/ü/s(1999) 141,22-28
Systematic status of the Black-collared Bulbul
Neolestes torquatus
ROBERT J. DOWSETT', STORRS L. OLSON2*, MICHAEL S. ROY^ & FRANÇOISE DOWSETT-LEMAIRE1
'Rue des Lavandes 12, Ganges F-34190, France
^Department of Vertebrate Zoology, National Museum of Natural History, Smitttsonian Institution,
Washington DC 21560, USA
^Zoological Institute, University of Copenhagen, Department of Population Biology, Universitetsparken 15,
Copenhagen DK-2100, Denmark
The Afro tropical monotypic genus Neolestes has been treated variously as a bulbul
(Pycnonotidae) or a 'shrike' (Malaconotidae, Laniidae or Prionopidae]. Recent field
observations and fresh laboratory material have enabled a multi-disciplinary review
incorporating biology, anatomy and DNA. The DNA analysis suggests that N. torquatus
represents a deep branch of the pycnonotid clade. The plumage oí Neolestes is unlike that
of any bulbul, although the juvenile plumage is more bulbul-like than shrike-like.
Evidence from the syrinx, skull, humérus and carpometatarsus eliminates a relationship
between Neolestes and the shrikes and mainly points to one with the Pycnonotidae.
Neolestes could be the primitive sister-group of the Pycnonotidae, or a member of some
larger group of which the bulbuls are but one manifestation. The vocalizations are not
shrike-like but are reminiscent of those of the African species of Pycnonoîus; other aspects
of its behaviour point in this direction. We believe there is no evidence for considering
Neolestes to be a 'shrike', and recommend treating it as a primitive member of the
Pycnonotidae, although as incertae sedis.
The Black-collared Bulbul Neolestes torquatus was
described by Cabanis [1875} as a monotypic species in
a monotypic genus. He found it difficult to identify the
nearest relatives of Neolestes, which he treated tentatively as an aberrant member of the Malaconotinae.
Gadow (1883] followed suit in placing Neolestes as an
unnumbered genus, at the end of the Malaconotinae,
but with a footnote (p. 103) stating that it 'does not
appear to be a Bush-Shrike, but to be allied to the
Bulbuls or Pycnonotidae', a conclusion based mainly
on bill morphology. Unfortunately the association
of Neolestes with the Malaconotinae in Gadow's
influential catalogue seems to have carried more
weight than his parenthetical opinion, which has
almost always been overlooked.
Shelley (1896], presumably swayed by the undeniable similarities in plumage pattern, went so far as to
include Neolestes in the now-abandoned genus
Pelidnius, which in his treatment included species that
'Corresponding autlior.
Email: Dowsette@aol.conn
© 1999 British Ornithologists' Union
are all now referred to the bush-shrike genus
Telophorus (type-species the Bokmakierie T. zeylonus],
which is now often included in Malaconotus (typespecies Grey-headed Bush Shrike M. blanchoti).
{Pelidnius is now considered a synonym oí Laniarius].
In his continuation of Shelley's work, Sclater (1912)
placed Neolestes in the 'Laniariinae', between
Telophorus and Malaconotus.
The most influential appraisal of the relationships of
Neolestes was that of Chapín (1921), who presented
evidence suggesting that it was in fact an aberrant
bulbul [Pycnonotidae]. He knew the species well in
the field; his later experience did not change his view
(Chapin 1953) but rather prompted him to rule out
completely any idea of a relationship with the
Malaconotinae. This was perhaps rather precipitate,
but has been the practice followed by most subsequent
writers. Sclater [1930] put Neolestes in an enlarged
Laniidae, between Nicator and Lanioturdus, but he
gave it the English name 'Black-collared Bulbul' and
mentioned the views of Chapin [1921] in a footnote.
Bannerman (1936) omitted Neolestes in his treatment
of the Pycnonotidae and thus had to include it with the
Systematic status of ttie Black-collared Bulbut
Laniidae in a later volume, apologetically citing Chapin
in a footnote (Bannerman 1939). Bannerman also
covered all fronts by giving it the English name 'Blackcollared Bulbul-Shrike'! Delacour [1943] questioned
the pycnonotid affinities of Neolestes, while believing it
not to be a shrike. Apart from the fact that Bannerman
[1953] did not trouble to make any changes to his
earlier account when he published his condensed
handbook, in the literature Neolestes has remained a
bulbul ever since, based mainly on the authoritative
position taken by Chapin [1953).
