). However the extent of mixing andfurther developments due to introductions of Siberian major birds deserve attention.
The bokharensis Sektor (of P. major sensu stricto) enters our area only marginally, innorthern Afghanistan (turkestanicus), and in northern and central Xinjiang Province ofnorth-west China iliensis and dzungaricus are involved (Delacour & Vaurie, 1950, Cheng,1987) 48. The complicated taxonomy of the bokharensis group was reviewed by Delacour& Vaurie (1950, see their map), Vaurie (1957a), Eck & Piechocki (1977: 131) and Eck (1977,
1992). This group has a range that covers the central Asian desert areas and bridges thegap between the south-west Asian and Siberian parts of the major group within P. major
sensu stricto. At the southern edge of this range, in ‘Transcaspia’, the bokharensis group isin contact with the cinereus Sektor (Fig. 5) (Kerimov & Formozov, 1986). As Vaurie (1957a)

48 Cheng (1987: 880) listed only turkestanicus but had dzungaricus in synonymy, mentioning the Junggar

Basin; he did not mention iliensis, but the type locality, now called Zharkent, is just across the borderfrom Yining and iliensis is very likely to occur in Xinjiang in the valley south of the Borohoro Shan.

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Fig. 6. Territorial song verses of Parus major sensu lato: a-b: Parus minor; c-f: Parus major sensu stricto, twoeach of the major Sektor (c-d) and of the bokharensis Sektor (e-f); g-h: Parus cinereus. - Recording localities:a) China, Yunnan, N of Zhongdian, 2002; b) Thailand, Chiang Mai, Doi Suthep, 1998; c) Kyrgyzstan, Issyk-Kol basin, SW of Kara-Kol, 1993; d) Iran, Mazandaran, Dasht-Nazir, 1978; e-f) Kyrgyzstan, Yarodar,
1993 (one verse each of two different males); g) India, vale of Kashmir, Tangmarg, 1976; h) Nepal, Bhojpur Distr., Arun valley W of Tumlingtar, 2001. - Recordings by J. Martens.

observed, the four subspecies of the bokharensis group are not only morphologically different, they also inhabit ecologically different areas. Though the differences between
these subspecies are only slight, it presently makes sense to recognize them all.
As suggested by Päckert et al. (2005), bokharensis group is here merged with the
major group of former P. major sensu lato to form P. major sensu stricto within a superspecies P. [major].
Colour images: Martens, 1994, fig. 5a: major, fig. c: bokharensis; Quinn in Harrap, 1996,
pl. 28, figs. 86a-e: bokharensis.
Parus [major] minor Temminck & Schlegel, 1848
Taxa included (10): P. minor wladiwostokensis Kleinschmidt, 1913 [Vladivostok]; P. minor dageletensis
Kuroda & Mori, 1920 [Dagelet I. (Ullungdo), Korea]; P. minor minor Temminck & Schlegel, 1848
[Japan = N Kyushu 49 see Morioka et al., 2005: 87] Synonym kagoshimae Taka-Tsukasa, 1919 [S Kyushu] (see Morioka, 2000: 315)]; P. minor artatus Thayer & Bangs, 1909 [Yichang chi/Hubei, China];
P. minor subtibetanus Kleinschmidt & Weigold, 1922b [Tatsienlu (= Kangding), Sichuan, China]; P.
minor tibetanus Hartert, 1905a [Chaksam, Tsangpo valley = Yarlung Zangbo, Tibet, China]; P. minor

49 Restricted by Momiyama (1927: 29).

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37

amamiensis Kleinschmidt, 1922 [Amami-Oshima I. 50]; P. minor okinawae Hartert, 1905a [Okinawa,
northern Riu Kiu Is.]; P. minor nigriloris Hellmayr, 1900b [Ishigaki, southern Riu Kiu Is. = S Nansei
shoto]; P. minor nubicolus Meyer de Schauensee, 1946 [Doi Pa Hom Pok, border between Thailandand southern Shan States].

