The avifauna of Flores (Lesser Sunda Islands)
G.F. Mees

Mees, G.F. The avifauna of Flores (Lesser Sunda Islands).
Zool. Med. Leiden 80-3, 25.viii.2006: 1-261, figs. 1-18. ISSN 0024-0672.
G.F. Mees, Nationaal Natuurhistorisch Museum, Leiden, The Netherlands. Present address: 31 West
Street, Busselton 6280, Western Australia, Australia.
Key words: Indonesia; Flores; avifauna; birds; zoogeography.
The avifauna of the island of Flores (Lesser Sunda Islands) is reviewed. Introductory sections, which
include a chapter on the history of ornithological discovery, are followed by the main part, a systematic
account in which each species and subspecies known from Flores is treated separately. A discussion of
the zoogeography, a gazetteer, a list of references and an index complete the volume. At present 214
forms (210 species) are accepted as having been reliably recorded. Of these, ca. 160 are, or may be assumed to be, residents, and of ca. 100 the eggs have been collected and are described. The balance consists of visiting sea and freshwater birds, migrants from the North and migrants from the South. It is
likely that few additions remain to be made to the list of residents, but migrants from the North andvisiting seabirds have obviously been under-recorded, and in these two categories another 20 to 25 species are to be expected as regular visitors. Lonchura punctulata insulicola subspec. nov. is described from
Mauritius, Réunion and Mahé (Seychelles Is.).

Contents
1. Introduction ........................................................................................................................................................ 1
2. Acknowledgements ........................................................................................................................................ 3
3. Topography ......................................................................................................................................................... 4
4. Adjacent smaller islands .............................................................................................................................. 5
5. History of ornithological exploration .................................................................................................. 9
6. The egg collection (see also pp. 254-261) ........................................................................................ 19
7. Forsten’s collection from Bima, Sumbawa .................................................................................... 22
8. Systematic account ....................................................................................................................................... 22
9. Unconfirmed and erroneous records .............................................................................................. 219
10. Species and subspecies with type-locality Flores .................................................................... 222
11. Zoogeography ............................................................................................................................................... 223
12. List of localities ............................................................................................................................................. 228
13. Postscript .......................................................................................................................................................... 230
14. Bibliography and references ................................................................................................................ 233
15. Systematic index .......................................................................................................................................... 250
1. Introduction
That the Lesser Sunda Islands, forming a connecting link between the strikingly
different faunas of Asia and Australia, belong to the zoogeographically most fascinating regions of the world, is no news. Important zoogeographical studies have been
based on their avifauna (particularly Rensch, 1928d, 1930, 1936; Stresemann, 1939;
Mayr, 1944a, 1944b). Yet, in spite of an early start, and of their acknowledged crucial

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Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

importance for an understanding of the zoogeographical relationship between Asia
and Australia, most islands are at present no more than moderately well-known ornithologically.
Collections have been made during (usually short) visits by professional collectors,
heavily relying on native helpers, rather than by ornithologists, and much of the collecting has been rather haphazard.
For Flores, this situation has now been changed through the efforts of two long-time
residents, the Fathers J.A.J. Verheijen, SVD, and E. Schmutz, SVD, who, during theirstay, devoted much time to ornithological investigations. It has been my privilege to
receive and study the collections resulting from their activities, and the paper here presented is the outcome of my involvement.
This paper is far less comprehensive than I should have liked it to be. Many species,
dealt with only superficially, require a revision before their zoogeographical significance in the fauna can be evaluated properly, but any work is a compromise between
time and means available, and the desired result.
As in the preparation of their admirable work on the birds of Wallacea, White &Bruce (1986) have had free access to all the material and notes brought together by the
Fathers Verheijen and Schmutz, one might wonder, and indeed I have wondered,
whether after the appearance of their publication, there was still any need, or even justification, for a separate paper on Flores. However, it will be evident that a paper exclusively on the birds of Flores, can go into much greater detail than a work covering thewhole of Wallacea, and therefore deserves publication. In addition, there was the obligation to the collectors, who were entitled to see the published result of all their work.
Finally, it seems worth publishing these collections for their own sake. Complaints haverepeatedly been made, that collections present in the Leiden museum remained unpublished. The truth is that, for the size and importance of its collections, the museum has
always been understaffed and that the complaint was justified. Catalogues of the collections were begun by Schlegel, and anew by Finsch, but they never got very far. To savethe Flores collections made by the Fathers from oblivion, a paper was evidently required.
Authors of faunistic and systematic works aspiring to scientific accuracy are increasingly confronted with the question of what to do with field observations. At
present the ornithologically more sophisticated countries even have panels of specialists sitting in judgement over observations made by others, at best a risky matter. Thereis a danger here of taking ourselves too seriously. When, however, birdwatchers and‘ticklisters’ from the above-mentioned countries, where they would not hesitate to subject their unusual observations to the whole rigmarole, travel abroad, they suddenly
shed this serious approach and seem to believe that a cross on their pre-printed ticklist
should be all the documentation required to have their observations accepted. Thus theacceptance of such records, for which there is no tangible evidence, becomes guesswork
or a matter of faith. When my interest in the avifauna began, in the mid-nineteen sixties,
the problem did not yet exist as the Lesser Sunda Islands, even Bali, were well outside
the range of ornithotourism, which at that time was only just beginning to develop.
This situation is now changing rapidly.
A comparatively recent development, of which the region has not remained entirely
free, is that of multi-author papers, combining observations made by different observers

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

3at different times. Superficially this may seem rational, but frequently it is impossible
to distil from these texts, who is responsible for certain records and observations, sothat in practice these are anonymous. The custom, also becoming popular, of referring
to unpublished notes and reports, irritates: one tries to check a statement made in the
text with a reference, and finds it to be based on unpublished notes, and therefore of
no help at all.
Another aspect has to be mentioned: several recent visits to the Lesser Sunda Islands
have been made under the auspices of conservation organizations. Nothing wrong with
that, but in a few instances I got from the text of their reports the impression that actually bird material had been collected, but is not clearly mentioned, or glossed over (asconflicting with the avowed purpose of these visits?). This means that important documentation is withheld: a report should clearly state what has been collected, and inwhich institution such material is held, so that other students will know where to go forfurther information.
Originally I had not intended to include in this report any species not based on collected material, but gradually this has been relaxed, with the inclusion of Fregata ariel,
Pandion haliaetus, Hieraaetus kieneri and Phalaropus lobatus. This relaxation has also induced me to list some (not all) of my own field observations made on Sumba and Sumbawa, as far as they are relevant to the avifauna and understanding of the zoogeographical position of Flores.
2. Acknowledgements
From the introduction, my debt to Fathers Verheijen and Schmutz will be evident. It
is on their collections, and often stimulating correspondence, that this paper has been
based.
Help with information about specimens in their care, loans, etc., was received from:
D. Amadon and Mrs M.K. LeCroy (American Museum of Natural History, New York:
AMNH); G. Cowles, M. Walters (British Museum (Natural History), Tring: BM); R.W.R.J.
Dekker (Rijksmuseum van Natuurlijke Historie, Leiden: RMNH); S. Eck (Staatliches
Museum für Tierkunde, Dresden, MTD); K. Größler (Leipzig); J.T. Marshall, Jr. (National Fish and Wildlife Laboratory, Washington, D.C.); the late S.A. Parker (South Australian Museum, Adelaide: SAM); R.A. Paynter, Jr. (Museum of Comparative Zoology,
Cambridge, Mass.: MCZ); C.S. Roselaar (Zoölogisch Museum, Amsterdam: ZMA); H.
Schifter (Naturhistorisches Museum, Wien, MV); R. Schodde (Australian National Collection, Canberra: ANC, CSIRO); S. Somadikarta (Museum Zoologicum Bogoriense,
Bogor: MZB); B. Stephan (Zoologisches Museum, Berlin, ZMB); E. Sutter (Naturhistorisches Museum, Basel: NMB); C. Voisin (Muséum National d’Histoire Naturelle, Paris,
MHNP). Mr T.J.G.M. van Oyen (RMNH) helped with the weighing of egg-shells on anelectric balance. Acknowledgements are also due to A. van Haasteren (Leiden University) and H.J. van Grouw (RMNH) for photographing the eggs and to E.J. Bosch (RMNH)
for drawing the maps.
As also mentioned in the Postscript, without considerable assistance from myformer colleague, J. van der Land, this paper could not have been completed. In recent
years, my old friend A.C. van Bruggen helped greatly in keeping things moving. C.R.
Trainor, a field worker in the Lesser Sunda Islands (at present in Timor-Leste), recipient4

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)of a copy of the manuscript, encouraged its publication, writing stimulating letters bothto me and to Leiden, thus helping to push through bottlenecks.
With sadness I must mention that three more of the persons acknowledged above(Amadon, Paynter, Sutter) have passed away. I gratefully remember their friendshipand help, not only in the preparation of the present paper, but on many previous occasions.
3. Topography
The island of Flores derives its name from its easternmost cape, which was namedby the Portuguese, the first European navigators to reach the Lesser Sunda Islands,
Cabo de Flores. The name has stuck and has been extended to include the whole island, although the name of the cape itself has in later years been translated to Tandjong
Boenga.
Until past the middle of the 19th century, Portugal maintained some posts on theeastern and southern coasts of Flores, but by the treaty of Lisbon, which was ratified on
20 April 1859, these posts (Paga, Sikka, and Larantoeka with the fort Posto), were transferred to the Netherlands, by which the whole island came, nominally, under Dutch
sovereignty. A garrison was established in Larantoeka. It is perhaps worth mention thatthe colonial yoke was not very heavy for at least another forty years: inland tribes continued to raid the coastal villages. When in July 1896, Everett arrived in Endeh, there
was even shooting down the streets, so that he was unable to use the place as a base for
exploration of the interior, as he had planned. In 1907, during a conflict between coastal
and mountain tribes, the whole of Endeh was burned down. This finally led to intervention by the Netherlands Indian Government, and a stronger colonial administration.

Rinca

Fig. 1. Flores and surrounding islands.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

5

Flores is situated between 119°48’ (Tg. Rasé) and 123°1’30” (coast south of Tg.
Matandoi) E., and between 8°4’ (Toro Kopondai or Floreshoofd) and 8°58’ (Ngaloe
Nageh, south coast of Ngada) S., its greatest length is ca. 354 km, its axis almost WestEast (fig. 1). The shape of the island is somewhat bizarre, perhaps best likened to thebody of a scorpion, with its heaviest, broadest part in the west, tapering eastwards to atail, terminating in a sting, curved upwards, at the eastern end. In the western part
(Manggarai), the width is over 60 km, but in the middle (Endeh), it is no more than 20km, and farther east, at the narrowest part (Maumere) only ca. 12 km. The total (flat)
surface area is ca. 15,175 km2. The country is strongly mountainous: extensive highlands, and the highest mountains, are found in the broad western part (Manggarai),
where several peaks attain over 2000 m, the highest one ca. 2400 m; in the middle andeast the mountains may be lower but are by no means negligible. The famous coloured
crater lakes of Geli-Moetoe are in the eastern part of the island, north-east of Endeh.
In the mountains the rainfall is high (fig. 2), especially in Manggarai: in the mountains south and west of Ruteng it averages over 4000 mm a year, and in the mountains
farther eastwards it is still over 3000 mm; this contrasts with some of the coastal lowlands, especially along the north-west coast, where the annual average is less than 500
mm. The gradient is very steep here, for the mountains with the highest rainfall are little
more than 25 km away.
4. Adjacent smaller islands
The islands of Endeh, in the Bay of Endeh off the south coast (visited by Rensch),
Palué or Rusa Radja in the Flores Sea north of Flores (visited by Weber and Verheijen),
and Groot Bastaard or P. Besar, off the north coast of Flores (visited by Ten Kate), havealways and quite correctly been included with Flores, ornithologically, as they are inthis paper.

0
100
0
200

30
20000
0

500

3000

10

1000
0
200

10
00

00

40

00

Fig. 2. Average annual rainfall in mm on Flores.

0
200
3000

00
10

0

0
100 0
200

200

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Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

It would make sense, geographically, to include the smaller islands to the west, viz.,
Rintja, Padar and Komodo, with Flores. Strait Molo, separating Rintja from mainland
Flores, is at its narrowest part less than a kilometer wide; the strait between Rintja andPadar, perhaps 2 km, and Strait Linta, separating Komodo from Rintja and Padar, about
5 km. Strait Sape, on the other hand, which separates Komodo from eastern Sumbawa,
is much wider and deeper, about 20 km, or 15 + 5 km when measured through the small
island of Kelapa.
The avifauna of these islands has become known through Hoogerwerf (1954b, 1956),
who visited all three of them in June 1953. He claimed that the total number of species
observed on the three islands combined, was 69. An unrecorded, but probably fairly
large, proportion of these were collected. It is a pity that Hoogerwerf’s published paper,
through its ecological arrangement (according to habitat), rather than systematic arrangement, makes it difficult to extract exact information from it, and has alreadycaused confusion. Moreover, as pointed out on a later page, some species included were
almost certainly misidentified. This would leave about 65 species for the three islands.
Especially because of doubt of some of them, a list of the collections made, split out toisland, would be useful. Hoogerwerf (1966) wrote a special paper on a few specimensfrom Komodo and Rintja, belonging to three species (Accipiter novaehollandiae, Coracina
novaehollandiae, Philemon buceroides), showing that a further study of the avifauna ofthese islands would be not without interest. Pfeffer stayed some weeks on Rintja, andvisited Komodo; on Rintja he collected seven species of birds, enumerated in the historical section (p. 16), one of which (Stiltia isabella) was not recorded by Hoogerwerf.
The eastern end of Flores is in equally close contact with adjacent islands. Opposite
Larantoeka, across the Flores Strait, Adonara is at most little more than a kilometer away,
and farther southwards, the distance from Solor is less than 5 km. Ornithologically,
Adonara and Solor are very inadequately known. Semmelink visited Adonara early in1862 and collected one bird (Pitta elegans), Colfs in August 1880 at least two (Arenaria
interpres, Terpsiphone paradisi), and Ten Kate spent a few days on Adonara and Solor in
May 1891. On Adonara, Ten Kate collected Halcyon chloris, Pachycephala fulvotincta, Anthreptes malacensis (cf. Büttikofer, 1892), and Oriolus chinensis (cf. p. 12). On his way to orfrom Flores, Allen must have landed on Solor, where he obtained a syntype of Treron
floris and the holotype of Gerygone sulphurea. I have not investigated what else he may
have collected on Solor: scanning of the pages of the “Catalogue of Birds” might yielda few more species. As mentioned above, Ten Kate visited Solor, but I am not aware thathe collected birds there.
From the western islands, Hoogerwerf (1954b, 1956) recorded eight species which
are not known from Flores, viz.:
Falco severus: island not specified
Charadrius peronii: Komodo and Padar; on Rintja heard but not seen; a specimen was collected
Numenius arquata: Padar
Tringa totanus: Padar
Ducula bicolor (with a query, see below)
Caloenas nicobarica: Komodo
Cypsiurus parvus (= balasiensis): island not specified
Dendrocopos macei: island not specified

The record of Ducula bicolor was, as Hoogerwerf (1954b: 130) clearly stated, basedon an observation by the photographer F. Huysmans, a non-ornithologist, who saw aMees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

7flock of white birds with black wing-tips come from the sea and disappear into a valley.
Hoogerwerf (who was obviously responsible for the identification) commented that theoccurrence of the species was of course very likely, but stressed the point that he himself
had not seen it. He mentioned Ducula bicolor again, with a query, but this time without
explanatory text, in his following publication (Hoogerwerf, 1956: 111). I consider theevidence too vague for acceptance.
Besides Ducula bicolor, the identification of at least three other species on the above
list may be questioned: Falco severus, Cypsiurus balasiensis and Dendrocopos macei havenever before been recorded from the Lesser Sunda Islands east of Bali. The casual way
in which Hoogerwerf mentions them, shows a complete lack of awareness that observations of these species were in any way unusual. In the case of Falco severus, one canthink of confusion with F. longipennis, not mentioned by Hoogerwerf. In the case ofCypsiurus balasiensis, it may be recalled that its occurrence in Sulawesi has recently
been established (Escott & Holmes, 1980): very likely a recent extension of its range.
The same could be happening in the Lesser Sunda Islands, for, as Hoogerwerf said,
there is no lack of suitable nesting trees. Hoogerwerf’s record of Caloenas nicobarica
from Komodo is acceptable, but the listing of Padar and Rintja by White & Bruce (1986:
192-193) is not.
Hoogerwerf was surprised by the number of migrant waders from the Northern
Hemisphere which he found still present in June. He observed Pluvialis fulva, Charadrius
leschenaulti, Numenius arquata, Numenius phaeopus, Tringa hypoleucos, Tringa totanus.
With the knowledge in mind that Hoogerwerf has made misidentifications, one
might legitimately wonder about the validity of the June records of these waders. Restricting myself to the two species not yet recorded from Flores, I have tried to find how
likely it is that they still occur in their winter quarters in June.
Material from the Archipelago in the RMNH collection, split out over the months, isgiven in the table below, which shows that June records of neither species are exceptional. When Junge (1936: 18) discussed the two June specimens of Tringa totanus from
Simalur, he was worried by their dates, and by their winter plumage, and speculated
that an error in labelling might have been made. This is very unlikely, because bothhave original labels. In the light of present knowledge, there is no reason to doubt theJune dates. One other point has, however, to be mentioned. Whereas in the western partof the Archipelago, Numenius arquata is a more common winter visitor than N. madagascariensis (cf. Hoogerwerf, 1948b: 54), farther east, east of the Sunda Shelf, the situation isreversed to the extent that there is only a single specimen record of N. arquata from thewhole of Wallacea (the specimen from Halmahera, listed above: , 20.xi.1861, Kaou,leg. Bernstein, RMNH cat. no. 17; cf. Schlegel, 1864c: 89, where it is cat. no. 29). One
might therefore think of confusion between these two, but Hoogerwerf certainly knew
the difference and moreover, in his first publication (Hoogerwerf, 1954b: 145) he states
expressly that the bird he had seen belonged to the large white-rumped species. Further
support is provided by recent field-observations from Timor (Andrew, 1986). Records
from Australia (Darwin and Point Peron) have not found general acceptance (cf. Blakers et al., 1984: 656; Higgins & Davies, 1996: 116-117), and a sight record from New
Guinea (Van den Assem, 1960) has been rightly ignored in later literature: it was almost
certainly due to confusion with N. madagascariensis.

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Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Material of Numenius arquata and Tringa totanus from the archipelago in the RMNH collection, arranged
by month.
s.d.
I
II
III
IV
V
VI
VII VIII IX
X
XI
XII
Numenius arquata
Sumatra
1
–
–
–
–
–
–
–
–
–
4
–
–
Bangka
1
1
–
–
–
–
–
–
–
1
–
–
–
Borneo
2
–
–
–
–
–
–
–
–
–
–
–
–
Java
6
–
3
–
3
–
1
1
2
5
–
6
1
Halmahera
–
–
–
–
–
–
–
–
–
–
–
1
–
Tringa totanus
Sri Lanka
3
–
1
–
–
–
–
–
–
–
–
–
–
Sumatra
1
–
–
–
–
–
–
–
–
–
4
–
3
Simalur
–
–
–
–
–
–
2
–
–
3
–
–
–
Bangka
–
–
–
–
–
–
–
–
–
1
2
–
–
Borneo
2
–
–
–
–
–
–
–
1
–
–
–
–
Bawean
1
–
–
–
–
–
–
–
–
–
–
–
–
Java
6
5
7
–
2
13
3
–
13
9
3
5
5

How these (and other) records were transferred to Rintja, and claimed by Pfeffer(1958) as his own, will be explained in the next section.
Before leaving the western islands, I want to mention that on 19.ix.1984, from theislet of Sabita, off the east coast of Komodo, I observed an individual of Numenius madagascariensis (very large, brown rump, calling), flying past over the strait separating Sabita from an even smaller islet to its south.
Of the few birds recorded from the eastern islands Adonara and Solor, only one,
Arenaria interpres, is not yet known from Flores proper.
The preceding evaluation leaves six species which would be added to the avifaunaof Flores if the satellite islands to the east and to the west were included: Charadrius
peronii, Numenius arquata, Numenius madagascariensis, Tringa totanus, Arenaria interpres,and Caloenas nicobarica.
Addendum.— A summary of the species added to the avifauna of Flores on thebasis of sight records, published by Verhoeye & Holmes (1998) is given in the Postscript. This leaves one form recorded by these authors from Komodo but not from
Flores, which therefore deserves mention in this chapter. It is Anas gracilis (which theauthors prefer to treat as a separate species, not as a subspecies of A. gibberifrons). Five
individuals were seen in a small pond behind the beach on 7.vii.1992. The record is notunlikely, but is the first for any of the Lesser Suna Islands. I regret the absence of any
information on how the birds were identified: females and juveniles of nominate gibberifrons do not have the high forehead of the drakes and must be very difficult todistinguish from gracilis in the field. This is also why I prefer to keep the two forms
together in one species.
Through a short visit (14/18.xii.2000) by Trainor (2002a) the island of Adonare has
become ornithologically better known. As was to be expected its avifauna is impoverished compared with that of Flores. Not only its small size as such (497 km2), moderate
elevations (although with a peak reaching 1659 m) and fairly low annual rainfall (10002000 mm) would be responsible but certainly not in the last place the dense humanpopulation (165/km2). The list of about fifty species is not entirely critical, some species
having been included on the basis of hearsay information from locals.