Thus practically throughout its history, the question
has been; 'Is Neolestes a bush-shrike (Malaconotinae)
or a bulbul [Pycnonotidae)?' More recently, Olson
(1990) examined a fluid-preserved specimen and
tentatively suggested placing Neolestes as incertae sedis,
nearest to the helmet-shrikes (Prionopidae). Although
Keith et al. (1992) illustrated this species with the
Pycnonotidae, they accepted the proposal of Olson and
intend treating Neolestes with the shrikes^ in a later
volume. Sibley and Monroe (1990) and Dowsett and
ForbesWatson (1993), however, provisionally retained
Neolestes with the Pycnonotidae, although as incertae
sedis.
Neolestes torquatus is a species of lightly wooded
savannas, occurring in a belt across central Africa,
mainly south of the equator. It is confined to Angola,
Congo, Gabon (of very local distribution), Zaire and
the extreme northwest of Zambia, a distribution that
agrees well with the Guineo-Congolian/Zambezian
transition zone of White (1983). The extreme limits of
its range are from Lubango [ex-Sa da Bandeira) in
Angola, where it is evidently rare [W.R.J. Dean, pers.
comm.), northwards to Tchibanga in Gabon (Malbrant
& Maclatchy 1949) and further north to the Odzala
area in Congo (Dowsett-Lemaire 1997). Eastwards it
occurs to extreme eastern Zaire on Idjwi Island, Lake
Kivu (Prigogine 1967). It ranges southward throughout the savannas of Zaire to the Marungu highlands
(Dowsett & Prigogine 1974). In Zambia, it occurs no
farther south than Mwinilunga district (Benson et al.
1971). Neolestes is common from about sea-level to
1750 m, and occurs in small numbers as high as 2060 m
in the Marungu area (Dowsett & Prigogine 1974).
METHODS
Olson (1990) stressed that more satisfactory skeletal
material was needed for a thorough study; with this
aim R.J.D. and F.D-L. collected a specimen in the
Odzala area of northern Congo (00°36'N,14°54'E), As
well as making available to S.L.O. the whole body
23
preserved in spirit, they also collected tissues for DNA
analysis by M.S.R.
Field studies
R.J.D. and F.D-L. (either singly or together) have studied Neolestes in Congo, Zaire and Zambia; R.J.D. has
handled live examples captured in mist-nets in all
three areas. Tape recordings were made by F.D-L. and
copies are available through C. Chappuis [France) and
the National Sound Archives (London).
Anatomical studies
The specimen was preserved whole in ethanol after
examination in the field, with the exception of some
organs which were extracted for separate DNA
analysis. Upon dissection, S.L.O. preserved the skin in
fluid, as well as the tongue, trachea and syrinx, while
the skeleton was prepared by using trypsin.
S.L.O. studied the syrinx, carpometacarpus, nasal
ossification and the structure of the skull and humérus,
comparing these (together with study skins) directly to
specimens in the National Museum of Natural History,
Smithsonian Institution, Washington DC, where the
specimen is preserved as USNM 620817.
DNA analysis
M.S.R. undertook the analyses at the Department of
Population Biology, University of Copenhagen. For the
analysis, 282 bp of cytochrome b sequence were
compared between Neolestes torquatus and the
following members of several passerine families:
Pycnonotidae
[Eastern
Mountain
Greenbul
Andropadus
nigriceps,
Yellow-streaked
Bulbul
Phyllastrephus flavostriatus and Western Nicator
Nicator chloris), Sylviidae [Green Warbler Phylloscopus
nitidus, genbank accession number 773489, Helbig et
al. 1995), Paridae [Great Tit Parus major, genbank
accession number D38314, K. Chikuni, N. Minaka &
H. Ikenaga unpubl. data, and Plain Titmouse Parus
inomatus, genbank accession number X60944,
Edwards et al. 1991), Prionopidae (White Helmet
Shrike Prionops plutnatus] and Malaconotidae [Bulo
Burti Boubou Laniarius liheratus, Smith et al. 1991),
Corvidae [Jungle Crow Corvus macrorhynchus,
genbank accession number D38313, K. Chikuni et al.
unpublished, and Blue Jay Cyanodtta cristata, genbank
accession number X74258, Cracraft 1993) and
Turdidae (Eye-browed Thrush Turdus obscurus,
genbank accession number D38323, K. Chikuni et al.
) 1999 British Ornithologists' Union, Ibis, 141, 22-26
24
R.J. Dowseíí et al.
unpubl. data). The South American suboscine Baron's
Spinetail Cranioleuca haroni was used as an outgroup
[J. Garcia-Moreno unpubl. data).