Taxonomy: there are several places across southern China and into Indochina where
birds occur that show similar characteristics. These are thought to be of hybrid origin
and Snow (1967) grouped them under the senior name: Parus major commixtus Swinhoe,
1868 [Tingchow = Dingzhou Mts., Fokien = Fujian]. The names fohkienensis la Touche,
1923a [Fujian], indochinensis Delacour, 1927 [NE Tonkin], and makii Momiyama, 1927
[Taiwan] are all treated as synonyms, but each needs further consideration.
On the border between north-east China and eastern Siberia P. minor is in contact
with P. major sensu stricto but, as discussed above, hybridisation on the middle Amur islimited. In southern China P. minor is in contact with P. cinereus, and there are several
locations where populations seem to be of hybrid origin (Fig. 5). According to Hartert(1923: 28, footnote), Delacour & Vaurie (1950) and Vaurie (1957a) Parus major “commixtus” can only be referred to hybrids from a contact zone between P. minor and P. cinereus.
The mapped contact points occur within an elongated area from coastal south-east China west, probably, to parts of northern Thailand. But there are no recent investigations
on population structure, extent of hybridization or vocalizations, let alone molecular
genetics. Consequently, the population structure is virtually unknown and their interactions need to be studied.
General characteristics: the extent of the white markings on the tail feathers and ofgreen on the back differs between subspecies.
Compared to nominate minor, wladiwostokensis is longer-winged and relatively
long-tailed. In general, P. minor is longer-winged in the high-altitude areas in the southof its range (the wing in tibetanus is up to 82 mm). Birds from the islands of Japan aresmall and dark, but nigriloris from Ishigaki is a small, blue-backed and remarkably distinct form (T6 51 is white-tipped; wing length 么 69 mm [type], 2 乆乆 62 and 65 mm),
resembling P. cinereus, rather than being greenish-backed as is normal in P. minor. We
recognize P. minor wladiwostokensis as it is longer-winged and has a greater relative taillength than nominate minor from Japan. Eck & Quaisser (2004) placed tschiliensis Kleinschmidt, 1922, in the synonymy of wladiwostokensis, which therefore reaches China.
One specimen from north-western Thailand (nubicolus, Mae Hong Sorn Prov., MTD)
and two from northern Yunnan (subtibetanus, Zhongdian; MTD) represent the same
cytochrome-b haplotype. This cluster is sister to all other specimens investigated from
northern China, the Amur valley in eastern Siberia and Japan (M. Päckert, pers. comm.,
see topology in Päckert et al., 2005). These two groups are separated by distance valuesof between 0.9 and 1.7%. These differences amount to the cytochrome-b distances between the major group and bokharensis group of P. major sensu stricto and to those between P. caeruleus and P. cyanus in the western and eastern Palaearctic (Salzburger et al.,2002a). A complex pattern of what is probably Pleistocene diversification seems likely
within the P. minor Sektor but this needs further study.
50 Contrary to Snow (1967: 108) Amami-Oshima is not strictly a part of the Riu Kiu Islands (Morioka etal., 2005: 24).
51 T6 = the sixth pair of rectrices, thus the outer ones.

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Territorial song of P. minor (Fig. 6e-f) largely differs from song of P. major sensu stricto
by the much broader frequency range and sharp frequency changes within single notesof the song verse (Martens, 1996, Päckert et al., 2005), but there are similarities to thesong of P. cinereus (q.v.).
When compared to the bokharensis group of P. major sensu stricto, P. minor is longertailed, with a relative tail-length (TWI) of up to 95% vs. 90% (Eck, 2006).
Distribution: from the middle Amur valley of Siberia across all China except for thebarren and treeless parts of Qinghai, Tibet, Xinjiang and Nei Mongol and for southern
China where it meets, and allegedly locally hybridizes with P. cinereus giving rise to“commixtus” (see above).
From Myanmar to Vietnam the distribution of the forms of P. minor and P. cinereusinterdigitates and forms a rather complicated mosaic pattern, with various local subspecies involved. Snow (1967) listed nipalensis (of P. cinereus stock) in north-west Myanmar, with tibetanus in the north and nubicolus in the east (both of P. minor stock) and thelargely disjunct form ambiguus Raffles, 1822, (of P. cinereus stock) in coastal Tenasserim.
P. cinereus is also represented in north-east Thailand and neighbouring southern Vietnam by templorum Meyer de Schauensee, 1946. This part of the range of P. cinereus isisolated from that of P. cinereus nipalensis in north-east Myanmar by the interposition of
P. minor nubicolus in north-west Thailand (Meyer de Schauensee, 1946, Vaurie & Delacour, 1950). In Thailand great tits occur only patchily (Lekagul & Round, 1991), and theexact distribution of the subspecies of P. minor and P. cinereus stock is yet not well documented. Lekagul & Round (1991) provided only one description although in addition
to nubicolus, which is of minor stock, there is templorum on the eastern plateau and ambiguus in mangroves in the extreme south, both of cinereus stock.
Colour images: Martens, 1993, fig. 5b: wladiwostokensis, Quinn in Harrap, 1996, pl. 27,
figs. 85f: minor (here given as similar to wladiwostokensis), i: tibetanus, j: nigriloris k: ‘commixtus’.
Parus [major] cinereus Vieillot, 1818
Taxa included (12): P. c. ziaratensis Whistler, 1929 [Ziarat, Baluchistan, Pakistan]; P. c. decolorans Koelz,
1939 [Jalalabad, eastern Afghanistan]; P. c. caschmirensis Hartert, 1905a [Gilgit, N Pakistan]; P. c. nipalenis Hodgson, 1837 52 [Nepal]; P. c. vauriei Ripley, 1950 [Chabua, NE Assam, India]; P. c. stupae
Koelz, 1939 [Sanchi, Bhopal, India]; P. c. mahrattarum Hartert, 1905b [Nuwara Eliya, Sri Lanka]; P. c.
templorum Meyer de Schauensee, 1946 [Wat Pa, Lomsak, “central Siam” = eastern Thailand 53]; P. c.
ambiguus Raffles, 1822 [Sumatra, Indonesia]; P. c. cinereus Vieillot, 1818 [Batavia, now Jakarta, Java,
Indonesia]; P. c. sarawacensis Slater, 1885 [Bungal Hills, Sarawak, Malaysia]; P. c. hainanus Hartert,
1905b [Hainan I., China].

