151

Proc. Linn. SOC.Lond., 177, 2, p . 151

With 2 platea and 2 text-figures
Printed i n Qreat Britain
July, 1966

Field characters as a guide to the specific relations of swiftlets

BY LORDMEDWAY,F.L.S.
University of Malaya, Kuala Lumpur
(Accepted for publication December, 1965)

INTRODUCTION

The swiftlets comprise a group of small swifts (Aves, Apodidae) extending in range half-
way round the world, from islands of the western Indian Ocean through southern con-
tinental Asia, the Philippines, and the Indo-Australian archipelago, to north Australia
and the west and south-west Pacific. Characteristically their plumage is dull blackish
brown to glossy black, relieved only by limited areas of w h t e or dull grey. Morphological
differences between species are unobtrusive and, in extreme cases, may even be undetect-
able in the prepared skin. As a result, determination of taxonomic relations among
swiftlets is difficult on the basis of museum skins alone, and the systematics of the group
have, for a long time, been confused.
All swiftlets nest in caves or cave-like situations. Breeding colonies occur on many
oceanic islands, often remote from neighbouring populations. Even on continuous land
masses, breeding caves may be separated by wide stretches of intervening terrain which
lack acceptable sites, and as a result, breeding colonies in continental regions may be as
effectively isolated as those on oceanic islands. Many of these topographically or actually
isolated populations have been described as distinct taxa. I n some cases the specific
relations of the named form are clear, but in others the limited morphological characters
available from specimens in the form of prepared skins and the lack of any clear criteria
may not permit the museum worker to make more than a tentative assignation to species.
Previous reviewers of the group have inferred relationships largely on the basis of
morphological characters, including notably the degree of feathering of the tarsus
(Oberholser, 1906), the furcation of the tail and relative lengths of wing and tail (Strese-
mann, 1932), the size and shape of the bill, the structure of the feathers of crown and
throat, and the presence or absence of white downy tips to the basal barbs of the feathers
ofthe back (Mayr, 1937).Mayr’s review of the swiftlets occurring east of (roughly)Wallace’s
line (Mayr, 1937) demonstrates very well that a satisfactory classification can be derived
from careful evaluation of all such characters. On the other hand, my own attention has
been &awn to the taxonomic value of certain characters of the living bird not normally
available to the museum worker. These are firstly the ability to orientate acoustically by
means of echolocation, and secondly the type of nest built.

Echolocation
The occurrence of acoustic orientation by echolocation among swiftlets was first
demonstrated by Novick (1959)in Ceylon, and myself (Medway, 1959a) in Borneo. When
in flight in darkness or poor light, those species possessing this faculty utter a penetrating
rattle-like call, composed of an irregular succession of brief clicks. This call is invariably
audible to man. Spectrographic analysis of the calls of three species (Medway, 1959a,
and manuscript in preparation) has demonstrated that all component frequencies are in
fact well within the range of normal human hearing.
152 Proceedings of the Linnean Society of London v77,
However not all swiftlets utter such a call, and those which lack it apparently lack also
the ability to orientate acoustically (Novick, 1959; Medway, 1 9 5 9 ~ )They
. nest only in
illuminated regions of caves, whereas echolocating species are able to penetrate to regions
of total darkness. If forced to fly in darkness, in caves, by night, or under laboratory
conditions, they are totally disoriented, and collide immediately and heavily with obstacles
that are easily avoided by species uttering the rattle call (Medway, manuscript in pre-
paration).
Other differences in behaviour are associated with the possession or lack of the power
to orientate by use of the rattle call, and field study has shown that these may have
repercussions on many aspects of the species’ biology (Medway, 1 9 6 2 ~ )Such
. differences
must inevitably increase the degree of genetic separation. Clearly the ability or lack of
ability to utter the rattle call is of taxonomic importance.

Type of Nest
Like most members of this family, swiftlets characteristically build a ‘ bracket-shaped’
nest (Lack, 1956),in the form of a hanging half-cup in which the extraneous nest material
is held together by a more or less copious application of ‘nest-cement ’. This nest-cement
has been shown to be a product of the paired sublingual salivary glands (Marshall t
Folley, 1956; Medway, 1962b). The typical bracket-shaped nest of swiftlets is in fact built
up from a basal crescent (which has been called the ‘hinge’ of the ne8t-e.g. Banks, 1935),
composed of almost pure nest-cement thickly applied to the cave wall, which serves to
support the weight of the rest of the nest proper (referred to below as the ‘nest cup’)
and its contents (Plate 1).
Among the various described forms of swiftlets, however, the structure of the nest and
the nature of its constituent materials are variable. Three main types may be recognized
(cf. Medway, 1959b).These are, firstly, the ‘vegetable’ nest, the bulk of which is composed
of more or less fibrous material, usually but not invariably vegetable in origin, collected
outside the nest site by the building birds, and normally held together by a moderate to
copious application of nest-cement which varies from being firm and dry to the touch to
being soft and permanently moist. Secondly, the self-supporting bracket-shaped nest
constructed exclusively (or almost so) of narrow concentric laminae of firm nest-cement,
without incorporating any significant amount of extraneous material. Thirdly, the self-
supporting bracket-shaped nest composed of firm nest-cement applied in thin concentric
laminae, between which are interspersed a large number of feathers apparently derived
from the plumage of the building birds themselves.
Nests of all categories may be commercially exploited in different parts of South-East
Asia as a source of the raw material of ‘bird‘s-nest soup ’, the chief constituent of which
is of course the edible nest-cement. Nests of the second category, composed predominantly
of the white or translucent nest-cement, intermixed with a few feathers or occasional
strands of vegetable material, are commonly referred to as ‘white’ nests. These require
minimal cleaning and preparation before being cooked, and are commercially by far the
most valuable type of nest throughout the region. Nests in which the nest-cement is
adulterated by the addition of vegetable material or feathers are often referred to col-
lectively as ‘ black’ nests. I n Malaya and Borneo this term is restricted to the type of nest
in which the extraneous matter is solely the feathers of the building birds, blackish brown
in colour (cf. Smythies, 1960).
The greatest variety of types of nest and the widest diversity of sympatric species of
swiftlets are found in the Indo-Malayan region. Here the morphological differencesbetween
species are in some cases very slight, and are consequently overlooked by the rural people
concerned in the commercial exploitation of swiftlets’ nests. Among these people it is
commonly believed that the same speciesof swiftlet may build nests of more than one type.
This misconception has in some instances been perpetuated in both popular and scientific
1965-661 Guide to the specific relations of swiftlets 153
literature. I n fact, it would seem more likely that such divergent types of nest structure,
each inevitably involving totally different patterns of behaviour, should be species
specific. Careful collecting a t the nest sites has shown that this is so, and that radical differ-
ences in nest structure can be correlated with consistent morphological characters
(Medway, 1959 b, 1 9 6 2 ~ )It. is now generally accepted that among the swiftlets, the type of
nest built is a good indicator of taxonomic relations (cf. Sims, 1961).
I n the following pages, the named forms of swiftlets are tentatively grouped in what
appears to be natural order. Assessments of relations are based primarily on morphological
characters, supplemented where available by field observations, firstly of the ability or
otherwise to utter the rattle call, and secondly of the type of nest built. SuEcient material
has not been available to reassess the validity of all subspecies described on morphological
grounds, and in general the judgement of previous authors on this subject has been
accepted. The purpose of this paper is to demonstrate that among swiftlets the field
characters discussed above do in fact provide a sound basis against which the taxonomic
significance of morphological differences can be evaluated, and are therefore useful
indicators of taxonomic relations a t the level of species and above.