Blood or tissue samples of A. nigriceps, P. flavostriatus, P. piumatus, N. chloris and N. torquatus were
washed three times in blood buffer (50 mM Tris pH
7.5, 400 mM NaCl, 100 mM EDTA, 0.5% SDS) and
subjected to standard proteinase K, phenol/chloroform
extractions [Arctander 1988). The primer pair LI4841
and HI 5149 [Kocher et al. 1989) was used to amplify
and sequence 282 base pairs of the cytochrome b gene.
Sequencing of both strands was accomplished by initially amplifying the target sequence with one of the
primer pair biotinylated. Dynabeads (DynalR) were
used to separate the strands of the amplified product
and dideoxy sequencing was carried out as normal
using the non-biotinylated primer.
Sequences were aligned visually using the program
SeqApp (Gilbert 1992). Aligned sequences were
analysed using PAUP [version 3.1 Swofford 1991) in
order to infer phylogenetic relationships; CS3
[Siegismund unpubl. program) was used to compare
substitution patterns.
We follow Dowsett and Forbes-Watson (1993) in
recognizing the Pycnonotidae (bulbuls), Malaconotidae [bush-shrikes), Laniidae (true shrikes) and
Prionopidae (helmet shrikes) as separate families. For
the sake of discussion at the generic level, it is more
convenient here to consider Tehphorus as a genus apart
from Malaconotus (following Olson 1990).
Malaconotidae in central and southern Africa is two
eggs; four is exceptional (R.J. Dowsett & J.F.R.
Colebrook-Robjent, unpubl. data). In the Laniidae,
however, four eggs are more common, while in
Prionopidae a clutch of four is the commonest, with
groups of these co-operatively breeding birds laying up
to seven eggs in one nest. In contrast to the large
parties formed by helmet-shrikes, Neolestes has never
been reported in any more than small groups of up to
four birds (presumably family groups). In this respect
the evidence from Neolestes points to it possibly
belonging to the Pycnonotidae and almost certainly not
to the Prionopidae. The young are born naked (Bowen
1983), which is a character shared by all Pycnonotidae
and at least some Malaconotidae, while the evidence in
Prionopidae is unclear (Markus 1972).
The species has a mixed fruit-insect diet (various
observers), the strong fruit element being typical of
bulbuls and rare or unknown in most Laniidae and
Malaconotidae. A nestling was almost invariably fed
fruit, both adults sharing this duty (Bowen 1983).
For a discreet species usually feeding low in dense
herbaceous vegetation, the song is the best way of
locating the bird, as it is given from the top of a bush
or small tree (F.D-L.). In northern Congo, song was
heard throughout the year, with some seasonal fluctuations. Both songs and call-notes are reminiscent of
those of the African species of Pycnonotus, especially
the Common Bulbul P. barbatus; they are quite unUke
the vocalization of any shrike.
RESULTS AND DISCUSSION
Plumage
Field studies
The nest and eggs have been well described (Chapin
1953 p.l56, Bowen 1983). They are similar to those of
savanna Pycnonotidae in all respects, with the nest
being a frail cup of slender grass and plant stems and
the eggs oval, coloured pinkish white with darker pink
and rufous speckles and shading. However, the nests
and eggs of true shrikes (Laniidae) and bush-shrikes
(Malaconotidae) are also similar to this. Lippens and
Wille (1976) considered the eggs to be very different
from those of other bulbuls, but did not explain
this and a comparative study would be needed to
confirm it.
There are at least four nests for which contents were
known with certainty; three of these contained two
eggs, the other contained two chicks (Bowen 1983,
Chapin 1953, Lippens & Wille 1976, Salvan 1972).
The most common clutch size in Pycnonotidae and
© 1999 British Ornithologists' Union, Ibis, 141, 22-28
Neolestes torquatus is a distinctively and boldly
patterned bird with whitish underparts, a broad
iridescent blue-black band across the breast that is
continuous with a stripe of similar colour extending
through the eye and over the auriculars to the neck.
The crown and nape are grey; the back, wings and tail
are green; and the underwing coverts and bend of the
wings are bright yellow.
Various combinations of such plumage can be found
elsewhere with the Telophorus and Malaconotus bushshrikes, although the whitish underparts are unusual,
occurring only in T. bocagei. This colour pattern is what
led to the former placement of Neolestes with the
shrike-like birds.