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9

Trichoglossus euteles is the only species on the Adonare list not known from Flores.
Considering the proximity of Adonare to Flores, where T. euteles is replaced by T. weberi,
the relationship between these two parrots requires further study. The occurrence ofHalcyon fulgida and Tesia everetti on Adonara constitutes a slightly eastward extension of
their known ranges. The same might be true for Zosterops palpebrosa, listed as common,without comment: it is a species I would not expect on Adonara, where Z. chloris, notmentioned by Trainor, is more likely to occur.
5. History of ornithological exploration
The first two collectors who worked on Flores: Semmelink for the Rijksmuseum van
Natuurlijke Historie, and Allen on behalf of A.R. Wallace, were both there in 1862. Unfortunately, very few specimens belonging to either of these early Flores collections aredated exactly. But Semmelink is known to have collected a bird on the second of Marchon the off-shore island of Adonara, so it may be presumed that he was present on Flores
for the whole calendar year. I have seen no dated specimen of Allen’s, but consideringthat he only stayed for four months on Flores, chances are that he started collecting
later than Semmelink. Therefore I have decided to place Semmelink ahead of Allen in
the chronological sequence of collectors.
J. Semmelink (1837-1912), Officier van Gezondheid (Military Surgeon), was stationed at Larantoeka, where there was a garrison, during 1862/1863. He collected material of various animal groups for the Rijksmuseum van Natuurlijke Historie, andpossessed a private shell-collection. According to our register, 55 bird skins were received on 19.viii.1863, and one more on 15.xii.1863. These birds were, with a few exceptions, not individually labelled, but Semmelink wrote that they had been: “Verzameld te Larantoeka en omliggende plaatsen in den loop van 1862”. Included in the collection was one specimen from Adonara, which is an island only a few kilometers off
Larantoeka (Pitta elegans concinna, 2.iii.1862, Adonara, RMNH cat. no. 4; cf. Schlegel,
1874a: 14). I have been able to find 45 species collected by him, 9 of which were notobtained by Allen. In spite of the small size of the collection, it contains several uncommon species: Ardea alba, Spizaetus floris, Circaetus gallicus, Ptilinopus regina flavicollis,
Dicaeum agile tinctum.
C. Allen (ca. 1838 - ca. 1900), spent almost four months of 1862 collecting on Flores(cf. Wallace, 1864: 480), but there is no exact information (dates and localities) on his
stay. It does not look as if Wallace had given his assistant very clear instructions, or as if
he was interested in proper data (sex, date, locality) of the collected material, for Weber(1890/1891: vii) wrote: “Auch brieflich konnte mir Herr Wallace keine Auskunft ertheilen, welche Gegend von Flores Ch. Allen besuchte”. Allen’s labels give merely theyear of collecting, and the name of the island.
Only speculation is therefore possible about Allen’s stay. I speculate, then, that
Larantoeka is the most obvious place for him to have visited: at that time it was the
only place with a garrison, and therefore the only place with regular connections withthe outside world. The fact that Allen also visited Solor, just off Larantoeka, supports
this speculation (cf. Wallace, 1869, II: 385). Rensch (1931a: 454) also supposed that
Allen collected: “wohl hauptsächlich im Osten der Insel”. On the other hand, if Allen
had stayed in Larantoeka for any length of time one would expect him to have met

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Semmelink, and to have been mentioned in one of Semmelink’s letters to Schlegel.
According to the list published by Wallace (1864), Allen collected 86 species of birds
on Flores. A few of these records are doubtful, being either based on misidentification,or on specimens erroneously labelled as having been obtained on Flores. For example,
Munia ferruginea (= Lonchura malacca ferruginosa) is in its natural distribution confined
to Java and is not known from any of the Lesser Sunda Islands. As Wallace also enumerates all four species of Lonchura known from Flores, the specimen may come in thesecond rather than in the first category. It is possible that the specimen still exists somewhere, but I have not traced it.
More serious is the case of Trichoglossus euteles, of which Allen is supposed to havecollected several individuals on Flores (cf. Hartert, 1898a: 43-44; Hellmayr, 1914: 78). T.
euteles is known from Timor and from the smaller islands to the east of Flores, from
Lomblen to Nila. Chances that it might be present on Flores are remote. This shows thatmaterial, forwarded from Flores by Allen, may have originated elsewhere. Therefore,
species which have been recorded from Flores only by Allen, and the occurrence of
which has not been confirmed by subsequent collectors, have to be considered very
critically. Besides the two species already mentioned, Lonchura malacca ferruginosa andTrichoglossus euteles, this concerns the following: Gallinula tenebrosa frontata, Loriculus
flosculus, Petrochelidon nigricans timoriensis and Lichmera indistincta limbata. The occurrence of none of these four species on Flores is a priori improbable, and therefore I havefound no reason to eliminate them from the list.
Wallace was a professional collector, whose material has been widely dispersed
through trade channels. At least a large proportion of his material, including types ofthe majority of species, has gone to the British Museum, either directly, or later, throughbequests of private collections. There is also material in the RMNH, and undoubtedly
in many other museums.
Very little is to be found in the literature about the person of Charles Allen. Wallace(1869, I: 72) says that, in May 1855, he was: “an English lad of sixteen”, which places hisdate of birth in 1838 or early 1839. Towards the end of 1862 (id., II: 385-386), “he obtained employment in Singapore, and I lost his services as a collector”. His name ismentioned a few times in Wallace’s published letters (Marchant, 1916: 40, 48, 49, 51, 6061, 79): Wallace and Allen apparently (but not quite certainly) travelled on the sameboat from England, and arrived together in Sarawak, in 1854, and when Wallace left,Allen stayed behind with the Bishop of Sarawak: “who wants teachers and is going to
try to educate him for one”. The most informing is a note inserted by L.V. Helmes(Bishop of Sarawak): “When Wallace left Sarawak after his fifteen months’ residence inthe country, he left his young assistant, Charles Allen, there. He entered my service, andremained some time after the formation of the Borneo Company. Later, he again joined
Wallace, and then went to New Guinea, doing valuable collecting and exploring work.
He finally settled in Singapore, where I met him in 1899. He had married and was doing
well; but he died not long after my interview with him. He had come to the East with
Wallace as a lad of 16, and had been his faithful companion and assistant during years
of arduous work”.
I cannot help feeling that Wallace has barely done justice to Allen who, young as hewas, worked independently on the Sula Islands, Morotai, Misool, Flores, evidence that
Wallace had full confidence in his abilities and integrity. From Allen’s Flores collection

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)11

alone 18 new species and subspecies of birds were described (mainly by Wallace andSharpe). There is an Accipiter fasciatus wallacii, occurring on Flores, and numerous otherspecies bear Wallace’s name. But Allen, the collector of many of these species, has notbeen honoured a single time in such a manner, and therefore his name is not mentionedin Wynne’s (1969) useful work.
Addendum.— Recently I came across another reference to Allen, taken from Wallace’s letters to his sister (Severin, 1999: 192-193). It is apparent that Wallace brought
Allen out with him from England as his assistant. Later, in Sarawak, Wallace considered the boy useless, so that he was happy to leave him behind with the Bishop of
Sarawak. I find it astonishing that Wallace, who was not a wealthy man, should havetaken the boy with him without first ascertaining his qualities (or the lack thereof). If
Allen was as incompetent as Wallace wrote, it is even less explicable that he re-entered
Wallace’s service later. In summary, I do not believe that my remarks given above areunfair to Wallace. I also realise that letters written by Wallace to his sister were notmeant for publication and that he may well have changed his opinion on Allen in later
years.
K.E. von Martens (1831-1904) was a zoologist of the “Preussische Expedition nach
Ost-Asien”. From 6-30 January 1863, he stayed in Larantoeka, as a guest of Dr Semmelink. In this period, he also made trips to Adonara and Solor, off Larantoeka, without
apparently collecting any birds. He did, however, collect a few near Larantoeka: Tinnunculus Moluccensis (= Falco moluccensis), Cacatua sulfurea, Scythrops Novae Hollandiae, Halcyon collaris (= Halcyon chloris), Tropidorynchus sp. indet. (= Philemon buceroides), Nectarinia
solaris, Gracula cf. venerata (= Gracula religiosa venerata), Treron sp. indet. (= Treron floris)
and Geopelia striata (cf. Martens, 1876: 244, 365-373). These nine species had also been collected near Larantoeka by Semmelink and (possibly near there) by Allen the year before,
so that Martens’s contribution to the ornithological knowledge of Flores is negligible.
In a letter addressed to Schlegel, dated 23.iii.1863, Semmelink mentioned this visit:“Dr Eduard von Martens uit Berlijn, die toen bij mij logeerde, was mij behulpzaam bij
het bestemmen en inpakken der zaken en stelde mij in de gelegenheid UEd. eene opgave van de vogels te kunnen zenden, die naar Uwe Handleiding tot de beoefening derDierkunde bestemd werden” 1).
P.F.A. Colfs (1853-1882) was a collector employed by Governor-General J.W. van
Lansberge. Colfs stayed in West Flores from 7.ii-25.iii and from 11.iv-15.vi.1880; on
16.vi.1880 he left by ship for Maumeri (Maoemere), where he arrived on the 19th. Helater sailed regularly between East Flores, Timor and Alor, although he did not visit
Flores (Larantoeka) after 29.ix.1880. In 1882 the RMNH received from Van Lansberge
143 bird specimens from Sumbawa, Flores, Ombaai (= Alor) and Timor. A list of thismaterial, dated 31.vii.1884, was prepared by Büttikofer; it includes 36 specimens in 23
species marked as being from Flores, but two of these are from Adonara, and of one,
Zoothera peronii, an error in labelling may be suspected. Vorderman’s (1888) little volume is less informative than one might hope, and throws no further light on Colfs’s
collecting activities.

1)

Dr Eduard von Martens of Berlin, who was my guest at that time, assisted me with identifying andpacking of the items and enabled me to send Your Honour a list of the birds, named according to yourHandleiding tot de beoefening der Dierkunde.

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Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

M.W.C. (Max) Weber (1852-1937) explored Flores from 21.xi.1888 to 9.i.1889. In the
introduction to his work, Weber (1890) gives an itinerary, but the only dates mentioned
in it are those of arrival and departure. I owe Mr Roselaar the following approximate
reconstruction of Weber’s itinerary: Reo (23/25 November), Bari (26/28 November),island Rusa Radja, now known as Paloë or Palué (29 November), Maumeri (1/18 December), Kotting and Mbawa (19/24 December), Sikka (25/31 December), Endeh (1/8January 1889), Rokka near Endeh (9 January). The ornithological results of Weber’svisit were quite important; they included the discovery of several undescribed species(Büttikofer, 1894). The greater part of the material (66 skins, 13 skeletons) was retainedin Leiden, but a considerable proportion (52 specimens) was returned to the Zoölogisch Museum in Amsterdam (of which Weber was Director). A total of 60 species was
obtained.
H.F.C. ten Kate (1858-1931) visited Flores in 1891, his itinerary being as follows:
Endeh (28/30 January); on the last-mentioned date he left for Timor, but he returned to
Flores in April: Sika and surroundings (13/24 April); from there, he walked to Koting,where he stayed three days until the 27th, and then walked on to Maumeri. On 1 May,he sailed from Maumeri to Groot Bastaard; on 5 May, he arrived at Hading, from where
he went overland to Larantoeka; on 11 May, he left for Adonara and Solor, returning toLarantoeka on the 17th. The next two days were spent writing and packing, and on 20
May, he travelled on to Koepang (cf. Ten Kate, 1894). In his first article on Ten Kate’s
birds, Büttikofer (1891), lists only three birds from Flores: Artamus leucorhynchus (1)and Philemon buceroides neglectus (2, syntypes of neglectus); in his following article, 11
species from Flores are enumerated (including Groot Bastaard, but without Adonara).
As one of the 11 was a species mentioned in the 1891 paper, Ten Kate’s total score was
12 (Büttikofer, 1892).
A.H. Everett (1848-1898), a famous professional collector in the service of Lord
Rothschild, visited Endeh in August 1896, but was not allowed into the interior because of unsettled conditions, and left almost at once (for Savu), after having shot only
a few common birds. On his way to Endeh, he seems to have called at Maumeri (cf. p.
86). Accompanied by his “native collectors”, he returned to Flores in October of thesame year, making his headquarters at Nanga Ramau (Nangaramoe, Nanga Roma,
Nanga Ramo, Nanga Ramut), a village on the south coast of Manggarai, in western
Flores. From his account (in Hartert, 1897b: 513-514), it is apparent that his hunterswere unable to penetrate very far inland, but did succeed in reaching levels of about
5000 ft. in the directions of Puchu Reah (not identified, but probably towards the Golo
Radjong, 1490 m) and Puchu Leoh (probably = Potjo-Léok with Golo Mompong, 1383m), all within a radius of ca. 10 km of Nanga Ramau. The material sent home by him
included a number of typical mountain birds, thus confirming this altitude. The exact
duration of his stay is apparently not known: his material is dated October and November 1896, and in December he was on Sumba, so that his stay cannot have lasted
more than about six weeks. In his various publications, Hartert is very reticent about
Everett’s exploits, and no exact information is provided by him. I have enquired
whether there are any letters in the BM-archives (into which the archives of the former
Rothschild Museum have been incorporated), which might provide more exact data,
but the reply from the Archivist was, unexpectedly, that the Rothschild correspondence for 1896 does not include any letters from A.E. Everett (Thackray, in litt., 11.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

13

x.1993). Subsequently, Mr Walters (in litt., 4.xi.1993) enlightened me: “I deeply regret
to have to say that letters to Rothschild/Hartert, if not among the Rothschild correspondence in London, are certainly destroyed. Both Rothschild in his lifetime, andsubsequently ‘officials’ of the BMNH, burned sackfull after sackfull of Rothschild’s
papers...even as late as the 1960s...Nobody knows what was destroyed...but it is fairly
certain that this included the catalogues of Rothschild’s mounted skins, and Max
Kuschel’s catalogue of Count Roedern’s egg collection. The loss of the Roedern catalogue rendered this fine collection almost valueless”.
The number of species collected was 114, of which some 15 species and subspecies
were new (mainly from the previously unexplored mountains): an impressive result for
a comparatively short stay. The material went to the Tring Museum, and most of it is
now in the American Museum of Natural History, but, soon after it was received inTring, some of it was sold to the dealer Rosenberg and became widely dispersed. Unfortunately, Hartert (1897a, 1897b, 1898a) only recorded the actual number of specimens
collected when they were few. His text abounds with expressions like: “a fine series”,“several skins”, etc. Rensch (1931a: 455), placed Everett’s collecting work much too far
east: “in Mittelflores (wohl hauptsächlich Gebiet des Keo- und Inerie-Vulkans)”.
G.A.J. van der Sande (1863-1910), a Naval Surgeon, collected 19 birds on the islandsof Flores and Sumba, at various times between December 1907 and March 1909. These
are in the Zoölogisch Museum, Amsterdam. Of this total, 11, belonging to 10 species,
are from Flores. Three species, Elanus caeruleus, Haliaeetus leucogaster and Esacus magnirostris, were new records for Flores. For biographical particulars of Van der Sande, I
refer to Pulle (1910).
J. Elbert (1878-1915), leader of the “Sunda-Expedition”, visited several of the Lesser
Sunda Islands as well as Sulawesi (Celebes), and fairly large collections of birds were
made. His stay on Flores was, however, short: 24.I-10.ii.1910. According to Van Bemmel& Voous (1951: 28), Elbert’s bird collections were divided between: “the Buitenzorg
Museum and the Senckenberg Museum at Frankfurt; some of the specimens are now inthe museums at Amsterdam and Leiden”. This does not look as if the authors were
aware that 105 specimens were presented to the RMNH directly (100 of which remain;
five were sent to the MZB). Rensch (1931a) discussed the material from the Lesser Sunda Islands in the Senckenberg Museum. He did not mention specimens from Flores. In
Leiden, there is only a single specimen from Flores: Eudynamys malayana. But there areseveral interesting species from other islands, such as two specimens of Dendrocygna
javanica from Lombok.
Endih, the anonymous “Javanese” preparator mentioned by Rensch (1931b: 400),
collected a few birds at Réo on the north coast in 1911. The number of species listed by
Rensch is nine, the number of specimens 13. It seems, however, that Rensch has missed
some of Endih’s specimens (cf. Somadikarta & Noerdjito, 1982). In spite of its small size,
this collection is important, as it contains a specimen of Falco cenchroides, the only record
from Flores.
B. Rensch (1900-1990), as participant in an expedition which included also his wife,
the botanist Ilse Rensch-Maier, the anthropologists G. Heberer and W. Lehmann, andthe herpetologist R. Mertens, stayed on Flores from 6.vi-25.vii.1927. Rensch arrived at
Endeh. A visit of two days was made to Poeloe Endeh; a motor vehicle was rented,
with which he travelled over the east-west road, which was at that time still under14

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

construction, to Badjawa (1200 m), Aimere (coast), Mborong (coast), Sita (700 m), Rana
Mesé (1200-1400 m) and Waë Renó (1000 m). After the return to Endeh, over the sameroad, a shorter excursion was made in an easterly direction, to the Geli Moetoe (ca. 1500m). Descriptions of the main collecting-localities are to be found in Rensch (1931a: 459460). Thanks to the help of four experienced preparators (“Fleißige und anstellige Leute”)
made available by the Zoölogisch Museum Buitenzorg (the senior one of whom was
Denin, the bird-preparator Darna), Rensch managed to bring together a collection of 91
species of birds in this short period. Several of these were new for Flores. He described
10 new subspecies from this collection, four of which are accepted here.
J.K. de Jong (1895-1972), at that time on the staff of the Zoölogisch Museum Buitenzorg, and assisted by two preparators from that Museum (one of whom was the mantri
Madzoed), made a collection of birds in western Flores (Mboera, Laboean Badjo, Wai
Sano), in October-November 1929. The list, of 69 species, was published by Rensch(1931b). Several species were new for Flores; one of these was described by Rensch as a
new subspecies, Collocalia francica dammermani (= Collocalia fuciphaga fuciphaga). It may
be assumed that most of the ornithological collecting was done by Madzoed, as De Jong
was a herpetologist, and his main interest was in Varanus komodoensis.
Jonkheer W.C. van Heurn (1887-1972), in the first half of May 1930, spent about tendays on Flores. He had no opportunity for ornithological collecting, and only in thesecond of his two articles (Van Heurn, 1931, 1932) are a few birds mentioned: Streptopelia chinensis, Geopelia striata, Cacatua sulphurea, Graucalus sp. (= Coracina sp.), Saxicola
caprata, Philemon buceroides. More interestingly, a group of 12 Ciconia episcopus was seenin the lowlands near Reo, and Gallus varius seemed to be particularly common near
Pota. Bee-eaters listed as Melittophagus leschenaulti would have been Merops ornatus.
Some unidentified herons, ducks and noisy green parrots, probably of more than one
species, complete the list of his observations.
E.R. Sutter (1914-1999), a member of the Swiss “Sumba-Expedition” spent, on the
return voyage from Sumba, in November 1949, some three weeks on Flores (cf. Sutter& Wegner in Bühler & Sutter, 1951: 214-215). In this period he collected only two birds,
both at Todabelu, and both on 13.xi.1949: one Turnix maculosa floresiana, and one Cecropis
striolata ( ad.) from the breeding-colony at the school. Notwithstanding the modest
ornithological results of this stay, it requires mention here, as one of the specimens has
been recorded in the literature (Sutter, 1955: 122). This would make readers wonder,whether more had been collected (as it did me), so that the point had to be cleared up.
J.A.J. Verheijen, SVD (1908-1997). Father Verheijen arrived on Flores as a missionary
in 1935 and, apart from a period of internment during the second world war, has remained there until 1993, when he retired to a monastery in the Netherlands. His interestin natural history in general and birds in particular was aroused and stimulated by
L. Coomans de Ruiter (1898-1972) during their stay in Japanese internment camps in
Sulawesi (Celebes) (cf. Verheijen, 1964: 194), with the result that after his return to Flores
he started observing birds and from 1952 onwards assembled a large collection of eggs,
later also of bird-skins.
Originally (ca. 1954-1960), the skins were sent for identification to the Museum of
Comparative Zoölogy, Cambridge, Mass. This first collection already contained anumber of additions to the known avifauna of Flores (cf. Paynter, 1963). The MCZ-collection consists of 335 skins, of which 34 are from Palué (Turnix suscitator, Amaurornis

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

15

phoenicurus, Ptilinopus melanospilus, Lalage sueurii, Rhipidura rufifrons, Monarcha cinerascens, Parus major, Nectarinia jugularis, and Zosterops chloris). The total number of species
in this collection is 88, of which 8 (3 from Palué) are not represented from Flores in the
RMNH-collections: Porzana pusilla, Gallinago stenura, Chlidonias hybridus, Tyto longimembris, Lalage sueurii, Rhipidura rufifrons, Monarcha cinerascens, and Lonchura pallida. In thisperiod, specimens were also sent to Bogor (cf. Jany, 1955). The later material (ca. 19681978), as well as some material, mostly common species, from the first collection, cameto Leiden and forms the main subject of this paper. The bulk of the skin collection isfrom Ruteng and its wider surroundings, in central Manggarai, as Father Verheijen lived
in Ruteng when it was assembled (figs 3-4).
Very important is the large collection of over 4000 eggs, belonging to ca. 100 species,
mainly brought together in the period 1952/1962. In Verheijen’s (1964) paper, based on
this collection, he acknowledges the help of several assistants and colleagues; amongst
the latter the Fathers J. Geeraeds (Rekas), J. Klizan (Borong), A. Mommersteeg (Soa),and P. Rehmet (Nanga Rema and other localities). The greater part of the egg collectionis also from Manggarai, but it includes material from the island of Palué (Verheijen,
1961), from Soa (received from Father Mommersteeg, as just mentioned), and from Mataloko (=Todabelu), where Father Verheijen worked in 1960 and 1961/62. Soa and Mataloko are outside Manggarai, in south-western Ngada. The collection was purchased by
the Rijksmuseum van Natuurlijke Historie in 1973, with the exception only of the eggs

Fig. 3. Kedaluans (districts) of Manggarai (Western Flores). 1. Nggorang; 2. Mburak; 3. Mata-Waé; 4.
Boléng; 5. Kempo; 6. Badjo; 7. Patjar; 8. Wélak; 9. Wontong; 10. Mules; 11. Berit; 12. Kolang; 13. Lelak; 14.
Todo; 15. Rého; 16. Ndoso; 17. Rahong; 18. Pongkor; 19. Nggalak; 20. Ruis; 21. Tjibal; 22. Ruteng; 23.
Potjo-Léok; 24. Lamba-Léda; 25. Ndé; 26. Sita; 27. Torok-Golo; 28. Riwu; 29. Tjo; 30. Manus; 31. Kepo; 32.
Rongga-Koé; 33. Biting; 34. Rembong; 35. Radjong.