SYSTEBIATIC SECTION

Colloculia esculenta (Linn) : White-bellied Swiftlet

Members of this group constitute a well-marked morphological division of the swiftlets,
characterized by small size (winglength not exceeding 107 mm.), and by generally blackish
upperparts strongly glossed with blue or green (in some cases interrupted by a white band
across the rump), and white or whitish edges to the feathers of the abdomen.
Descriptions of the nests of most named forms are summarized below. They are
characteristically of the self-supporting bracket-shaped type, in which a firm translucent
nest-cement is applied in the form of a network of fine threads to bind together the material
of the nest cup, and is copious only a t the hinge. The materials of which the nest cup is
constructed may be very diverse. Strands of bryophyte or lichen, straw or dry blades of
grass, pine needles, casuarina foliage, fibres from palm spathes, the mycelium of ‘horse-
hair ’ fungus, little twigs of angiosperm trees, and aerial rootlets, are among the vegetable
materials commonly utilized. Human hair is one of the more unusual materials that have
been recorded. The selection of the chief constituents appears not to be taxonomically
significant, but to depend partly on the local availability of suitable material, and partly on
individual preferences of each building pair, so that neighbouring nests in a single colony
may be made of entirely different materials (Medway, 1962 c).
The typical race, C.e. .esculentu (Linn.) occurs in Celebes, the Moluccas, Western
Papuan Islands, islands in Geelvink Bay, New Guinea (lowlands), Aru Islands, D’Entres-
casteaux and Louisiade Archipelagoes (Peters, 1940). The structure of its nest has been
discussed in some detail by Stresemann (1914a))who also quotedoriginal details from the
writings of Rumphius. In addition there are descriptions from Buru (Siebers, 1930: 135) ;
and from Indrulamen, southern Celebes : ‘ The nests consisted of moss, rootlets, lichens
and little twigs, agglutinated and fixed to the walls of the cave with saliva, clutch 2, the
eggs measuring 17-16 x 10-11-1 mm.’ (Hartert, 1696a); and from St Aignan, Louisiade
Archipelago : Nests composed of small grasses and fibres, fixed to the rocks with saliva
and loosely held together by the same material, eggs 15.6 x 10.6, 16 x 10.5, 16 x 10.6 mm.’
(Hartert, 1699: 211).
In the American Museum of Natural History, there are several collections of nests of
closely related forms of this group from the general region of the south-west Pacific. These
include nests attributed to C.e. stresemanni Rothschild & Hartert, as follows: two (un-
numbered) from near Kandrian, New Britain, collected by E. T. Gilliard, 21 January, 1959,
which are deep, thick walled, and rounded or cup-shaped rather than truly bracket-
shaped nests, built of vegetable material including bryophyte and ‘horse-hair fungus ’;
154 Proceedings of the Linnean Society of London P77,
no. 14711, from New Britain, collected by W. F. Coultas, 1 August, 1932, a similar deep
somewhat cup-shaped nest, loosely woven from fibrous strands by bryophytes, with nest-
cement conspicuousonly a t the hinge; no. 5342, a cluster of three adjoined bracket-shaped
nests, made of fibrous vegetable material cf. dead grass stems and blades.
There are also one nest of C.e.nu;ckiren.aieMayr, no. 6344, from Barro Islands, Solomons,
collected by R. R. Beck & F. P. Drowne, 4 April, 1927, which is a deep cup of grass,
apparently attached to a dead tree by the characteristic nest-cement, and a number of
nests of C.e. uropygialis Gray from the New Hebrides. The latter vary from the true
bracket-shaped type, constructed of vegetable material with very little nest-cement in
evidence (e.g. no. 5341, from Aurora Island, collected by R. H. Beck, 22 January, 1927),
to more or less rounded nests, again constructed almost exclusively of strands of moss or
blades and stems of grass (e.g. 14733, from Inakahi, E. Tanna, collected 16 October, 1935,
by L. M a om an ).
From elsewhere south and east of the range of typical esculenta, the only named forms
placed in the esculentu group on morphological grounds for which no details are known of
the construction ofthenest are C.e. erwini Collin & Hartert, occurring in the high mountains
of southern New Guinea, C.e. becki Mayr, from the northern and central Solomon Islands,
and C.e. desiderata Mayr, from Rennel Island (distributions from Peters, 1940).
From the South West Islands, nests of C.e.perneglecta Mayr have been described (under
C,e. neglects) by Hartert (1904: 83): ‘Two nests from Wetter are entirely composed of
a kind of olive greyish lichen, while one from Roma consists of black hair-like palm fibre
with a few pieces of yellowish green lichen’. From the eastern Lesser Sunda Islands, nests
of C.e.neglectu Gray have been described from Savu and from Alor. From Savu (Hartert,
1897 : 268) : ‘The nests . . . are built of grass and fibre, with a few feathers and fragments of
dry leaves, just held together by a small amount of saliva’; from Alor (Hartert, 1898:
460) : ‘ . . . composed of rootlets, fibres and lichens, held together by sometimes very little,
sometimes a great amount of saliva, by which they are also fixed to the rocks . . .’, eggs
17 x 11 and 16-6x 11.1 mm.
I n the Museum Zoologicum, Bogor, Indonesia, there are three nests taken by P. F.
Frank, 1 May, 1925, from K. Mao Maroe (=Maru), Sumba. These are attributed to
‘ Collocalia linchii’ i.e. C. esculentu group, and although not associated with skins may be
referred to C.e. sumbawae. These nests are uncharacteristic in being thick-walled and
rounded. It is evident that they were originally supported to a great extent by a sloping
rock face. The nest-cement had dried and deteriorated a t the time I examined them (in
1961), but had apparently been only sparsely applied a t the time of building. The shape
of the nests may be related to the vegetable material utilized in their construction, which
was not strand-like but consisted exclusively of the pappaceous seeds of a grass, kindly
identified by staff of the Herbarium, Bogor, as Penisetum sp.
The representative of the esculenta group on the western-most Lesser Sunda Islands,
Lombok and Bali, and on all Java, the Kangean Islands and Bawean is linchi Horsfield &
Moore. There are several published descriptions of nests from different parts of its range,
including detailed observations by Spennemann (1928), Hoogerwerf (1949) and Medway
(1962~) in Java, and Hartert (1896b) on specimens from the island of Lombok.
With the exception of the mountains of northern Borneo and apparently also of northern
Sumatra, where the esculenta group is represented by dodgei Richmond (the nest of which
is not known), in the lowlands of Borneo, the Natuna Islands, the Riau-Lingga archipelago,
Malaya and (?) eastern Sumatra, the species is represented by cyanopttila Oberholser.
Bracket-shaped nests of the type characteristic of the esculenta group have been described
in detail from lowland Borneo (Medway, 19593, 1962a; Smythies, 1960), and Malaya
(Gibson-Hill, 1948).
On Christmas Ialand, Indian Ocean, the esculenta group is represented by C.e. natalis
Lister. Gibson-Hill(l947: 150-152) has described nests in limestone caves built of lichen
and fibres of sago palm; the fibres formed a ‘sagging sling’4.e. of typical bracket-shape.
1965-661 Guide to the speci$c relations of swiftlets 155
Northwards from Malaya, on continental Asia cyanuptila is replaced by C.e.elachyptera
Oberholser, which builds nests of the characteristic type (Smythies, 1953) and on the
Andaman and Nicobar Islands by C.e. afinis Beavan. Hume very explicitly described
typical bracket-shaped nests of afinis, from Jolly Boy, Macpherson’s Straits, Andaman
Is., built of strands of vegetable material ‘firmly agglutinated with saliva; only along the
line of junction with the place of attachment is there a thickish film of unmixed in-
spissated saliva . . .’ (Oates, 1890: 34, under the name Collocalia linchi). I myself have
seen nests of afinis collected on South Andaman Island together with the associated birds
and kindly made available for examination by M i Humayun Abdulali (specimensnow in the
collection of the Bombay Natural History Society). They are of the characteristic bracket-
shape, consisting of fibrous strands of moss and other bryophytes, bound together with a
firm translucent nest-cement, copiously applied only a t the hinge.
I n the Philippines C.e. isonata Oberholser is found on Luzon, Mindora, Mindanao and
Bongao (Peters, 1940) ; from the alpine summits of Mt Apo, Mindanao, a distinct form C.e.
bagobo Hachisuka has been described. The nest of the latter is not known, but Delacour &
Mayr (1946) have described the nest of isonata as the typical bracket-shaped type. A very
aberrant structure is described by McGregor (1909): ‘Nest composed of lichens and placed
on the ground in the protection of grass or ferns’. It is very uncharacteristic for a swiftlet
to build a nest on the ground, and it is unlikely that this type of nest is typical of isonata.
None of the subspecies of C . esculenta, as listed above, is known to utter the rattle call.
Both Bornean and Malayan populations of C.e. cyanoptila fly silently, without uttering
the rattle call under any circumstances, and are apparently unable to orientate in darkness
(Medway, 1959a; personal observation in Malaya). I n Malaya, we have recently operated
a Holdgate Ultrasonic Receiver ( M k . Iv) in a nesting colony of C.e. cyanoptila, and have
verified that there are no ultrasonic components to any calls uttered in flight within the
range 20-160 kc./s. (Cranbrook & Medway, 1965). It has aho been found that Javanese
representatives of C.e.linchi not only always fly silently, but also when in darkness cannot
detect obstacles in their flight path which are successfully avoided by species uttering the
rattle call (Medway, manuscript in preparation).