Another curious feature is the tuft of stiff, plush,
chestnut feathers behind the ear, which Olson (1990)
noted was similar to the dark tuft of feathers in
helmet-shrikes of the genus Prionops sensu stricto
(although, as noted below, S.L.O. is now convinced
Systematic status of the Black-collared Bulbul
that the differences in cranial osteology rule out any
relationship between Neolestes and Pnonops).
Chapín [1921) remarked that the black eyestripe
passes over the ear-coverts in Neolestes but under them
in those shrikes that have such markings, whereas in
fact in several species of the latter the stripe may pass
through the auriculars. Chapin (1921) thought that
there was only a 'superficial resemblance' between
Neolestes and bush-shrikes, and later he went further
(Chapin 1953) in saying that it 'has no real resemblance to a shrike'. What he failed to stress is that
Neolestes certainly bears no resemblance whatsoever to
any bulbul in plumage pattern. No bulbul has so much
as a black breast band, the closest approach being the
blackish pectoral patches in the Red-whiskered Bulbul
Pycnonotus jocosus, which nearly reach across the
breast. Given that there is nothing in the plumage
pattern alone to suggest a relationship oí Neolestes with
anything other than the Malaconotidae, it is little
wonder that early museum taxonomists placed the
genus among the shrikes.
Filoplumes
One of the characters long associated with the
Pycnonotidae by skin-taxonomists is the presence of
well-developed filoplumes on the nape. This condition, as determined by Olson from examination of the
Smithsonian collections, is indeed widely distributed in
the species attributed to this family. Filoplumes are
present in some species in virtually every genus, with
exceptions including genera of doubtful affinity such as
Nicator. Nevertheless, elongate filoplumes in the nape
could not be found in certain species that otherwise
seem certainly to be correctly identified as bulbuls.
Not only is this character thus not invariably present; it
also occurs in other families. The fact that Neolestes
lacks such filoplumes, as also noted by Chapin (1921),
seems to have little taxonomic significance.
The juvenile plumage (Chapin 1953, pers. obs.] is
like a duller version ofthat of the adult; but the crown
and nape are greenish (like the upper back], not grey
as in adults - and not brownish, as illustrated by Keith
et al. (1992). The black breast band develops early in
the juvenile, but is dull, not glossy. The juvenile wing
differs in the buffy tips to some of the greater coverts.
As mentioned by Chapin, there is no sign of the fine
barring present in the first plumage of Laniidae sensu
lato. The specimen collected was a juvenile as shown
by the pale buff tips remaining on some wing coverts
and by the small fleshy rictal flanges. The skull was
nevertheless well ossified, although the parietals were
largely unpneumatized (untrabeculated).
25
In contrast to such genera of Pycnonotidae as
Phyllastrephus and Bleda, there is no sexual dimorphism in size (but neither is there in such genera as
Pycnonotus). Bowen (1983) summarizes wing lengths
of sexed specimens: male 67-76 mm (mean 72.9 mm,
n = 17), female 68-76 mm (mean 72.7 mm, n= 12).
Morphology
Syrinx
The syrinx in Neolestes was similar to that examined in
a specimen of the Sombre Bulbul Andropadus importunus, but markedly different from the syrinx in the
Helmet Shrike Pnonops plumatus (Olson 1990) or of
the Bush Shrike T. zeylonus [examined in this study).
That of P. plumatus is unusual in that the ventral
musculature is broad and elongate, and is asymmetrical, with fibres from the left side crossing the
midline to the right side of the trachea, which is
possibly twisted. The syrinx in one individual of T.
zeylonus was similar to that of Prionops in having the
elongate ventral musculature, but was not asymmetrical.
Syringeal morphology of bulbuls, shrikes and
possible allies has not been subjected to detailed study,
in contrast, for example, with thrushes and their
possible relatives (Ames 1975). However, it would
seem that theirs is otherwise a very generalized
syringeal morphology that is probably similar to that in
a great many oscines. Although the syrinx neither
confirms nor refutes a pycnonotid relationship for
Neolestes, it does appear to rule out any affinity with
Prionops or the Malaconotidae.
Carpometacarpus
In Neolestes there is a distinct, reasonably welldeveloped dentiform process ('process D'; Pocock
1966) on the leading edge of the major metacarpal
above the tendinal groove, curving over this last. This
character is present in many oscines. In Pocock's
(1966) initial study examining the taxonomic utility of
this character, he noted that the process was absent
in the Laniidae sensu lato (including Malaconotidae),
but present or weakly developed in the Pycnonotidae.
