Systematic notes on Asian birds. 44.
A preliminary review of the Corvidae
E.C. Dickinson, S. Eck & J. Martens

Dickinson, E.C., S. Eck & J. Martens. Systematic notes on Asian birds. 44. A preliminary review of theCorvidae.
Zool. Verh. Leiden 350, 26.xi.2004: 85-109.— ISSN 0024-1652/ISBN 90-73239-95-8.
Edward C. Dickinson, c/o The Trust for Oriental Ornithology, Flat 3, Bolsover Court, 19 Bolsover
Road, Eastbourne, East Sussex, BN20 7JG, U.K. (e-mail: asiaorn@tiscali.co.uk).
Siegfried Eck, Staatliche Naturhistorische Sammlungen Dresden, Museum für Tierkunde, A.B. Meyer
Bau, Königsbrücker Landstrasse 159, D-01109 Dresden, Germany. (e-mail: Siegfried.Eck@snsd.smwk.
sachsen.de).
Jochen Martens, Institut für Zoologie, Johannes Gutenberg-Universität, Saarstr. 21, D-55099 Mainz,
Germany. (e-mail: martens@uni-mainz.de).
Keywords: magpies; treepies; jays; nutcrackers; crows; Corvidae; type locality; vocalizations.
Certain Asian species of corvids, mainly those that have been reviewed since 1962, are discussed andattention is drawn to recommended changes in subspecific or specific status. Problem areas remain
and research is suggested, some morphological, some molecular and some into voice. The rationale
behind the ongoing revision of Corvus macrorhynchos is explained, but it is recommended that it beconsidered hypothetical and further developments awaited so that all relevant taxa can be assigned
safely to species.

Introduction
Our preliminary reviews are arranged to coincide with parallel articles on types
and are preparatory to a planned ’Synopsis of the Birds of Asia’ (see Introduction to’Systematic notes on Asian birds’: Dickinson & Dekker, 2000). In Peters’s Check-list ofBirds of the World Blake & Vaurie (1962) dealt with the Corvidae. Prior work by Vaurie(1954a; 1954b; 1955; 1958; 1959) makes detail available on much of the thinking relevant
to Asia. Blake dealt only with New World species. Vaurie’s arrangement in 1962 is
examined and more recent treatments are noted, especially the ’monographs’ of
Goodwin (1976) and Madge & Burn (1993). Goodwin’s study benefitted from his greatinterest in behaviour for he had often previously published about corvids1, sometimes
based on his own aviary studies. We include here revisions from these and othersources that we would expect to be accepted in the planned synopsis. In this paper weretain the sequence of species used by Blake & Vaurie (1962), although Goodwin (1976)
recommended changes and these should be re-examined when sufficient species havebeen sampled for molecular evidence.
The family is large with about 130-140 species attached to about 25 genera, but
relationships within the assemblage called core corvidans by Cracraft et al. (2003) are
not yet well defined. Molecular sampling has been limited and there are conflicts withearlier evidence from DNA hybridization. The family, as accepted by Blake & Vaurie
1 In this paper the term ’corvid’ is used for any member of the family Corvidae.

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Dickinson et al. A preliminary review of the Corvidae. Zool. Verh. Leiden 350 (2004)(1962), is worldwide in distribution, except for the Polar Regions. Generic limits are
not all settled. The affinities of several species remain insufficiently understood andneed focussed field studies to describe their behaviour. Others require serious revision,
for which more acoustic evidence is needed. The recognised behaviour traits of thefamily were listed by Amadon (1944) as ’long-continued courtship feeding; nestbuilding and feeding of young by both male and female; incubation and brooding byfemale only; burying or hiding of food; breaking of food with the bill while the food isheld in the feet’; ’omnivorous and more or less predatory feeding habits; bold andinquisitive nature’. A few Asian species have not been studied to determine how wellthey fit this general pattern, which is not one in which all species share every trait.
Corvids have loud and harsh voices with extensive call repertoires and there are also
accounts of local dialects in crows (Goodwin, 1976) and it is also thought that there areoften sexual differences in voice. Gathering and understanding the acoustic evidence
will not be easy (Laiolo & Rolando, 2003).
Within the genus Corvus, where morphology is relatively less helpful than in mostgenera and size variations within currently recognised species seem to suggest overlumping, Martens & Eck (1995) have provided evidence of the utility of vocalizations.
Indeed this is a more general issue (Martens et al., 2003). Scattered, and still limited,voice samples of Corvus species. gathered by JM from Kashmir to the lower Ussuri
show significant homogeneity and offer considerable hope that evidence will become
available to complement that from other sources.
In their preliminary study of molecular evidence Cibois & Pasquet (1999) had
samples from 15 species of 11 genera and although this dataset could be interpreted as
supporting monophyly, they rightly noted that several key species were missing2. They
mentioned the Asian species Platysmurus leucolophus (Temminck, 1824) as problematic.
Since then James et al. (2003) have shown that Pseudopodoces is not corvine. A further
Asian species that may not be a corvid is Platylophus galericulatus (Cuvier, 1816) as
discussed by Goodwin (1976) and below. Cibois & Pasquet noted that this species
does not have the unspotted juvenile plumage found in corvids.
No new Asian taxon has been named since Blake & Vaurie (1962) except from Asia
Minor.
Nomenclature
One generic name needs comment. Blake & Vaurie (1962) used the generic name
Cissa Boie, 1826, for the green magpies. Some previous authors, including Amadon(1944) and Ripley (1961), used Kitta Temminck, 1826a, but as footnoted by Blake &Vaurie (1962: 242) this is founded on the bird he called Kitta holosericea and depicted in
Pl. Col. 395, a picture not of a corvid, but of what we now call Ptilonorhynchus violaceus(Vieillot, 1816)3, a bowerbird.
The name of one species has needed change (see Corvus pectoralis Lesson, 1831,
below).

2 Jollie (1978) examined the phylogeny of the crows before current methodologies came into use.
3 Not an Asian name and not in our list of references.