Collocalia marginah Salvadori

C . marginata is sympatric with C . esculenta isonota over much of its range, and must
therefore be recognized as a distinct species. It is none the less clearly related, linked by
common morphological features, an identical type of nest, and the inability to utter the
rattle call.
The nest has been described by McGregor (1909) as ‘composed of blackish brown hair-
moss, and held together by the characteristic glutinous saliva’, with a clutch of two eggs,
and by Manuel (1937 : 383) as largely consisting of ‘ dark mossy materials held together
by a scanty gelatinous substance’.
Novick has recorded that while in Luzon, Philippine Islands, he found many colonies
of a swiftlet which did not utter the rattle call, and was apparently unable to enter the
total darkness of true caves (Novick, 1959: 500-501). I n his report, the species is not
identified, However, a specimen was deposited a t the Museum of Comparative Zoology,
Harvard, where i t has been identified by R. A. Paynter as Collocalia marginatu (Mayr,
1961).
Collocalia troglodytes Gray
This species is morphologically very close to the esculenta group (including marginatu)
with which it is sympatric in the Philippines. It is a small bird, wing length 86-96 mm.
(Oberholser, 1906), with glossy black upperparts broken only by a white band over the
rump, and a white abdomen.
156 Proceedings of the Linneun Society of London u77,
The nest is said to be ‘edible’, by Delacour & Mayr (1946),perhaps following McGregor
(1909): ‘Builds edible nests. Nests always in caves and composed chiefly or entirely from
secretions from the mouths of the birds. When the birds are persistently robbed, however,
they seem t o become discouraged and mix in grass, stems of small plants, moss etc.’
On the other hand, the only nests of troglodytes that I have seen, nos. 28993-2901 1 in the
United States National Museum, from Guimaras, Philippine Islands, collected by Bourns
& Worcester, January 1888, are bracket-shaped nests constructed of fibrous vegetable
material, apparently including seaweed, held together by strands of firm nest-cement.
Nests of troglodytes are also described by Kutter (1882): ‘Bei betrachtlich geringerer
Grosse, bestehen dieselben durchweg aus einem Netzwerk erharteter Speichelleimf aden,
zwischen welchen nur an der vorderen Nestwand einzelne Halme & Blattreste einge-
flochten sind’. As already indicated, in circumstances when the demand is strong enough,
such apparently incomestible nests may be collected for the sake of the small proportion
of nest-cement incorporated into them, and could then be considered ‘edible’, as by the
authors quoted above.
There are no observations whether or not this species utters the rattle call.

Colloculia slpodiopygia (Peale) : Pacific White-rumped Swiftlet

This species comprises a number of named forms occurring east of Wallace’s line, all of
which agree in being larger than any of the esculenta group, marginata or troglodytes, with
median wing size in the range 115-120 mm., and having upperparts black, more or less
glossed although not so glossy as the preceding forms, interrupted by a pronounced
whitish or grey band across the rump, and underparts more or less uniformly pale grey
with darker shaft lines. There are a few inconspicuous concealed white barbs a t the bases
of the feathers of the back. The nest, where known, is more or less bracket-shaped, the cup
composed principally of strands of vegetable material bound together with a relatively
copious application of nest-cement which characteristically takes the form of laminae
between which the extraneous vegetable material is sandwiched.
The typical race, C.S.spodiopygia,occurs on the Samoan Islands (Peters, 1940). There
are several nests in the U.S. National Museum from different Samoan islands, collected by
R. H. Beck in 1923, including no. 5336 from Ovalau Is., completely bracket-shaped, con-
structed of black fibre-like strands of moss bound together with a copious application of
nest-cement; no. 14673 from Kalawanga Is., in which the extraneous vegetable material
consists predominantly of stiff racemous flowering shoots ; nos, 5360-5367, composed of
strands of bryophyte together with some other materials, including a grass leaf-blade and
feathers, loosely bound with a rather sparser application of nest-cement in the form
of sheets rather than threads. The last group vary in shape from approximately bracket-
shaped to completely rounded, in relation t o the degree of support provided by the site
in which each nest had been placed.
The structure of the nests of the following named forms, recognized by Peters (1940) as
subspecies of C . spodiopygia on morphological grounds, is not known: C.S. assimilis
Stresemann, from the Fiji Islands, C.S.eichorni Hartert from the Bismarck archipelago,
C.S.ceramensis van Oort from Ceram and Bum, C.S.infuscuta Salvadori from Morotai,
Ternate and Halmahera, and C.S.sororum Stresemann, from Celebes. However, nests
resembling those of the typical race are known to be built by a t least three other forms
united with spodiopygia on morphological grounds, as follows. Nests of C.S.townsendi
Oberholser from the Tongan Islands are described by Finsch & Hartlaub (1870).Nests of
(7.8. leucopygia Wallace from the Loyalty Islands are in the collection of the American
Museum of Natural History, nos. 14769-14771, collected by L. MacMillan, 21 December,
1937. They are truly bracket-shaped, constructed of casuarina twigs with the addition of a
few feathers and grass stems, interspersed with copious nest-cement. A nest of the same
subspecies from New Caledonia, evidently incorporating a smaller proportion of nest-
1965-661 Guide to the specific relations of swiftlets 157
cement, has also been described :‘Eine halbkreisformige Schiisse1,ganzausMooagebautmit
nur sehr wenig verfestigendem Speichel ’ (Sarasin, in Stresemann, 1925).
Finally, the nests of C.S.terraereginue (Ramsay) are evidently also of the same type.
Nests from Dunk Island, Rockingham Bay, near Cardwell, have been described: ‘All were
adherent to the roof by a semi-transparent white substance resembling isinglass, with
which also the grass tendrils and bark composing the nest were consolidated ’ (Banfield,
in Campbell, 1909). Nests from Bedarra Island were described by Busst (1956, under the
name Collocalia francica) as consisting chiefly of casuarina foliage. Through the kindness
of Mr John Scott, I was able to examine a nest from this colony. It consisted entirely of
casuarina twiglets interspersed between laminae of firm, translucent nest-cement (now
deposited in the British Museum (Natural History) ).
Observations on the ability to orientate in darkness are generally lacking. Only in the
case of C.S.terraereginae does a description of the bird’s vocalization leave no doubt that i t
utters the rattle call : ‘ . . . a blending of cheeping, clicking and chattering, yet metallic,
and not very unlike the quick winding up of a clock’ (Banfield, in Campbell, 1909).

C’ollocaliafrancica (Gmelin): Grey-rumped Swiftlet

This is the only species of swiftlet represented on the Indian Ocean islands of Mauritius
and RBunion, where it is widespread (Rountree, GuBrin, Pelte & Vinson, 1952). It is also a
middle-sized swiftlet, with wing 112-114 mm. (Oberholser, 1906). The general colour of
the upperparts is a dark blackish brown, not heavily glossed with green or blue, broken
only by a whitish band across the rump in which there are inconspicuous dark shaft stripes.
Concealed white barbs a t the bases of the feathers of the back are present. The underparts
are pale brownish grey, again with relatively inconspicuous dark shaft stripes.
From RBunion, the nest of this swiftlet is described as being composed of vegetable
matter, primarily filaments of lichen, agglutinated with salivary product (Berlioz, 1946 :
52). No unambiguous description of nests from Mauritius has previously been published,
although Meinerzhagen (1912 : 93) has implied that extraneous materials are utilized,
with the words: ‘ . . . though it is cemented together by a gelatinous substance, I have
never heard of its being used as a food.’
There are however specimens in the Museum of Comparative Zoology, Harvard, nos.
1327-1333, from Black River, Mauritius, collected by Colonel A. Pike, October, 1872.
These are true bracket-shaped nests, composed exclusively of strands of fibrous lichen,
bound together by a sparse application of nest-cement, which is copious only at the hinge.
Two undamaged specimens measured externally (no. 1330) 6 x 5.5 cm. by 2.5 cm.deep,and
(no. 1332) 6.5 x 5.5 by 3.5 cm.
I have also been able to examine two fresh nests very kindly collected by M. Jean
Vinson, from the Caverne des hirondelles, La Louise, on 6 March, 1963. The first is a true
bracket-shaped nest, composed of fibrous lichen, bound together with fine strands of a firm
translucent nest-cement, which is copious and agglutinated only a t the hinge; it measures
5 cm. in diameter externally, 4.2 cm. in diameter internally, greatest external depth of the
nest cup 3 cm. The second is a shallow nest, more or less circular but definitely asym-
metrical and incipiently bracket-shaped, evidently having been resting on a large pile of
old and rotten vegetable material (presumably the material of earlier nests on that spot) ;
this nest is composed principally of conifer needles bound together by sparse strands of
firm white or translucent nest-cement which is only slightly more copious a t the hinge;
measurements a t the rim: external diameter 5.5, internal 4.5, external depth 1.7 cm.
These nests have now been deposited a t the British Museum (Natural History).
There is no evidence whether or not C.francica is capable of echolocation.
The namefrancicu is one of the older available among Colloculia and has a t times been
in use as a specific epithet for large groups of swiftlets. Oberholser, who placed much
stress on the feathering of the tarsus in the taxonomy of Collocalia,united under the specific
11
158 Proceedings of the Linnean Society of London [177,
name francicu all middle-sized swiftlets (wing 110-124 mm.) with unfeathered tarsi and
with a pale band across the rump a t the base of the tail (Oberholser, 1906). This grouping
however is obviously not natural, and was not widely followed. Subsequently Stresemann
(1925) suggested that francicu was conspecific with a large number of middle-sized pale-
rumped swiftlets extending from the Seychelles to Australia and the south-west Pacific,
including forms listed above under C . spodiopygia. He later modified this arrangement,
and restricted the use of francica t o forms occurring in South-East Asia, including in-
expectata and germani. Later Mayr (1937) separated francicu from South-East Asian swift-
lets. Mayr was followed by Peters (1940) who united the South-East Asian group under the
name inexpectutu. Sims (1961) examined topotypes of francicu, inexpectata and germani,
concluded that they were conspecific, and reintroduced the use of francica as the specific
name for this group (cf. Medway, 1961). However, francicu is now shown to builda very
different type of nest to the ‘white’ nest of inexpectata and its allies (below),and the two
forms cannot be considered to have close affinities.