16

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)of the parasitic cuckoos, which had previously been sold separately to J. Ottow. Thismeans that eggs of the following species, although collected by Verheijen, are not represented in the collection acquired: Scythrops novaehollandiae, Eudynamys scolopacea, Cacomantis variolosus. Some eggs are shown in figs 12-18 on pp. 254-261.
P. Pfeffer (*1927) visited the islands of Komodo, Rintja and Flores between 18.iv and14.vii.1956. He spent five weeks of this period on Flores: in the South at Endeh and‘Mont Keli-Mutu’ (one day), and in the West around Labuanbadjo, Soknar and Lenteng.
His report (Pfeffer, 1958) is very confused; he gives an ecological classification of vegetation-types, listing the birds occurring in each, but there is not a single date or locality;
frequently, it is not even clear whether Komodo, Rintja, or Flores are meant. Many
records of birds are so vague that one suspects them not to be based on personal observations, but copied from the literature. This suspicion becomes a certainty when one
compares his article with Hoogerwerf’s (1956) paper on the birds of Komodo, Padar
and Rintja; Pfeffer’s paper is little more than a translation of Hoogerwerf’s contribution, except that for Flores he has added a few species highly unlikely to have beenfound in the coastal lowlands around Soknar, where he stayed most of the time, e.g.
Accipiter virgatus, Cuculus poliocephalus (= C. saturatus lepidus) and Brachypteryx floris.
Much of Hoogerwerf’s (1956) paper was based, almost verbatim, on the ornithologicalchapters of a report with a limited distribution, issued two years earlier (Hoogerwerf,
1954b) and Pfeffer has apparently used the report rather than the publication. Very

Fig. 4. Localities in Western Flores. 1. Labuanbadjo; 2. Mburak; 3. Renari; 4. Soknar; 5. Look; 6. Tjeréng;
7. Paku; 8. Naga; 9. Nisar; 10. Waé Radja; 11. Sésok; 12. Nunang; 13. Waé Sano; 14. Goang; 15. Waé Wako;
16. Tado; 17. Rekas; 18. Waé Djamal; 19. Nanga-Lili; 20. Bari; 21. Orong; 22. Léma; 23. Déngé; 24. Lamba;
25. Réo; 26. Rua; 27. Tjantjar; 28. Wangkung; 29. Tjumbi; 30. Ruteng; 31. Kumba; 32. Rua; 33. Dampék; 34.
Nanga-Rema; 35. Nanga Rema; 36. Waé Tua; 37. Waé Rambung; 38. Béntég Djawa; 39. Nunuk; 40. Peta;
41. Lawir; 42. Rotjong; 43. Mano; 44. Pongkor; 45. Rana; 46. Potjo Rana-Ka; 47. Méné; 48. Rembong; 49.
Sita; 50. Puntu; 51. Waé Reca; 52. Léwé; 53. Mborong; 54. Kisol; 55. Badjawa; 56. Toda-Belu.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)17

likely he was not even aware of the existence of the latter. The impractical ecologicalarrangement is also taken from Hoogerwerf.
In Pfeffer’s paper, there are no acknowledgements, and there is no literature list.
Hoogerwerf is mentioned twice, the second time: “M. Hoogerwerf signale avoir rencontré Ducula bicolor et le Pigeon à crète, Caloenas nicobarica, que je n’ai jamais eu la chanced’apercevoir”. This is apparently to emphasise that Pfeffer himself did observe all theother species mentioned in his paper (which is, of course, not so). Generally, Pfeffer’stranslation is a good one, but occasionally he has gone astray. Discussing Haliaeetus
leucogaster, Hoogerwerf notes: “Wij zagen van deze soort eens 5 exemplaren bijeen in delucht, maar 1 of 2 stuks werden dagelijks gezien, zowel volwassen exemplaren als nog
in het jeugdkleed gestoken vogels, ofschoon vaker op Komodo dan op Padar en Rintja.
Die zee-arend werd ook in het binnenland van Rintja aangetroffen bij een zoetwaterreservoir waarin nogal wat visjes leefden. Zulk een verblijf was ons van deze grote
roofvogel volkomen onbekend, want een dergelijke plaats is bij uitstek het biotoop van
die andere arend, Ichthyphaga ichthyaetus en - maar in mindere mate - van Pandion, devisarend” 2). There is no mention of the occurrence of Ichthyophaga ichthyaetus here; only
the statement that Haliaeetus leucogaster occurred here in a habitat where (elsewhere)
one would rather expect Ichthyophaga ichthyaetus and Pandion haliaetus.
Pfeffer, after discussing Haliastur indus and Haliaeetus leucogaster, birds so common
and conspicuous that he was able to give some personal observations, comes with this:“Moins nombreux que les deux espèces précédentes sont Pandion haliaetus et Ichtyophaga
ichtyaetus, cependant on peut être assuré de rencontrer au moins un individu ou un couple de chacune de ces deux espèces en une matinée d’observation”. Pfeffer even copiedfrom Hoogerwerf the erroneous spelling Haliaetus for Haliaeetus, and from Hoogerwerf(1954b: 217) the erroneous spelling Ichtyophaga ichtyaetus for Ichthyophaga ichthyaetus, for
Hoogerwerf’s (1956) error (Ichthyphaga) was a different one. Bruce (in White & Bruce,
1986: 118) noted about Ichthyophaga ichthyaetus: “From Rinca it was also recorded by
Hoogerwerf, perhaps the source of Pfeffer’s error”. Bruce was correct in pointing out thesource of Pfeffer’s error, but did not realize that Hoogerwerf’s “record” was not a record
at all. Verhoeye & Holmes (1998: 12) not only transferred Hoogerwerf’s discussion of
Pandion haliaetus from Rintja to Komodo, but moreover seem certain that he observed
there two individuals of the species he stated clearly that he had not seen!
Pfeffer’s observations of Numenius arquata and Tringa totanus in June, allegedly on
Rintja, which were rightly questioned by White & Bruce (1986: 170, 173), were presumably also copied from Hoogerwerf, and were from Padar, an island not visited by Pfeffer. Hoogerwerf is also responsible for the record of Cypsiurus parvus (= C. balasiensis):“de Apodidae Collocalia esculenta, Collocalia sp. en Cypsiurus parvus ... maar deze laatste
soort kwam hier toch in geen verhouding voor tot de prachtige nestel-gelegenheid,

2)

Of this species we once saw 5 individuals together in the air, but 1 or 2 were noted daily, adults aswell as birds in juvenile plumage, although more frequently on Komodo than on Padar and Rintja. Thissea-eagle was also found inland on Rintja, at a freshwater reservoir containing numbers of small fishes.
Such a habitat was completely unknown to us for this large bird-of-prey, for such a place is especiallythe biotope of that other eagle, Ichthyophaga ichthyaetus and - to a lesser degree - that of Pandion, theOsprey.

18

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

welke de lontarpalmen haar konden bieden” 3). Pfeffer: “Les Apodidés sont bien représentés aussi: Collocalia esculenta, Collocalia sp. (francica?) et Cypsiurus parvus, qui
nichent tous à la base des palmes de lontar”. The statement that the Collocalia species
nest in Lontar Palms (Borassus flabellifer) is another mistranslation.
The supposed common occurrence of Dendrocopos macei originated also with Hoogerwerf. As Hoogerwerf’s article deals with the islands of Komodo, Padar and Rintja,
and not with Flores, there is no need here for a discussion of his paper and for speculation as to whether he has actually seen Cypsiurus balasiensis and Dendrocopos macei (Iregard it as very unlikely).
The observation that Ninox scutulata is: “commune à Flores et Rintja” is not derived
from Hoogerwerf, but is Pfeffer’s own, and is unexpected, both because otherwise only
three specimens are known from the island, and because only in the first one or two
weeks of Pfeffer’s stay might this northern migrant have been likely still to be present.
Pfeffer continues, however: “d’après autopsie d’un individu, il semble que cet oiseau
serait plutôt insectivore que franchement carnivore”. This certainly suggests that a specimen was collected. In reply to my request for a loan from the Paris Museum, I received
a Pfeffer specimen of Ninox scutulata, but it was from Borneo, not from Flores, and theenumeration of Pfeffer’s skins from Flores and Rintja, given below, shows that he did notcollect the species there. More about the specimen from Borneo will be said in the maintext. Enough about Pfeffer’s publication: in the rest of this paper, it shall be ignored.
The preceding notes were necessary to clear up the record, but it would be unfair toPfeffer not to mention that he has published a very readable book about his visit to
Flores and Rintja (Pfeffer, 1965). Throughout, the book has the ring of truth, and someof its stories (like the addiction to “soentik”) look very familiar, being similar to my own
experiences in the East. The 17 bird species mentioned in the text (of which the scientific names are given in an appendix) are common lowland birds, of which there can be
no reasonable doubt that he has observed them.
During his stay on the islands, Pfeffer collected a small number of birds for theParis Museum. The acquisition-register of the Museum shows the following specimens
to have been received. From Flores: Halcyon chloris (2), Pachycephala fulvotincta, Philemonbuceroides (2) and Dicrurus hottentottus. From Rintja: Stiltia isabella, Chalcophaps indica,
Halcyon sancta, Anthus novaeseelandiae (2), Coracina floris, Nectarinia jugularis and Poephila guttata.
E. Schmutz, SVD (*1932) arrived on Flores in 1963 and, mainly from 1968/1978, but
again in 1982/1983, collected birds, of which the great majority was sent to Leiden. Like
Father Verheijen, with whom he cooperated closely, his main field of activity was in
Manggarai, West Flores. He lived in Nunang, and his collections are mainly from thesouth-western corner of Manggarai. Especially in the earlier years, material collectedby him was labelled and numbered in the Verheijen collection, which makes it in somecases difficult to be sure whether a bird was collected by Schmutz or by Verheijen. Perhaps it does not matter much: for example, interesting material from the locality Poco3)

the Apodidae Collocalia esculenta, Collocalia sp. and Cypsiurus parvus . . . but the latter species did notoccur here in proportion to the beautiful nesting opportunities offered it by the Lontar Palms.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

19

Nernancang (near Ruteng) was actually obtained by the native collector Lazarus, andwas forwarded by either Verheijen or Schmutz. Father Schmutz has been particularly
successful in collecting a number of rare or little-known birds such as Podiceps novaehollandiae, Pernis ptilorhynchus, Columba vitiensis, Caprimulgus macrurus, Coracina novaehollandiae. From his collections, I was able to describe one new species (Monarcha sacerdotum) and one new subspecies (Accipiter virgatus quinquefasciatus). During a visit to Leiden in 1995, I was shown another small collection of birds from Flores (ca. 80 specimens), received from Father Schmutz after my retirement. I took notes of the more interesting specimens and included them in this paper.
In recent years bird collections have been made in the Lesser Sunda Islands (including Flores) on behalf of the Western Australian Museum, Perth. Mainly for the sake ofcompleteness, I should have liked to include these, but my request for access to thismaterial was declined.
Several authors (e.g. Junge, 1954: 311; White & Bruce, 1986: 71) list Flores amongst
the islands visited by the great Italian botanist and ornithological collector O. Beccari(1843-1920), but that is hardly correct. On his first expedition, with L.M. d’Albertis(1841-1901) to New Guinea, the route was from Makassar through Strait Flores to Timor(Koepang and Dilli). Departure from Makassar 24 February 1872, arrival Koepang the
28th. The voyagers must have seen the eastern end of Flores from their ship, and they
may even have spent a few hours in the roads of Larantoeka, but it could not have been
more than this (cf. d’Albertis, 1880: 5).
6. The egg collection
Until Father Verheijen started his investigations, practically nothing was known of
the nidification of birds of the Lesser Sunda Islands. Rensch noted the condition of the
gonads of the birds he collected, and photographed the nest of a sea-eagle Haliaeetus
leucogaster with a chick on the island of Endeh. The title of a short article by Schönwetter(1934) promises more than it gives, as eggs of only five species of birds from Timor aredescribed in it, and all are widely-distributed species, the nidification of which was
already well known in other parts of their range (Podiceps ruficollis, Anas superciliosa,
Rallus philippensis, Porphyrio porphyrio, Chalcophaps indica). And that is about all, as far as
published evidence goes. It may truly be said that Verheijen did pioneer work, and hisarticle on breeding seasons of birds on Flores is a contribution which deserves more
attention than it has received.
Verheijen (1964) lists 101 species which have been found breeding, of which 9 witha query:
Ixobrychus cinnamomeus
Accipiter virgatus
Rallus pectoralis
Porzana cinerea
Porzana fusca
Streptopelia bitorquata
Ptilinopus melanospilus
Pnoepyga pusilla
Culicicapa ceylonensis

Of these nine, I have been able to confirm four: Ixobrychus cinnamomeus, Rallus

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Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

pectoralis, Porzana cinerea, and Culicicapa ceylonensis. With much hesitation, I have also
accepted Ptilinopus melanospilus, which will be further discussed below. Moreover, thefollowing species have to be eliminated from Verheijen’s list: Otus alfredi = Otus silvicola(as suggested by Verheijen in a footnote), Collocalia inexpectata (there are no eggs that
could be ascribed to this species, now C. salangana, in the collection; see p. 219), Geocichla interpres (no eggs in the collection), Erythrura hyperythra (identification doubtful).
The case of Rhipidura diluta is interesting: at first rejected (the eggs had been misidentified), it could subsequently be re-instated, as a clutch ascribed to its congener R. rufifrons was found to belong to it. On the other hand, the following species can be added:
Accipiter novaehollandiae, Falco longipennis, Otus silvicola, Caprimulgus macrurus, Merops
philippinus, Orthotomus cucullatus, and Passer montanus.
When these corrections are carried out: 8 species subtracted, and 7 added, the total
number of species of which the eggs have been collected is exactly 100 (of which 97 in
the RMNH; the eggs of three parasitic cuckoo-species in the Coll. Ottow). Consultationof the register will show that this is close to two-thirds of the total number of species
which may reasonably be assumed to be breeding birds. This means that, rich and important as the collection is, it is far from being complete. This was pointed out by Verheijen (1964: 194): “Hardly anything has been collected on the coastal islands, along thebeach and in the coast vegetation, nor above 1300 m”. Even keeping these limitations in
mind, the absence of eggs of some species, particularly Lophozosterops superciliaris andLichmera lombokia is surprising, seen in the light of the large numbers of skins of these
species that have been obtained, and show them to be common.
The collection contains a number of unidentified clutches. Further study of these willprobably lead to the addition of a few species to the list of 100. A much greater problem
than unidentified eggs, are misidentified eggs. Inevitably, large collections brought together with the help of natives, must contain misidentified eggs, and the collection from
Flores is no exception. In a few instances (such as the eggs ascribed to Accipiter virgatus,
Rhipidura diluta) such errors were easy to detect. Real problems are caused by the whiteeggs of the Columbidae: unless the birds are seen attending the nest, there seems no possibility of separating eggs of Streptopelia bitorquata from those of S. chinensis; the problem
is not confined to within a genus, for the eggs of Ptilinopus melanospilus also agree inmeasurements although perhaps not in weights. In this and several other cases, the original identification by the finders has just to be accepted, as a verification on the basis of
the eggs alone is not possible. The same difficulties apply to the pair Zoothera interpres /Zoothera dohertyi, the Lonchura species, and several others.
Father Verheijen has been generous with his eggs. He presented several clutches to
Coomans de Ruiter and to Hoogerwerf, who described some of them in a paper on theoology of Java (Hellebrekers & Hoogerwerf, 1967: 53, 122, 135, 136, 149, 158). The eggcollections of both these gentlemen have since been incorporated into the RMNH collection; included is an egg of Chrysococcyx minutillus, which was already in Hoogerwerf’s possession before the eggs of the parasitic cuckoos were sold to Ottow. I am
personally indebted to Father Verheijen for presenting me with clutches of Zosteropidae,
then undescribed, for inclusion in my revision (Mees, 1969: 203, 211, 280).
As regards seasonality, Verheijen’s (1964) enumeration of the combined totals of
nests found each month, shows that some breeding activity takes place throughout
the year, but that there are clear maxima in April and May, and minima in December,Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

21

January, and February. The period of minimal breeding activity corresponds to the wet
season (heavy rainfall), whereas the peak period, in April and May, corresponds to theend of the rainy season, and the beginning of the dry season. Of course, Verheijen’s tableshows only a rough approach and undoubtedly there is some bias in it. For example,
Verheijen visited the island of Palué from mid-April to the first week of May, 1960, andduring this short period, large numbers of clutches of a limited number of species wereobtained (15 of Turnix sp., 17 of Nectarinia jugularis, 65 of Zosterops chloris). Although the
Palué records have certainly boosted the figures for April and May, I believe that they
have accentuated rather than distorted the general picture.
When comparing Verheijen’s table with the numbers of clutches now in his collection, it will be noted that the former in many cases contains more. This is only as was tobe expected, as the table was based on all the nests and eggs found, not all of which
would have been collected. On the other hand, some clutches have been added to thecollection after the manuscript was closed (in June 1962). There are, however, several
discrepancies that cannot be explained in this way. An extreme example is Nectarinia
jugularis, of which Verheijen lists 36 clutches (or nesting records) divided over themonths as follows: March (4), April (3), May (23), June (6). His collection, however,contains clutches collected in March (4), April (19), May (11), June (7), and July (3).
Perhaps more interesting than the general (main breeding activity in April-May atthe beginning of the dry season), is the particular: the species that do not conform. Verheijen (1964: 197) already mentioned Saxicola caprata as an example of a pronounced
dry-season breeder, which begins breeding only towards the end of July, and stops
abruptly in the beginning of November. Other dry season breeders are Caprimulgus affinis, Merops philippinus, Cecropis striolata, Terpsiphone paradisi, Dicrurus hottentottus, Artamus leucorhynchus, Corvus florensis and Corvus macrorhynchos, and of course, perforce,the brood-parasites of these species: Cacomantis variolosus, Eudynamis scolopacea andScythrops novaehollandiae. None of these other cases is based on such an impressive material as that of Saxicola caprata, and one seems dubious: Verheijen listed nesting of Terpsiphone paradisi in October (1), November (2) and December (1), which certainly issuggestive, but his collection contains, in addition, a clutch taken in March.
Are there also birds with a preference for the wet season? Not unexpectedly, thedata for some of the Rallidae (Rallina fasciata, Amaurornis phoenicurus) suggest this,and one would expect it of the Anatidae (about the breeding of which there is little
information).
Finally, there is the category of species that have been found breeding in all monthsof the year. There are not many: Turnix suscitator/maculosa, Rallus philippensis. Anthus
novaeseelandiae, Tesia everetti and Dicaeum annae have also a remarkably extended season(11 months in Verheijen’s table), although the egg-material of A. novaeseelandiae available to me, covers only 9 months, but including May, the month for which no breeding
was listed by Verheijen. Of Tesia everetti, material collected in 10 months is present. On
the other hand, eggs of Philemon buceroides are represented in the collection from 10
months, although listed by Verheijen from 8 months only. In many other species, there
is a clear and predictable relation between the number of clutches present, and the extent of the breeding season that can be deduced from them. It is apparent that these
species, although they have a distinct and fairly short peak in their breeding, do occasionally nest in other months. As observed by Verheijen, Philemon buceroides appears to22