52 Not 1838 as in Snow (1967); this is in Vol. 2, part [1] comprising pages 1-74, within which on p. 52 thereis the date Apr. 15, 1837.
53 Meyer de Schauensee (1946) described Wat Pa as ‘about 75 miles east of Pitsanulok’, and Deignan(1963) placed this, the type locality, in Phetchabun province. Lomsak is a crossroad town where the roadeast from Pitsanulok meets a north-south road on the eastern side of the Dong Phaya Fai range. This
places it on the eastern plateau and Deignan noted it from Ubon which is on the eastern side of the plateau. Snow (1967: 107) mentioned “western and central” Thailand, which does not agree, and included
birds from southern Vietnam.

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Taxonomy: the correct taxonomic rank for the cinereus Sektor of the Parus major complex is rather more difficult to establish. It is separated from the minor Sektor by a notable molecular genetic difference of 2.1 to 2.9% of the cytochrome-b gene (Päckert et al.,2005). These values, in themselves, are not a reason for a split at the species level. However, it is a level of the same magnitude as that between the major and minor Sektoren
between which isolating mechanisms are established (see above).
Vocalizations of the cinereus Sektor (Fig. 6 g-h) are similar to those of the minor Sektor(Martens, 1996, Päckert et al., 2005) and considerably less different – both as evidenced
by sonagrams and in song parameters – than those between the major and minor Sektoren, and while differences exist (Päckert et al., 2005) their meaning in a population
context is unknown. External morphology, especially the flight feather proportions between these Sektoren is also similar. However, we must admit that morphological constraints, population structures (including hybridisation) and vocalizations in the south
Chinese contact zone between the minor Sektor and the cinereus Sektor (where “commixtus” occurs) are little known. Outside our area at the north-western limit of therange of the P. cinereus Sektor, in north-eastern Iran and south-western and southern
Turkmenistan, there is contact with the major group and the bokharensis group of P. major
sensu stricto.
As suggested by Päckert et al. (2005), we give specific rank to the cinereus Sektor, as
well the others, but substantial further research in zones of secondary contact is needed.
See Helbig et al. (2002) for comments on the general taxonomic treatment of populations in areas of contact and (limited) hybridisation.
Distribution: the P. cinereus Sektor occurs across an enormous area from the central
Afghan mountains and the subtropical foothills of the north-western Himalayas
through India and deep into the Indo-Malayan archipelago. Conspecificity of cinereus
from Java, and of the populations of Sumatra and Borneo, with those from the Asian
mainland has not yet been investigated using modern techniques.
Although birds of the major Sektor carry lipochromes those of the bokharensis Sektor
and of the P. cinereus Sektor do not, so that cinereus and bokharensis are somewhat similarin outward appearance, although differing in flight feather proportions. A detailed investigation of the great tits of Kopet Dag, the contact zone of these three taxa, will thus
be particularly challenging and will be extremely interesting. To this area is ascribed“Parus bocharensis var. intermedius“ Zarudny, 1890, [’valleys of Karguy-Sou, Firouse, andGujarmaou rivers, and hills and forests of Soumbar & Chandyr’ = along Kargy-su, Firyuza, Gyaur, Sumbar, and Chandyr rivers, Kopetdag Mts., SW Turkmenistan; extralimital]. It was suggested by Hartert & Steinbacher (1933: 176) that this name had been
given to a hybrid population and this view was accepted by Delacour & Vaurie (1950),
Vaurie (1957b), Mishchenko (1982) and Kerimov & Formozov (1985). However intermedius although said to have a greenish back and whitish underparts (similar to minor
stock) may yet be shown to be best placed in the cinereus Sektor on non-morphologicalgrounds. An understanding of this population will contribute importantly to our overall understanding of the allospecies treated here, as will a better knowledge of the commixtus zone (Fig. 5) and whether populations to the south of that (e.g., hainanus) truly
belong with the cinereus Sektor.
For a fuller overview of western contact zones of Great Tits see Fig. 5 and Formozov
et al. (1993).

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Colour images: Martens, 1993, fig. 5c: nipalensis, Quinn in Harrap, 1996, pl. 28, figs.
85p-q: caschmirensis
Parus monticolus Vigors, 1831
Taxa included (4): P. m. monticolus Vigors, 1831 [Himalayas = Simla54, N India]; P. m. yunnanensis la
Touche, 1922 [Milati = now Zhicun, SE Yunnan, China]; P. m. legendrei Delacour, 1927 [Djiring, Langbiang Mts., S Annam, Vietnam]; P. m. insperatus Swinhoe, 1866 [mountains of S Taiwan, China].