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Taxonomy above the species level
We have referred earlier to recommendations made and adopted by Goodwin(1976). Goodwin followed Amadon in treating the colourful Asian ’magpies’ in one
genus. By contrast Vaurie in Blake & Vaurie (1962: 240-244) split the species between
Urocissa and Cissa. In Goodwin’s view the degrees of difference displayed within
Urocissa by U. caerulea Gould, 1863, and U. whiteheadi Ogilvie-Grant, 1899, argued forlumping all members of Urocissa in Cissa. Goodwin (1976: 204), also like Amadon(1944), submerged Dendrocitta and Temnurus in Crypsirina. While we see logic in submerging Temnurus in Crypsirina we are less convinced by the submergence of both.
We are content to leave Vaurie’s generic treatment in place until molecular evidence
can be brought to bear.
As mentioned earlier Pseudopodoces will need to be removed from the Corvidae.
Taxonomy at and below species level
Platylophus galericulatus Cuvier, 1816
This crested tropical forest ’jay’, which is usually found in small noisy flocks (Sheldon et al, 2001), was long treated as a relative of the shrikes (Amadon, 1944). Amadon
accepted the very limited evidence of Bartels & Stresemann (1929) as sufficient for thetransfer of this monotypic genus to the crows. He did not mention that Chasen (1935,
1939), who would have known the bird quite well in the field, did not accept this.
Goodwin (1976: 343) noting the atypical juvenile plumage remarked ’it is not certainthat it is now correctly considered a member of the Corvidae’. Smythies (2000) reported
that Amadon had written about removing the species to the Prionopidae.
The name applied to birds from the Malay Peninsula may need to change. The
type of ardesiacus Bonaparte, 1850, may not be from mainland Asia. It is thought tomatch Sumatran birds, thus making it a synonym of coronatus Raffles, 1822, and thename malaccensis Cabanis, 1866, may have to be used instead.
There is some disagreement whether there are two valid forms in Borneo. Madge& Burn (1993) did not recognize lemprieri Nicholson, 1883. We follow Smythies (2000)
who maintained two Bornean forms and noted an unstable intermediate situation in
Sarawak.
Platysmurus leucopterus Temminck, 1824
Goodwin (1976: 206) thought this species likely to belong close to the genus Crypsirina. Vaurie’s treatment suggests the species is an early or basal corvid and we do not
feel that this view needs correction until molecular studies are available. In behaviour
and in movements both on the ground and in the air its habits are not convincingly
corvine, nor perhaps is the voice (see Smythies, 2000)4.

4 Sheldon et al. (2001) report participation in mixed feeding flocks.

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Garrulus glandarius Linnaeus, 1758
In some areas, especially in Europe, northern forms of the jay are migratory or
irruptive and the specimen record needs careful interpretation when determining thebreeding ranges and whether apparent ’hybridisation’5 is between neighbouringforms or between residents and overstaying irruptive forms. Variation in Europe was
examined by Voous (1953) who detailed the clines perceptible in the colour of theupperparts and underparts. There is also considerable individual variation (Voous,
1953), which may result from interbreeding with irruptive birds. Whatever the casemay be in particular locations, there seems to be general agreement between reviewersthat the entire complex across Eurasia should be treated as one species since the groups
seem to be connected by intermediate populations.
Reviews of all or major parts of this species can be found in Kleinschmidt (1893),
Hartert (1918), Kleiner (1940) reviewed by Stresemann (1940a), Voous (1945, 1953),
Vaurie (1954b), Kuroda (1957), Keve (1973, 1974) and Roselaar (1994). Necessarily
these authors generally focussed on breeding specimens.
In the case of species with such extensive ranges it is not unusual to be able toperceive groups of subspecies sharing common features. For Garrulus glandarius
drawing on Kleiner, Stresemann (1940a) listed eight such groups: glandarius, brandti[sic], bispecularis, leucotis, japonicus, atricapillus, hyrcanus and cervicalis6. Nor is it unusualthat authors disagree and see differing group configurations. Vaurie (1954b: 3-10)
accepted the groups of Kleiner/Stresemann, but dealt with only six groups: glandarius,
cervicalis, atricapillus, brandtii, bispecularis and japonicus. The other two were not
Palaearctic. However, Kuroda (1957), paralleled by Goodwin (1976: 222-223), acceptedjust five groups: glandarius, brandtii, japonicus, bispecularis and leucotis. Kuroda andGoodwin included both the atricapillus and cervicalis groups in their nominate group.
Four of these groups are represented in our region. The brandtii group is associatedby Kuroda (1957) with the Manchurian refugium, the japonicus group with the Japanese
refugium and the bispecularis group with the south China refugium. This third groupextends into the eastern and central Himalayas where it co-exists with G. lanceolatus
Vigors, 1831; but lanceolatus alone then occurs west as far as the Afghan border(Whistler, 1944) and into the oak forests of Nuristan (Paludan, 1959).
The fourth group, leucotis, which is found south of the other groups, has received
less attention. Vaurie in Blake & Vaurie (1962: 233 footnote) wrote ’The populations of
Burma and Assam belong to two types connected by intermediate forms. The puretypes are leucotis (Hume, 1874) which is essentially uniformly black on center andposterior of crown, and white on forecrown, ear coverts, and throat; and persaturatus(Hartert, 1918) which is strongly vinous throughout and not black or streaked oncrown.’ He then listed three forms that he considered to be intermediate between
these types and wrote that he had not seen specimens of oatesi Sharpe, 1896 (from theChin Hills) or from the Lushai and Mishmi Hills and that these required further
5 Better termed intergradation to distinguish between inter-breeding between species.
6 Were these names used as subspecific names the authors and dates would be included according toour usual practice. In the discussion of subspecies groups the addition of such names and dates would
make the text much more difficult to follow.

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study, as did the distribution of the intermediate forms. Ali & Ripley (1948) did not
obtain Garrulus glandarius in the Mishmi Hills and Ripley (1961, 1982) and Ali & Ripley(1972: 201) submerged persaturatus Hartert, 1918, and azureitinctus Koelz, 1951, in interstinctus Hartert, 1918. Abdulali (1980) had just one specimen from the Mishmi Hills andlisted it as interstinctus. We tentatively accept the views of Ripley, but the examinationof fuller series may eventually sustain the views of Hartert (1918) and show that there
are constant differences between the birds north and south of the Brahmaputra.
In Japan Vaurie (1954b) wrote that hiugaensis Momiyama, 1927a, ’required confirmation’ and he later listed it as a subspecies (Vaurie, 1959; Vaurie in Blake & Vaurie,1962), but it was not accepted by the Ornithological Society of Japan (OSJ, 2000: 324),
which we follow.
It is perhaps appropriate to discuss the type locality of japonicus Temminck &Schlegel, 1847. The OSJ (1958) cited the type locality as ’Japan’, not mentioning earlier
restrictions. Blakiston & Pryer (1878: 233) restricted it to ’south Japan to the Strait ofTsugaru’. This is no more than recognition that a quite different form occupies
Hokkaido. The birds of northern Honshu were described and named kakes Momiyama, 1927b, at much the same time as Momiyama named those of southern Kyushu
hiugaensis, but apparently he did not seek to determine whether the type material in
Leiden matched either of these birds. Given that Decima, the small ’island’ linked toNagasaki, where von Siebold was based, is in southern Kyushu and thus must be alikely, but not certain, provenance for the type-specimens, this omission is of some
importance. As it happens the three syntypes now present in Leiden are all labelled’Kioe Sioe’ [= Kyushu]. We consider it likely that these birds were obtained in thegeneral vicinity of Nagasaki, but it has not yet been possible to make comparisons
between the types and specimens known to be from northern and southern Kyushu.
Once this has been done it will be possible to restrict the type locality of Garrulax glandarius japonicus Temminck & Schlegel, 1847, appropriately, placing one or the other ofMomiyama’s names in synonymy. Until then we recommend that G. g. japonicus
should be treated as if it is from the same type locality as hiugaensis Momiyama, 1927.
Pending evidence of this, and only if it is proposed to recognise past splits in japonicus
and to recognise the birds of Honshu or at least northern Honshu we recommend that
future workers would do better to resurrect kakes Momiyama, 1927b, from Iwate Prefecture. This name is available and is much less likely to find itself in synonymy once
the true provenance of G. g. japonicus has been identified.
Garrulus lidthi Bonaparte, 1850
In view of our general introductory comments about a lack of information on
voice in Asian corvid species, and the comments of Goodwin (1976), mention must be
made of information on this species provided by Bruce (1979) on voice and behaviour
when calling and foraging for food.
Urocissa ornata (Wagler, 1829)
Ali & Ripley (1972: 203-212) were not comfortable with the treatment of the morecolourful magpies in Blake & Vaurie (1962) and preferred to place them all in one