This was confirmed by P.L. Ames (pers. comm.) who
found the process to be absent in 29 species in 11
genera of Laniidae sensu lato, but variable at the
generic and specific level in the Pycnonotidae.
Because the process occurs in many other families of
birds, its presence in Neolestes cannot be taken by itself
to imply relationships with the Pycnonotidae, but it
strongly suggests that the genus does not belong among
the shrikes.
© 1999 British Ornithologists' Union, Ibis, 141, 22-28
26
R.J. Dowsette\a\.
Nasal ossification
Examination of a fluid-preserved specimen oí Neolestes
had shown no ossification of the nostril; however, that
examination was regarded as inconclusive without a
proper skeletal preparation (Olson 1990). In the
present specimen of Neolestes, it can be confirmed that
the posterior margin of the bony nostril was entire,
with no trace of pycnonotid-like ossification of the
posterior margin.
One of the best characters defining the Pycnonotidae is this ossification of the margin of the nostril,
which is present in every genus of the family except a
few whose relationships had been questioned (Olson
19903. This ossification may range from the most
advanced state, in which there is a distinct sheet of
bone covering the posterior third or more of
the nostril that is perforated by neural or nutrient
foramina, to a few^ ragged bony outgrowths along the
margin of the nostril, the rest of the sheet presumably
being cartilaginous.
Thus, what would be the most satisfactory evidence
for a pycnonotid relationship of Neolestes is lacking in
the genus. This is not necessarily proof that Neolestes
is not a bulbul. S.L.O. has examined all available
skeletons of African Pycnonotidae in the USNM
collections.
All of the genera are represented and all of the available species of these genera have the nasal ossification,
except for Nicator (Olson 1990) and for some
individuals of the genus Andropadus (which are often
included in Pycnonotus^. The nostril was strongly
ossified in all 11 individuals of Little Greenbul A.
virens examined and two of Yellow-whiskered
Greenbul A. latirostris. It was less developed in three
Stripe-cheeked Greenbul A. milanjensis and two
A. nigriceps.
In the Cameroon Sombre Greenbul A. cunnrostris,
the condition was well developed in two individuals
and absent in three others; in Slender-billed Greenbul
A. gradlirostris it was slightly developed in one and
absent in four; ossification was absent in two
specimens each of Little Grey Greenbul A. gradlis and
A. importunus.
Whether the absence of this nasal ossification in
these bulbuls is primitive or a secondary loss cannot be
ascertained. What is certain, however, is that Neolestes
also lacks this character, but in plumage it bears no
similarity whatever to the 'sombre bulbuls'.
Skull structure
The skull of Neolestes was compared particularly with
that of Telophorus and also Malaconotus sensu stricto,
© 1999 Brilish Ornithologists' Union, Ibis, 141, 22-28
among the bush-shrikes. Because Chapin (1921) considered Neolestes to be closest to Pyctionotus itself, skull
comparisons were made with African species of that
genus, including Andropadus. On this basis, the overall
similarities of the skull of Neolestes are decidedly with
the bulbuls and not with the shrikes.
As might be expected from external morphology,
the rostrum of Neolestes is not heavy and shrike-like,
but it also differs from that of Pycnonotus in being
shorter, broader and more decurved. As in bulbuls,
there is no nasal septum - a character well developed
in Telophorvs but absent in Malaconotus sensu stricto
The maxillopalatines are long, thin and slightly
expanded at the tips, as in bulbuls, and quite unHke the
short, wide and heavy processes in shrikes. The
palatines are expanded and truncate posteriorly, lacking transpalatine processes, and in lateral view they
descend ventrally below the jugal bar, unlike those of
shrikes. The transpalatine processes may be present or
absent in Pycnonotus sensu lato and when absent the
palatines are truncate, as in Neolestes, though not quite
as expanded. In the shrikes, the transpalatine processes
are much stronger. The postorbital process is vestigial
in Neolestes and Pycnonotus, and well developed in
Malaconotus.
Humérus
All the taxa in question here have a single pneumatic
tricipital fossa of the humérus. However, in Neolestes
and Pycnonotus there is an incipient accessory fossa
partially undercutting the head, whereas this is absent
in Malaconotus. Several other differences between
Pycnonotus and the shrikes were noted in the proximal
end of the humérus; in all of these Neolestes agreed
with the bulbuls, although a larger survey will be
needed to determine the taxonomic distribution of
these characters.
Molecular results
Substitution patterns
Analysing the 12 in-group species across 282 bp of
cytochrome b revealed 1481 transitions (ts) and 1140
transversions (tv), with an average ts/tv ratio of 1.36.