Taxonomy: the Green-backed Tit is smaller than the Great Tit complex (P. major sensulato), with coloration similar to parts of it, but with differences in the pattern of the innervanes of outer tail feathers T4-6 (cf. Eck, 1980b: 388, fig. 3). Coloration increases in intensity from west to east (Vaurie, 1957a). It is a relative short-tailed species, TWI 79.9%, sd
2.38 (n = 9). In the cytochrome-b topology (Päckert et al., 2005) P. monticolus is sister to thewhole P. [major] complex. The presence of lipochromes causing yellow and greenish coloration of belly and back in this complex thus seems to be a plesiomorphic trait, and its
absence in P. cinereus and in the bokharensis group of P. major sensu stricto a secondaryloss.
The highly disjunct legendrei has a conspicuously broad black ventral stripe. Vocal
and molecular data for this form would be particularly interesting.
Distribution: the Himalayas, west and central China; disjunct populations in southern Vietnam and Taiwan.
Colour images: Martens 1993, fig. 5e: monticolus; Quinn in Harrap, 1996, pl. 29, figs.
87a-b: monticolus, c: legendrei
Parus nuchalis Jerdon, 1845
Taxon included (1): Parus nuchalis Jerdon, 1845 [Eastern Ghats, India]. Monotypic.

Taxonomy: the White-naped Tit is endemic to two apparently disjunct parts of India;
locally present in the northwest though scarce, and rarer still in the south. This drycountry species, which is morphologically close to the P. major sensu lato complex and to
P. monticolus, has not been sequenced for molecular genetic study and its voice has notbeen analysed.
It may form a superspecies with P. monticolus (for details see Eck, 1988: 115, footnote).
Recent distributional and ecological information of this little known species were
presented by Hussain et al. (1992), Tiwari & Rahmani (1997), and Lott & Lott (1999).
Colour images: Quinn in Harrap, 1996, pl. 29, figs. 88a-c.

54 Type locality restricted by Baker (1920b). Snow (1967: 111) dated this from the 1923 reprint; in fact

this should be cited from J. Bombay Nat. Hist. Soc. 27 (2): 228-247 (for the date see Pittie, 2003). Notethat Ticehurst & Whistler (1924) restricted the type localities of two other titmice (P. melanolophus and P.
xanthogenys) from the same collection to Simla-Almora.

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41

Parus xanthogenys Vigors, 1831
Taxa included (3): P. x. xanthogenys Vigors, 1831 [Himalayas = Simla-Almora 55, India]; P. x. aplonotus
Blyth, 1847 [Mts. of C India = Chaibasa, Singhbhun, Bengal 56, India]; P. x. travancoreensis (Whistler& Kinnear, 1932) [Mynali 57, S India].

Taxonomy: Rasmussen & Anderton (2005: 529) split the Black-lored Tit based on vocalizations and elevated aplonotus of peninsular India to specific rank, with travancoreensis as a subspecies. The peninsular population is certainly disjunct from nominate
xanthogenys of the western Himalayas, however there is no detailed information on territorial song, except for some sonagrams, and no acoustic playback experiments havebeen carried out yet. In addition no molecular genetic data seem to be available. The
differences in colour in the wing bars of northern and southern populations may be no
more than subspecific variation. For the time being we refrain from adopting this decision, believing more detailed studies are needed to support this split.
Distribution: endemic to India; two disjunct populations, widely separated by theGangetic plain.
Colour images: Quinn in Harrap, 1996, pl. 31, figs. 89a: xanthogenys, b: aplonotus, c-e:
travancoreenis.
Parus spilonotus Bonaparte, 1850
Taxa included (4): P. s. spilonotus Bonaparte, 1850 [Darjeeling, N Bengal, India]; P. s. subviridis Blyth,1855 [Tenasserim, SE Myanmar]; P. s. basileus (Delacour, 1932) [Thateng, Bolovens Plateau, S Laos];P. s. rex David 1874 [Kuatun, Fokien = Guadun, Fujian, E China].

Taxonomy: in the Yellow-cheeked Tit nominate spilonotus meets P. xanthogenys, theBlack-lored Tit, in easternmost Nepal; but their interactions are so far unknown. A
cytochrome-b difference of some 5.1% (Gill et al., 2005) strongly supports their validity as separate species; however the samples sequenced came respectively from “Nepal” and “Vietnam”. These are very distant populations, and if the sample from Vietnam came from the southern part of that country it may relate to a form totally disjunct from the Himalayan range. Further sampling is needed from closer to the contact area in eastern Nepal.
Distribution: mountainous subtropical areas from east Nepal through northern partsof Myanmar and Thailand to Laos, Vietnam and southern China, where perhaps the
population of Fujian/Jiangxi is widely separated from more westerly Chinese birds
now associated with it.
Colour images: Quinn in Harrap, 1996, pl. 31, figs. 90a-b: spilonotus, c-e: rex.

55 Type locality restricted by Ticehurst & Whistler (1924).
56 Type locality restricted by Whistler & Kinnear (1932).
57 Not Mynall as in Snow (1967) and elsewhere. About 20 miles NNE of Trivandrum, in the shadow of

two peaks greatly revered in South Travancore, Agastyamalai (6132 ft.) and Champunjimalai (5621 ft.).

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Parus holsti Seebohm, 1894
Taxon included (1): Parus holsti Seebohm, 1894 [Taiwan]. Monotypic.