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genus (Cissa) in which Delacour (1929) had included U. ornata but not the otherspecies treated in Urocissa by Blake & Vaurie. Nor was Goodwin (1976: 188) who
referred to the group as ’the blue, green and Whitehead’s magpies’. He reasoned that
U. ornata and U. whiteheadi Ogilvie-Grant, 1899, were almost as different from the typespecies (erythrorhyncha Boddaert, 1783) as they were from Cissa and that Urocissa, as
employed by Blake & Vaurie (1962), was probably polyphyletic. He considered but
rejected the idea of treating each of these two in monotypic genera7. As mentioned
earlier we await molecular studies on the group. Goodwin (1976: 189) considered
Urocissa caerulea Gould, 1863, part of a superspecies (erythrorhyncha).
Urocissa flavirostris (Blyth, 1846)
Roberts (1992: 420) has drawn attention to the fact that the key to subspecies in Ali& Ripley (1972: 206) is wrong. The Punjab race does not have the underparts ’almost
pure white’ but has them pale primrose in fresh plumage. Goodwin (1976) noted thatsome birds from Punjab ’have the pale head patch more extensive than usual’ and thatin this they match the type. This seems to be an individual trait. Ali & Ripley (1972: 208)
suggest that cucullata Gould, 1861, has the upperparts and wings bluer than nominate
flavirostris. Further study seems needed to clarify whether this is merely clinal along
the Himalayas and to examine whether there are other constant distinctions.
Cissa hypoleuca Salvadori & Giglioli, 1885
Delacour (1929) treated Cissa hypoleuca as five species and admitted two subspeciesof hypoleuca: the nominate in southern Annam and Cochinchina and chauleti Delacour,
1926, in central Annam. Both these have yellowish underparts while allopatric C. concolor Delacour & Jabouille, 1928, of northern Annam, with green underparts, occurredjust to the north of chauleti. His other three species were insular endemics: C. thalassina(Temminck, 1826b) from Java, C. jefferyi Sharpe, 1888, from Borneo and C. katsumatae
Rothschild, 1903, from Hainan. Of these five, two, jefferyi and katsumatae he consideredhighland birds and two, concolor and hypoleuca he reported from low altitudes. A further
race jini Delacour, 1930a, was described from the Yaoshan Massif as a form of concolor.
Kuroda (1933: 32) united chinensis (Boddaert, 1783) and thalassina and included
minor Cabanis, 1850, believing it to originate from Sumatra alone, as well as jefferyi
from Mt. Kinabalu. It should be noted that Kuroda’s information showed thalassina to
be a montane form. Chasen (1935: 310) followed but added the Bornean lowlands to
the range of minor. Judging from Smythies (2000)8, the two Bornean taxa overlap on
Mt. Kinabalu where minor ranges from 300 m to 1220 m (once 1800 m) in mixed dipterocarp forest and lower montane forest, and jefferyi from 900 m to 2440 m keeping tomontane forest.
7 One generic name is available: Cissopica Delacour, 1927, for whiteheadi, but no generic name has been

proposed with Urocissa ornata as the type species. For this Wolters (1977) used a monotypic subgenus,of Cissa not of Urocissa, but did not name that.
8 Sheldon et al. (2001) give slightly narrower altitudinal ranges suggesting overlap largely confined to
between 900 and 1000 m.

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Delacour (1940) united concolor with hypoleuca. Later, disagreeing with Kuroda
and with Chasen, Delacour (1947) united the Indochinese lowland forms, the Javannominate form and the highland form of Mt. Kinabalu (jefferyi) in a broad species Kitta
thalassina. This is not incompatible with sympatry with chinensis in Borneo andIndochina. As noted by Goodwin (1976) these taxa share a common character in differing from chinensis in the colour of the inner secondaries.
The logic of combining two highly disjunct montane forms from the Greater Sundas
with lowland populations from Indochina is surprising and yet Vaurie in Blake &Vaurie (1962) accepted Delacour’s treatment without apparent demur, as recently has
Smythies (2000). However Goodwin (1976: 190), without discussing the relevance ofaltitudinal ranges (explained here and now admittedly better known), noted that thetwo insular montane forms differed from hypoleuca and felt that the Indochinese birds
should not be combined with them. He treated jefferyi as a race of thalassina. We accept
Goodwin’s view for the moment, but few montane species from Mt. Kinabalu are
represented by subspecies in Java. It is our expectation that careful comparison willshow the Javan and the Bornean montane forms deserve to be treated as separate
species, as suggested by Wolters (1977: 226).
Dendrocitta vagabunda (Latham, 1790)
Blake & Vaurie (1962: 284) added bristoli Paynter, 1961, in their Addenda. They
did not mention the re-restriction by Paynter of the type locality of D. v. pallida (Blyth,1846). The earlier restriction to Simla by Ticehurst (1922) did not fit with the bird
described by Blyth. Blyth’s bird was smaller and could have been a more southern
specimen or a bird from further east in the Himalayas. Instead Ticehurst’s restriction
placed the type locality of pallida within the range of the larger of the two north Indian
forms. Paynter (1961) suspected that Blyth’s specimen was not from the Himalayas atall, but believed that, for the sake of nomenclatural stability (and in the absence of a
type specimen), he should give Blyth the benefit of the doubt. Hence his re-restriction
is to a location in the range of the smaller of the two northern forms. Ripley (1982)
followed Paynter.
Dendrocitta formosae Swinhoe, 1863
Blake & Vaurie (1962) treated occipitalis (Müller, 1836) and cinerascens Sharpe,
1879, as conspecific and separated formosae with a footnote recording the view that it
may be conspecific with them. Deignan (1963) preferred to take the line suggestedby the footnote (although he did not state clearly that he brought in cinerascens).
However, Deignan did not provide any information to rebut the treatment by Blake& Vaurie. Goodwin (1976: 205) considered that occipitalis and cinerascens should betreated as separate species and van Marle & Voous (1988) accepted occipitalis as a
Sumatran endemic species. Eck (1996) recommended a superspecies formosae grouping
this species with occipitalis and leucogastra. Smythies (2000) also treated cinerascens as
an endemic.
As regards India, Biswas (1964) considered that the disjunct population sarkari
Kinnear & Whistler, 1930, of the Eastern Ghats did not merit recognition.