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

have a double peak in its breeding-activity, in April/May, and September. A second
example of a double peak, Gerygone sulphurea, is less convincing. This brings me to aconcluding remark about the value of Verheijen’s egg collection. Originally I was inclined to judge it by the number of species represented in it, and to regret as unnecessary the large series of some of the commoner birds. However, only these large series
can show where the peak period of breeding activity lies, and indeed, without them theinformative value of the collection would have been very much less.
7. Forsten’s collection from Bima, Sumbawa
In the following pages, mention will repeatedly be made of specimens from Bima,
received from Dr A.E. Forsten (1811-1843) in 1842. It is the first collection known from
Sumbawa, and there can be no doubt of its provenance, as it contains several species
and subspecies of birds endemic to the central Lesser Sunda Islands. Some of thesewere described from 1850 onwards (Trichoglossus haematodus forsteni, Zosterops aurifrons= Zosterops wallacei, Gracula religiosa venerata, Oriolus chinensis broderipi, Dicrurus hottentottus bimaensis), but others remained undescribed, and were only described many years
later from other collections (Treron floris, Pitta elegans concinna, Terpsiphone paradisi floris,
Philemon buceroides neglectus, Lonchura pallida). There remains, however, an enigma
around this collection, as Forsten did not visit Sumbawa. On his way out from Java to
Sulawesi (Celebes), he passed in sight of Sumbawa, but did not go ashore. I have consulted Forsten’s diary in the archive of the Rijksmuseum, and it confirms that he neverset foot on Sumbawa. Junge (1954: 307), confronted with the same problem, suggested
that Forsten had sent hunters to Sumbawa, which seems possible, but very unlikely, as
Forsten never worked anywhere near Sumbawa, his activities being confined to Sulawesi (Celebes) and the Moluccas. The possibility that after Forsten’s death his hunters,on their way back to Java, might have called at Bima and done some collecting there,
has occurred to me, but that conflicts with the fact that the collection was received inLeiden in 1842, when Forsten was still alive.
Of the early collectors, only C.G.C. Reinwardt (1773-1854) visited Bima, but he is notknown to have collected birds there, and his collections were received in Leiden twenty
years earlier (as far as they were not lost by ship-wreck). Moreover, he did apparently
not label his specimens, which later has caused no end of confusion. Therefore a collection definitely from Bima, is not likely to have originated from him. That leaves theriddle of who was the collector of the Bima material unsolved; the solution is probably
still buried in an archive: after all, not that many people interested in birds and able tocollect them, can have visited Sumbawa around 1840. In 1839/1840, just before leavingon his journey to Celebes, Forsten was for a short time director of the natural history
museum in Batavia. It is likely that during this period he either received the specimensfrom Bima, or found them already present in the collection, and that he arranged theirtransport to the Netherlands.
8. Systematic accountAll measurements are in millimetres; weights of the eggs in grammes. It will beclear that these are the weights of the empty shells, not of the full eggs.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

23

Fig. 5. Podiceps ruficollis; right and middle specimens from Java, with little black on chin, left specimen
from Flores with large black throat patch, as discussed in the text.

Podiceps ruficollis vulcanorum Rensch
Podiceps ruficollis vulcanorum Rensch, 1929, Journ. f. Orn. 77, Ergänzungsb. II: 205 – Kratersee Segare
Anak (2000 m), Lombok.
Collectors.— Allen, Rensch (6), Schmutz.
Material.— 2 (?), vii.1969, Nunang (crater lake), 650 m (Schmutz, RMNH nos. 66087, 66088); , 11.
viii.1969, Nunang (Schmutz, RMNH no. 85142); , 28.x.1971, Nunang, 650 m (Schmutz, RMNH no.
81084).

Notes.— Father Schmutz found this a common species on the crater lake Nunang.
The subspecies vulcanorum is characterized by a large black throat-patch, although
individual variation in extent of the black makes this a less distinct subspecies than one
would wish. Unexpectedly, two specimens from East Java, where this species is rare
and its status uncertain, have very little black on the throat and are quite typical tricolor(, , 22.x.1939, Klakah, leg. Kooiman, RMNH nos. 22951, 22953) (fig. 5). In lateralview, the bill of P. ruficollis vulcanorum/tricolor, is thicker than that of P. novaehollandiae.
Measurements:
2
2(?)

wing
98, 100
101, 101

exposed culmen
22.8, 2123, 27bill/wing index
22.3, 21.0%
22.8, 26.7%

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Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Podiceps novaehollandiae novaehollandiae Stephens
Podiceps Novae Hollandiae Stephens, 1826, in Shaw, General Zool. 13 (1): 18 – New Holland.
Podiceps novaehollandiae javanicus Mayr, 1943, Emu 43: 6 – Rakukak, 4000’, Java (= Rakoetak, see Notes).
Podiceps novaehollandiae timorensis Mayr, 1943, Emu 43: 7 – Sumul, Timor.
Collector.— Schmutz.
Material.— , 17.i.1969, Nunang,Waé Sano, 650 m (Schmutz, RMNH no. 65167). Iris golden yellow, bareskin around the eye citrine yellow.

Notes.— This specimen does not agree with the description of P. n. timorensis (about
which more will be said below), but it fits well into a series from Java. Many years ago,
I compared specimens from Java with Australian material, and was unable to confirm
the existence of differences that would justify recognition of javanicus. P. n. timorensis is
not recognized by White & Bruce (1986: 62, 89); these authors did not discuss P. n. javanicus, which is outside the geographical region treated by them.
The type-locality of P. n. javanicus was given as “Rakukak” by Mayr (1943). No locality of that name exists, but according to Greenway (1973: 217) it appears on the label
to be Rakoetak. The Rakoetak (7°09’S, 107°43’E) is a mountain to the south-east of Bandoeng, West Java. This would also be the locality referred to as Raketak by Ripley (1952),
who honestly states: “I have been unable to identify this locality”.
Although van Oort (1910) already wrote: “All the little grebes from Java in the collection of the Leiden Museum (15 specimens) belong to novae hollandiae”, this was forsome reason overlooked by later authors, such as Bartels & Stresemann (1929: 90: P.
ruficollis philippensis; this is especially strange, since specimens in the Bartels collection
are correctly labelled P. novaehollandiae), Chasen (1935: 50: P. ruficollis vulcanorum) andKuroda (1936: 587), until Mayr (1943) drew renewed attention to it.
The status of this species on Flores is probably that of a rare visitor: the present
specimen is the only one ever found by Father Schmutz. P. ruficollis vulcanorum, on theother hand, is a resident. There are now records of P. novaehollandiae from Bali (Wiegant& van Helvoort, 1987), Flores, Alor and Timor, and in each case it concerned single individuals, without any suggestion of breeding. The Bali bird might conceivably have comefrom Java, whereas the birds from the eastern Lesser Sunda Islands are more likely tohave been stragglers from Australia. Mayr’s (1943: 4) assertion that P. ruficollis: “lives
side by side with novaehollandiae on a number of places without interbreeding”, remains
unconfirmed. It does certainly not apply to the Lesser Sunda Islands, but may be valid
for northern New Guinea (Lake Sentani), where the situation requires further study.
P. n. timorensis was based on a single specimen, supposedly distinguished fromnominate novaehollandiae by having the: “Facial pattern and white on secondaries as innovaehollandiae, but underparts darker than in the other races of the species and distinctly washed with tawny (chestnut), resembling tricolor in this respect. General size
and relative size of bill large”. Being aware of the presence of a specimen from Timor in
the ANC, I asked Dr Schodde for information about it. On the basis of his reply, I feeljustified in placing timorensis also in the synonymy of nominate novaehollandiae, inagreement with White & Bruce.
Particulars are: , gonads 3.0 × 3.0 mm, 14.vi.1974, 10 km west of Lautém, East
Timor (I.J. Mason, CSIRO no. 30240), weight 170 g. “The specimen is white-breasted

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)25

and -throated with vestiges of white mottling on sides of head and a faint russet cast to
the mid grey of the lower throat, and sides and back of the neck. This suggests to me
that it is an immature male moulting into a nearly completed first non-breeding plumage .... In comparison with T. n. novaehollandiae from Australia in equivalent plumage I
can find no difference other than some russet feathering in the crown which could be
individual. The bird is as white on the breast, belly and upper throat as typical Australian specimens” (Schodde, in litt., 11.vi.1993).
Measurements:
wing
exposed culmen
bill/wing index
Flores105
22
21%
Timor104
21+
ca. 22.1%
The bill of the Timor specimen is damaged; the index is calculated from the assumption that ca. 2 mm at
the tip are missing.

Pelecanus conspicillatus Temminck
Pelecanus conspicillatus Temminck, 1824, Recueil d’Ois. 5 (livr. 47): pl. 276 – des Terres Australes = Swan
River, Western Australia (cf. Stresemann, 1951).
Collector.— Verheijen (a wing only, ZMA).
Material.— None.

Fig. 6. Pelecanus conspicillatus captured on Flores (Watuneso).

26

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Notes.— The wing preserved by Verheijen, was from a bird taken near Sika, about
September 1960 (cf. Voous, 1967). In 1978, an invasion took place in southern Indonesia.
Somadikarta & Holmes (1979) listed records from the Moluccas, from Java and Sulawesi(Celebes), and from the Lesser Sunda Islands of Bali, Lombok, Sumbawa and Timor.
The largest flock reported was one of ca. 300 birds in West Sumbawa. Flores, although
not specifically mentioned by these authors, was not passed by. Local newspapers recorded 26 pelicans flying above and fishing in Rana Loba, a small lake ca. one kilometerfrom Mborong, Manggarai, on 26 April. On 10 July, four birds were captured near Watuneso (fig. 6). As an explanation for the invasion, adverse conditions in Australia, following several good breeding seasons and a build-up of the population, has been suggested
by Somadikarta & Holmes.
Sula leucogaster plotus (Forster)Pelecanus plotus Forster, 1844, Desc. Anim. (ed. Lichtenstein): 366 – Ternate.
Collector: – Verheijen.
Material.— (?) ad. 16.x.1970, Nanga Ramut (Verheijen, RMNH no. 65168). Iris light greenish grey, billbluish slate, its tip almost white, legs spotty greenish blue.
Measurements:
(?)

wing
ca. 390

tail
198

tarsus53

culmen from forehead feathers
93

Phalacrocorax melanoleucos melanoleucos (Vieillot)
Hydrocorax melanoleucos Vieillot, 1817, Nouv. Dict. Hist. Nat. 8: 88 – l’Australasie.
Collector.— Verheijen.
Material.— , 5.vii.1969, Look (Verheijen, RMNH no. 65169).

Notes.— Mathews’s (1912: 241) substitution of “New South Wales” for the type-locality as originally given by Vieillot, was, as far as I can judge, purely arbitrary. I haveno objection to a type locality restriction being arbitrarily made, when this is needed forthe study of geographical variation, and the original type material cannot provide thisinformation, or is no longer available. I do object, however, to arbitrary restrictions andsubstitutions being made (as in the above case), without any mention that they are restrictions. According to Berlioz (1929: 65) the type is “d’Australie”, and was broughtback by the Expedition Baudin in 1804.
I cannot find clear published evidence of breeding of this species in Sulawesi(Celebes), so that it is worth recording the presence of two eggs in our collection: 1,
11.viii.1863, Ayer Pannas (Rosenberg, RMNH cat. no. 1); 1, 13.ix.1863, Panybie (Rosenberg, RMNH cat. no. 2), measurements 46.0 × 30.3 and 48.2 × 30.4 mm, weights 1.5802
and 1.8488 g.
Cat. no. 1 has on the shell, in what I believe to be Schlegel’s handwriting: “s. n. Ibis
peregrinus, v. Rosenberg, Celebes”. On the oldest label Plegadis falcinellus, (on which isadded in pencil a word which I believe reads “falsch”, in Schönwetter’s handwriting),
and then one with Phalacrocorax javanicus (Horsf.). The third label reads Phalacrocorax

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)27

spec., with the last word crossed out and, in the hand of Hellebrekers changed to melanoleucos and on the last label, the current one: Phalacrocorax melanoleucos melvillensis.
Cat. no. 2 has written on the egg: “s. n. Ardea picata, v. Rosenberg”. On the oldest label,bearing the name Notophoyx picata, is added in pencil: “ist Phalacrocorax spec.”, signed
Schönwetter. The next label reads Phalacrocorax javanicus, then Phalacrocorax spec.,changed by Hellebrekers, and the final one of P. melanoleucos melvillensis.
To the impressive list of island records provided by White & Bruce (1986: 94), Alor
(Ombaai) can be added: (?), viii.1880, leg. Colfs, RMNH cat. no. 21.
Phalacrocorax sulcirostris (Brandt)
Carbo sulcirostris Brandt, 1837, Bull. Acad. Sci. St. Pétersb. 3: col. 56 – Terrae australes (Südsee).
Collector.— Weber.
Material.— (?), i.1889, Endeh (Weber, RMNH cat. no. 14).

Notes.— Bruce & White (1986: 94) state: “Breeds in E. Java (Hoogerwerf 1953) andnow also W. Java (McKinnon MS)”. But Hoogerwerf’s (1953 = 1954a) paper, listed correctly in their bibliography, is about breeding on Pulau Dua in West Java, in 1952. Ourcollection contains nine clutches of eggs from Pulau Dua, collected by Hoogerwerf in
1953, 1954, 1955 and 1956. Hoogerwerf (1969/1971: 448-449) found it also nesting on thenorth coast of the Udjung Kulon peninsula, the western tip of Java, in March 1955. As
regards East Java: when Hoogerwerf (1935) visited the tambaks near Soerabaja, he wasassured that the species had been breeding there for as long as could be remembered. I
do not know on what McKinnon’s statement is based that Phalacrocorax sulcirostris “hasbeen extending its range westward in the last century and now nests in E. Java andPulau Dua in West Java”: correct for West Java, but in East Java it might, for all that is
known, have been a resident for centuries.
There is no definite record of breeding in Sulawesi (Celebes). White & Bruce (1986:94) state: “Breeds in E. Java..... presumably also in Wallacea”. Meyer & Wiglesworth(1898: 891), like Blasius (1883: 127-128, and 1886: 174) before them, dismissed Rosenberg’s (1881) claim of the occurrence of P. sulcirostris on Lake Limbotto. Yet, our collection contains two clutches, identified as P. sulcirostris, c/1, 21.vi.1863, Gorontalo (v.
Rosenberg, RMNH cat. no. 1) measurements 53.7 × 33.3 mm, weight 2.7708 g; and c/4,
11.viii.1863, Ayer Pannas (v. Rosenberg, RMNH cat. no. 2) measurements 49.2 × 37.1,
50.4 × 37.0, 50.4 × 37.2, 53.3 × 36.0 mm, weights 3.2220, 3.5772, 3.8336, 3.7391 g. Cat. no.
2 has written on the shells: “Carbo sulcirostris”, v. Rosenberg, Celebes”, and therefore
was correctly identified from the beginning. Cat. no. 1 was originally marked Ardea nigripes, re-identified as Phalacrocorax on a label written by van Oort, after that as Anhinga
melanogaster, and finally as Phalacrocorax sulcirostris by Hellebrekers.
Fregata ariel ariel (G.R. Gray)
Atagen ariel G.R. Gray (ex Gould MS), 1845, Gen. Birds 3: 669, pl. 183 – no locality; designated type
locality Raine Island (cf. Mathews, 1914: 121).
Collectors.— None.

28

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Notes.— Frigate birds were first recorded from Flores by Rensch (1931b: 505), who
observed them, high in the air, near Endeh and Mborong. He was unable to give a specific identification, and listed them as Fregata sp. Verheijen (1961) reported F. ariel as:“often seen above the coastal areas of Flores”. Schmutz (MS, 1977) observed frigate
birds at Nanga-Lili, as F. ariel subsp. Although, as far as I know, no specimens have beencollected, the evidence is strong enough for acceptance.
Fregata ariel is certainly the commonest frigate bird in the region, but F. minor may
also occur. There is a specimen of F. minor from Timor in our collection, x.1880 (leg.
Colfs, RMNH cat. no. 5), and Haniel collected a specimen of F. minor off the north coastof Samau, on 21.vi.1911 (cf. Hellmayr, 1914: 109; Stresemann, 1922). Goenoeng Api inthe Banda Sea, a well-known breeding station of F. minor (Hoogerwerf, 1939; Van Bemmel & Hoogerwerf, 1940), is ca. 500 km north-east of Flores. Schmutz says of the birds
he observed, that the white of the breast may be entirely absent, but he says nothing
about the presence or absence of white patches on the flanks, the character by which
adult males of the two species may be distinguished.
F. minor is a common breeding bird on Christmas Island (Indian Ocean). White &Bruce (1986: 93) also claim breeding of F. ariel on the island, but surely in error: until
recently, F. ariel had not even been recorded as a visitor to Christmas Island, but in thelast few years birds have been observed there, their behaviour suggesting that they may
be about to settle (cf. Stokes, 1988: 32).
Ardea sumatrana Raffles
Ardea Sumatrana Raffles, 1822, Trans. Linn. Soc. Lond. 13: 325 – Sumatra.
Ardea sumatrana mathewsae Mathews, 1912. Novit. Zool. 18: 230 – Cooktown, North Queensland.
Collectors.— Allen, Verheijen (egg only).
Material.— Egg: c/1, iii.1961, Nanga Rema (RMNH no. 70047). The egg is very pale greyblue.
Measurements and weight:

RMNH no. 70047

68.5 × 47.4

7.24

Notes.— Measurements and weight of this egg fit nicely into the range of variation
given by Schönwetter (1960: 87) for a series of 11 eggs from northern Australia.
There is no general agreement on the question of whether the species shows geographical variation, in particular of whether there is a distinguishable Australian subspecies. That this question has not yet been solved to everybody’s satisfaction is certainly due to the large size of the bird: very few museums have adequate material fromall parts of its fairly extensive range, and their large size leads to a reluctance to ask for,or to provide, loans, merely for the solution of what is, after all, only a very minor
problem. As far as I can make out, in the past fifty years, only Amadon & Woolfenden(1952) have actually compared material. They concluded that the subspecies describedfrom Australia, mathewsae, was not valid, although they admitted that their material
was somewhat inadequate. Also, their material suggested that Australian birds are
smaller. Many authors have accepted this conclusion, and have treated Ardea sumatrana as a monotypic species (cf. Condon, 1975; 52; Payne, 1979: 201; Dickinson et al.,
1991: 101), but others have continued to recognize subspecies, either implicitly, byMees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

29

using a trinomial for the birds from the northern and western parts of the range (forexample Smythies, 1957: 560 and later publications; duPont, 1971b: 21), or explicitly. To
the latter category belong Hancock & Elliott (1978: 280), who claim Australian birds tobe: “noticeably browner”, and Hancock & Kushlan (1984: 74), who state that: “TheAustralian subspecies mathewsi [sic] is very noticeably browner, almost bronze, in colour”. Recently, the race: “mathewsii [sic] restricted to the Australian subcontinent”[why sub?] was also accepted by Lansdown (1992).
Clearly, there was reason for a further examination of the problem. The RMNH collection is not exactly poor in specimens, but is very unbalanced: 24 specimens, but only
one of these from Australia. The Australian bird is a mounted specimen from ClarenceRiver, NSW (received in 1862 from the Australian Museum, Sydney); the other specimens are from: Continental India (2), Sumatra (1), Simalur (2), Java (3), Sulawesi(Celebes) (6), Morotai (1), Halmahera (1), Batjan (1), Obi Major (1), Kelang (1), Waigeo(1), Misool (1), Salawati (1), and Sorong, Vogelkop, New Guinea (1). In this material,
there is a considerable variation in plumage. Very distinctive is the bird from Waigeo, ajuvenile: it is warm brown, almost chestnut, with black stripes. Other brownish birds,
specimens having brownish edges along the feathers, may be assumed to be immature(cf. Amadon & Woolfenden, l.c.). Adult birds, however, do not differ in any way fromthe Australian specimen, which, contrary to descriptions in the publications cited
above, is quite dark. I cannot see any significant difference in plumage between an
adult bird from Continental India (admittedly an old, mounted specimen of not very
precise provenance, type of Ardea typhon Temminck, RMNH cat. no. 1), several adults
from intermediate localities, and the Australian bird. The wing-length of the Australian
bird is 485 mm, that of the bird from India 480 mm, clear confirmation that Australian
birds are not smaller.
It is not surprising that the subspecific name mathewsae bestowed by Mathews hascaused some confusion (mathewsi, mathewsii). Mathews did not explain the name, so wemust assume that, somewhat artlessly, he named the bird after his wife. I suppose that,with the naming of some 560 species and subspecies in this one publication, his phantasy was heavily taxed.
Ardea purpurea manilensis Meyen
Ardea purpurea var. manilensis Meyen, 1834, Nova Acta Acad. Caes. Leopold.-Carol. Nat. Cur. 16, Suppl.:
102 – Manila (reference not verified).
Collectors.— De Jong (1), Schmutz.
Material.— , 12.vii.1969, Kenari (Schmutz, RMNH no. 65170).