Taxonomy: the Yellow Tit is endemic to Taiwan. Monotypic with a peculiar and very
distinctive morphology. The cytochrome-b topology places it as sister to P. spilonotus
and P. xanthogenys (Gill et al., 2005).
Colour images: Quinn in Harrap, 1996, pl. 18, figs. 91a-c.
Parus cyanus Pallas, 1770
Taxa included: A. White-breasted Azure Tits (1, extralimital forms listed for convenience: 3): P. c.
koktalensis (Portenko, 1954) [Koktal, Ili basin, Kazakhstan] EL; P. c. hyperriphaeus (Dementiev &Heptner, 1932) [Kustanai, Turgai region, Kirghiz steppe, Kazakhstan] EL; P. c. yenisseensis Buturlin,
1911 [Konnyi near Krasnoyarsk, C Siberia, Russia] EL?; P. c. tianschanicus (Menzbier, 1884) [typelocality restricted to Tian Shan; synonym apeliotes Meise, 1934, Tschen 21 km SE of Harbin, Heilongjiang, NE China)]; B. Yellow-breasted Azure Tits (2, extralimital forms listed for convenience:
1): P. c. carruthersi Hartert, 1917 [Samarkand, Uzbekistan] EL; P. c. flavipectus Severtsov, 1873[‘Uylgum’, not located, in Fergana basin apud Vaurie (1957a: 15)]; P. c. berezowskii (Pleske, 1893)[Guide, upper Chuan-che = Guide (36°04’N, 101°20’E) on upper Hoang-ho, NE Qinghai, China].

Taxonomy: once acoustic and molecular information was added to the taxonomic
decision process species limits in the ‘Cyanistes’ complex changed substantially. Both in
the west Palaearctic (blue tits) and in the east Palaearctic (azure tits) species limits are
now drawn differently.
Traditionally, the split between Blue Tits (P. caeruleus sensu lato) and Azure Tits (P.
cyanus sensu lato) had been thought to be older than that between the European Blue
Tits (caeruleus group) and the North African/Canary Islands Ultramarine Tit group(based on teneriffae Lesson, 1831) (Martin, 1991). However, apparently the opposite istrue and the “similar” African/Canary Islands and the European Blue Tits are theproduct of an older radiation and specific separation. These two “blue tit” groups, now
P. caeruleus sensu stricto and P. teneriffae, were elevated to independent species by Salzburger et al. (2002a), based on remarkably high cytochrome-b difference values anddistinct vocalisations. The average distance between the African and the European
clade amounted to 4.9%. By contrast, distances were remarkably small (1.6-1.9%) between the European Blue Tits and the subspecies of the Azure Tit that have been analyzed. The value of 4.9% is quite enough to sustain specific rank, although the two BlueTit groups are strictly allopatric (north and south of the Mediterranean) and no zone ofcontact is thought to exist. The 1.6-1.9% value is considerably closer to a subspecies
level of distinction than to species rank. Equally the acoustic differences are more pronounced between the two “blue tit” species than those between the European Blue Tit
and the Azure Tit (Becker et al., 1980; Martens & Schottler, 1991, Martens 1996, Schottler,
1993a, b, 1995).
P. caeruleus sensu stricto and P. cyanus are not strictly allopatric, and hybridisation
occurs frequently; but that differs over time and may be dependent on population size
(Johansen, 1952; Vaurie, 1957a; Meise, 1975; Portenko et al., 1982). Judging from territorial song and cytochrome-b distance values of below 2%, these two groups might beEck & Martens. Review of the Aegithalidae etc. Zool. Med. Leiden 80 (2006)

43

seen as only subspecifically distinct, but no constant area of intergradation seems toexist and due to their different ecological preferences there may be no constant area ofsyntopy or even sympatry. We need to know their ecological requirements more precisely, as well as their behaviour in the area of potential sympatry.
The split between P. caeruleus sensu stricto and P. cyanus seems to be phylogenetically recent; and, in addition, no notable cytochrome-b distance is shown between
white-breasted and yellow-breasted Azure Tits (see group members above), whicheven share the same haplotype cluster (Salzburger et al., 2002a). Premating isolating
mechanisms may exist, but this seems very unlikely. Allegedly, mixed pairs and hybrids have ‘often’ been observed (Beaman, 1994), but the breeding area where theymeet seems to be unknown. Because the species is supposed to be sedentary, one
would expect such hybrids to occur in the Tian Shan at about 76°E, but Beaman gave
no localities. Dolgushin et al. (1972) enumerates localities for P. cyanus west to Almaty
and on the NW shore of Issyk Kul (Toraygur), for P. flavipectus east to SW Zailiskiy
Alatau and upper Naryn River, with an occasional December find at Talgar, just east of
Almaty. Dolgushin does not seem to list hybrids. In summary we cannot identify any
sound reason to split white-breasted and yellow-breasted Azure Tits at the species
level despite earlier suggestions and arrangements (Dolgushin et al., 1972, Stepanyan,
1990; Harrap, 1996, Inskipp et al., 1996, Sibley & Monroe, 1990, Dickinson, 2003) and
we treat them as conspecific.
More information on the disjunct berezowskii from north-western China is especially needed. Within the last 100 years it has not been found within its presumed
minute range.
General characteristics: white-breasted subspecies: crown and back vary, crown between white and lavender-grey, back between light grey and dark grey-blue, and theamount of white in the tail feathers varies (Meise, 1934: 31). So also does the thicknessof the bill, which is remarkably thick in koktalensis. The underparts of young yenisseensis and tianschanicus are light yellow which may be further evidence of a link to flavipectus (Harrap, 1996), but juveniles of most white-breasted parids are pale yellowish
below.
In the yellow-breasted subspecies the amount of white in the tail feathers varies, as
does wing length; comparing flavipectus and carruthersi one finds the latter is shorterwinged. In disjunct berezowskii the crown is light grey, and there is no eye stripe behind
eye, the neck collar is broken, and the belly lacks the median stripe. Wing length: up to
67 mm (very few skins available).
For measurements of all azure tit subspecies and comments on them see Eck(2006).
Distribution: white-breasted forms occur in the central Palaearctic from eastern Europe to eastern Asia, yellow-breasted forms from the Tian Shan to the Pamirs plus awidely disjunct tiny area in north-east Qinghai, China (see map in Portenko et al., 1982;
Mauersberger, 1976).
Colour images: Quinn in Harrap (1996), pl. 30, figs. 93c: hyperrhiphaeus, d-e: tianschanicus, figs. 94a-c: flavipectus, d: carruthersi, e: berezowskii; Pleske (1890), pl. VII, fig. 3
and 4; berezowskii.