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Dendrocitta frontalis Horsfield, 1840
The treatment of Dendrocitta frontalis as a monotypic species by Blake & Vaurie(1962) lumped the population of NE Tonkin with the Himalayan population. This
species does not occur in northern Thailand (Deignan, 1963). Blake & Vaurie (1962:249) did not list the species for China, but Cheng (1976: 504; 1987: 541) mentioned its
occurrence in westernmost Yunnan on the border with Burma, citing ’Stanford et al.’.
However, he gave no suggestion that the distribution of this taxon extends along thesouthern Yunnan border with Indochina and it seems pretty certain that the Tonkin
population is an isolate. We have not traced a specific rebuttal of the validity of kurodae
Delacour, 1927. It was not placed in synonymy by Delacour (1940), Delacour &Greenway (1941) or Delacour (1951). Ali & Ripley (1972) treated the Indian bird under
the nominate trinomial suggesting they agreed that it was valid. We have made asuperficial comparison of kurodae with the nominate form, but Tring has only three
kurodae, all females. Derek Goodwin left a note with these saying that the race is a’poor’ one, but might just be valid. Without seeing more specimens, including males,
this cannot be confirmed and it is probably better left in synonymy where Blake &Vaurie put it.
Temnurus temnurus (Temminck, 1825)
This species was discovered in west central Thailand in May 1990 (P.C. Round, pers.
comm.)9 where breeding has also been reported (Robson, 2001). We are not aware of
any reported differences in these birds from Indochinese stock. Given the probability
that the Thai population is small it may be best that no specimens are taken to explore
this unless distinctions are observed.
The type locality originally provided was Cochinchine. It was not recorded fromsouthern Annam by Blake & Vaurie (1962). There is no reason to suppose a population
survives there, and indeed the attribution may not have been reliable, but then until
recently no population was known from Thailand.
Pica pica (Linnaeus, 1758)
It is often asked whether the Tibetan population bottanensis Delessert, 1840, is adifferent species. There is indeed a degree of difference, but its does not appear to beat specific level. Goodwin (1976: 11) suggested that the principal differences, relatively
long wings and short tail, might be accounted for by the open habitat in Tibet requiring
it to fly much longer distances than is usual for European magpies. He suggested thatthe short wings of European magpies confer an advantage in terms of manoeuvrability
in dense cover. Recently, Eck (1997) compared measurements of bottanensis with
Mongolian leucoptera Gould, 1862, which have equally long tails, but shorter wings.

9 And included by Robson (2000).

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Pseudopodoces humilis (Hume, 1871)
The distinctness of this genus was discussed by Li & Cheng (1965). James et al.
(2003), who noted two earlier anatomical studies that suggested that this species is nota crow, have recently presented molecular and osteological studies showing that thisis a parid. They recommended the transfer of the genus Pseudopodoces to a position
within the Paridae.
Nucifraga caryocatactes (Linnaeus, 1758)
Hartert (1897) accepted three ’species’ of nutcracker in Eurasia (caryocatactes, multipunctata Gould, 1849, and hemispila Vigors, 1831), and then described japonica as new.
During revisions at the time that subspecies were becoming accepted Hartert (1903)
united these in one species. However, Baker (1922: 67) treated multipunctata as a separate species and suggested that the distribution of this taxon overlapped with N. c.
hemispila ’in many places’. Meinertzhagen (1927) examined the details of his claim andrejected it, as recounted by Vaurie (1954b). Vaurie (1954b) examined the specimen
record and found the Pir Panjal range consistently separated the two populations,
except for one stray individual, and treated the two forms as subspecies.
In Europe the species is highly irruptive and until this was recognised the breeding
distribution of European forms was not understood. Quite apart from the potential for
confusion due to individual variation, which Meinertzhagen suspected, there would befurther difficulties identifying vagrants of different origins due to varying conditionsin the annual cone crop in their home ranges. Irruptive behaviour, incidentally, is not
restricted to Europe. Blake & Vaurie (1962) mention irruptive migrant macrorhynchos
Brehm, 1823, in Korea and northern China, and Morioka in OSJ (2000: 325) reiterated a
1923 record of macrorhynchos from as far south as Kyushu.
Madge & Burn (1993) recently suggested that there may be two species, but nofresh evidence was put forward and Inskipp et al. (1996) did not endorse it. Wolters(1977) agreed with Vaurie’s treatment as, pending fresh information, do we.
Biswas (1950) advocated the subspecific recognition of yunnanensis Ingram, 1910,
but he did not compare it with macella Thayer & Bangs, 1909, in the synonymy ofwhich yunannensis is now found.
Corvus enca (Horsfield, 1821)
Two species of Asian corvid seem to us to be particularly deserving of urgent fieldstudies with the careful collection of acoustic evidence. This is one; the other is Corvus
macrorhynchos Wagler, 1827.
Blake & Vaurie (1962) followed Meinertzhagen (1926) and listed a broad species
taking in the birds of Malaysia (enca, and compilator Richmond, 1903), of Sulawesi
and nearby islands (celebensis Stresemann, 1936, unicolor Rothschild & Hartert, 190010,
and mangoli Vaurie, 1958), the southern Moluccas (violaceus Bonaparte, 1850) and thePhilippines (pusillus Tweeddale, 1878, samarensis Steere, 1890, and sierramadrensis Rand
& Rabor, 1961). Vaurie (1958) gave details of the forms he united, but said little about
why he rolled them all into one species. He considered them all to have very rounded