Notes.— Previous records are of a bird collected at Mboera, 26.x.1929 (cf. Rensch,
1931b: 395), and a sight record from Mborong in the beginning of July 1927 (Rensch,
1931a: 497). Evidently A. purpurea is a resident on the Lesser Sunda Islands, althoughproof of breeding is not yet available. I do not know on what evidence, if any, breeding
records from Flores in March and from Sumbawa late April by White & Bruce (1986: 99)
are based, and presume them to be erroneous.

30
Measurements:

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)wing
330 (band)

tail
115

tarsus122½

exposed culmen
123

Ardea alba modesta J.E. Gray
Ardea modesta J.E. Gray, 1831, Zool. Misc.: 19 – India.
Collector – Semmelink
Material.— (?), 1862, Larantoeka (Semmelink, RMNH cat. no. 16).

Notes.— When Schlegel (1863a) completed his catalogue of the Ardeae in April
1863, the above specimen had not yet reached him, so that it remained unpublished.
That is why the species was only added to the avifauna of Flores seventy years later, on
the basis of a sight record (of a single bird) by Rensch (1931a: 501).
To the island records listed by White & Bruce (1986: 100), I can add Khoor (= Koer,
Kur), where v. Rosenberg obtained a specimen (, 2.ix.1865, RMNH cat. no. 32).
Since the revision by Payne & Risley (1976), this species has frequently, but by nomeans universally, been placed in the genus Ardea (rather than in Egretta or Casmerodius).
The reason why I did not follow this, was a certain scepticism (cf. Mees, 1982a: 14) andmainly a reluctance (due to conservatism) to abandon the binomen Egretta alba, that has
been familiar to me and that I have used in publications for over fifty years. Records ofinterbreeding in the wild between Egretta alba and Ardea cinerea in the Netherlands, theraising of viable young from such pairs, and especially the fact that a hybrid pairedwith an Egretta alba, again produced viable young, proving its fertility (Van der Kooij &Voslamber, 1997), indicate that the two species are fairly close and thus support theirunion in one genus. That interbreeding is not exceptional or confined to a single locality is shown by a further record from Latvia, where in 1997 a mixed pair succesfullyraised four hybrid young (Baumanis in Anon., 1998: 39, 56 pl. 4).
McCracken & Sheldon (1998), although highly critical of the osteological work ofPayne & Risley, nevertheless agreed in placing alba close to Ardea and remote from
Egretta garzetta, the type species of Egretta. In the light of all this evidence I feel compelled, still reluctantly, to transfer alba to Ardea.
Ardea novaehollandiae Latham[Ardea] novae Hollandiae Latham, 1790, Index Orn.: 701 – nova Hollandia.
Collectors.— Weber, Rensch (1), Verheijen (5, MCZ), Verheijen.
Material.— (?), xi-xii.1888, Endeh (Weber, RMNH cat. 20);  juv. nestling), x.1968, Kuwu, Rahong (Verheijen, RMNH no. 65172); (?)juv. (nestling), 29.x.1968, Lawir (Verheijen, RMNH no. 65175);  ad., (?)juv.
(nestling), 31.x.1968, Rahong (Verheijen, RMNH nos. 65171, 65174);  ad., 14.xi.1968, Ruteng (Verheijen,
RMNH no. 65173).
Eggs: c/1, vii/viii.1957, Bénténg Djawa (RMNH no. 70048); c/1, 11.vii.1958, Dampék (RMNH no. 70049).
The eggs are plain light blue.
Measurements and weights:

RMNH no. 70048
RMNH no. 70049

44.1 × 34.2
47.6 × 33.5

1.691
1.940

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)31

Notes.— Breeding of Ardea novaehollandiae on Flores was recorded by Verheijen(1964: 198) and Mees (1975a). Unfortunately, these records were overlooked by Hancock& Kushlan (1984: 113-116), who mention it as merely a migrant visitor or a vagrant to
the Lesser Sunda Islands.
Egretta intermedia (Wagler)A[rdea] intermedia Wagler, 1829, Isis: 659 – Java.
Collector.— Schmutz.
Material.— , 20.ix.1969, Kenari, west coast (Schmutz, RMNH no. 65179). Iris light yellow, bill yellow,
its tip brownish black, legs black.

Notes.— For measurements and a discussion of this species, cf. Mees (1982a: 16).
Egretta garzetta nigripes (Temminck)
Ardea nigripes Temminck, 1840, Manuel d’Orn. (2. éd.) 4: 376 – l’Archipel des Indes = Java.
Collectors.— Semmelink, Allen (cf. Wallace, 1864: 487, s. n. Egretta immaculata).
Material.— (?), 1862, Larantoeka (Semmelink, RMNH cat. no. 7).

Notes.— Egretta immaculata in Wallace’s list stands for this species, so that it musthave been collected by Allen. The records by Semmelink and Allen seem to have been
overlooked by later authors.
In the description of A. nigripes, Temminck first gave its range as “les îles de laSonde et les Moluques”, and a few lines further down, as “l’Archipel des Indes”. The
type locality was restricted to Java by Mathews (1927: 195), followed by Chasen (1935:56), but Peters (1931: 111) restricted it only to the Sunda Islands, and so did Payne(1979: 213). Our collection contains four specimens which would have been available
to Temminck in 1840: , not dated, but 1821/1823, Java (van Hasselt, RMNH cat. no.
1); , not dated, but see the date of the following specimen, Soerabaja (S. Müller,
RMNH cat. no. 2); , ii. “1827”, but 1828, Soerabaja (S. Müller, RMNH cat. no. 3); (?),
presumed juvenile, not dated, but 1821/1823, Java (van Hasselt, RMNH cat. no. 4). See
also Schlegel (1863a: 14), where these specimens are listed under the cat. nos. 18-21. As
all four specimens are from Java, Mathews’s restriction was perfectly justified. The twocollected by Van Hasselt would be from West Java, whereas Soerabaja is, of course, in
East Java. I see no need for a further restriction or a lectotype designation.
The year of collecting given for a Soerabaja specimen, 1827, cannot be correct, as S.
Müller did not visit East Java in that year. Members of the Natuurkundige Commissie,
including Müller, did, however, collect at Soerabaja in February 1828, when on their
way to Makassar (cf. Mees, 1994: 6, 52).
Egretta sacra sacra (Gmelin)[Ardea] sacra Gmelin, 1789, Syst. Nat. (ed. 13) 1 (2): 640 – Tahiti.

32

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Collectors.— Semmelink, Rensch (3), Verheijen/Schmutz.
Material.— (?), 1862, Larantoeka (Semmelink, RMNH cat. no. 13); , 22.ii.1969, Nisar (Schmutz, RMNH
no. 66070); , 11.v.1969, coast between Nanga-Lili and Sésok (Schmutz, RMNH no. 66071). Iris yellow.
Eggs: c/3, 14.vii.1958, Nanga Rema (RMNH no. 70050). The eggs are dull white, faintly tinged withbluish grey.
Measurements and weights:

RMNH no. 70050

45.1 × 34.9
45.3 × 34.1
47.3 × 34.0

1.87791.9422
1.8270Notes.— The bird collected by Semmelink belongs to the grey morph (cf. Mees,
1986: 23), as does one bird collected by Schmutz, whereas the other one belongs to thewhite morph.
Bubulcus ibis coromandus (Boddaert)
Cancroma Coromanda Boddaert, 1783, Table Planches Enlumn.: 54 – Coromandel (based on Crabier deCoromandel, Buff., Pl. Enlumn. 910).
Collector.— Semmelink.
Material.— (?), 1862, Larantoeka (Semmelink, RMNH cat. no. 46).

Notes.— Semmelink’s specimen, as well as the birds obtained in 1861 by Wallace at
Dilli, Portuguese Timor, provide proof that the Cattle Egret already inhabited the Lesser Sunda Islands before its recent expansion. On Flores it is apparently not common, forbesides the specimen listed above, there is only a field observation by Rensch (1931a:
500, 633). Father Schmutz has never seen it.
There is little doubt that the species is a resident on the Lesser Sunda Islands, although I do not know of any definite breeding records. White & Bruce (1986: 103) comment on the general absence of breeding-records from Wallacea; they could only mention a record of a nestling, not yet able to fly, from Paré-Paré, southern Sulawesi(Celebes), by Coomans de Ruiter (1948b: 75). However, in our collection there is a c/2,
6.vi.1863, Gorontalo, Celebes (Rosenberg, RMNH cat. no. 1). In evaluating the scarcityof nesting-records of this and other common birds, it should be kept in mind that reporting on their nesting would not have had priority with ornithologists of the 19th
century: breeding was taken for granted, and required no documentation. Incidentally,
Van Marle & Voous (1988: 62) record a c/2 from the Padang Highlands, Sumatra, in the
RMNH collection. This clutch, originally identified as being of Bubulcus ibis, was reidentified and published by me as Egretta intermedia (cf. Mees, 1982a: 18). Unfortunately
I failed, at that time, to correct the entry in the card-catalogue of the egg collection, although I did so on the label of the clutch.
Ardeola speciosa speciosa (Horsfield)
Ardea speciosa Horsfield, 1821, Trans. Linn. Soc. Lond. 21: 189 – Java.
Collector.— De Jong (1).
Material.— None.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)33

Notes.— De Jong’s specimen was recorded by Rensch (1931b: 395), but appears tohave been overlooked by all later authors. Rensch gives for this bird a wing-length of
213 mm, which is large for a member of the nominate subspecies.
Butorides striatus javanicus (Horsfield)
Ardea Javanica Horsfield, 1821, Trans. Linn. Soc. Lond. 13: 190 – Java.
Butorides striatus steini Mayr, 1943, Emu 43: 10 – Dilly, Timor.
Collectors.— Allen, Everett (2), Schmutz.
Material.— , 7.vii.1969, Nunang (Schmutz, RMNH no. 66072); , 15.vii.1969, Nggoér (Schmutz,
RMNH no. 66073). Iris yellow.

Notes.— Both specimens were marked as having large gonads, which suggests local
breeding, but there is no definite record.
The subspecies steini was not based on a great amount of material: Flores (2, presumably the two collected by Everett), Alor (1), Sumba (2) and Timor (1). For comparison, Mayr (1943) had only a single topotypical specimen of javanicus from Java, and one
from Bali (additional specimens from various other islands). In the large series fromJava available in Leiden, the characters stressed by Mayr as diagnostic of steini show agreat deal of variation. I found it impossible to separate the Flores specimens in any
way from birds from Java and therefore it is with some confidence that I assign them tojavanicus. With only two specimens from Timor (one of which is mounted, v.1829,
Koepang, leg. S. Müller, RMNH cat. no. 42), I am not in a good position to judge thevalidity of steini as such, but then, Mayr had only one specimen from Timor. At any rate,
I feel reasonably confident that these birds are also javanicus.
In uniting steini with javanicus, I am in agreement with White & Bruce (1986: 104105), but as other recent authors have retained steini as a valid subspecies (cf. Payne,
1979: 224; Hancock & Kushlan, 1984: 173), it still seemed worth mentioning my conclusions, based on material different from that examined by White & Bruce.
Measurements:



wing
194
185

tail
74
64

tarsus55
52

exposed culmen
64
62

Nycticorax nycticorax nycticorax (Linnaeus)[Ardea] Nycticorax Linnaeus, 1758, Syst. Nat. (ed. 10) 1: 142 – Europa australi.
Collector.— Weber.
Material.— (?) ad., xii.1888, Maumeri (Weber, RMNH cat. no. 52).

Notes.— Hitherto, the above specimen constituted the only published evidence forthe occurrence of N. nycticorax in the Lesser Sunda Islands east of Bali (where it is a
resident; cf. Ash, 1984), but Van der Sande collected one on Sumba ( ad., ii.1909, southwest coast of Sumba, ZMA no. 26011): I have examined the bird, which is in perfect
adult plumage.
The small-scale map in Hancock & Kushlan (1984: 191) is inaccurate, in that it shows34

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

all the Lesser Sunda Islands, as far east as Timor, included in the breeding-range. The
inaccuracy is based on an equally inaccurate map in Cramp & Simmons (1977: 263). On
the other hand, Sulawesi (Celebes) can definitely be added to the breeding-range, on
the basis of the evidence supplied below.
There has been, and still is, much confusion about the status of this species anywhere east of the Line of Wallace. White (1973b), partly on the basis of information
supplied by me, rejected earlier records of breeding in northern Sulawesi (Celebes), as
the two specimens in juvenile plumage in our collection ( juv., 21.viii.1863, Ayer Pannas, RMNH cat. no. 39 and  juv., 7.ix.1863, Panybie, RMNH cat. no. 43, both collectedby v. Rosenberg) were fully grown and would have been well able to fly. Juvenile dispersal over great distances is one of the characteristics of the species.
Rosenberg is supposed to have found both species in northern Sulawesi (Celebes).
Our collection contains six specimens of N. nycticorax obtained there by him, and two
specimens identified as N. caledonicus. Hitherto, everybody has been expecting N. caledonicus to be the common species in Sulawesi (Celebes), and N. nycticorax the unlikely
one. Therefore I had not, previously, bothered too much about Rosenberg’s two juvenile specimens ascribed to N. caledonicus. Now I did so, and was surprised to find thatboth specimens have actually been misidentified and are N. nycticorax. Both are juveniles ( juv., 3.ix.1863, Limbotto, RMNH cat. no. 28 of caledonicus and  juv., 7.ix.1863,
Panybie, RMNH cat. no. 29 of caledonicus). Cat. no. 28 is not fully fledged (wing-length
ca. 200 mm; nest-hairs on the head) and therefore provides definite proof of breeding.
Note that the information about these two specimens, which I forwarded to White(1973b) is erroneous. Cat. no. 29 is not only of the same date and place, but also of thesame size and appearance as specimen cat. no. 43 which has always been correctly
identified as N. nycticorax.
Rosenberg (1881) never claimed to have found more than one species of night-heronin northern Celebes, which he listed under the name of Ardea caledonica, and of which
he stated to have taken 18 specimens. With the re-identification of the two specimens,
there are now 8 specimens of N. nycticorax from him in Leiden, and not a single one of
N. caledonicus. A specimen that found its way to Brüggemann (1876: 98, s. n. Nycticorax
aegyptius) was also N. nycticorax. Evidently, Rosenberg had misidentified the species.
Nycticorax caledonicus hilli Mathews
Nycticorax caledonicus hilli Mathews, 1912, Novit. Zool. 18: 233 - North-West Australia (Parry’s Creek).
Collectors.— Verheijen (1, MCZ), Verheijen/Schmutz.
Material.—  juv., 26.vi.1969, Nunang (Schmutz, RMNH no. 65180);  ad., 3. xii.1969, Waé-Mésé, Kandang, Nisar (Schmutz, RMNH no. 66074);  juv., 28.vi.1976, Wangkung, Rahong (Schmutz, RMNH no.
85147). Iris of adult yellow, bare facial skin and basal two thirds of the mandible green-yellow, remainder of bill black, legs green-yellow.

Notes.— The nos. 65180 and 85147 are in the streaked juvenile plumage. Juvenilesof this species are easily distinguishable from juveniles of N. nycticorax in corresponding plumage, by the strongly rufous tinge of the remiges and rectrices. The literaturetends to be hazy about these differences; for example, Hancock & Kushlan (1984: 193)
only say: “The immature is brown with much buff and white spotting and streaking”,

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)35

without any reference to the young of N. nycticorax. I cannot confirm the claim by Meyer & Wiglesworth (1898: 848) that the two species would differ in the relative proportions of tarsus and middle toe.
My reasons for continuing to use the subspecific name hilli for birds from Australia
and adjacent islands, instead of novaehollandiae as propagated by Schodde & Mason(1980: 18), have been given elsewhere (Mees, 1982c).
There is no evidence yet of breeding of this species on any of the Lesser Sunda Islands, although it is quite likely that it does. I have been unable to find on what theclaim of breeding on Flores by White & Bruce (1986: 106) is based.
Ixobrychus sinensis (Gmelin)
Ardea Sinensis Gmelin, 1789, Syst. Nat. (ed. 13) 1(2): 642 – Sina.
Collectors.— Allen, Verheijen (2, MCZ), Schmutz.
Material.— , 26.xii.1982, Lemboi, 250 m (Schmutz, RMNH no. 81026). Iris yellow.