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Parus varius Temminck & Schlegel, 1845
Taxa included (8): P. v. varius Temminck & Schlegel, 1845 58 [Japan = Honshu’ 59]; P. v. sunsunpi Kuroda, 1919 [Tanegashima, Osumi Is., S Japan]; P. v. namiyei Kuroda, 1918 [Niishima, Seven Is. of Izu,
Japan] 60; P. v. owstoni Ijima, 1893 [Miyakeshima, Seven Is. of Izu, Japan]; P. v. orii (Kuroda, 1923b)[Minami-Daitojima, Borodino Is. = Daito shoto, Japan]; P. v. amamii (Kuroda, 1922) [Amami Oshima];
P. v. olivaceus (Kuroda, 1923a) [Iriomote I., S Riu Kiu Is. = Nansei shoto, Japan]; P. v. castaneoventris
Gould, 1863 61 [Taiwan, China].

Taxonomy: because the Varied Tit occurs both on the Asian mainland and on theJapanese islands and Taiwan its relatively small and isolated populations show considerable geographic variation. Consequently, numerous, indeed too many, subspecies
were erected (Kuroda, 1927), so that for decades infraspecific nomenclature was rather
unstable. Vaurie (1957a) revised the group and accepted nine subspecies. Snow (1967)
largely agreed, but synonymised yakushimensis with sunsunpi 62. We have retained
Snow’s arrangement.
Gill et al. (2005) found a cytochrome-b distance between nominate varius (based on
one specimen from Korea) and castaneoventris of Taiwan of 6%. This difference needs
validation through molecular sequencing of the Japanese populations from the mainislands south to Iriomote before species limits for P. varius can be seriously reconsidered. However, C. Roselaar informs us that castaneoventris stands well apart in its smaller measurements and remarkably short tail compared to the other geographical representatives of P. varius. See also the comments by Chikara (2002).
General characteristics: the main variable features are the overall colour pattern, theextent of the rufous on the nape and on the upper back, the wing length (longest in
owstoni, 84 mm) and bill length. The light parts of the face may be whitish (in nominate
varius) or dark chestnut (as in owstoni).
Between 1830 and 1966 six hybrids between P. varius and P. montanus are known
to have been found in Japan. The most recent one was intermediate in colour, but invoice was close to varius (Mishima, 1969). Does this perhaps hint at an affiliation of P.
varius to the Poecile group? Gill’s et al. (2005) cytochrome-b phylogeny placed P. varius
close to the basis of Poecile. The possible relationship to the rather similar P. semilarvatus of the Philippines is unresolved because its cytochrome-b has not yet been determined. In morphology it is somewhat similar to P. varius as it has a white frontal
patch. But the Philippine species has a strongly down-curved, and thus very differently shaped, bill.

58 Not 1848 as used by Snow (1967: 117). See Morioka et al. (2005).
59 Type locality restricted by Hartert (1905a: 354).
60 Morioka (2000: 315) observed that this might not justify recognition.
61 Not 1862; see Duncan (1937).
62 A position not taken by Orn. Soc. Japan (2000: 239), but Morioka (op. cit. p. 315), although he had notseen specimens from Tanegashima (sunsunpi), implied that probably both yakushimensis and sunsunpi
should be subsumed in nominate varius. That he did not act on this view in the Checklist itself suggeststhat his material was too scanty to be relied upon.