10 Described in the genus Gazzola.

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wings11 and a relatively short and slightly rounded tail (about half the length of thewing). They have the base of the culmen bare12. Fig. 1 in Vaurie (1958), a photographof the bills in five forms he treated as C. enca, makes clear that Vaurie regarded billlength, and probably depth, as of subspecific not specific value13. Vaurie also rejected
size as a basis for specific separation in this group (p. 6). The taxa assembled are said
to have in common black plumage with a purplish lustre14 (but not very glossy) withwhite or whitish bases to the feathers of the nape, breast and abdomen and to have’somewhat’ elongated throat feathers15, but not true hackles (of the kind seen in otherspecies of Corvus). His table of measurements reveals that his sample sizes weremodest to negligible and so they give no more than a rough idea of relative size. It
should not be supposed that Vaurie neglected the issue of sample size. His otherstudies, especially that on the drongos (Vaurie, 1949), show that he was willing tomeasure large series. The correct conclusion is probably that past collectors simply
neglected the collection of such obvious and apparently similar birds
Vaurie (1958) restated the evidence of a very distinctive voice reported by Heinrich
for the allied species C. typicus (Bonaparte, 1853) that Stresemann (1940b) considered
important enough to warrant generic separation. Apart from this Vaurie made no
mention of voice differences.
More recent information on voice needs to be collected. McGregor (1910) had notedthat in the Philippines the Palawan form (pusillus) and the Samar form (samarensis)
had ’entirely different’ voices. This was repeated by Goodwin (1976: 68) who could findno suggestions of differences in voice between compilator, nominate enca and violaceus.
Smythies (2000: 632-633) provides the fullest comparative information on the voices ofC. enca and C. macrorhynchos.
The Slender-billed Crow is not typically a bird of open country. Chasen (1939)
wrote, of the Malay Peninsula, that it replaced C. macrorhynchos ’in the less open country
and in the inland districts’ and that it was ’a bird of the forest rather than the coasts,
but it does nevertheless visit the mangrove belt where this is backed by forest’. In
Borneo Smythies (1960) said ’I have met with it usually in small flocks along the rivers
and forest edges’, and yet Corvus macrorhynchos was almost unknown. More recently
Smythies (2000) suspected it was now only scarce and was perhaps increasing, andreported that C. enca was adapting to open secondary growth. In Sulawesi, however,C. macrorhynchos is absent and C. enca is more evident. Raven in Riley (1924) notedthat C. enca seemed to prefer coconut groves. Subsequent writers of privately published

11 Oortwijn (1987) added that the outer primary is relatively short.
12 This refers to a lack of the nasal bristles that normally conceal the culmen in the C. coronoides group(Meinertzhagen, 1926), which includes Corvus macrorhynchos. This was the grouping proposed by Stresemann (1916).
13 Of course the species is known as the Slender-billed Crow and the bill is proportionately more slender than that of C. macrorhynchos.
14 Meinertzhagen (1926) mentioned ’a distinct violet tinge on the under-parts, which is invariably
absent in the C. coronoides-group’ (a group including C. macrorhynchos).
15 Meinertzhagen (1926:70) referred to the throat feathers as less lanceolated. It will be rememberedthat full adult crows generally have relatively more lanceolated and elongated throat feathers than
younger birds (Meinertzhagen, 1926: 80).

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reports agree that C. enca is reasonably common around the villages. In the Philippines the birds of Luzon (Dickinson, pers. obs.), Mindoro (Ripley & Rabor, 1958: 54)
and Samar (McGregor, 1910; Rand & Rabor, 1960: 413) are definitely forest birds. In
Palawan, where C. macrorhynchos is reportedly absent16, C. enca pusillus seems to be
widespread.
There have been three reviews since Blake & Vaurie (1962). Goodwin (1976: 68-69),
like Rand & Rabor (1961), rejected the separation of violaceus in Dorst (1947) where thePhilippine forms were associated with it, and retained the treatment of Blake & Vaurie,
except that he perceived a relationship between unicolor and typicus, both having greybases to the feathers not white. Goodwin thus detached unicolor from C. enca andsuggested it be seen as forming a superspecies with typicus. We tentatively accept thistreatment, as did Wolters (1977) and White & Bruce (1986).
Oortwijn (1987), whose material was limited to that in collections in Amsterdam andLeiden and did not permit him to study Philippine populations, had more specimensfrom the Malay Peninsula, Sumatra, Java, Bali and Sulawesi than Vaurie (1958) andconcluded that birds from Sulawesi seemed insufficiently different from the nominate
form to deserve separation. Although Eck (1975) had confirmed that Taliabu birds
should be placed with mangoli and recognised that form (still on limited material),
Oortwijn, with another small sample from the Sula Islands also submerged this race innominate enca. And with ten specimens from Borneo, no doubt from Kalimantan, he
concluded that their affinities lay with compilator. We accept the latter finding as regards
Borneo, but as the Wallacean forms have been retained by White & Bruce (1986: 321)
we prefer to retain them too. Like Oortwijn we keep an open mind on whether violaceus
is a separate species and whether Philippine forms are any part of that. It seems moreprobable to us that if violaceus is accepted as a distinct species then the Philippine
forms will deserve separation into one, possibly two other species.
Amadon (1944: 15) emphasised the ability of the genus Corvus to ’exploit many
secondary habitats produced by human activities’. Rand & Rabor (1961) suggested thatthe absence of enca from parts of the Philippines might be ’due to ecological factors’.
This, taken in conjunction with the loss of forest habitat, makes sense. Equally the
relative absence, at least historically, of macrorhynchos from Borneo, Palawan and
Sulawesi seems likely to have been due to the high proportion of original forest cover
within which only enca could survive. White & Bruce (1986: 25) noted that deforestation
had left much of the Lesser Sundas covered with grassy hills and little forest and it isperhaps unsurprising that only C. macrorhynchos is present.
Corvus corone Linnaeus, 1758
This species, or species complex, is mainly Palaearctic. The debate on whether
this is best treated as one, two or three species continues; see Eck (2001) and Knox etal. (2002).

16 Its absence from Palawan now may or may not be true. Neither Rand’s 1970 MS note that it appeared

to be absent, nor the footnote by McGregor (1910: 722) triggered a cautionary word in Dickinson et al.
(1991) or in Kennedy et al. (2000) as either should have.

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Corvus macrorhynchos Wagler, 1827
The problems here are even more complex than those discussed under Corvus
enca. Here we have a ’species’ in the eyes of virtually all authors from 1930 to 1995
where two populations are now known to overlap in the Himalayas; apparently
secondary contact among populations that evolved in isolation. The taxonomic issues
involved have been particularly apparent since Martens & Eck (1995) detailed the altitudinal ranges of what they labelled Corvus japonensis intermedius Adams, 1859, andCorvus levaillantii Lesson, 1831. This contrasted with the traditional view in Ripley(1961, 1982) and reflected in Grimmett et al. (1998), that a broad species macrorhynchos
is represented by four subspecies of which these are two. But Grimmett et al. well
portrayed the experience of those who have travelled from the Indian lowlands into
the Himalayas and have noted the considerable size difference - the large Himalayan
bird seeming raven like, even to the extent of having a wedge-shaped tail - and thedistinctness of their calls, which are higher pitched in the lowlands. The situation was
known before 1995. Sibley & Monroe (1990) flagged it by treating levaillantii as anallospecies, but as the appropriate placements of all the subspecies in macrorhynchos
was not explained readers either assumed that levaillantii consisted of just that form or
remained confused.
Martens & Eck (1995) noted that intermedius seemed to occur in the range 18504200 m and that levaillantii was found from the lowland up to 2660 m. They provided
sonagrams of numerous vocalizations (their Fig. 116) of ’C. japonensis’ (including someof C. j. intermedius) and of levaillantii (their Fig. 117). The calls of the former werecharacterised as ’harsh and coarse’ and those of the latter as ’brighter and clearer’.
They noted that the difference in voice had already been observed by Biswas (1964)17.
Significantly, they did not associate the name macrorhynchos with either species. The
signal here is that they did not consider macrorhynchos to be conspecific with either of
the two taxa discussed. The only direct comment was to suggest that Vaurie (1954b)
had lumped levaillantii with macrorhynchos ’without particular reason’. But importantly
they pointed to Hartert (1929) and buried in his paper about types in the Tring Museum
one finds a review that does indeed seem to have escaped Vaurie and other post-war
writers.
Hartert (1929) provided a useful historical perspective, drawing on his initialproblems with Australian corvids. He noted the paper by Stresemann (’1914’ = 1916 18)that to him had seemed satisfactory in ’1921 to 1923’, and the critique of this, and of theviews of Mathews19, by Meinertzhagen (1926). He concluded that Meinertzhagen had
posed useful questions, but had been wrong to place the many forms of ’raven’ orcrow listed by Mathews in just two subspecies of one very broad species coronoides
Vigors & Horsfield, 1827, namely in bennetti North, 1901, and nominate coronoides.
Hartert also felt that Meinertzhagen had undervalued the distinction between whitebases to the feathers and grey bases. Hartert thought that there were two species, a