Notes.— Further records from the Lesser Sunda Islands are: 2  ad., 17 and18.vi.1932, Dilly, Timor (Stein, cf. Mayr, 1944a: 132) and (?), 23.iii.1925, Kambera, Soemba(Dammerman, cf. Rensch 1931b: 373).
Material from other islands, arranged by month is given in the following table.
Material from other islands, arranged by month:
I
II
III
IV
Sumatra
–
–
3
4
Java
6
5
4
1
Bali
4
–
–
–
Borneo1–––
Sulawesi (Celebes)
–
–
–
2

V
–
3
–
–
–

VI
–
1
–
–
–

VII
3
–
–
–
–

VIII
1
–
–
–
5

IX
–
–
–
–
1

X
–
1
–
–
–

XI
–
6
–
2
3

XII
–
20
–
–
–

Supplementary information on birds from Java: the June date is 1 June 1902 (Pangerango, leg. Bartels); the earliest autumn date is October 1827, without exact date(leg. S. Müller); the first subsequent autumn date is 6 November 1925 (Tjisaroeni, leg.
Bartels).
White & Bruce (1986: 108) claim breeding in Java, with a reference to Hancock &Elliott (1978: 60), where Java is listed without any explanation amongst the breeding
places. I have no idea on what this is based.
The table suggests breeding in Sumatra rather than in Java, cf. Robinson & Kloss(1924: 219): “Females (July and August) had developed ovaries”. Sumatra is only referred to as a vague possibility in the publications mentioned above. Hancock & Elliott(l. c.): “Java and possibly Sumatra”, and this is repeated without alteration by Hancock& Kushlan (1984: 246). Further to breeding in Sumatra, there is the definite statement by
Van Marle & Voous (1988: 64): “Female and eggs collected 18 Dec 1914, Perbaungan,
Deli (coll. Waldeck, ZMA)”, which would seem to clinch the matter. Because of the importance of the record, I asked for more particulars, which Mr Roselaar (in litt., 23.
x.1995) provided in great detail. There is a female, with the date cited above (ZMA no.
26034), but collected by De Bussy, not Waldeck. Apparently, De Bussy used to pass on

36

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

duplicate eggs to Waldeck. There are several clutches (and skins) of I. cinnamomeus from
Perbaoengan/Perbaungan in the collection, but none of I. sinensis. Moreover, the specimen was originally identified as I. cinnamomeus, and only in the nineteen seventies reidentified as I. sinensis (by Roselaar). There is no clutch dated 18.xii.1914 in Amsterdam,
but an apparently large part of Waldeck’s egg collection was lost in 1917, when the ship
on which it was transported, the ‘Koningin Emma’, was torpedoed and sunk in themouth of the Thames (Voous, 1995: 537). Mr Roselaar concludes that the available
records do not make clear whether the specimen was actually found in association withthe eggs. A weak point is also the original misidentification as I. cinnamomeus. In theabsence of the eggs, proof either way is impossible, but it seems at least likely that the
specimen of I. sinensis was a visitor and just had the misfortune of being collected at aplace where its congener I. cinnamomeus was nesting.
On the basis of the information provided by the table, breeding in Sulawesi (Celebes)
is also a distinct possibility, although Stresemann (1941: 86-87) called the species onlya “Spärlicher Wintergast” there. Lansdown (1987) records cases of nesting in Malaya(from where it was already known), and North Borneo (“Sabah”), new, in 1986. Johnstone et al. (1996: 165) report the find of a nest with one nestling of this species on Sumbawa, in May 1988; they seem not to have considered the possibility that it was I. cinnamomeus, which is much more likely.
Our collection gives no support to the suggestion by Beehler et al. (1986: 59) of nesting in New Guinea; our dated material is from October (2) and November (16), cf. Junge(1953: 5).
Ixobrychus cinnamomeus (Gmelin)[Ardea] cinnamomea Gmelin, 1789, Syst. Nat. (ed. 13) 1 (2): 643 – Sina.
Collector.— Verheijen (eggs only).
Material.— Eggs: c/3, 20.vi.1955, Dampék (RMNH no. 70751). The eggs are white.
Measurements and weights:

RMNH no. 70751

35.5 × 28.4
36.2 × 29.1
36.3 × 28.0

0.708
0.714
0.758

Notes.— According to White & Bruce (1986: 109): “the only Flores record (Verheijen1964) perhaps a vagrant”, but Verheijen (1964: 195, 198) mentioned no specimen, but
only this clutch, which he provided with a query. In my opinion the identification iscorrect. Both measurements and weights are within the (upper part of the) range ofvariation of I. cinnamomeus and outside (= above) that of I. sinensis as presented by
Schönwetter (1960: 93-94).
Hancock & Kushlan (1984: 254-255), both in their text and in the map, have excluded Sumatra and Java from the range, thus giving a completely misleading picture of the
distribution of this species. It is a common breeding bird in both islands. Our collection
contains 6 clutches of eggs from Sumatra, and 77 from Java, the earliest one of which
was collected in 1827 (Tjikao, leg. Boie & Macklot, RMNH cat. no. 1). See also the description and photographs of nest, eggs and young by La Bastide (1941).
Marin & Sheldon (1987) claim several nests of this species found in 1982 and 1983 inMees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)37

Sabah (North Borneo) as first nesting records for Borneo, commenting: “It is surprisingthat nests of this common padi bird had not been discovered previously in Borneo”. But
there had. In our collection, there is a c/3, 28.ii.1931, Peniti, Pontianak (Coomans deRuiter, RMNH no. 73536), a find which precedes those of Marin & Sheldon by over fifty
years. Coomans de Ruiter (1948a: 62) clearly refers to clutches (in the plural) collectedby him in West Borneo, all consisting of three eggs, but only the one clutch listed above
remained when his private collection was incorporated in the RMNH-collection.
Ciconia episcopus neglecta (Finsch)
D[issoura] neglecta Finsch, 1904, Orn. Mber. 12: 94 – Java, Sumbawa, Lombok, Celebes, Philippinen,
restricted to Java by Rensch (1931a: 502).
Collector.— Verheijen.
Material.— (?) , nestling, 16.x.1970, Pong Nggéok (Verheijen, RMNH no. 65182).
Eggs: c/2, 27.ii.1956, Bénténg Djawa (RMNH no. 70052); c/4, ca. 20.iii.1956, Bénténg Djawa (RMNH no.
70053); c/2, ca. 25.vi.1961, Soa (RMNH no. 70054); c/2, 7.iii.1962, Mengé-Ruda (RMNH no. 70055); c/2,
17.ii.1963, Mongu Luwa (RMNH no. 70056); c/2, 14.vii.1963, Mongu Luwa (RMNH no. 70057). The eggs
are plain white, dull, with a not entirely smooth surface; most are slightly dirty, and the eggs of one
clutch (no. 70053) are strongly stained brownish.
Measurements and weights:

RMNH no. 70052
RMNH no. 70053

RMNH no. 70054
RMNH no. 70055
RMNH no. 70056
RMNH no. 70057

62.0 × 44.6
63.0 × 44.2
60.3 × 42.5
61.4 × 44.2
61.9 × 44.3
62.2 × 44.2
61.6 × 45.6
62.4 × 45.6
63.2 × 45.1
63.7 × 44.0
61.9 × 45.3
62.3 × 45.0
61.4 × 43.6
62.5 × 45.1
63.0 × 44.8
65.2 × 44.7

6.74086.74235.28756.8001
6.72336.12536.47786.48716.7561
6.1062
6.21416.64795.67356.48366.25736.4246

Notes.— This species was previously included in the avifauna of Flores on the basisof field-observations by Rensch (1931a: 503) and Van Heurn (1932).
The validity of the subspecies neglecta is questionable (cf. White, 1974). When Finsch
described neglecta, he did not indicate a type or a type-locality, but only what he believed to be the range of the subspecies, without making clear on what material thesubspecies was based, but a year later (Finsch, 1905: 152) he mentioned that his material consisted of specimens from Java (5), Celebes (5) and Sumbawa (1); he states expressly, that he had not examined birds from the Philippines, and only speculates thatthey belong to the new subspecies. An examination of the material from before 1904 in
our collection, showed the five specimens from Celebes and the one from Sumbawa stillpresent, but from Java I could only find four specimens, without any evidence that a
fifth one would ever have been present. Subsequently, I found in Schlegel (1865a: 10),

38

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

specimen no. 1 (“Adulte, Sennaar, présenté en 1859 par Mr Ruysenaers”), crossed out
with, in Finsch’s handwriting, in pencil, added: “Zweifellos falsch und von Java (D.
neglecta F.)”. The specimen has remained catalogued as C. e. microscelis and, unlike
Finsch, I see no reason to question its identity and provenance. It would, however, explain why Finsch recorded the presence of five specimens from Java. This leaves four
specimens from Java, the restricted type locality of neglecta. They are mounted birds,
cat. nos. 1 and 2, merely labelled “ad., Java, Cabt. Temminck” (which means that theydate from before 1820), Cat. no. 3, , xii.1826, Buitenzorg, leg. H. Boie, and cat. no. 4, ,iii.1870, Java, without locality or name of collector. Only cat. no. 3 is properly labelled,
and in addition, it has a type-label attached to it by my predecessor as curator of thebird collection, G.C.A. Junge. There is every reason, therefore, to accept it as lectotype.
Although in their text, Hancock et al. (1992: 82-83) give the known range in theLesser Sunda Islands correctly as Bali, Lombok, Sumbawa, and Flores, in their distributional map all the Lesser Sunda Islands, including Sumba, Timor and Wetar have beencoloured in.
As Van Marle & Voous (1988: 66) knew of only one specimen record from Sumatra,
I mention here the presence in our collection of the following specimen: , 27.vi.1920,
Soekarami, Palembang, leg. J.C. Batenburg (original label, RMNH no. 20405). Soekarami
is a little west of Sekajoe on the Moesi.
Platalea regia Gould
Platalea regia Gould, 1838, Synops. Birds Austr. 4, app.: 7 – East coast of New South Wales.
Collector.— Schmutz.
Material.— , head and one wing only, 1.viii.1969, near the mouth of the Waé Djamal, opposite NangaLili (Schmutz, RMNH no. 85037). Iris red, bill, face and legs black. Stomach contents small crabs.

Notes.— This bird was obtained from a flock of nine. Presumably, these were stragglers from Australia. There has been speculation, whether this bird should be considered a separate species or a subspecies of P. leucorodia. Most recent authors (with theimportant exception of Steinbacher, 1979: 267) have treated it as a separate species, following arguments presented by Amadon & Woolfenden (1952). However, lately it has
been suggested that not P. leucorodia, but P. minor is its closest relative (Hancock et al.,
1992: 261). These authors did not go so far as to treat P. regia and P. minor as conspecific,
and I follow them in this cautious approach.
Pandion haliaetus cristatus (Vieillot)
Buteo cristatus Vieillot, 1816, Nouv. Dict. d’Hist. Nat. (nouv. éd.) 4: 481 – la Nouvelle Hollande.
Pandion haliaetus melvillensis Mathews, 1912, Austral Av. Rec. 1: 34 – Melville Island, Northern Territory.
Pandion haliaëtus microhaliaëtus Brasil, 1916, Rev. Fr. d’Orn. 4: 201 – Nouvelle Calédonie.
Collectors.— None.

Notes.— Schmutz (MS, 1977) repeatedly observed Ospreys along the coast, in the
mangrove and along the beach. Near Look, a feather was found floating in the water(Vo 309a). This widely-distributed species is present throughout the archipelago, andMees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)39

there can be no doubt that it occurs regularly on Flores and the adjacent smaller islands.
This does not mean that the present records from Komodo, Padar and Rintja are acceptable. White (in White & Bruce, 1986: 113) correctly rejected them, as Pfeffer records. But
on the next page, they are re-instated by Bruce, with a reference to an FAO list (probably
unpublished, and not available to me); however, it is a safe assumption that the FAO list
was based on Hoogerwerf’s (1954b) report. The source of the records of Pandion haliaetus from these islands would therefore be the same as Pfeffer’s (and based on the same
mistranslation).
Within a year after its description, the subspecies melvillensis, based on “whiter headand smaller size” (no measurements given), was withdrawn by its author (Mathews,
1913: 113; 1916: 302), “as the characters are not constant and I find, moreover, the Australian birds, as a whole, constitute only an ill-defined subspecies”. Mathews occasionally had these flashes of insight. The matter was further complicated by the introduction of a second subspecies of supposedly small size in the region: P. h. microhaliaëtus,originally described from, and thought to be confined to, New Caledonia. Because of
the geographical distance separating the type-localities, its author did not consider it
necessary to compare microhaliaetus with melvillensis. Next came Swann (1922: 233),
who extended the range of microhaliaetus to “New Caledonia; Celebes”, compared withcristatus: “Smaller; wing (New Caledonia), 392-412, wing (Celebes),  383,  430, tail175 mm”.
Meise (1929: 479-480) measured the material in the Berlin and Tring (Rothschild)museums, and arrived at the very definite conclusion: “daß auf Grund der Größe keine
weitere Einteilung dieser von Celebes bis Australien verbreiteten Subspecies möglich
ist .... Es mag sich wohl ein größerer Durchschnitt für den Flügel der Australischen
Fischadler ergeben, aber 3/4 oder mehr aller dortigen Stücke könnte man nach den
Maßen nicht bestimmen. Ein besonderer Name ist also für die Inselbewohner nicht
angebracht”.
One might have considered this conclusive, but it was accepted by Stresemann(1940a: 487) only with qualifications: “Die subspezifische Stellung der Celebes-Vögel ist
noch umstritten, ihre Identität mit australischen zweifelhaft”.
The next to study the problem was Amadon (1941), who made no mention of
Meise’s publication, although for a large part he examined the same material as Meise,that in the mean time had been sold from Tring (Rothschild) to New York. Nevertheless, he arrived at an opposite conclusion. He measured an impressive number of specimens from the tropical part of the range, including northern Australia and MelvilleIsland, and opposed them to five specimens from “Central and southern Australia”,
which were clearly larger. These last five he listed individually, thus showing that certainly one of them is misplaced: a large  from Point Torment, Western Australia, for
Point Torment, near Derby (ca. 17°30’S) is well up into the tropics. The two other Western Australian birds, from Lewis Island and Point Cloates are also from within thetropics, of which, indeed, Amadon was aware. His two remaining specimens werefrom Port Mackay, Queensland, and New South Wales. Port Mackay is also in thetropics, so that the New South Wales bird remains as the only genuine non-tropical
Australian specimen. It is indeed the largest of the whole series (, wing 480 mm).
Amadon also refers to the supposed type of P. leucocephalus from Tasmania in Philadelphia, more about which will be said below (, wing 455 mm - very likely mis-sexed,40

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

but to Amadon it supported his opinion that southern birds are large).
Amadon included New Caledonia into the range of the smaller tropical subspecies
P. h. melvillensis, placing P. h. microhaliaëtus in its synonymy.
Probably following Amadon, most subsequent authors (Mayr, 1941b: 18; Delacour,
1947: 52; van Bemmel, 1948: 349; Hoogerwerf, 1949a: 36; van Bemmel & Voous, 1951:
82-83; Vaurie, 1965: 144; Stresemann & Amadon, 1979: 279) recognized melvillensis. Also
Rand & Gilliard (1967: 86), who, however, extended the range of melvillensis to Tasmania, thus usurping the presumed type-locality of cristatus. The same error is found in
Simpson (1972: 100).
Gyldenstolpe (1955: 361) doubted the validity of melvillensis, but accepted it as helacked the material for a proper evaluation.
In the ensuing years, Amadon seems to have lost some confidence, for Brown &Amadon (1968: 196) regarded melvillensis as: “Doubtfully distinct; very similar to P. h.
cristatus, differing in being smaller” (this is followed by a summary of the measurements published by Amadon in 1941).
Frith & Hitchcock (1974: 127) considered it: “most unlikely that more than one raceof this world-wide species is represented in Australia”, and commented on the inadequacy of Amadon’s Australian material. Via Condon (1975: 76), who neither then nor in
his earlier publications accepted more than one Australian form, we come to White(1975b), who “assembled [does that mean, personally took?] wing measurements ofthirty-one specimens from Wallacea. The wings of twelve males are 413-439 mm, rather
longer on average than the range for melvillensis, which is given by Brown & Amadon
as 384-428 but about the same in length as those of cristatus males, given as 426-431. I do
not regard melvillensis as recognisable”. But Dickinson et al. (1991: 119) again recognized melvillensis. Marchant & Higgins (1993: 225, 233) admit only one Indo-Australiansubspecies, for which, without explanation, they use the junior synonym leucocephalus
instead of cristatus.
The latest word about geographical variation of the Osprey in Australia is by Olsen& Marples (1993). These authors introduced a new character, size and shape of the eggs,
and distinguish again two subspecies in Australia, one of which is confined to theSouth-West of the Continent. For this subspecies, they use the name subcristatus; thename confused me considerably, as it seemed to indicate a Mathewsian creation that I
had overlooked. It took some time, before I realised that the subcristatus of these authors
is no more than a lapsus for cristatus. Note that, without comment, cristatus has beenshifted away from its traditionally- accepted type-localities (Tasmania and New South
Wales), and that eastern Australia has been included into the range of melvillensis.
The continuing doubt about the validity of P. h. melvillensis, naturally made me eager to form my own opinion. To this purpose I examined the material from the melvillensis/cristatus range in the RMNH, and took wing-measurements. These were takenwith a tape, following the bend of the wing, to the nearest 5 mm (see table).
Pandion haliaetus cristatus (Vieillot): specimens examineddateisland
7.ix.1906Java
 juv.
9.ix.1906“

27.xii.1912“

6.x.1920“

locality
P. Lang“
Moeara Angke
P. Bokor

wing-length
430
430
470
430

notes

1

)

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

41


11.viii.1927“
P. Lantjang435

ca. 1841
Bawean430

2.ix.1844
BorneoPagattan
430
2)

10.x.1863
Sulawesi (Celebes) Poë450

14.vi.1914
“
Mara
435

9.x.1939“
Likoepang420
30.vi.1861
Ternate425

9.vi.1910
Ceram
P. Kasoeari
435
30.ix.1873
Ceram/Amboina
410
2
)

17.vi.1898
Babar
445

19.ix.1948
MisoolFakal
425

19.vi.1865
Aru Isl.
435

26.i.1869
Mefoor (= Numfor)430

25.vi.1903
New Guinea
L. Sentani
425
3
(recd. 1861)
Australia
460)
 juv.
7.ix.1906
Java
P. Lang460
15.vi.1912“
P. Bokor465
(1841)
Bawean–455
(1883)
Sulawesi (Celebes) Menado500

20.vi.1914
“
Mara
465

18.x.1939“
Bangka Isl.
470
(1866)
Siao475
1

4.viii.1865
Sanghir425)

3.xi.1865“455

24.i.1866“455

24.vi.1862
Ternate450

2.x.1863
Moti
460
1

31.xii.1861
Morotai420)
4

11.i.1861
Batjan475)
25.vii.1923
Buru
Djikoe-Merasa
470(1842)
Ceram
Kaibobo470

27.ii.1897
New Guinea
Sekroe4805)
22.vi.18(78)“Doktien
445

1875
“
Doreh
470
6

8.xii.1953
Biak
470
)

1.iii.1881
Duke of York460
(1865)New Caledonia
450

(1841)
Australia
490
1) Almost certainly mis-sexed.
2) Possibly mis-sexed.
3) Provenance questionable, and probably nominate race, see text.
4) Probably nominate race, see text.
5) Locality not traced; from A. A. Bruijn.
6) Junge (1956) recorded for this specimen a wing-length of 451 mm. He must have used a method of
measuring very different from mine, unless the figure is a misprint.

Unfortunately, the series is even more deficient in Australian material than Amadon’s: only a single specimen, which is large (wing 490 mm), but being without locality,
has little value. In spite of this deficiency, the list of measurements is most instructive.
It shows that birds from Sulawesi (Celebes) are not small: indeed, one specimen fromthere is the largest of the series. A bird from New Guinea (wing 480 mm) is as large as
Amadon’s largest Australian bird, from New South Wales. None of my specimens has

42

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

a wing-length of less than 400 mm, as recorded by several previous authors (perhaps adifference in the method of measuring?). In the following paragraphs, I shall try to showthat the Osprey does not occur in south-eastern Australia and Tasmania, the supposed
type-locality of cristatus, which eliminates this region as the epicentre of a special, large
subspecies. Measurements from Western Australia presented by Johnstone & Storr(1998: 128), some of which would be of extra-tropical birds (wing 425-475 mm, sexes
combined) are within the range of variation of the birds measured by me. Little is
known of the isolated South Australian population, but as no name has been based onit, and as Condon, probably the only ornithologist who had access to material, did not
accept more than one Australian subspecies, it can safely be left out of discussion. The
egg-evidence provided by Olsen & Marples (1993) is intriguing, but requires confirmation, especially as the Osprey has an almost unbroken range from the south-west into
the tropics of Western Australia. Even if confirmed, it would only doubtfully qualify forrecognition in nomenclature. Therefore I join with confidence those authors who reject
melvillensis and place it in the synonymy of cristatus.
The type locality originally given for Pandion haliaetus cristatus was “la Nouvelle
Hollande”. Mathews (1912: 254), without explanation, misquoted this as New South
Wales. The following year, Mathews (1913: 113) substituted Tasmania, again without
mention of the originally-given type locality, and without stating that this was a restriction. Later, he corrected the omission, quoting both the originally-given type locality,
and the restriction to Tasmania, true, without an explanation of the basis for this restriction (Mathews, 1915/1916: 254; 1927: 267). The restriction is not an obvious one, asPandion haliaetus is of problematical occurrence in Tasmania (Green, 1977: 17). In the
19th century, however, Gould is believed to have personally taken a specimen in the
Recherche Bay, southern Tasmania (Gould, 1848, 1865: 22). The specimen still exists andwas thought to be the type of Pandion leucocephalus Gould (a junior synonym of P. h.
cristatus), by Stone & Mathews (1913: 147). De Schauensee (1957) has not followed this,and rightly, for the original description of P. leucocephalus was communicated at themeeting of the Zoological Society of 26 December 1837, and was first published in April
1838. Gould only arrived in Tasmania (or Van Diemen’s Land as it was then called), in
September 1838 (cf. Whittell, 1954: 88). In both descriptions of P. leucocephalus, the type
locality is Australia, without any indication that would justify a restriction.
In spite of Gould’s (1848) definite statement (made some ten years later): “I myself
shot it in Récherche Bay, at the extreme south of Van Diemen’s Land”, and the presenceof a mounted specimen so labelled, I must admit to having a slight but gnawing doubt
about his record of Pandion haliaetus from Tasmania; one reason is that if the species
does occasionally visit Tasmania (as I am quite prepared to believe), Recherche Bay on
the south coast is about the last place one would expect it to reach. The other is that
Haliaeetus leucogaster is common there, and presumably was so in Gould’s time, but was
not especially mentioned by him. Even Australians are not always fully aware of thetotal lack of reliable records from Tasmania. For example, Sharland (1958: 88) has thisto say about its occurrence there: “Uncommon...The Osprey is so little known in Tasmania that odd birds which appear along the coast could be passed off for Sea Eagles
if not carefully examined...If it does inhabit Tasmania and is not merely a visitor fromthe mainland, then it must keep to remote parts of the coast, because it is seldom seen.
Occasionally a bird turns up on Flinders Island”. Simpson (1972: 100) claims that:

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

43“Although they range right around the Australian and Tasmanian coastline...”.
Even in Victoria, the Osprey is a surprisingly rare visitor. Wakefield (1958) stated
that: “There is no authentic record for Victoria”, and several of the few subsequent
records have later been queried (Cooper, 1975: 86).
Nowhere does Mathews say on what his fixation of Tasmania as the type localityof P. h. cristatus was based, but it is apparent that this was because he (mistakenly) believed Tasmania to be the type locality of the synonym P. leucocephalus. Even had hebeen right about Gould’s Tasmanian bird, there is no logic here: the fact that Gould may
have obtained a bird in Tasmania in 1838/1839, can have no bearing on the place in “laNouvelle Hollande” where, some thirty years earlier, a French expedition collected thetype of P. h. cristatus. Hartert’s (1929) correction of the type-locality of cristatus: “Australia, not Tasmania”, was published in too succinct a way to have had much impact onlater revisers. Amadon (1941) accepted the restriction to Tasmania, and rejected Hartert’s correction, on the assumption that the French expedition (the Expédition Baudin)which may have brought the type of P. h. cristatus home: “did most of its work in Tasmania”. This is not entirely correct, for considerable time was spent along the coast ofWestern Australia, and visiting coastal islands, where P. haliaetus is still a regular breeding-bird. It illustrates well the arbitrary way in which some ornithologists made type
locality restrictions and designations in the first quarter of the 19th century.
The examination of the material from the Archipelago leads to some further comments. Smythies (1957: 585) mentions that Coomans de Ruiter had found the species
breeding on the Karimata Islands, off West Borneo, but Smythies has misread this publication, for in it, Coomans de Ruiter (1936: 49) expressly states that he never found thespecies breeding and had never obtained or received its eggs. Smythies must have become aware of his error very soon, for three years later he did not mention breeding andeven stated that the resident subspecies melvillensis or cristatus was not known from
Borneo, all records of Ospreys from Borneo to date being of migrants of the nominate
race (Smythies, 1960: 160). Twenty years later the position was still the same (Smythies,
1981: 57), albeit with the addition of some possible sight records. However, a mounted
, 2.ix.1844, Pagattan, Borneo, collected by Schwaner (RMNH cat. no. 5), belongs undoubtedly to the subspecies cristatus. It was already recorded by Schlegel (1862: 23).
Considering that P. h. cristatus is not uncommon and ranges throughout the Archipelago, it is surprising how few breeding records there are. The only definite records I
know of are from Madu and Kalao tua (v. Plessen, in Meise, 1929: 480), and West Java
(Hoogerwerf, 1969/1971: 462). White & Bruce refer to breeding on Lombok, of which I
have failed to find the reference, unless it is Rensch’s statement that a bird collected
there had large gonads. There is no evidence that the population is augmented in thesouthern winter by migrants from Australia, as has sometimes been suggested (cf.
Dickinson et al., 1991: 120). Towards the southern part of the range in Western Australia, the adult birds are sedentary, and the young disperse, rather than migrate(Holsworth, 1965).
Of the two birds labelled as being from Australia in the RMNH collection, I haveno reason to doubt the provenance of one (RMNH cat. no. 1, purchased from Frank in1841). The other one, however, has the feathers of the head conspicuously brown andI suspect strongly that it is a mislabelled specimen of the nominate race. This bird (RMNH
cat. no. 2), was also purchased from Frank, in 1861, and bears a note on its origin:44

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)“voyage de Gardner” (see also Schlegel, 1862: 23, where it is cat. no. 10). In the remarkably complete work of Whittell (1954), I have failed to find a Gardner who could have been
active in Australia around 1860, and this compounds the doubt I have about the specimen. Gardner is more likely not to have been a traveller, but the London natural history
dealer of that name, who was in business around the middle of the 19th century andtraded in bird-skins from all continents (Salvin, 1882: xi). With such a background, there
must have been ample opportunity for a mix-up in labelling. Brown & Amadon (1968:
196) differentiate P. h. cristatus and P. h. melvillensis on the basis of small size and by having: “a pure white head, not streaked on crown”. White (1975b) qualified this: “It should
be noted that the crown in cristatus is not always pure white and some dark brown spots
may be present”. But all specimens examined by me have at least a narrow brown median band, longitudinally, from the hind-crown to the nape. Some specimens fromwithin the range of the nominate race have heads almost as white as cristatus, and support Mathews’s opinion, quoted above, that cristatus is only an ill-defined subspecies.
The statement by Dickinson et al. (1991: 119) that this species: “fishes inland, andmore rarely coastal”, is certainly not applicable to P. h. cristatus, which is almost entirely
coastal in distribution and feeds mostly on marine and estuarine fish.
Addendum.— There are still no recent records from Tasmania (Green, 1993: 14) andmy doubt about Gould’s record increased so much that in my opinion the species
should be eliminated from the Tasmanian avifaunal list. Davison (in Smythies, 1999:
209) repeated the erroneous claim first made by Smythies (1957), but soon withdrawn
by that author, that Coomans de Ruiter would have found this species nesting in West
Borneo. He cannot have read the paper by Coomans de Ruiter, as is also evident fromthe fact that he omits mention of nidification of Accipiter virgatus in lowland West Borneo, reliably recorded in the publication (Coomans de Ruiter, 1936) and further discussed by Mees (1981: 381-382.
To the meagre records of nesting in the Archipelago, Komodo can be added (Bishop
in Verhoeye & Holmes, 1998: 13). In this publication the error is repeated, based on a mistranslation by Pfeffer, that Hoogerwerf would have seen Pandion haliaetus on Komodo.
Aviceda subcristata timorlaoensis (A.B. Meyer)
Baza timorlaoënsis Meyer, 1893, Abh. Mus. Dresden, 1892/93 (3): 5 – Timorlaut.
Collectors.— Weber, Everett.
Material.— , xii.1888, Maumeri (Weber, RMNH cat. no. 1).

Notes.— It is perhaps surprising that, since Everett obtained “a series” in 1896, thisconspicuous bird has not been recorded again from Flores. The use of the name timorlaoensis for the Flores population follows tradition and current usage, but is by no means
definitive. Lack of adequate material prevents me from going into this. A few notes on
the adjacent subspecies pallida, described from the Kei Islands, are, however, perhaps
not out of place.
The original description of pallida reads: “In der Färbung wie die vorigen Formen,
aber kleiner”; the two preceding forms are the nominate one, and timorlaoensis. Peculiarly, Stresemann says nothing about differences in plumage, and therefore it is difficult
to understand why he regarded the name pallida as appropriate. Actually, the meticulous

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)45

Siebers (1930: 249) observed that some specimens of topotypical pallida had very pale
under tail coverts. This was followed by the descriptions of pallida by Brown & Amadon(1968: 210) as “very pale ... pale grey upper breast”, and by White & Bruce (1986: 115):“paler above and on the breast, ventral barring less heavy”. I can confirm that the two
specimens in Leiden also show the character of paleness, and thus support the validityof pallida.
Measurements:


wing
303

tail
207

tarsus33

exposed culmen
26

culmen from cere
19.8

Pernis ptilorhynchus orientalis Taczanowski
Pernis apivorus orientalis Taczanowski, 1891, Faune orn. Sibérie Orientale, Mém. Acad. Imp. Sci. St.-Petersbourg (7) 39: 50 – Koultouk sur le Baïkal méridional; l’embouchure de l’ Oussouri au 48° L. N., et....
l’îlot Askold au 43° L. N.
Collector.— Schmutz.
Material.— , 2.xii.1971, Lo Kong, 900 m (Schmutz, RMNH no. 81111).

Notes.— This is only the third specimen of P. ptilorhynchus orientalis to be recorded
from anywhere east of Java, the previous records being from Saleyer and Kisar (cf.
White, 1973a). The measurements of the specimens from Flores and Kisar prove thatthey belong to the subspecies orientalis, a migrant from north-east Asia, as was to be
expected. According to Stresemann (1940b: 163), the specimen from Saleyer is alsoorientalis.
The subspecies wintering in Java has generally been considered also to be orientalis
or its synonym japonicus (cf. Bartels in Hartert, 1922: 2207; Bartels, 1923; Chasen, 1935:
79; Stresemann, 1940b), but Vaurie & Amadon (1962) decided (on the basis of one immature specimen from that island, and a re-interpretation of a series of measurements
published by Stresemann but provided by Bartels), that it is ruficollis and not orientalis
which winters in Java. This was repeated by Vaurie (1965: 149), but Brown & Amadon(1968: 221) and Stresemann & Amadon (1979: 287-288) have quietly and without explanation reverted to excluding ruficollis and including orientalis in the avifauna of Java. In
the meantime, Wells & Medway (1976) had demonstrated convincingly that two winterbirds from Perak were orientalis (a specimen from Trang showed somewhat mixed characters) and concluded that the occurrence of ruficollis in the Malay Peninsula remained
unconfirmed. In these later publications it is also suggested that ruficollis, which breeds
in south-eastern Asia, is not or at most slightly migratory. Nevertheless, the series fromJava in the RMNH (mostly from the Bartels collection), deserves further study, forwhich, unfortunately, I did not have the time before leaving Holland.
In the original description of this species, the spelling of the name was Falco ptilorhyncus (cf. Temminck, 1821: pl. 44). It was emended to Pernis ptilorynchus in the text,
and to Pernis ptilonorynchus on the caption of the plate, by Stephens (1826: 44, pl. 35).
Subsequently to Falco ptilonorhynchus by Temminck (1839: 3) himself. Later, another
emendment, P. ptilorhynchus, came into general use (for example, Stresemann, 1940b),so that there was a choice of five different spellings. As ptilorhynchus is now the mostcommonly used spelling, I am also using it, although I realise very well that under the46

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Rules the original spelling, ptilorhyncus would have to be retained. Cases like this one
have been sanctioned in the new edition of the Code (ICZN, 1999: art. 33.2.3.1).
Measurements:

wing

tail

tarsusFlores 
Kisar ca. 470
ca. 470

257
280

63
65

bill from forehead feathers
40(bill damaged)

bill from
cere
26

wing
tip
147
151

Both specimens are in moult. The bird from Flores shows moult in wings and tail;in the wings, on each side, primary 5 is very short, primary 6 is short; curiously, thefollowing primaries (7-10) are not clearly new. The bird from Kisar is in the last stage ofprimary moult, with only the outer (1st) primary not yet fully grown; its tail moult iscompleted. Vaurie & Amadon (l.c.) have illustrated the tails of adult and juvenile individuals, showing the adult female with three weaker bands between the dark terminal
and middle bands, the juvenile female with four. The Kisar bird, although its tail cannot
have the juvenile pattern (as it has just been moulted), has six of these bands, pointing
to a considerable amount of individual variation in this character, as indeed Vaurie &Amadon mentioned.
Elanus caeruleus hypoleucus Gould
Elanus hypoleucus Gould, 1859, Proc. Zool. Soc. Lond. 27: 127 – Vicinity of Macassar, Celebes.
Collectors.— Van der Sande (1, ZMA), Verheijen (1, MCZ), Schmutz.
Material.— (?), 18.iii.1908, flat coastal region east of Maumere (Van der Sande, ZMA no. 13729); ,
1.vii.1969, Tjeréng (Schmutz, RMNH no. 66085); ? nestling, 14.vii.1969, Tjeréng (Schmutz, RMNH no.
66086). Iris of adult red, of nestling dull brown.
Egg: c/1, 2.viii.1955, Bénténg Djawa (RMNH no. 70064).
Measurements and weight:

RMNH no. 70064

40.7 × 32.5

1.785

Verheijen (1964) listed breeding in August and October, but the October-record was based on misidentified eggs, and has to be withdrawn. The nestling provides evidence for eggs in late May/June, and sodoes one recorded by Paynter (1963)

Notes.— The two adult specimens have been discussed, and their measurements
provided, elsewhere (cf. Mees, 1982a: 24-34).
Stresemann (1939: 318) has given a reconstruction of the distributional history ofthis subspecies: “E. c. hypoleucos [sic] hat folgendes Wohngebiet: Philippinen, Celebes,
Kalao, Sumba, Lombok, Java, Sumatra, Süd-Borneo. Bei dieser Art vermute ich die Ausbreitung von den Philippinen aus nach Süden über Celebes-Flores, von dort ostwärts
bis Sumba, westwärts bis Sumatra”. Since this was written, the range of E. c. hypoleucus
has been filled out further, in particular by its discovery in Timor and New Guinea.
Interestingly, on 28.ix.1984 I observed an individual on the islet of Goeang, off the west
coast of Saleyer, another link between Sulawesi (Celebes) and Flores. Nevertheless, inthe light of present knowledge, Stresemann’s reconstruction is no longer the obvious
one. Some years ago I measured a fair amount of material of this form (Mees, l.c.) andfound that continental specimens (from Thailand and Malaya, poorly represented inMees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

47

collections) average about 5% smaller in linear measurements than most insular populations. Specimens from Sumatra, however, average some 2% smaller than birds from
Java, Sulawesi and the Philippines; in other words they are intermediate between continental birds and birds from Java, etc. It seems more logical to consider the Sumatran
birds an intermediate link, than to believe that first the Philippines were colonized fromthe Asiatic mainland, with an increase in size, and that the subspecies subsequently
extended over Sulawesi (Celebes) and Flores to Java, and finally to Sumatra. where its
size became autonomically reduced when it approached the continental populations.
Another point is that in Stresemann’s reconstruction one would expect the species tohave come by way of Taiwan, rather than with a direct long jump from southern China
to the Philippines, but that at present it does not occur on Taiwan. Finally, the conceptof E. c. hypoleucus coming from Asia and spreading comparatively recently to the Philippines and Indonesia, seems somewhat simplistic in the light of the knowledge that thegenus Elanus must be a fairly old resident in Australia, as is apparent from the presenceof two species there. As I have pointed out before, E. c. hypoleucus is as well-differentiated as other forms of its genus, currently regarded as distinct species. It may be assumed to be an old resident rather than a new colonist in the region, but if an expansion
in the sense understood by Stresemann has taken place, it is likely to have been throughThailand/Malaya (small) and Sumatra (intermediate) to Java, etc. (large). The as yet
insufficiently-known New Guinea population may be the largest of all, which fits into
this concept.
The genus Elanus, with its limited number of well-differentiated forms, distributed
over six continents, would make an excellent subject of a comprehensive zoogeographical study or a monograph.
Haliastur indus intermedius Blyth[Haliastur] intermedius Blyth, 1865, Ibis (n. s.) 1: 28 – Java.
Collectors.— Semmelink, Allen, Everett, Verheijen/Schmutz.
Material.— (?), 1862, Larantoeka (Semmelink, RMNH cat. no. 16); , 29.ix.1968, Nunang (Schmutz, RMNH
no. 65186); , 17.i.1969, Nunang (Schmutz, RMNH no. 65185); (?), 17.iii.1976, Nunang (Schmutz, RMNH
no. 81037).
Eggs: c/1, not dated, Soa (RMNH no. 70077); c/1, 3.vi.1954, Potjong (RMNH no. 70078); c/2, 29.iii.1955,
Tado (RMNH no. 70079); c/2, 12.v.1955, Montjok, Tado (RMNH no. 70080); c/2, 18.vi.1956, Bénténg
Djawa (RMNH no. 70081); c/2, 3.v.1957, Potjong (RMNH no. 70082); c/2, 17.v.1957, Mano (RMNH no.
70083); c/2, 17.vi.1959, Pongkor (RMNH no. 70084); c/2, 20.vi.1959, Poéng (RMNH no. 70085); c/2,
29.xii.1959, Poéng (RMNH no. 70086); c/2, 5.iii.1961, Soa (RMNH no. 70087); c/2, 13.iii.1961, Soa (RMNH
no. 70088); c/2, 21.iii.1961, Mataloko (RMNH no. 70089); c/2, 4.v.1963, Soa (RMNH no. 70090). The eggs
are dull white, sparsely marked with red-brown dots.
Some measurements and weights:

RMNH no. 70079
RMNH no. 70084
RMNH no. 70086
RMNH no. 70090

51.7 × 43.1
52.8 × 43.6
50.0 × 42.4
50.1 × 42.4
51.8 × 42.2
53.9 × 42.9
51.1 × 41.9
53.9 × 41.9

3.78233.63633.93044.02033.85044.03033.5951
3.9355

48

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Haliaeetus leucogaster (Gmelin)[Falco] leucogaster Gmelin, 1788, Syst. Nat. (ed. 13) 1: 257 – no locality.
Collectors.— v.d. Sande (1), Rensch (1).
Material.— , 24.vi.1908, Celebesbaai (v.d. Sande, ZMA no. 13596).

Notes.— The skeleton from Bima, listed without comment amongst the birds from
Flores by Büttikofer (1894: 290), was of course from Bima, Sumbawa. It was returned to
Amsterdam, where it cannot now be found.
Circaetus gallicus gallicus (Gmelin)[Falco] gallicus Gmelin, 1788, Syst. Nat. (ed. 13) 1: 259 – Gallia, rarior in reliqua Europa.
Collectors.— Semmelink, Everett (1), Verheijen/Schmutz.
Material.— (?), 1862, Larantoeka (Semmelink, RMNH cat. no. 12);  juv., 3.x.1971, Ruteng (Verheijen,
RMNH no. 85133); , 20.i.1973, Nunang (Schmutz, RMNH no. 81114).

Notes.— Previous records of this species in the Lesser Sunda Islands are from Lombok, Sumbawa, Flores, Roti and Timor (cf. Mees, 1975b: 120-123). Westwards it is now
known to reach Bali, and even the extreme East of Java (van Balen & Compost, 1989).
On 16.ix.1984, I observed an individual near Tg. Penitie, eastern Sumba, thus addingthat island to its range. In order not to be accused of doing exactly what I condemn inothers: making ex cathedra statements about observations new to islands, I translatehere a passage from my diary about this observation: “walking along an interesting,although not particularly well-wooded, ravine, my attention was drawn by a not very
long, whistling call: ‘kyüü … kyüü … kyüü …’, and after a moment an eagle appeared
and flew rather low overhead. What drew my attention, apart from its huge size, wasthat the WHOLE under wings as well as the under surface of the body showed darkbrown bars on cream. In these regions there is no other eagle showing this colour pattern”, The voice further supports the identification.
In spite of the increased number of specimens now available, I am still unable to distinguish the isolated populations of the Lesser Sunda Islands satisfactorily from the nominate race. Therefore I feel obliged, still somewhat reluctantly, to keep these birds under
the name of the nominate race. The reluctance is due to the fact that these populations
must have been isolated for thousands of years, and that Hieraaetus fasciatus, which presumably has a similar history in these regions, has developed an endemic subspecies. The
fact that they have not differentiated does, of course, by no means decrease the zoogeographical interest these isolated populations have. I consider it likely that the originalcolonisation of the Lesser Sunda Islands took place through an open country corridor
from south-eastern Asia, in the Late Pleistocene, rather than that this isolated population
would result from northern migrants that have remained behind in their winter quarters.
In India the species is still widely distributed in open country (cf. Tyto longimembris).
Measurements:

wing

tail

tarsus

 ad.
(?)

530
ca. 525

268
290

96
93

exposedculmen42
50

culmenfrom cere
32
35

depth
culmen17
19.2

middletoe
49
45

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

49

Accipiter soloensis (Horsfield)
Falco Soloënsis Horsfield, 1821, Trans. Linn. Soc. Lond. 21: 137 – Java (by inference: Solo).
Collectors.— Everett (1), Schmutz.
Material.— , 18.x.1971, Tjeréng-Look (Schmutz, RMNH no. 84863); , 28.xi.1978, Tjeréng-Look(Schmutz, RMNH no. 81044); , 2.xii.1982, Nunang (Schmutz, RMNH no. 81043); , 9.xii.1982, Nunang(Schmutz, RMNH no. 81047); , , 14.xii.1982, Nunang (Schmutz, RMNH nos. 81051, 81046); , 14.
i.1983, Nunang (Schmutz, RMNH no. 81316); , (no date)Nunang (Schmutz, RMNH no. 81048).