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Distribution: somewhat scattered. Found in north-eastern China in Liaoning and
Jilin Provinces (Cheng, 1987, Meyer de Schauensee, 1984), Korea, S Kuril Islands,
throughout Japan, Taiwan, and also Sakhalin where it seems likely to have been a vagrant(but is perhaps a sporadic breeder) 63.
Colour images: Quinn in Harrap (1996), pl. 18, figs. 95a-b: varius, c: owstoni, d: castaneoventris.
Parus semilarvatus (Salvadori, 1865)
Taxa included (3): P. s. snowi Parkes, 1971 [N Luzon]; P. s. semilarvatus (Salvadori, 1865) [Mt. Arayat,
Luzon]; P. s. nehrkorni (Blasius, 1890) [Mindanao].

Taxonomy: if the genus Parus is to be satisfactorily divided into genera this species
needs molecular sampling so that we can determine its generic assignment. It was notincluded by Gill et al. (2005).
Colour images: Quinn in Harrap (1996), pl. 22, figs. 96a-c: semilarvatus, d-e: snowi, f:nehrkorni.
Melanochlora sultanea Hodgson, 1837
Taxa included (4): M. s. sultanea (Hodgson, 1837) 64 [Nepal]; M. s. flavocristata (Lafresnaye, 1837) [Sumatra]; M. s. seorsa Bangs, 1924 [Yenping, Fujian, China]; M. s. gayeti Delacour & Jabouille, 1925
[Bana, C Annam = Vietnam].

Taxonomy: these three isolated populations show significant colour differences. For
example, gayeti from Vietnam has a black crest rather than a yellow one.
In the Paridae cytochrome-b topology Melanochlora is placed close to the base of thetree, the only other clade that is sister to all other parids being ‘Cyanistes’ (Gill et al.,2005). Löhrl (1997) examined Melanochlora behaviour in a general parid context andfound many behavioural traits were conspicuously unlike parids. This seemed especially true in its marked ‘panic’ in reaction to unusual noise or unexpected encounterswith other species inhabiting the same pen.
Distribution: from east Nepal to south-east Asia including peninsular Malaysia, withdisjunct and local forms in Sumatra, S China, and Vietnam.
Colour images: Quinn in Harrap (1996), pl. 31, figs. 102a-c: sultanea, d: gayeti.

63 Based on 2 birds observed 21 Sep 1976 at Pugachevka river basin, Sakhalin (Bardin, 1987 apud Harrap,

1996).
64 Dickinson et al. (2006) re-examined the old question of whether the specific name should be sultanea
or flavocristata and showed that as far as can be determined sultanea has a couple of months priority
over flavocristata.

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Sylviparus modestus Burton, 1836
Taxa included (3): S. m. simlaensis Baker, 1917 [Simla]; S. m. modestus Burton, 1836 [Himalayas = Nepal 65]; S. m. klossi Delacour & Jabouille, 1930 [Langbian Mts., S Annam = Vietnam].

Taxonomy: local differences are small. The greenish feathering which camouflages thebird in the canopy of broad-leaved forests, as in the case of Phylloscopus warblers, apparently does not allow much variation. Vaurie (1957a) and Snow (1957, 1967) concluded
that although numerous names are available, only three subspecies should be accepted.
Martens & Eck (1995: 318) agreed adding that “it seems more appropriate merely to callattention to the differences [of local populations] that can exist, rather than finding a newsubspecies for each of them”. However, in view of apparent gaps in the range it will beinteresting to learn about possible molecular differences between populations.
Distribution: Himalayas and many isolated populations in southern China and innorthern parts of tropical SE Asia.
Colour images: Quinn in Harrap (1996), pl. 12, figs. 101a-b: modestus, c: simlaensis.
Pseudopodoces humilis (Hume, 1871)
Taxon included (1): Pseudopodoces humilis (Hume, 1871) [above Kichik-Yailagh at 36º47’N, 78º16’E,
near Sanju Pass, SW Xinjiang, China]. – Monotypic.

Taxonomy: Dickinson et al. (2004a, b) discussed this species within the preliminary
review of the Corvidae and signalled in a footnote the proposal that this species be reassigned. It is now fully apparent that this species of the Tibetan highlands belongs to theparids. This new position is based on morphological and anatomical studies (James etal., 2003), on molecular genetics (James et al., 2003, Gill et al., 2005) and even on biochemical traits of the uropygial glands (Gebauer et al., 2004).
In the cytochrome-b topology Pseudopodoces is not, as one would expect, a sister tothe bulk of ‘true’ Parus, but it is placed as sister to the species around P. major thoughwith low support (Gill et al., 2005), thus in a broad genus Parus it would need to be included there.
Geologically the high plateau of Asia is due to relatively recent uplift, but it would
appear to have been invaded twice by parids. This species, due to its distinctiveness,
is assumed to be the first of the invaders, or did it evolve on the spot despite the orogenic disturbances? P. superciliaris which also occurs at high-altitude and occupies thesame novel niche is thought to be a younger immigrant to the high plateau. See also
Gill et al. (2005).
Colour images: Anderton in Rasmussen & Anderton, 2005: pl. 155 fig. 6 (head only),
James et al. (2003, fig. 1b).

65 Type locality restricted to Nepal by Baker (1920b). Snow (1967: 111) dated this from the 1923 reprint;

in fact this dates from the J. Bombay Nat. Hist. Soc. 27 (2): 228-247.