17

Citing it as 1963, which is the volume date (see Pittie, 2003).
Hartert (1929) correctly referred to 1916 at other points in his paper.
19
Various works on Australian birds from 1911 to 1920: for details see Meinertzhagen (1926).
18

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crow and a raven, as Mathews had said, differing in size and that there were ’nointermediates’ and that neither of these two species could include bennetti. The crow
had white feather bases, except, he wrote, in young birds, and the ’raven’ grey; bennetti
also had white feather bases. As far as this went we would now say that Hartert wasright (however, two other ravens with restricted ranges were not then picked out, no
doubt due to his limited material)20. Leaving bennetti aside Hartert placed the crow ina broad species macrorhynchos and the ’raven’ in a restricted species coronoides, thelatter being of no concern to us in the context of Asia.
Hartert then assembled the large crows with white bases to the neck feathers in a
species macrorhynchos, including the races philippinus (Bonaparte, 1853), mengtszensis
La Touche, 1923, orru Bonaparte, 1850 and three other regional forms that for us are
extralimital. Of these forms, all of which Hartert claimed had snowy white bases tothe feathers, mengtszensis is a distant outlier surrounded by populations that Hartert
assigned to C. levaillantii21. In levaillantii he placed japonensis Bonaparte, 1850, mandshuricus [sic]22 Buturlin, 1913, hassi Reichenow, 1907, colonorum Swinhoe, 1864, hainanus
Stresemann, 1916, connectens Stresemann, 1916, and intermedius apparently accepting
the views of Stresemann (1916) and Meinertzhagen (1926) both of whom accepted aseparate monotypic species in Sri Lanka (anthracinus Madarász, 1911).
This treatment came too late to influence Baker (1922). However, Baker (1930)
acknowledged Hartert’s review and removed all Indian forms from coronoides andplaced them in levaillantii (including macrorhynchos despite its prior name). It came toolate for Robinson (1927, 1928) as well23. Change to Robinson’s nomenclature either
escaped the attention of Chasen (1939), although he had used the name macrorhynchos
in 1935, or, more likely, was deliberately left alone to minimise confusion with what
was in the prior volumes by Robinson. But Corvus macrorhynchos was used by Delacour(1930b) who discussed colonorum from Indochina and wrote ’… but the Jungle-Crowsof Asia cannot be divided into two species according to white or grey feather-bases.
There are only races of one and the same bird and also intermediates …’. So while
Baker (1930) tried to reunite everything South Asian as levaillantii, Delacour began toreunite eastern populations, correctly using the older name macrorhynchos.
Having read Hartert (1929), Whistler & Kinnear (1932) wondered whether thejungle crows might not be forms of Corvus corone. They re-examined the arrangementof Indian races and accepted just three forms of coronoides from the sub-continent:

20 A comment based on the treatment by Schodde & Mason (1999). However it should be noted that
the Australian raven that Hartert (1929) called C. macrorhynchos ceciliae Mathews, 1912, was separated
along with orru and other forms by Stresemann (1943).
21 There are also unresolved problems here both as regards to the colour of the bases of the neck feathers
and as to whether the name was applied to ’hybrids’ (see Mayr, 1940) - the reason, presumably, why
there is a ’?’ before this entry in Blake & Vaurie (1962). Vaurie (1954b) had not seen specimens (the ’cotypes’ are at the MCZ, Harvard).
22 Often emended to mandschuricus as it was by Hartert (1929).
23 Robinson used Corvus coronoides macrorhynchos. Rather earlier Kloss (1921) had reviewed the treatmentof Stresemann (1916) and clearly believed that Stresemann, when visiting Malaya, had mistaken the
open country macrorhynchos for C. enca and seemed to suspect that Stresemann had muddled his
specimens too. Kloss set out the evidence available from the Malayan collection.

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intermedius across the west and central Himalayas, culminatus in the peninsula (treating
Sri Lankan birds as this too, sinking the endemic anthracinus), and macrorhynchos inthe east, both high in the Himalayas and down through Assam to the sea. The namelevaillantii disappeared into the synonymy of nominate macrorhynchos.
Mayr (1940) did not agree. In examining collections from northern Burma, andstating that these belonged to an undescribed form24, he assembled enough material
to make some judgements about the birds found from Bengal and Assam south to the
Malay Peninsula, as well as those in the Andamans. His comparative southernmost
population, restricted to Java and islands to the east of it, he called macrorhynchos, thus
removing this name from contention in the sub-continent. His remaining four forms
were from: 1) Bengal and Assam, which has been taken to be levaillantii back out ofsynonymy; 2) a lowland form in Burma and northern Tenasserim, 3) a slightly different
form in peninsular Thailand and the Malay Peninsula, and 4) the Andamans birds
which differed from all the others in having the ’nape white’. In the years that followed
Deignan (1945) considered that northern Thai birds belonged to population 2 (butrather than create a subspecific name employed levaillantii for it) and retained this’label’ when he applied macrorhynchos to form 3 from Prachuab south (Deignan, 1963),
here following the lead of both pre-war authors on Malaya and of Gibson-Hill (1949).
In Indochina, where Delacour’s treatment was repeated (Delacour, 1940) andremained unchanged (Delacour, 1951), the only oddity appears when considering thejuxtaposition between nominate macrorhynchos ’sensu’ Delacour, stretching throughmost of Indochina, and its absence in neighbouring Thailand until around Prachuab in
the north of the peninsula, replaced through northern, eastern and central Thailand bylevaillantii ’sensu’ Deignan (in fact Mayr’s ’population 2’). Presumably this is evidence of
gradual intergradiency between birds of Bengal in the west and those of the whole ofmainland South-East Asia except where the south Chinese form colonorum occupies
north-east Laos and northern Vietnam (Delacour, 1940). When account is taken of thereported extent of individual variation, the irrelevance of first year birds to an evaluation, and the modest and scattered collections of specimens it is not surprising that noreal attempt has been made to examine in more detail the populations listed by Mayr
(1940) and the eastward extension of them into Indochina.
Ali & Ripley (1948) resurrected the name tibetosinensis Kleinschmidt & Weigold(1922), which Meinertzhagen (1926) had put in synonymy, using this for the birds
with ’large raven-like bills’ quite different from those of intermedius, that they foundin the Mishmi Hills and that Mayr (1940) had reported from North Burma and had
considered were unnamed25. Ripley (apparently just he, for the pronoun ’I’ is used)stated that this was pending the direct examination of the type and other specimensof tibetosinensis from SE Tibet. Vaurie (1954b) endorsed this view, but did not actually
examine Himalayan specimens east of Simla and did not mention whether he had
examined the type or not26. Ripley (1961) united Andamans birds27 with those of Bengal
24 ’They are large, are very black and glossy, have dusky bases to napes, and medium long, heavy