Collocalia elaphra Oberholser ; Seychelles Swiftlet

A somewhat larger swiftlet than francica, wing 120-121 mm. according to Oberholser
(1960) (wing of the type measured by me as 124 mm.) and generally similar in appearance
except that the rump is hardly paler than the back. On the type specimen there is much
white on the concealed barbs of the plumage of the back. The dark shaft lines of the pale
grey abdomen are inconspicuous.
This swiftlet is undoubtedly resident in the Seychelles. There are reports that nesting
colonies were found in caves up in the hills of Mah6 Island in 1934 (Loustau-Lalanne,
19Gl). Unfortunately no description of the nest was given, and it has not yet proved
possible to relocate the site.
There are no observations on its ability or otherwise to utter the rattle call. On mor-
phological grounds, elaphra is close to unicolor of Ceylon (below), but without further
field details I hesitate to unite it with either unicolor or with francicu, its nearest geo-
graphical relative.

Colloculia uniwlor (Jerdon) : Indian Edible-nest Swiftlet

A medium-sized swiftlet, wing 112-120.5 mm. (Oberholser, 1906), with upperparts
uniformly blackish brown, the rump concolorous with the back. There are concealed
white tips to the basal barbs of the plumage of the back. The underparts are pale brownish
grey, with poorly marked dark shaft lines.
Nests from the type locality, the Nilghiri hills, are described in Oates (1890: 28) as
‘small shallow semi-saucer-like structures glued to the rock ’, always consisting ‘chiefly
of a long grey thread-like lichen (so common on trees in the Nilghiris) firmly agglutinated
together with saliva. . . . The normal number of eggs appears to be two.’ Similar nests
are described from all parts of the range of uniwlor in southern India in first hand reports
by observers including the contributors to Oates (1890), Abdulali (1942),and as summed
up by Stuart-Baker (1927).
In normal circumstances there is apparently quite a degree of variation in the amount of
nest-cement incorporated in the nest cup. I have seen two nests (no. 1242, in the Museum
of Comparative Zoology, Harvard) from a cave in Wellatte, Badulla district, Ceylon.
I n both the hinge was pure nest-cement, but whereas in one the cup was constructed of
distinct laminae of nest-cement between which strands of vegetable material were sand-
wiched, in the other nest-cement was much less copious, being restricted to a network of
fine threads serving to bind together the fibres of moss which comprised the bulk of the
nest. In some districts of south India and Ceylon, the quantity of edible nest-cement in
nests of the first type makes them commercially attractive, and rights of collection are
leased annually.
1965-661 Guide to the specgc relations of swiftlets 169
Novick (1959, under the name Collocalia brevirostris unicolor) trapped swiftlets of this
species and found that they were able to utter the rattle call, and to orientate in darkness
apparently by echolocation.
This swiftlet has been considered conspecific with fuciphaga of Java by Oberholser
(1906) and Ripley (1961; but see also Ripley, 1964) and with brevirostris by a number of
authors including Peters (1940).Neither attribution appears correct. Firstly, the type of
nest built by true fu,ciphuga as shown below is very different. Secondly, although records
of unicolor from the Himalayas, where it was once thought to be sympatric with brevirostris,
have been denied by Whistler (1936) and Ripley (1961), the two forms appear to be
morphologically quite distinct. On geographical grounds, the relations of unicolor might be
suspected to be with francica or ebphra, but since there are no observations on the
echolocating ability of either, nor the nest of eluphra, the exact taxonomic position of
these two forms must remain in doubt. Apart from the lack of intervening congeners,
unicolor might be considered most closely related to, and probably conspecific with the
group of swiftlets united under the name C . vanikorensis (below).
Collocalia vanikorensis (Quoy & Gaimard)
The forms united under this name are again all medium-sized swiftlets, with wing
length in the range 115-126 mm. The upperparts are blackish brown, usually somewhat
glossed, with concealed white barbs a t the bases of the feathers of the back. The underparts
are pale grey, with darker shaft stripes.
The range of the species is extensive, and a large number of subspecies have been
described. The nests of all are not known, but such examples as have been described or
collected are somewhat rounded, although tending to the characteristic bracket-shape,
composed largely of strands of vegetable material bound together with a moderate t o
sparse application of a firm nest-cement. I n some cases nests are noted as resting on a
supporting surface, rather than being suspended as is the true bracket-shaped type.
A nest of topotypical C.V.vanikorensis from Vanikoro Island, preserved in the American
Museum of Natural History, no. 5345, collected by J. G. Correia, 20 September, 1926, is
damaged but still:clearly somewhat rounded in shape, constructed of vegetable materials.
A better preserved specimen in the same museum from Goodenough Island, Papua,
attributed to the same species (presumably C.V. granti Mayr), no. 17343, collected by
H. M. van Deusen, 7 October, 1953, is also a distinctly rounded vegetable nest composed
of plant material, largely bryophyte, in the form of both tufts and strands, with very little
~ a l i v aapparent.
A nest collected on St Aignan Island, Louisiade Archipelago, attributable t o C.V.
tclgulae Mayr, has been described as ' composed of only one kind of long, wiry fibre, of a
dark colour, but strongly agglutinated and fastened to the upper surface of some rocks,
viz. resting on the rock, not hanging on the sides of the rock, as Collocalia nests usually
do' (Hartert, 1899). The nest of C.V.moluccarum Stresemann is described as being largely
composed of vegetable matter (Stresemann, 1925) and is probably similar.
I can find no records of the nests of C.V. yorki Matthews, C.V. steini Stresemann &
Paludan, C.V.waigeuensis Stresemann & Paludan, C.V.lihirensis Mayr, C.V.coultasi Mayr,
C.V. aenigm Riley, or C.V. heinrichi Stresemann, all forms assigned to the species
vanikorensis on the basis of morphology and distribution (Mayr, 1937; Peters, 1940).
Another form tentatively included in C . vanikorensis on the basis of morphology alone,
without any data on the type of nest built, is capnitis Thayer & Bangs from south China;
the type specimen (examined in Museum of Comparative Zoology) has a wing of 118 mm.
by my measurement, tail 53 mm. (central feathers 45 mm.), tarsus heavily feathered,
white tips to the concealed barbs of the feathers of the back, and rump only very slightly
paler than the back. The form has however been synonymizedwith C .brevirostris innomi-
nata (see below) by Deignan (1955b), and may just be an abnormally small example of
this group.
160 Proceedings of the Linnean Society of London P77,
Other morphologically similar forms may also be assigned to C. vanikorensis on the
basis of nest type. The first is bartschi Mearns, from Guam and other islands in the
Marianne group. Nests of bartschi are preserved in the Zoologisches Museum, Berlin, and
in the U.S. National Museum (no. 40195). These are rounded in shape and composed
largely of vegetable materials, conforming to the descriptions of Oustalet (1895) and
Stresemann (1925). Mayr (1937) has noted that pelewensis Mayr is close to bartschi, and
this form too is probably best considered conspecific with vanikorensis rather than with
inexpectatu (see Peters, 1940), even though its nest has not been reported.
Two others are amelis Oberholser and mearnsi Oberholser, sympatric on certain islands
in the Philippines, which again have previously been linked to the ‘white’ nest group on
morphological grounds. However, both are stated t o build nests ‘ made entirely of moss
and a sparse admixture of plant fibres, glued together with hardened saliva’ (Ripley &
Rabor, 1958). The type specimens of both have concealed white basal barbs in the plumage
of the back, indicating affinity with C . vanikorensis. Although the degree of tarsal feather-
ing is shown not in fact to be a diagnostic criterion, Ripley & Rabor (1958)have maintained
the separation of the two forms on other morphological characters. If the two types are
not in fact representatives of one variable population (for which amelis would be the
prior name), they provide the only suggestion that the vanikorensis group, as treated here,
has evolved reproductively isolated sympatric forms (cf. C. fuciphaga, below).
There are no observations on the ability or otherwise of any member of the species C.
vanikorensis, as outlined above, to utter the rattle call.