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47

Abbreviations and acronyms
HI
IZAS
MTD
m.y.
sd
TWI
WTI
YIO
ZFMK
ZISP

Handflügel Index (or wing-tip index)
Institute of Zoology, Academy of Sciences, Beijing.
Staatliche Naturhistorische Sammlungen Dresden, Museum für Tierkunde,Dresden.
million years
standard deviation (omitted when sample too small)
Tail wing index
Wing tip index
Yamashina Institute for Ornithology, Abiko City
Zoologisches Forschungsinstitut und Museum A. Koenig, Bonn
Zoological Institute, Russian Academy of Science, St. Petersburg.
Acknowledgements

Over the decades the extended field expeditions to many parts of Asia by J.M. andhis co-workers were sponsored by Deutscher Akademischer Austauschdienst, Deutsche Forschungsgemeinschaft, Feldbausch-Stiftung and Wagner-Stiftung, the latter two
at Fachbereich Biologie of Mainz University. Vereinigung “Freunde der Universität
Mainz”, Deutsche Ornithologen-Gesellschaft (East Asia grants to J.M., A. Gebauer, and
M. Kaiser), and Gesellschaft für Tropenornithologie provided travel funds. E.C. Dickinson has kindly shared his notes and provided literature excerpts. Y. Red’kin helpfully
provided his views on Far Eastern parid taxonomy. Special thanks go to Hiraoka Takashi from the Yamashina Institute for Ornithology for providing information on thetype specimen of Parus major bargaensis and several digital images of this specimen (onwhich helpful comments were also received from A. Nazarenko), to M. Päckert for information on Yunnan P. minor cytochrome-b haplotypes and for helpful criticism of themanuscript. Mrs K. Rehbinder with her unrivalled skill provided the meticulous drawings of titmice. M. Päckert, D.T. Tietze and T. Töpfer helped with editorial corrections
and searching for references, some of which were obtained by M. LeCroy. Christiane
Quaisser helped with a discussion of the Dresden type catalogue. The help of Aasheesh
Pittie and his uncle allowed the correction to one type locality from ‘Mynall’ to Mynali
and is appreciated. Kees Roselaar refereed an earlier draft of this paper which he carried out with the utmost care and added a number of important references previously
unknown to us. Many and sincere thanks are due to all those mentioned.
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66 The 1923 reprint (240 pp.) was entitled: A Handlist of genera and species of birds of the Indian

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67 We accept 1850 for pp. 1-106 (see Dickinson et al., 2004: 138); the subsequent pages are best dated 1851.

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68 The author is given as “Hauptmann Thom. Conr. von Baldenstein in Graubünden” but see Conti(1947) for reason not to accept the title ‘von Baldenstein’.

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71 Spelled Hachlor in the original and doubtless a typographical error for Hachlov or Hachlow which isthe normal German spelling; Khakhlov in Russian.

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73 This appears as a separate paper following a longer one in Japanese (pp. 313-321) that is differently

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74 Issue date: 26 January 1904.

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Final draft received: 29.vii.2006
Accepted: 03.viii.2006
Edited: E.C. Dickinson

75 Often cited as Messager Ornith.

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Plate I: A series of Aegithalos concinnus from China (talifuensis and/or concinnus) in different plumage
states, from newly fledged juveniles to the adult state. Note the grey crown in the youngest birds (left),changing to chestnut in the older/adult ones (right). – In a. and b. the same series is shown from below
and from the left side. Based on material in the collection of the Chinese Academy of Sciences, Beijing(IZAS). – Photo: J. Martens.

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Plate II: Himalayan representatives of Parus rufonuchalis and Parus rubidiventris. – a) P. rufonuchalis,
Nepal, Dolpo Distr., Phoksumdo Lake (ZMFK 71.1055); – b) P. r. rubidiventris, Nepal, Myagdi Distr.,
Dhorpatan valley (ZMFK 71.1064); – c) P. r. beavani, Nepal, Solukhumbu Distr., Pare (ZMFK 71.61). – P.
r. rubidiventris and P. r. beavani from Nepal belong to the same haplotype cluster. All drawings to scale.
– Originals by K. Rehbinder (first published in Martens & Eck, 1995).

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䊴 Plate III: European, central Asian and Himalayan subspecies of the Coal Tit, Parus ater. – a) ssp. abietum,
Austria (MTD C 52967); – b) ssp. rufipectus, Kazakhstan or Xinjiang/China; east ‘Turkestan’ (ZFMK
GX1f.b.gamma); – c) ssp. melanolophus, ‘normal’ morph, Nepal, Dolpo Distr., Phoksumdo Lake (ZFMK
95.026); – d) ssp. melanolophus, cinnamon-bellied morph, Nepal, Myagdi Distr., Dhorpatan valley; – e)ssp. martensi, holotype, Nepal, Mustang Distr., Chadziou Khola valley (ZMFK 95.024); – f) ssp. aemodius,
Nepal, Solukhumbu Distr., Imja/Phunki Drangka valley (ZFMK 95.023). All drawings to scale. – Original by K. Rehbinder (first published in Martens, 1993 and Martens & Eck, 1995).