As expected when comparing distant taxa using
cytochrome b, third codon positions were almost saturated with respect to transitions (average ts/tv
ratio = 1.28; Edwards et al. 1991J. For this reason
the phylogenetic analysis disregarded transitions at
third positions as well as first position C-T leucine
transitions (a frequent silent substitution prone to
saturation; P. Arctander pers. comm.).
Systematic status of the Black-collared Bulbul
Phyiogenetic inference
Six most parsimonious trees were generated, which
differed only in the terminal branching order of the
monophyletic clade containing the Laniidae and
Corvidae, All trees placed N. torquatus as a sister taxon
to the Pycnonotidae as represented by A. nigriceps and
P. flavostriatus. This association is also supported when
bootstrap resampling analysis is applied. In addition,
the Corvidae and Laniidae present a monophyletic
clade that is supported over 1000 bootstrap replicates.
This analysis also suggests that Nicator chioris is
not allied to the Malaconotidae as recently suggested
[Keith et al 1992], Instead it groups within a
monophyietic clade together with Pants major and
P. inomatus, although this association is not supported
by bootstrap analysis [Fig. 1). The same overall clade
membership is derived when the outgroup is omitted
and an unrooted phylogeny constructed [data not
shown]. Using this regime, higher bootstrap values for
major clades including the close association between
M torquatus and the pycnonotid clade were produced.
Conclusions
Despite limited data due to the removal of third codon
position transitions, the DNA analysis provided good
support for the grouping of N. torquatus together with
other Pycnonotidae in a monophyletic clade. Within
this clade, Neolestes was found to be basal, although its
exact evolutionary position cannot be ascertained until
a phyiogenetic analysis considers all members of the
Pycnonotidae together with additional outgroups.
In its biology, Neolestes exhibits an impressive
number of differences from the shrikes, and similarities
to the Pycnonotidae. Breeding behaviour [clutch-size
and the absence of co-operative breeding), diet
[frugivory), vocalizations and juvenile plumage all point
to it probably being a bulbul and certainly not a shrike.
The only real divergence is in the plumage of
Neolestes, which is quite unlike that of any bulbul; it
most resembles [at least superficially) the pattern seen
in some of the malaconotine shrikes. Evidence from
the syrinx, skull, humérus and carpometatarsus
provide no further support for a relationship of
Neolestes to the shrikes, however. The same elements
show no important points of departure of Neolestes
from the conditions obtaining in the Pycnonotidae.
Many or all of these similarities may, however, simply
be those of some primitive group of primarily frugivorous oscines and may well occur in other taxa.
Neolestes lacks the nasal ossification of most bulbuls,
which, had it been present, would have been the most
27
Neolestes torquatus
55
Ptïyiiastrephus flavostriatus
79
Andropadus nigriceps
41
Nicator ctiloris
35
Parus major
50
Parus ir\ornatus
Ptiylloscopus nitldus
Laniarius liberatus
Cyanocitta cristata
57
52
• Corvus macrorftynchus
Prionops plumalus
Turdus obscurus
Cranioleuca baroni
Figure 1. One of six most parsimonious trees (PAUP) generated
from the comparison of 232 bp oí cytocfirome b from 13 species
of passerines. Otfier trees differed only in branchiing order of thie
terminal nodes of the corvid/lanlid clade. This tree has a ler>gth
of 143 steps and a consistency index excluding uninformative
characters of 0.490. Bootstrap percentages over 1000 replicates are indicated above branches.
satisfactory evidence of its relationships to the
Pycnonotidae. This absence is probably primitive.
Neolestes could be the primitive sister-group of the
Pycnonotidae or a member of some larger group of
which the bulbuls are but one manifestation.
We recommend, following Dowsett and ForbesWatson [1993], the retention of Neolestes with the
Pycnonotidae, although as incertae sedis.
R.J.D. and F.D-L. undertook their research in Congo while
employed by the ECOFAC project (funded by the European
Union].
REFERENCES
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© 1999 British Ornithologists' Union, Ibis, 141, 22-28
28
R.J. Dowsett et a\.
Bonn. Zool. Beitr. 26: 107-134.
Arctander, P. 1988. Comparative studies of avian DNA by restriction
fragment length polymorphism analysis: convenient procedures
based on blood samples from live birds. J. Ornithol. 129:
205-216.
Bannerman, D.A. 1936-39. The Birds of Tropical West Africa, Vols
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Received 10 February 1997; revisions accepted
7 October 1997