Collomlia inquieta (Kittlitz): Carolines Swiftlet

The swiftlets from the Caroline Islands included in C. inquietu are morphologically little
differentiated from C. vanikorensis. Mayr (1937) has noted that the two groups will
probably prove to be conspecific.
Nests of C.i. inquieta from Kusaei Is. are preserved in both the U.S. National Museum
(no. 41760) and in the American Museum of Natural History (no. 17375), collected simul-
taneously from the same colony by J. H. Brandt, 10 May, 1959. These are distinctly
rounded, suppressed bracket-shaped nests, composed of moss and other soft vegetable
matter with nest-cement in evidence only a t the point of attachment to the cave wall.
The collector’s field notes record, ‘ I n cave in clusters stuck to walls and ceiling ’ ; but both
specimens from their structure had clearly been partly resting on irregularities in the rock
wall. The clutch is recorded as one egg. From Ponape Island there are in the American
Museum of Natural History (no. 17374)’ two adjoined nests of C.i. pompensis Mayr,
also collected by J. H. Brandt, 8 May, 1959. These nests are again distinctly rounded,
with entirely circular central depressions, 2 cm. deep and 4 cm. wide internally, having
apparently been built onto and subsequently been partly invaded by a termite nest. There
is a thin deposit of nest-cement round the outer margins of the nest cup and near the point
of attachment to the cave wall. The collector’s notes read: ‘ I n a small cleft of rock-stuck
to wall-20 ft. up-nests in clusters’. The clutch is recorded as one egg.
From Moen Island in the Truk Atoll, there are nests of C.i. rukensis Kuroda in both
American museums (U.S.N.M.41761,41762; A.M.N.H. 17376),collected by J. H. Brandt,
4 April, 1959. These nests too &remore rounded than truly bracket-shaped, with thick
walls composed of short lengths of fibrous vegetable material including seedless grass
heads, (?) fibres from the spathe of coconut palms, and short lengths of woody twigs,
bound together with nest-cement which is copious only along the line of attachment to the
rock wall. Collector’s notes for no. 17376 read: ‘In small man-made cavern-attached to
wall-8 ft. u p - s t u c k to small projection’. The clutch is recorded as two eggs. A general
description of nests from Truk by Brandt himself indicates that in fact both rounded nests
of this type and also more pronouncedly bracket-shaped nests are commonly built: ‘The
nest is a well cupped affair constructed of grass stems and small tendrils; and occasion-
1965-661 Guide to the speci$c relations of swijtlets 161
ally small sticks, fern stems, and more rarely, mosses are employed. A few feathers are
generally found embedded in the nest which is glued together with copious secretions of
saliva and firmly attached to the cave rocks. No lining is used. Some nests are entirely
supported from underneath by stone, whereas others protrude and are fastened only a t
the back of the nest’ (Brandt, 1962: 427).
There are no observations on whether or not inquieta and allies utter the rattle call.

Colloculia salangana Streubel

C. sahngana is a slightly larger swiftlet of the medium-sized group, with wing measure-
ments in the range 120-128 mm. The upperparts including the rump are uniformly
blackish brown ; the concealed barbs a t the bases of the feathers of the back are invariably
a uniform greyish brown throughout their length and do not have white tips. The latter
character is important in so far as it serves to distinguish C. salangana from sympatric
subspecies of C. fuciphaga (below).The underparts are pale grey, with inconspicuous dark
shaft stripes.
For many years, this swiftlet has been known as Colloculiafuciphuga, and it is under this
name that the nest of the nominate subspecies, C.S.sahngana of Java, was first described
by fitresemann (1926a: 107). It is built of vegetable materials, held together by a sparse
amount of nest-cement which is transparent when fresh, and does not harden (as does the
nest-cement of all species discussed so far) but remains permanently soft and moist to the
touch. Apparently because this nest-cement is structurally weak, the nests always depend
a t least for partial support on a ledge or on some small irregularity in the cave wall, and the
truly self-supporting bracket-shaped type is not found (cf. Medway, 1962 c).
The nest of C.S.natunae Stresemann from Borneo and the Natuna Islands is similar
(Medway, 1959b; Sims, 1961 ; both under C. fucipimga natunae; Medway, 1962a) (Plate
2 (a)).
There are two other named forms, aerophih Oberholser from Nias, other islands off the
west coast of Sumatra, and perhaps the Sumatran mainland itself, and maratua Riley
from the Maratua archipelago, off north-east Borneo, which can be associated with
C‘. salangana on morphological grounds, particularly on the presence of all-dark concealed
barbs a t the bases of the feathers of the back of the type specimens of both. No nests have
been attributed to either.
The biology of C.S. natunae has been studied in some detail a t Niah cave, Sarawak,
(Medway, 1962a) and it has been shown to utter the rattle call. C.S.salungana from West
Java is also known to utter the rattle call, and under experimental conditions individual
birds have been shown capable of oriented flight in darkness (Medway, manuscript in
preparation).
Although the distinctness of this species has been recognized for many years (Strese-
mann, 1914b), its nomenclature has been confused until quite recently (Medway, 1961).
The absence of white tips to the concealed barbs a t the bases of the feathers of the back is
a useful character, first emphasized in swiftlet taxonomy by Mayr (1937), which serves to
distinguish C . salangaizcc from sympatric C. fuciphaga and allies (see below).

C’olloculia hirundinacea Stresemann

C. hirundinacea is sympatric with C. uanikorensis in New Guinea and certain neighbour-
ing islands. Although the two species have been confused by some authors, they are
separable on morphological grounds. The differences have been summarized by Mayr
(1937: 12), who noted that C. hirundinacea can be distinguished inter alia by having the
tarsus feathered, by having the crown distinctly darker than the back, by not having the
silvery grey throat contrasting with a much darker abdomen, and by having the under
wing-coverts and feathers of the superciliary pure black not with pale edges. Wing length
within three recognized subspecies varies from 114-131 mm. (Mayr, 1937).
162 Proceedings of the Linnean Society of London P77,
The type of nest built is apparently quite distinct. There are two records available. The
nests of C.h. hirundinacea have been described by Rand (1942) as, ‘bulky, solid affairs,
flattened on the bottom and back where they rested against the ledge, otherwise just
masses of material with a depression a t the top. The nests were composed of mosses,
lichens, filmy ferns and some rootlets, the lining being similar but finer material was packed
and felted together with no salivary glue apparent.’ A similar nest attributed on the
label also to C . hirundimmu, is held in the American Museum of Natural History, collected
by E. T. Gilliard, 20 May, 1950, from Mt Kubor, New Guinea, c. 6,000 ft. This is also a
deep cup-shaped structure, built principally from fibrous lichen (‘ Spanish moss’), and
lacking all signs of nest-cement.
Similar nests constructed without nest-cement, have been found in a Sarawak cave,
where they were attributed to C. salangana (Medway, 1959b; under the name C .fuciphugu
natunae).
There is no evidence whether or not C . hirundinaceu utters the rattle call.

Colloculia leucophaea (Peale)

A middle-sized swiftlet, with wing 117-5-127 mm. (Oberholser, 1906), characterized
by soft, dull coloured, grey brown plumage, a long tail, reduced supkaloralspots, and black
under tail-coverts (Mayr, 1937).
A nest from Tau, Mariana Is., no. 14780 in the American Museum of Natural History,
collected by R. H. Beck, 26 December, 1923, attributed to this species (presumably C.2.
leucopham) is distinctly rounded, composed of vegetable material, largely fern fronds.
Collector’s notes record that i t was sited ‘ on a little ledge, 4 ft. above floor of low lava
tunnel.’ There are also seven nests of C.1.ocistu Oberholser, also collected by Beck, in 1922,
from Marqueses Is., nos. 5325-5331 in the American Museum of Natural History. These
again are deep, rounded nests constructed of strands of moss with the application of very
little nest-cement.
There is no evidence whether or not this species utters the rattle call.