bills.’ (Mayr, 1940: 694).
25 Vaurie (1959: 168) says they are glossy whereas intermedius is duller.
26 The holotype of tibetosinensis has been lost (see Dickinson et al., 2004; this issue). There is a paratype
in Budapest with which comparisons could be made.

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under the name levaillantii. We have not attempted to determine whether this treatedAndamans birds correctly, but it would appear to bring together birds with different
basal colours to their neck feathers. Further study is probably warranted.
Although Stresemann (1943) gave the Philippine form specific rank, stating that it
shared with C. m. macrorhynchos brown eyes, and feather bases that were ’schmutziger
weiss’ [= dirtier white], but had a more slender bill and a significantly different wingtail index28, Delacour & Mayr (1946) treated philippinus as a race of macrorhynchos andBlake & Vaurie (1962) and Goodwin (1976) both maintained subspecific treatment. So
too did Dickinson et al. (1991) and Kennedy et al. (2000), but the paper by Stresemann(1943) was unknown to them as was the evidence from parasitic mallophaga (Klockenhoff, 1969 a, b). No one has doubted that the birds of the Lesser Sundas are nominate
macrorhynchos.
To the north-east, in China and Japan, Hartert (1929) had listed one form (mengtszensis) in his species macrorhynchos and six forms in his species levaillantii, two Japanese(japonensis and connectens) and four Chinese (mandschuricus [sic]29, hassi, colonorum andhainanus).
In Japan, in a book that did not include Tsushima or the Ryukyus, Austin & Kuroda (1953) listed C. levaillantii japonensis alone. Two forms named by Momiyama(1927c), one from Honshu and one from Kyushu, were treated as synonyms. Due tothe scope of the book the presence of mandshuricus in Tsushima30, and of the two
smaller forms, connectens and osai Ogawa, 190531, in the Ryukyus was not mentioned,
but these were included by OSJ (1958), where the name levaillantii was still employed,
although this changed to macrorhynchos in later check-lists. Cheng (1958) listed macrorhynchos with four Chinese races: tibetosinensis, mandschuricus [sic], colonorum andhainanus. He did not include intermedius; perhaps Chinese territory did not theninclude its range, but this was added later (Cheng, 1976, 1987), by which time he hadplaced hainanus in the synonymy of colonorum, and had moved hassi from the synonymy
of mandshuricus to that of colonorum, where mengtszensis had figured all along. Cheng,
who probably never saw the types and may not have questioned the colour of thefeather bases, presumably accepted Greenway’s view as given by Mayr (1940).
Vaurie (1954b) said that he had not examined the types of mengtszensis and Vaurie(1959: 168-170) omitted all mention of it. The basis upon which he determined to list it
as a separate form in Blake & Vaurie (1962) seems to be based on his speculation in
1954 that it ’probably represents the end of a cline of decreasing size fromtibetosinensis’. Although it is evident that the syntypes of mengtszensis need to be reexamined, above all to re-examine the colour of the bases to the feathers, we feel that
we should follow Cheng in treating this as a synonym of colonorum. Obviously if the
27 Delacour (1930b) reported examining those in the British Museum (Nat. Hist.) and finding whitebases to the feathers, which led him to submerge andamanensis in nominate macrorhynchos.
28
A wing-tail index of 64.3-64.7 in philippinus versus 55.1-57.2 in nominate macrorhynchos (no othersubspecies cited).
29
Because the spelling is as used here by Hartert but not in the original.
30
Brazil (1991) thought that the population of Tsushima might actually be japonensis. OSJ (2000) treated
it as mandshuricus.
31
Not included in Hartert (1929).

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feather bases differ from those found in colonorum there will need to be further consideration of the suggestion that the name mengtszensis was given to hybrids.
With this background set down it is helpful to sketch Hartert’s arrangement on amap. Several observations can be drawn from it. The first is that levaillantii as Hartert
defined it was a species of south and east Asia, with one disjunct oddity – mengtszensis.
This, of course, would not be odd if it has feather bases that are not white as hethought (but although he may not have seen the types at the MCZ he must be presumed to have seen Yunnanese birds with such colouring). Secondly, if mengtszensis is
put aside, the populations with grey or brownish bases to the neck feathers of adults
evidently meet nominate macrorhynchos. This reaches north to Prachuab (Deignan,
1963) or up into all of mainland South-East Asia, except for northernmost Burma andnorthern Indochina, if, when dealing with Mayr’s intermediate populations, you
adopt Delacour’s nomenclature32. Thirdly, there remains the issue of the Andamans
with conflicting claims about the colour of the basal feathers, Delacour (1930) foundthem white, but Hartert (1929) grouping andamanensis with levaillantii no doubt influenced Ripley (1961, 1982) to place the former name in synonymy.
We must now return to the specific name Corvus japonensis used by Martens & Eck(1995). This is fundamentally a signal, initiated by Wolters (1977)33, that the species isoverlumped, not just because of overlapping forms with great differences in the
Himalayas, but also because there is a case to re-examine over the combination of thesouthern forms with the northern. The oldest names in the broad species are macrorhynchos 1827, levaillantii 1831, culminatus 1832, japonensis 1850, and philippinus, 1853.
The first and last belong with the southern birds, the second and third are clearlyattached to smaller lowland birds of the Indian subcontinent. What is needed is theoldest name for a northern species overlapping with levaillantii and it must include thelarge Himalayan forms and those of China and Japan. So for this assemblage the binomial japonensis was the correct name to use.
The case however remains hypothetical. There are several questions to be answered.
What does acoustic evidence tell us about the various populations that might be contained in such a species? After all as proposed it must include both large highland forms(intermedius, tibetosinensis) and lowland forms (e.g. colonorum), and it may not beappropriate to include the two forms from the Ryukyus which Brazil (1991) describedas ’very small and very small-billed’. Once the north is settled there will remain theneed to reassess the status of andamanensis and of philippinus.
Martens et al. (2000) noted that acoustic evidence ’clearly separates’ levaillantii,
japonensis and splendens in the Himalayas and they proposed a ’complex’ (superspecies)
allowing a four way split, of Corvus macrorhynchos s.l., in which 1) all Himalayan,
32 Logically the distinctions should be clear: white basal feathers in macrorhynchos and grey in levaillantii.