Collocalia brevirostris (McClelland): Himalayan Swiftlet

This swiftlet is characterized primarily by the marked furcation of the tail, in which the
difference in length between the long outer feathers and the short central feathers generally
exceeds 15% of the length of the former, and by its large size, with wing over 123 mm. in
most subspecies and reaching 142 mm. in some. The tarsus is lightly feathered or un-
feathered, and the rump colour variable.
The nominate race, C.b. brevirostris, in which the rump is brownish grey, clearly differ-
entiated from the blackish brown of black and mantle, ranges through the Himalayas to
northern Burma (Deignan, 1955b).Nests from the Himalayan region have been described
by Glennie (1944))from caves a t 9000 ft. in Chakatra District, U.P., India (under the name
C. fuciphugu).The nests are evidently of the self-supporting bracket-shaped type : ‘made
of moss gummed in layers by means of the viscid saliva which is also used to stick the nest
to the vertical rock wall, no use being made of rock ledges to support the nests.’ The clutch
is given as two eggs, measuring 20 x 13.8 mm. The skin of a swiftlet taken from this colony
is preserved in the collections of the Bombay Natural History Society, and was kindly
made available by the Hon. Secretary. The wing, only 119 mm. long, is shorter than the
minimum for C.b. brevirostris given by Deignan (1955b), 123-132 mm., or Whistler (1936),
121-1316 mm. On the otherhand, thereare twoother skins in the BritishMuseum (Natural
History) nos. 87.8.1.262 and 76.2.12.292, also of wing length 119 mm. and on other
characters all three conform to brevirostris. No other species is recorded from this area
(Ripley, 1961), and for the time being these very small specimens must be attributed to
abberant brevirostris. It still remains possible that in fact they represent a smaller species
related to the unicolor group.
 1965-661 Guide to the specijc relations of swiitlets 163
C. brevirostris is also resident in Sumatra, where nests have been described from a cave
a t Buo, Padang Highlands, as being ‘made of moss and other vegetable matter, fixed to the
wall with the well-known slimy secretion from the buccal glands ’ (Jacobson, in Robinson
& Kloss, 1924: 243). The subspecific attribution of this population, with wing length
125 mm., is uncertain; it is likely to be close to C.b. vulcanorum Stresemann, which is
resident in Java and apparently also builds vegetable nests (Medway, 1 9 6 2 ~ ) .
From Malaya, C. brewirostris is known only as a winter visitor (Gibson-Hill, 1949).
Further north populations distinguished as rogersi Deignan, breeding in the Southern
Shan States, west Thailand and Laos, and inopina Tliayer & Bangs and pellos Thayer &
Bangs, breeding in south-central and western China, are considered conspecific on mor-
phological grounds although their nests are not known (cf. Deignan, 1955b). Deignan
(19556 ) has assumed that the type of innominuta Hume, taken on the Andaman Islands
from which no other specimen has subsequently been collected, represents a winter
straggler from a northern locality, and has synonymized with it capnitis, inopinu and
pcllos. Of these, capnitis (a smaller swiftlet, with wing length of the type measured by
myself as 118 mm.) has already been mentioned above, with the suggestion that the unique
type specimen may represent a mainland congener of the micolor-vanikorensis group.
Smaller members of populations a t present united under the name rogersi, of which the
wing length is recorded as 116-128 mm. (Deignan, 1955b),may also ultimately be found to
be more properly related to vnnikorensis. Furthermore, since the species brevirostris is
resident in Sumatra and Java (of which Deignan was evidently unaware), and since these
Indonesian populations are little if any differentiated from the type of innominuta, the
position of the unique type of innominuta is left in some doubt. On available evidence, it is
certainly difficult to be sure that i t represents a migrant straggler from south-west China,
and I am inclined to conserve a local name for populations known to breed in that
region.
Of the other two Chinese swiftlets, both pellos and inopina (types examined) are un-
doubtedly conspecific with brevirostris on morphological grounds. Wing lengths of topo-
types of pellos from Szechwan in the Museum of Comparative Zoology, the Zoologisches
Museum, Berlin, and the British Museum (Natural History) vary from 128-136 mm.
(eight measured) ; they are further distinguished from C.b, brevirostris by a darkened rump,
which is virtually concolorous with the back. The holotype of inopina which was col-
lected further east, in Hupeh Province, more closely resembles nominate brewirostris in so
far as the rump is noticeably paler than the back. But this character is approached by
some topotypes of pellos. If the two are considered to represent one variable taxon, the
name inop’nu has priority as subspecific epithet.
In the collections of the British Museum (Natural History), the skin of a n immature
nestling is associated with adult specimens from Omi-hsien, Szechwan. Despite this con-
firmation that C.b. inopinu breeds in Szechwan, no unequivocable description of nests
from this region can be found, apart from a casual reference t o ‘inedible’ nests (Guide
Madrolle, quoted by Medway, 1964).
Specimens attributed to C.b. inopina in the British Museum also include skins from
Yunnan and from Tonkin (the latter collected 24 February, 1924). Morphologically these
are very close to the equally dark-rumped form whiteheudi Ogilvie-Grant from Luzon,
Philippine Islands, extending to New Guinea, of which the female ‘co-type’ (B.M. (N.H.)
97.5.13.290) has wing 136 mm. (my measurement, cf. 5.4 in. of Ogilvie-Grant, 1895).
C.b. inopha may be distinguished most easily by the possession of moderately well-
feathered tarsi, as opposed to whiteheudi of which the tarsi are entirely devoid of feathers.
The nest of whiteheudi is apparently similar to the nest of Sumatran and Himalayan repre-
sentatives of brewirostris: made of green moss and placed in a waterworn cave ’ (McGregor,
1909); ‘made of green moss fastened with saliva t o the wall of caves’ (Delacour & Mayr,
1946). Previous authors up to and including Peters (1940) have considered whiteheadi a
distinct species. However, in view of the intermediate position of inopina and the
164 Proceedings of the Linnean Society of London P77,
similarities in nest structure, there seem to be no valid grounds for separating whiteheadi
from the species C . brevirostris.
The remaining forms of C. brevirostris inhabiting the islands east of continental Asia
include four further subspecies, all dark-rumped and morphologically close to white-
headi. Of these, there are nests attributed to origenis Oberholser, from Todaya a t 4,000 ft.,
Mindanao, nos. 32092-32096 in the United States National Museum, collected 12 July,
1904, by E. A. Mearns, which are rounded vegetable nests constructed of green bryophytes
together with some fibrous plant material, and apparently not incorporating nest-cement.
The nests of a p e n s i s Hachisuka and orientalis M a p , listed under whiteheadi by Peters
(1940), are not known.
Finally palawanensis Stresemann, from Palawan Island, must also be placed in this
species, rather than with lowi (i.e. C . maxima, see below) as a subspecies of which i t was
described. Ogilvie-Grant (1895) differentiated carefully between specimens of these
two large swiftlets from Palawan, distinguishing those he attributed to whiteheadi as,
‘all having the tail distinctly forked, the middle pair of feathers being a quarter of an inch
shorter than the outer; also the tarsi entirely naked.’ This as opposed to lowi, of which
‘thc tail is practically square, the middle and outer feathers being subequal, and the tarsi
distinctly feathered.’ As shown below, the dense feathering of the tarsus is a character of
all subspecies of C . m a x i m ( =low*).At the date he described palawanensis, Stresemann
in fact considered low‘ and whiteheadi conspecific (see Stresemann, 1926b). Nonetheless
the tarsus of palawnensis is stated unequivocably t o be naked (Stresemann, 1914b), and
this form must accordingly be attributed t o the brevirostris-u~hiteheadigroup, now re-
cognized to be separate. As noted by Peters (1940), tsubame Hachisuka therefore falls
as a synonym of palawanensis.
Of the many subspecics of brevirostris detailed above, there are positive records of the
rattle call being uttered only by C.b. v u h m r u m in Java (Medway, 1962~).I n addition,
Jacobson noted that the cave a t Buo, Sumatra, from which nests were collected, is a
natural tunnel several miles long, the walls of which were covered with the nests of these
swiftlets : It is quite incomprehensible how the birds can find their way in the absolute
dark and among the bewildering noise caused by the twittering of hundreds of birds’
(Robinson & Kloss, 1924). From this description it can be inferred that these swiftlets were
certainly capable of orientation in darkness, presumably by echolocation.