One might have expected this to be discussed by Robinson & Kloss (1924), but this part of their paper
appeared before Meinertzhagen (1926) and Hartert (1929) and the colour of the bases of the feathers is
not mentioned. However, Riley (1938) is also silent on the subject. Sadly he wrote two years before
Mayr raised awareness of the need for study. Tellingly, and as an example of an underlying problem,
according to Riley, Hugh Smith apparently had no crows at all in his 6459 bird skins from Thailand!
33 As explained by Martens et al. (2000) six forms were identified by Wolters (1977) that he consideredappropriate for a hypothetical species C. japonensis.

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Chinese and Japanese birds are united in a species C. japonensis; 2) C. levaillantii
engulfs the lowland forms of the subcontinent, Burma and most of Thailand; 3) C.
macrorhynchos of the Greater and Lesser Sundas34 is seen as reaching further norththan Prachuab and east through south-east Thailand into Indochina, and 4) C. philippinus – a rather more tentative split in their text than in their map (for understandable
reasons), mainly dependant on the striking but incomplete mallophagan evidence of
Klockenhoff (1969a, b) for support. This hypothesis needs refining, with inputs from
specimens, acoustic evidence and molecular studies.
Enough adult specimens from Burma, Thailand and Indochina should be re-examined to determine where macrorhynchos becomes levaillantii. The map in Martens et al.
(2000) is based on standard works whose authors, without recorded close re-examination, chose different conventions to label the populations discussed but not namedby Mayr (1940). It should be noted too that Mayr saw at least his lowland forms asintergrading and he did not envisage that this intergrading ceased where nominate
macrorhynchos was met35 - unless it was south of the Malay Peninsula. If there is intergradiency then it is particularly important that species definitions explain where thisoccurs and why specific range borders are chosen that cut across this. The situation in
Sakhalin is instructive. Here Nechaev (1991) reported widespread overlap between
mandshuricus and japonensis with no apparent hybridisation.
A molecular study by Isawa et al. (2002) provided support for the separation ofjaponensis but the forms in the Ryukyus seem not to belong with japonensis. More of theforms in this whole complex require molecular evaluation.
As yet the amount of acoustic evidence that has been collected is limited. Several
taxa require such study, not least the small birds in the Ryukyus and the Philippine
isolate. Probably Sri Lankan birds should be reassessed too.
Here we defer the break up envisaged in Martens & Eck (1995) and by Martens etal. (2000), although we have suggested that in the accompanying paper on types theraces be sequenced in accordance with the four groups suggested. This deferment is
not because such action is unwarranted. Clearly some action is, and we suggest that
separating the Philippine form may be a relatively simple first step if voice recordings
can be obtained there. This, of course, will be just a step along the way. As regards
mainland Asia there remain too many loose ends. Martens et al. (2000) wrote ’supplementary analyses of their vocalizations may help to reveal additional species delimitations that we suspect exist within the macrorhynchos complex’. It is thus just a matter

34 Not actually absent from Borneo as their map suggests although admittedly still uncommon.
35 Martens et al. (2000) suggested that Deignan (1963) considered that ’two very different’ forms

encounter one another in southern Thailand. If this were so then implicitly the Mekong would represent the boundary between the same two forms around the edge of eastern Thailand and their map
except for cutting corners assumes this. In ECD’s view Deignan simply listed two allopatric subspecies,
with no indication that he saw them as very different and the treatment of Mayr (1940) suggests thatintergradation should be expected and not hybridization. Intergradation would presumably more easily
pass unnoticed against the background of individual variation. The mosaic pattern of mallophaganparasite speciation does tend to suggest that there is some interdigitation or sympatry and of two
distinct species, right up into northern Thailand, but this needs to be supported by voucher specimensof the birds as well as the parasites.

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of time. We anticipate however that from three to seven species should be expected;
the mallophagan evidence suggests more still might be expected.
Corvus pectoralis Gould, 1836
Blake & Vaurie (1962) and Wolters (1977) missed the comment by Kleinschmidt(1940) to the effect that Corvus torquatus Bechstein, 1791, is a prior name to Corvus
torquatus Lesson, 1831, and is not applicable to this species. Eck (1984: 26-27) has
provided a reminder.
Goodwin (1976: 66) suggested that torquatus Lesson [= pectoralis] was probably thegeographical representative of Corvus corone but that some overlap in breeding range
may exist. Maps 412 and 413 in Cheng (1987: 556-557) suggest that the two probably
breed allopatrically.
Corvus ruficollis Lesson, 1831
There are, or have been, mixed views about whether this is a species distinct from
Corvus corax Linnaeus, 1758. Vaurie (1954b) wrote, ’the two are sharply differentiatedmorphologically and ecologically’ and sustained this treatment in later works (Vaurie,
1959; Blake & Vaurie, 1962). Ripley (1961) did not accept this but changed his mind atsome point after 1972 (Ali & Ripley, 1971; Ripley, 1982).
Goodwin (1976) felt that specimens from Quetta and Kandahar might be specimens of subcorax that had been misidentified especially if missexed or possibly evenexamples of interbreeding with C. corax. These may still require reassessment.
Acknowledgements
We are most grateful for the support of the library staff at our various institutions,and especially to the Natural History Museum, Tring and South Kensington for help
with the more obscure literature. Further encouragement and help has been received
from René Dekker. Murray Bruce, Steven Gregory, Alan Peterson and Aasheesh Pittie
have kindly shared bibliographic information. Tony Holcombe generously helped withtranslations. JM greatly appreciates travel funding by the Feldbausch-Stiftung and
Wagner-Stiftung at Fachbereich Biologie of Universität Mainz as well as by Gesellschaft
für Tropenornithologie. Finally we should like to thank Kees Roselaar, our referee,particularly for references to two important papers we had missed.
References
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Ali, S. & S.D. Ripley, 1948. The birds of the Mishmi Hills.— J. Bombay Nat. Hist. Soc. 48(1): 1-37.
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Comp. Zool. 109(4): 277-637.

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36 The dates applicable to the volume are not certainly known, see Zimmer (1926) and Browning &Monroe (1991). Cissa minor (on p. 86) appears to be from December 1850.

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37
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Endre Kleiner in Hungarian, Andreas Kleiner in German. Later changed his name to Keve.
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39 This series is compiled largely from ’unconfirmed reports’ and the compiler adds a caution to that

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40 On sale April, 2000.

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Received: 5.viii.2004
Accepted: 27.viii.2004
Edited: C. van Achterberg