Colloculia gigas (Hartert & Butler) : Giant Swiftlet

C .gigas is the largest and rarest of swiftlets, known from a very small number of speci-
mens collected in Malaya, Sumatra and Java, and believed also to occur in Borneo
(Smythies, 1960). It is distinguished by large size, with a wing of 157-102 mm. (Strese-
mann, 1932), and uniformly dark upperparts. The nest is described by Hoogerwerf
(1949) as a vegetable nest, with thick walls, consisting mainly of aerial rootlets and
fibrous material.
Colloculia fuciphaga (Thunberg) : Edible-nest Swiftlet
A member of the medium-sized group of swiftlets, with wing in the range 110-128 mm.
The rump in most subspecies is greyish brown, more or less distinctly paler than the back
and tail, marked with dark shaft stripes. I n the plumage of the back there are always
white tips to a t least some of the concealed downy barbs a t the bases of the feathers. The
tarsi may be lightly feathered or unfeathered.
The nests of all named forms united by these common morphological characters are
constructed almost exclusively of concentric laminae of firm nest-cement. I n most nests a
few small swiftlet feathers are found adhering to and partially incorporated in the cup.
It is assumed that these feathers derive from the ventral region of the building birds, and
that they adhere to the nest-cement accidentally during construction (Plate 2 ( a ) ) .
1965-661 Guide to the specific relations of swiftlets 165
There are also reported instances from coastal locations of strands of marine algae being
incorporated into nests of this species. There are no observations of birds actually collect-
ing such materials for the nest, and i t is not clear whether their incorporation is deliberate
or accidental.
The ‘white ’ nest of the nominate race from Java, C .f.fucipllaga was described and figured
in the original description (Thunberg, 1812), and has since been redescribed several times,
generally under the specific name francicu. The inclusion of strands of marine algae in a
few nests has been recorded from a colony nesting among sea cliffs on the south coast of
Java, overlooking the Indian Ocean. It was reported a t the time by the nest collectors that
the same algae grew on the rocks t o which the nests adhered, and it was assumed that some
of these had been incorporated into the nest unintentionally (Medway, 1962c). The
process of nest building a t an inland site has been observed in some detail (Franck, 1926)
and neither in the account of this author, nor of others including for instance Kuroda
(1936), nor in my own relatively wide experience, has vegetable material been observed
incorporated into the main structure of any nest a t an inland site. On the other hand both
as reported by Franck (1926), and in my own experience (in Java, unpublished; in
Sarawak, see Medway, 1959b) nests of fuciphaga subsp. may sometimes be found built
onto a vegetable base or ‘hinge’. Such nests occur only in sites where other species of
swiftlet which build vegetable nests are also present (C. esculenta or C . salangana),and on
examining such nests, I have no doubt that they represent normal ‘white ’nests of fuciphaga
which have been built onto the abandoned remains of a vegetable nest originally con-
structed by the other species.
Specimens of C.f.fuciphaga taken from inland localities in Java cannot be separated
from those of coastal breeding populations and all must be included in a single sub-
species. The type of bartelsi Stresemann with wing 122 mm. (Stresemann, 1927), is larger
than the mean but still within the range of the species; i t is unlikely to represent a distinct
population and, with javensis Stresemann (already synonymized by Peters, 1940), may be
reduced to the synonymy of fucipllaga.
Populations attributable to C.f. fuciphaga also occur on Billiton (skins in the Zoological
Museum, Bogor) and the Kangean Islands. Other populations closely related to C.f.
fuciphaga, in which the overall coloration of the rump is only very slightly paler than the
back, are represented by dummermani Rensch from Bali and the Lesser Sunda Islands east
to Flores, which builds ‘good white edible nests’ (Hartert, 1896), and micuns Stresemann
from Sumba, Savu and Timor, recorded on the label of the type (from S a m ) as: ‘This is
the bird which constructs the white edible nests on this island’ (Stresemann, 1914b).
I n Sumatra as in Java there is apparently only one form of C . fuciphaga, which nests
both in inland sites and on coastal cliffs and in sea caves. This subspecies, vestitu Lesson, is
distinguished by a darkened rump which is concolorous with the plumage of the back.
Specimens from Sumatra cannot be differentiated from the ‘white ’-nest builders from
inland caves in Borneo, including those a t Tamaluang in south-east Kalimantan,
Gomantong and elsewhere in eastern Sabah, and the Baram district, Sarawak; these
populations have all been referred to vestitu (cf. Stresemann, 1932; Smythies, 1960).
As noted above, a somewhat paler-rumped swiftlet attributable to C.f. fuciphuga occurs
on the island of Billiton intervening between the dark-rumped populations of Sumatra
and Borneo. From this island northward there is a progressive cline of paling rump colora-
tion. In southern Malaya a slightly paler-rumped form is known to build white nests in
Singapore, Malacca and on islands of the Tioman archipelago (Gibson-Hill, 1948) ;this has
been assigned to amechanu Oberholser, type locality the Anamba Islands. On the coasts
and offshoreislands of Thailand, southern Burma (i.e. the Mergui Archipelago), Cambodia
and Vietnam, a still paler-rumped form, C.f. germani Oustalet, is again known to build
‘white’ nests (Gibson-Hill, 1948; Smythies, 1953; Giles, 1936; Sallet, 1930).
I n the Philippines, commercially valuable ‘white’nests are recorded from Palawan and
the Sulu archipelago (cf. Medway, 1964). These have been attributed to amelis by authors
166 Proceedings of the Linneun Society of London [177,
including Delacour & Mayr (1946). Oberholser (1906) recorded a specimen of amelis from
Palawan, but as listed by Peters (1940) its range is exclusive of the areas in which white
nests occur. Moreover, amelis apparently builds a vegetable nest, and has accordingly been
assigned to C. vanikmensis (above). The ' white'-nest swiftlet of the southern Philippines
may be identical with germani, or if separable may be closer t o perplexa Riley (below).
On the Andaman and Nicobar Islands, 'white' nests are built by inexpectata Hume
(specimens examined from the collections of the Bombay Natural History Society), cf.
Osmaston quoted by Stuart-Baker (1927) describing nests of ' pure white semitranslucent
inspissated saliva, half cups stuck up against the sloping roofs of small caves round the
coast '.
Off the west coast of Borneo, 'white' nests are found on the islands of south-west
Sarawak, built by a pale-rumped swiftlet, clearly separable by this character from the
dark-rumped populations of inland localities, and attributed t o germani (Smythies, 1960 ;
supported by personal observation). Off the north-east coast of Borneo, on Berhala
Island, Sandakan, another swiftlet building ' white' nests occurs, distinguishable from
vestita of inland sites by a less marked but none the less distinct paling of the rump. This
population is morphologically close t o birds from the Maratua archipelago, north-east
Kalimantan (Borneo),which have been described as perplexa Riley; the nest is not known.
Stresemann recognized the close afinity between vestita, the subspecies on Java (which
he knew as javensis and bartelsi), and germani, and up to his paper of 1932, considered
them conspecific. Chasen (1935) then drew attention to the existence of these dark-
rumped and grey-rumped populations virtually sympatric in northern Borneo, and
separated the former under the specific name C. vestita. This separation was subsequently
maintained by Mayr (1937),Peters (1940) and Sims (1961).
However in northern Borneo, the dark-rumped and the pale-rumped populations are
not strictly sympatric in so far as they are segregated in the breeding sites. All known nest
colonies of dark-rumped swiftlets are located in inland, landlocked caves, whereas the
pale-rumped forms are restricted to sea caves or offshore islands. Elsewhere within the
range of the ' white '-nest swiftlets, only one form occurs in each locality. I n Javafuciphqa,
in Sumatra vestita and in southern Malaya amechuna are all tolerant of a wide variety of
nest sites, favouring localities far inland as well as a t the coast or on offshore islands. On
the other hand the more northerly geographical representatives of the group, germani on
continental Asia and the subspecies on the Philippines (which is perhaps related to per-
plexa), are apparently confined to the coast and offshore islands, as they are in Borneo,
and are presumably better adapted t o such habitats. Among other species of swiftlet, it
has been reported that mated pairs copulate a t the nest site (Medway, 1 9 6 2 ~ )If. copulation
occurs only here, and if mate selection also takes place only at or near the nest colony,
then these different subspecies of C.fuciphuga will be effectively isolated by their divergent
preferences of breeding sites.
The situation in Borneo can thus be interpreted as a circular cline, in which the terminal
populations remain distinct and reproductively isolated although sympatric, yet inter-
grade completely through a series of intervening allopatric populations. The dark-rumped
vestitu presumably derives from Javanese C.f. fuciphuga (with which as noted above it
was for long considered identical) and colonized Borneo from the south; since the Bornean
population is separated from the morphologically identical Sumatran population by
typical fuciphuga on Billiton, these two dark-rumped populations may not in fact be
monophyletic. The pale-rumped germni may be a recent invader to the coast of Borneo,
where it is in fact geographically separated from any known vestitu colony. Finally perplexa
in the north-east may be derived from the population in the Philippines, or from the
Lesser Sunda Islands (see Text-fig. 1). Rather than erect an artificial division by separating
vestita, it seems preferable to unite all the forms discussed above in a single species.
As already noted, a number of previous authors have used the name fuciphaga to
designate the species correctly known as C. salangana. At the same time the' white'-nest
1965-661 Guide to the speci$c relations of swiftlets 167
builders have been united with francica (e.g. Sims, 1961). As shown above, C .francica in
fact builds a very different type of nest and cannot be considered conspecific with this
group. Mayr (1937))on different grounds, has in fact already separated the white '-nest
swiftlets of south-east Asia from francica. Since the namefuciphaga was not a t that time
considered to be available, Mayr used inexpectata as specific epithet and was followed by
Peters (1940)) Gibson-Hill (1949) and Smythies (1960) among others. There is now

C.ir

1000 105' 110° 115O I 20° I25O

Text-fig. 1. Distribution of recognised subspecies of Collocalia fuciphnga, the Edible-nest Swiftlet

however no doubt that the older name fuciphaga is correctly applied t o the white'-nest
swiftlet of Java (Medway, 1961) and therefore has priority as the specific name of this
group. It has already been shown that bartschi and pelewensis which were listed under
inexpectatu by Peters (1940),and mearnsi listed under vestita, are races of C . vanihrensis,
and aerophila and maratua, also listed under vestitu, are probably correctly referred to
C. salanganu.
I have personally ascertained that members of the following recognized subspecies of
C. fuciphaga utter the rattle call: vestita, perplexa, germani, nmechanu and fuciphaga.
168 Proceedings of the Linneun Society of London
Collocalia maxima Hume : Black-nest Swiftlet

Alarge swiftlet, with the wing 122-136 mm., in most subspecies over 125 mm. Over
most of its range C . maxima is sympatric with C . brevirostris-from the eastern Himalayas
to the Philippine Islands and south to Sumatra and Java, excluding only Borneo where
brevirostris is absent. It may be distinguished from brevirostris in all regions firstly by
the heavily feathered tarsus, which bears a distinct row of a t least six or seven small feathers
on the outer side and a second row of a t least four small feathers on the inner side ;secondly
by the more or less square tail, in which the difference in length between the long outer
pair of retrices and the short central pair normally does not exceed 12% of length of the
former. The rump colour is variable.
Use of the specific name maxima, which antedates low‘ Sharpe is, has been validated
by Deignan ( 1 9 5 5 ~ The
) . nominate subspecies extends from Tenasserim south to Malaya
and north-west to Bhutan and southern Tibet (Medway, 1 9 6 2 ~ )Nests . of the Malayan
population (formerly known as C. lowi robinsoni Stresemann) have been described by
Chasen (1939)as ‘black-nests’, with the comment, ‘I cannot distinguish these nests from
those of lowi taken in Borneo’. Similar nests are built by C.m. lowi Sharpe, northern
Borneo (Medway, 19593; Smythies, 1960)) and C.m. tichelmani in south-east Borneo
(Stresemann, 1932; specimens examined in Zoologisches Museum, Berlin), as well as by
the intermediate population in western Java ascribed to maxima 3 lowi (Medway, 1962 c ) .
All are bracket-shaped, incorporating feathers from all parts of the plumage of swift-
lets, and held together by a firm translucent nest-cement without the inclusion of any
vegetable materials.
It is clear from the remarks of Ogilvie-Grant (1895) that the species is also found on
Palawan, Philippine Islands, although as shown above palawanensis Stresemann is a
member of the brevirostris group. It has also already been noted that vulcawum Strese-
mann, assigned t o loui (i.e. maxima) by Hoogerwerf (1949)and others, is again correctly
a subspecies of brevirostris.
The rattle call is known to be uttered by C.m. bun’(Medway, 1 9 5 9 ~the ) ~ Javan repre-
sentative of the species (Medway, 1962c), and C.m. maxima in Malaya (personal
observation).
DISCUSSION

1 . Ecological Separation
From the preceding systematic section i t is clear that, with one exception, the forms of
swiftlet sympatric in any given region are separated either by relatively marked nior-
phological differences such as size or coloration, or by physiological differences expressed
as the type of nest built. I n Java for instance six species occur. Among these, four size
groups are recognizable: the very large C . gigas, large C. brevirostris and C. maxima,
medium C.fuciphaga and C. salangana, and small C. esculenta linchi. Of these, each of the
four species building vegetable nests (gigas, brevirostris, salanganu and esculenta) falls
into a different size group, and the two remaining species are each distinguished from the
builder of vegetable nests which they match in size by nests of very different types.
Only in northern Borneo is there a group of very similar swiftlets, separable by no more
than minor morphological characters, which build the same type of nest. This group
comprises the ‘white’-nest swiftlets vestita, germani and perplexa. The first named
colonizes inland caves, the latter two are restricted t o offshore islands of, respectively, the
west and the north-east coasts. I n view of the fact that the dark-rumped vestita is virtually
sympatric in different parts of its range with one or other of the paler-rumped forms,
previous authors have given it specific status. It has been suggested above that this
situation is more satisfactorily explained if the three forms are classified as races of a single
species, between which genetic isolation is maintained by their divergent ecology.
1965-661 Guide to the specific relations of swiftlets 169

2 . Evolution Within the Genus

From the data presented above, it is possible to draw up a scheme of evolutionary
relations within the genus as defined. There are still a number of species for which observa-
tions are lacking on the ability, or otherwise, to utter the rattle call indespensible for
echolocation. But among those for which definite observations have been made, two
distinct morphological groups can be recognized ; these correspond exactly to the group-
ing by ability or inability to echolocate. The swiftlets that are known not to utter the rattle
call, and to be unable to orientate in darkness, comprise the forms listed above under C .
esculenta and C . marginuta, all of which are among the smallest swiftlets, further distin-
guished by a blue or greenish gloss to the dorsal plumage, and by white edges to the
feathers of the abdomen. On the other hand, all those which are known to utter the rattle
call are larger ; the general colour of their upperparts is dull blackish brown with only a
slight gloss, with plain grey underparts lacking white markings. Such a division, into
‘glossy ’ and ‘grey ’ swiftlets, respectively, has already been tacitly recognized by previous
authors on purely morphological grounds (e.g. Stresemann, 1932). It is worth noting here
the importance of discovering whether or not troglodytes utters the rattle call, since in
other respects its position is intermediate.
Assuming that all ‘grey ’ swiftlets for which observations are lacking will ultimately
prove to be able to echolocate, it appears that the ‘glossy’ swiftlets are closest to the primi-
tive or ancestral stock of the genus. The type of nest built by these swiftlets likewise con-
forms to the primitive pattern within the genus and is built also by the smaller species of
‘grey ’ swiftlets : by spodiopygia and by the eluphra-francicu-unicolor-vanikorewis-
inquietu group. This type of nest consists of fibrous vegetable material arranged in more or
less concentric laminae and bound together by a firm translucent nest-cement. Normally
these nests are bracket-shaped, but as shown above, in conditions where the extraneous
substrate offers basal support, the bracket-shape may be suppressed and the final form of
the nest is more or less rounded with a circular central depression. Both the shape of the
nest and the proportion of nest-cement to vegetable matter are variable, both between
different populations of one species and between different nests in any given colony.
There is one rather poorly supported instance of a nest attributed to C . esculentu being
built without nest-cement a t all.
Within the morphological group of ‘glossy’ swiftlets, there is little if any taxonomically
significant advance in nest construction. Within the ‘grey’swiftlets, four main lines can be
recognized, three of which are clearly anticipated in the variations in the primitive type
already discussed. The evolutionary relationships suggested by comparison of these nest
type are illustrated diagrammatically in Text-fig. 2.
Firstly the line of simple increase in body size, without apparently any change in methods
of nest construction, represented by brevirostris and gigas (although the nest of the latter
is poorly known). Secondly the line of reduction in either the proportion or the mechanical
properties of the nest-cement (or both) resulting in nests which are obligatorily more
rounded and less bracket-shaped in final form. The species C . leucophaea represents a
morphologically distinct offshoot of this line. Less distinct morphologically, C . salangana
secretes a permanently moist, mechanically inefficient nest-cement. At the evolutionary
extreme C . liirundinacen apparently builds nests of a type in which nest-cement is not
utilized, thus differing not only from other members of the genus, but also from the
family of swifts in general (Lack, 1956).
The third line is represented by the converse trend, towards a great increase in the
proportion of nest-cement (to 100%) in the nest of C .fuciplmga. The occasional records of
vegetable matter incorporated in the nests of this species may represent a residual tendency
to utilize such materials, but as suggested above may equally well be accidental.
The final line is that represented by C .maxima in which the nest cannot easily be derived
from a modification of the ancestral vegetable type. Although the nest of unicolor in
170 Proceedings of the Linnean Society of London u77,
India is said normally to include a t least a small proportion of feathers (Stuart-Baker,
1927),there are no observations on the method by which these feathers are collected, and
no indication that they are deliberately incorporated in the nest structure. Similarly,
nests of fuciphaga subspecies also often have adherent a few small contour feathers; these
are thought to be incorporated accidentally. The feathers found in the nests of C . maxima
include not merely small feathers from all parts of the body, but also large remiges or
retrices, and there is no doubt that their incorporation in the nest is intentional. It must
be assumed that C. maxima represents an early divergence from the main evolutionary
line of echolocating swiftlets.
Further field observations are needed in order to establish whether or not all species of
the morphological group of ‘grey’ swiftlets are capable of echolocation. Observations of
the behaviour of C. troglodytes are also important. If such observations support the separa-
tion into a ‘grey’ group and a ‘glossy’, it may be desirable t o divide the genus formally.

NO echolocation ? echolocation Kmwn echolocation _ --1

I C. troglodytes
. . . I
C. francico ,r I
C. spodiopygio
I
I
;.’ .:.’
,.
--;.. .
,‘a
*;,

Presumed common ancestor

I
Text-fig. 2. Diagram to illustrate the postdated evolutionary relationships of recognized
species of the genus CoUocaZia.

I n this case it should be noted that esculentu Linn. is by designation the type species of the
genus Collocalia, which would accordingly be restricted to the Lglo~sy’ group. Earlier
names already proposed for supraspecific taxa within the genus (AerodromusOberholser
1900; Zoonava Mathews, 1920) do not accommodate the ‘grey’ g o u p as a whole, and for
this a new name would be required.

ACKNOWLEDOEMENTS

The field work on which these observations were based was supportedin Sarawaklargely
by the Sarawak Museum, t o which I was then attached, and in Indonesia by the Jajasan
Siswa Lokantara, under whom I held a research fellowship. The opportunity to visit
museums in the U.S.A. was made possible through a travel grant from the B.P. Bishop
Museum, Honolulu.
1965-661 Guide to the speci$c relations of swiftkts 17 1
I am grateful to the directors and curators of ornithological collections in all the museums
or other institutions mentioned in the text, for permission to examine material in their
acrc; and particularly to two leaders in the field of swiftlet taxonomy, Dr Ernst Mayr
who kindly made available his notes on the subject put together over many years, and
Dr Erwin Gtresemann, with whom I had the privilege of discussing several of the problems
raised. I am also grateful to Mr Tom Harrisson for permission to use the photographs of
Plat,es 1 and 2 ( b ) .
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EXPLANATION OF PLATES
PLATE
1
(a) A group of typical self-supporting bracket-shaped nests of Collocalia maximu built
onto a smooth rock surface, at this point concave and slightly over-hanging, in the interior of
Niah Cave, Sarawak. Photographed by flash.
( 6 ) Nests of C . mazima lo& Sharpe, photographed by flash in the interior of Nicth Cave,
Sarawak. The numbers on the cave wall relate t o a study of the breeding biology of the species.

PLATE
2
( a ) A cluster of nests of Collocalia fuciphaga uestita (Lesson), photographed by flash in a
narrow chimney in a cave in the Baram District, Sarawak.
(b) Nests of Collocalia salangana natunae Stresemann photographed by flash in the interior
of Niah Cave, Sarawak. Both nests are partially supported by irregularities in the cave wall.
Proc. Linn. Soc. Lond. Vol. 177, No. 2 Plate 1

MEDWAY (Facing p. 172)

 Platt. 2

MEDWAY