Notes.— Previously, Accipiter soloensis was known on Flores from a single immature
male only, obtained by Everett in 1896. The new records show it as a regular wintervisitor. For further notes on this species, see Mees (1981: 398-400).
Accipiter novaehollandiae sylvestris Wallace
Accipiter sylvestris Wallace, 1864, Proc. Zool. Soc. Lond. (1863): 487 – Flores.
Collectors.— Semmelink, Allen, Colfs, Weber, Everett, Endih (2), Rensch (1), Verheijen (1, MCZ), Verheijen/Schmutz.
Material.— ,  juv., 1862, Larantoeka (Semmelink, RMNH cat. nos. 2, 3); , 1880, Flores (Colfs, RMNH
cat. no. 4); , xii.1888, Sikka (Weber, RMNH cat. no. 5); , 2.vi.1969, Nunang, 650 m (Schmutz, RMNH
no. 65188; taken from nest with eggs);  juv., 7.vii.1969, Nunang (Schmutz, RMNH no. 66081);  juv., 
juv., 3.ix.1969, Nunang (Schmutz, RMNH nos. 66082, 66079); , 18.ix.1969, Rekas (Schmutz, RMNH no.
84860);  juv., 5.xi.1969, Nunang (Schmutz, RMNH no. 84859);  med., 26.ii.1976, Nunang (Schmutz,
RMNH no. 81041); , 2.xi.1982, Nunang (Schmutz, RMNH no. 81050). Iris  and  dark brown, cere,
eyelid and legs golden yellow, bill and nails black. An adult bird collected in November is in heavy
moult, birds collected in April and September show no moult, the former is in moderately, the latter inmore strongly worn plumage.
Eggs (fig. 12a): c/1, ix.1948, Mataloko, ca. 700 m (RMNH no. 70069); c/2, 4.iii.1956, Bénténg Djawa(RMNH no. 70070); c/2, 28.iii.1958, Tado (RMNH no. 70067); c/1, 20.vi.1959, Poéng (RMNH no. 70068);
c/1, 9.v.1959, Potjong (RMNH no. 70071); c/2, 3.vi.1960, Potjong (RMNH no. 70072); c/2, 27.iii.1961,
Mataloko (RMNH no. 70073); c/2, 2.vi.1969, Nunang (Schmutz, RMNH no. 75299, taken with skin no.
65188).
The eggs nos. 70067 and 70068 were originally identified as A. virgatus (see the discussion of that species).
Measurements and weights:

RMNH no. 70069
RMNH no. 70070
RMNH no. 70067
RMNH no. 70068
RMNH no. 70071
RMNH no. 70072
RMNH no. 70073
RMNH no. 75299

42.0 × 34.7
41.9 × 34.0
42.8 × 34.1
38.9 × 31.6
41.6 × 31.5
40.4 × 31.6
41.6 × 33.6
40.6 × 33.2
43.1 × 33.6
42.8 × 35.2
42.9 × 34.7
39.2 × 32.4
42.2 × 32.5

1.95611.8545
1.86751.65281.718
1.62371.69471.5872
1.7026
2.226
1.967- (large holes, evidently
heavily incubated)

50

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Notes.— The widely-distributed, strongly polytypical species A. novaehollandiae, as
at present understood, is another relict from a period that the species concept was
stretched to, and frequently beyond, its possible limits.The position of sylvestris as asubspecies of A. novaehollandae has been questioned, rightly in my opinion, for the Australian nominate A. novaehollandiae is strikingly different from the many forms occurring on islands to the north. A revision is required to establish more realistic species
limits and therefore I retain for the moment the established trinomen A. n. sylvestris.
A. n. sylvestris is perhaps not primarily a bird-hunter, for the stomach of no. 65188
contained a frog, that of no. 66082 a lizard.
Measurements:

wing

tail

tarsus4

174-188(181.5)197-208(203.4)132-141(137.3)152-157(154.2)49-54(51.6)
53.7-58(55.7)

7

entireculmen20-22(21.0)
22.5-24.6(23.3)

culmenfrom cere13.4-14.1(13.9)
15.5-17(16.3)

middle toe
25-27(26.0)
27-31.7(29.5)

Accipiter fasciatus wallacii (Sharpe)
Astur wallacii Sharpe, 1874, Cat. Birds Brit. Mus. 1: 95, 128, pl. V – Lombock, Bouru; by inference restricted to Lombok by Stresemann (1914c: 381).
Collectors.— Allen, Weber, Everett, v.d. Sande (1), Rensch (2), de Jong (2), Verheijen (2, MCZ), Verheijen/Schmutz.
Material.— , xii.1888, Endeh (Weber, RMNH cat. no. 1); , 13.ii.1969, Wangkung, Rahong (Verheijen,
RMNH no. 65187);  juv., 25.vii.1969, Nisar (Schmutz, RMNH no. 66078); , 19.ix.1969, Sok, Ruteng (Schmutz, RMNH no. 66077);  juv., 19.ix.1969, Sok-Rutung (Schmutz, RMNH no. 66080);  juv., 24.x.1971,
Nunang, 650 m (Schmutz, RMNH no. 84866); , 11.x.1975, Nunang, 650 m (Schmutz, RMNH no. 81022).
Iris  and  golden yellow, juvenile yellowish olive to sulphur-yellow, cere olive-yellow or olive; bill
black, its base slate; legs yellow. Adult birds collected in September, October and December are undergoing their main moult; an adult bird collected in February shows no moult, it is in fresh plumage.
Eggs (fig. 12b): c/2, 4.v.1957, Potjong (Verheijen, RMNH no. 70066); c/2, 24.vi.1969, Nunang, 650 m(RMNH, Schmutz 135). These eggs are a little longer and broader than the eggs of A. n. sylvestris, which
they otherwise resemble; they are also heavier.
Measurements and weights:

RMNH no. 70066
Schmutz 135

43.3 × 37.7
43.4 × 36.5
45.3 × 35.2
47.0 × 33.6

2.53212.44782.25002.1921

Notes.— Of the clutches listed by Verheijen (1964: 198) under this name, only theone mentioned above belongs to this species, all others belong to A. n. sylvestris.
Weber’s specimen (collector’s no. 89), was identified by Büttikofer (1894: 289-290)
as a “wahrscheinlich altes Weibchen” of Astur sylvestris. It was re-identified by Finsch as“wohl ” of Astur torquatus, and on Finsch’s label is in pencil added the name Accipiterfasciatus wallacii  ad. by Stresemann.
Brown & Amadan (1968: 501) describe this subspecies as small (wing  210,  237;
one of each sex?), and A. f. tjendaenae from Sumba as larger (wing  207-219,  247-252).
The specimens of wallacii measured by me are clearly larger; they support Wattel (1973:

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

51137) and White & Bruce (1986: 122-123), who show the two subspecies as practically
identical in size.
The inclusion of Sulawesi, Halmahera and Ceram in the range of A. fasciatus by
Marchant & Higgins (1993: 136, 137) is incomprehensible: even taken in its widest possible sense, the range of the species never extended to these islands.
Measurements:
2
4

wing

tail

tarsus229,[220]250-257(253.5)

161,166
185-192(188.3)60, 62
66.7-68(67.3)

entireculmen19.6, 2122.7-27(24.6)

culmenfrom cere15, 16.2
17-18(17.4)

middle toe
29.5, 3133-35.2(33.8)

Accipiter virgatus quinquefasciatus Mees
Accipiter virgatus quinquefasciatus Mees, 1984, Zool. Meded. 58: 314 – mountain forest above Ruteng,
Flores, ca. 1500 m.
Collectors.— Everett (1), Schmutz.
Material.—  juv., xi.1896, Mt. Repok, Mangarai, above 3500 ft. (A.H. Everett’s native collector, AMNH
no. 533861); , 26.vi.1978, mountain forest above Ruteng, ca. 1500 m (Schmutz, RMNH no. 81024, holotype).
No eggs (see below).

Notes.— This subspecies remains known from the two specimens listed above.
Verheijen (1964: 198) recorded, with a query, breeding (nest or eggs found). Twoclutches of eggs from his collection, identified as A. virgatus, are too large to belong tothis species, and are evidently referable to A. n. sylvestris.
Measurements:

wing

tail

tarsus juv.

153
151

118
113

47
48

entireculmen-culmenfrom cere
10.5
10.5

middle toe
28
27.5

Accipiter gularis (Temminck & Schlegel)
Astur (Nisus) gularis Temminck & Schlegel, 1844, Fauna Japonica, Aves: 5, pl. II – Japon.
Collectors.— Verheijen/Schmutz.
Material.— , 18.xi.1968, Nunang (Schmutz, RMNH no. 65190); , 15.iii.1969, Nunang (Schmutz, RMNH
no. 65189); , 16.xii.1970, Ruteng (Verheijen, RMNH no. 84857); , 16.x.1971, Tjeréng (Schmutz, RMNH
no. 84858); , 31.iii.1976, Ruteng (Verheijen, RMNH no. 84862); , 13.xii.1977, Tjeréng (Schmutz, RMNH
no. 81045).

Notes.— Although it has not been recorded by previous authors (indeed, there is
only one previous record from the whole chain of Lesser Sunda Islands, a subadultmale from East Timor, collected by Wallace in 1861), the material listed above showsthat this is a regular winter visitor to Flores. There are recent records (presumably sightrecords) from Sumbawa (Johnstone et al., 1996: 166). No details are provided, not eventhe name of the observer. The dates of these observations: 27 July (one bird) and 28 July
(two birds), seem unusual (cf. Mees, 1981: 394), but are given without comment.

52

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Hieraaetus fasciatus renschi Stresemann
Hieraëtus fasciatus renschi Stresemann, 1932, Orn. Mber. 40: 78 – Sumbawa: Wawo 500 m.
Collectors.— Verheijen/Schmutz.
Material.— , 21.viii.1971, Ruteng (Verheijen, RMNH no. 66076); , 8.xi.1971, Siru, 150 m (Schmutz,
RMNH no. 81113).

Notes.— The following specimens of this subspecies are now known: Sumbawa (,type), Flores (, ), Timor (), Wetar (), Luang (2 ). Assuming all sexing to be correct, it would seem that the female sex was hitherto unknown. However, the bird from
Wetar, for which Hartert (1904: 189) recorded a wing-length of 495-500 mm, seems verylarge for a male, and I presume that it is the same specimen listed without comment asa female, with a wing-length of 493 mm, by Brown & Amadon (1968: 677). On the other
hand, published measurements of the nominate race show so much overlap between
the sexes, that caution is indicated.
Measurements:



wing

tail

tarsus

450
470

225
264

100
107

exposedculmen45
45

culmenfrom cere
31.3
33

depth
of bill
23
25

middle toe
without nail60
64

Hieraaetus kienerii formosus Stresemann
Hieraaëtus kieneri formosus, Stresemann, 1924, Orn. Mber. 32: 108 - Nord-Celebes.
Collectors.— None.

Notes.— Individuals of this species were reported as having been seen on four occasions, in 1986 and 1989, by Verhoeye & King (1990). As Doherty collected a female on
Satonda, an islet off the north coast of Sumbawa, in May 1896 (cf. Hartert, 1896: 575), theoccurrence on Flores is not unexpected.
Spizaetus (cirrhatus) floris (Hartert)
Limnaëtus limnaëtus floris Hartert, 1898, Novit. Zool. 5: 46 – Flores.
Collectors.— Semmelink, ten Kate, Everett (2), Verheijen (2, MCZ), Verheijen/Schmutz.
Material.— (?), 1862, Larantoeka (Semmelink, RMNH cat. no. 1, syntype); (?), April 1891, Sika = Sikka(ten Kate, RMNH cat. no. 2); (?)pullus, 14.x.1970, Orong (Verheijen, RMNH reg. no. 65184); , 30.i.1976,
Waé-Wako, 180 m (Schmutz, RMNH reg. no. 81112); (?), without data, received in 1974, Flores (Verheijen, RMNH reg. no. 66257).
Eggs (fig. 12c): c/1, 20.vi.1954, Todo (RMNH reg. no. 70074); c/1, 9.vi.1955, Potjong (RMNH no. 70075);
c/1, 15.v.1955, Mano (RMNH reg. no. 70076).
Measurements and weights:

RMNH no. 70074
RMNH no. 70075
RMNH no.70076

69.8 × 56.3
73.0 × 55.1
71.0 × 53.4

9.27
9.51
weight not taken, as a large
piece of the shell is missingMees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

53

Notes.— Ten Kate’s specimen from Sika was recorded by Büttikofer (1892: 193) andby ten Kate (1894: 27) himself, but this has apparently escaped the attention of later
authors (cf. White & Bruce, 1986: 131).
Systematically, the genus Spizaetus is still very insufficiently known. The current
trinomial for the Flores population, S. cirrhatus floris, dates from a period, not so long
ago, that it was custom to stretch the species concept as widely as possible. The validityof this combination may now be questioned at two levels. The first is whether S. limnaeetus from south-eastern Asia, east to Java and Borneo, has properly been included asa subspecies in S. cirrhatus. Doubt about this has been expressed several times (cf. Amadon & Bull, 1988: 324). The second is whether, when limnaeetus is separated specificallyfrom cirrhatus, the form floris would still be regarded as conspecific with limnaeetus, orwhether the logical next step would be to give it also specific status. I incline strongly tothe last-mentioned view.
Brown & Amadon (1968: 694) give for S. c. floris the unexpectedly-large wing-measurements of 485-495 mm. As demonstrated below, the largest bird examined by me hasa wing-length of 470 mm. Hartert (1898a: 46) recorded for two males 437 and 450 mm,
Rensch (1931a: 512) for a male 438 mm (Rensch believed his bird to be a juvenile, but infact it was an adult). Rensch’s second specimen was misidentified and later became thetype of Hieraaetus fasciatus renschi. Hartert (1904: 189) recorded under the name S. c.
floris a bird from Wetar with a wing-length of 495 mm. Later, Mayr (1941a) re-identified
this bird as Hieraaetus fasciatus renschi. It occurred to me that the measurements for S. c.
floris presented by Brown & Amadon could have been taken from such misidentified
specimens, and I wrote to Dr Amadon to ask, whether he could throw light on the matter. From his reply (in litt., 15.iii.1988) I quote: “In any case ignore them! I suspect, as
you say, there was some sort of mix-up with the local race of Hieraaetus fasciatus”.
Measurements:(?) cat. 1(?) cat. 2
 81112(?) 66257

wing
450
470
460
460

tail
260
286
277
290

tarsus
116
121
123
115

entire culmen
54
52
49
48

culmen from cere
36½
35
35
34

Falco moluccensis subsp.
Collectors.— Semmelink, Allen (at least 3, cf. Sharpe, 1874: 430), Martens, Weber (4, of which 2 in ZMA),
Everett, Rensch (5), de Jong (2), Verheijen (2, MCZ), Verheijen/Schmutz.
Material.— [], 1862, Larantoeka (Semmelink, RMNH cat. no. 17); , 19/24.xii.1888, Koting (Weber,
RMNH cat. no. 1); , 1/8.i.1889, Endeh (Weber, RMNH cat. no. 2); , 18.i.1969, Nunang (Schmutz,
RMNH no. 66083); , 16.vii.1969, Sok-Rutung (Schmutz, RMNH no. 66084);  juv., (?)juv., 27.viii.1969,
Rahong (Verheijen, RMNH nos. 65196, 65195); , 27.ix.1969, Ruteng (Verheijen, RMNH no. 65197); ,
12.xii.1975, Nunang 650 m (Schmutz, RMNH no. 81049). The birds collected in December (2) and January (2) are undergoing their main moult.
Eggs (figs 12d, 13a): 23 clutches of 1 (6 ×), 2 (7 ×), 3 (5 ×) and 4 (5 ×) eggs, collected in the months April(10), May (6), June (3), August (3), and September (1). The short-oval eggs are white or pinkish white, so
heavily spotted and dotted with red-brown, that this dominates the ground colour.
Measurements and weights of some clutches:

RMNH no. 70091

35.0 × 30.9 1.4063
36.1 × 31.1 1.4137
37.8 × 31.2 1.2786

54

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)
RMNH no. 70092

RMNH no. 70093

RMNH no. 70094
RMNH no. 70095

RMNH no. 70096

RMNH no. 70098

RMNH no. 70099

37.3 × 31.7
37.9 × 31.6
38.2 × 31.1
39.2 × 31.4
38.2 × 31.3
38.4 × 31.7
39.3 × 31.5
39.8 × 31.9
37.5 × 31.6
37.6 × 31.0
37.1 × 31.8
38.0 × 31.7
38.1 × 31.6
39.3 × 31.6
39.7 × 32.0
39.8 × 32.3
42.6 × 31.4
36.3 × 30.6
36.9 × 30.5
37.3 × 30.3
37.4 × 31.2
38.5 × 31.0
38.5 × 31.1–
1.40171.5284
1.53881.6362
1.73621.656––
1.51701.40541.4844
1.37691.6484
1.47831.7433
1.73521.498
1.50151.47161.54481.65491.6341

Notes.— The four specimens collected in December/January are in the last stage ofprimary moult; two collected in July and September show no moult and are in rather
worn plumage. Compare this with the egg-dates.
There is little doubt that F. moluccensis has been oversplit at the subspecific level, but
the way White & Bruce have disposed of subspecies, was not justified by the meagrematerial they actually examined. A revision on the basis of a large material is required.
Actually, the Leiden collection would form a good basis for it, but it is one of the problems I did not find the time to solve before leaving Leiden. I did note, however, thatbirds from the North Moluccas in the Leiden collection are darker than those from thecentral islands and suggest that F. m. bernsteini is a valid subspecies. In view of the meticulous care Siebers used to take over the description of subspecies, I would also hesitate to reject F. m. renschi from Sumba, without a renewed investigation.
Measurements:
4
3

wing
202+-224(212)
218-238(228)

tail
142-150(146)
146-156(151.3)tarsus33-36(34.8)
35-38(36.3)

culmen from cere
14-16
(14.8)
15-16(15.5)

Falco cenchroides cenchroides Vigors & Horsfield[Falco] Cenchroides Vigors & Horsfield, 1827, Trans. Linn. Soc. 15: 183 – Australia = Parramatta, N.S.W.,
where Caley, the collector of the type material, lived.
Collector.— Endih (1).
Material.— None.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

55

Notes.— The single bird collected at Reo by Endih, in 1911, remains the only record.
On the islands north of Australia and west of New Guinea, the species appears to be no
more than a rare straggler. It did, however, colonize Christmas Island (Indian Ocean)around 1950, and was already common there when I visited that island in 1961.
Falco longipennis hanieli Hellmayr
Falco longipennis hanieli Hellmayr, 1914, in Haniel, Zool. Timor 1: 100 – Bonleo, Westtimor.
Collectors.— Allen (1), Endih (1), Rensch (1), Verheijen/Schmutz.
Material.—  juv., 1.iii.1969, Sésok, 900 m (Schmutz, RMNH no. 65192);  ad., 15.xi.1969, Paku, 400 m(Schmutz, RMNH no. 65195);  im., 22.ii.1970, Ruteng (Karot) (Verheijen, RMNH no. 65191);  subad.,25.x.1971, Nunang, 650 m (Schmutz, RMNH no. 84856);  ad., 12.xi.1971, Nunang (Schmutz, RMNH no.
84853);  ad., 9.xii.1971, Nisar (Schmutz, RMNH no. 84855);  ad., 7.vi.1972, Nunang (Schmutz, RMNH
no. 84854);  ad., 7.vi.1972, Nunang (Schmutz, RMNH no. 85430);  juv., 9.vi.1972, Nunang (Schmutz,
RMNH no. 84852);  ad., 12.xii.1975, Nunang, 650 m (Schmutz, RMNH no. 81040). Adult birds collectedin October, November (2) and December (2) were undergoing their main moult.
Egg (fig. 13b): c/1, 5.ix.1958, Potjong (RMNH no. 70097). The egg was identified as belonging to F. moluccensis, but is too large; I consider it a safe assumption that it belongs to the present species.
Measurements and weight:

RMNH no. 70097

43.3 × 31.9

1.803

Notes.— There is a conspicuous difference in size between the sexes, the female being much larger than the male. There is little or no difference in plumage: possibly the has slightly more extensive and darker brown on the under surface than the , but
the difference is dubious. Juvenile birds of both sexes differ from the adults by having
light brown edges to the feathers of the mantle; the central pair of rectrices is black withlight brown bars (in adults, black with grey bars), and the black feathers of the forehead(behind the white frontal band) have tiny brown tips.
Rensch’s (1931a: 517)  juv. from Dompoe, Sumbawa, with wing 217, tail 117, culmen 13 mm, has obviously been mis-sexed and must be a .
All specimens examined belong to the subspecies hanieli. The nominate subspecies
has been recorded from Flores (Stresemann & Amadon, 1979: 417), but in error, as waspointed out by Bruce (in White & Bruce, 1986: 135). I know of no evidence to support
the claim by Marchant & Higgins (1993: 278) that some Australian birds winter in theLesser Sunda Islands and suppose its basis to be this same erroneous record from
Flores. Small-scale distribution maps in handbooks are rarely accurate; therefore it is
not really surprising to see in Cade (1982: 187 fig. 21) the range of F. longipennis extended westward to cover the whole of Java. In contrast, the map in Marchant & Higgins (1993: 270) seems to suggest that the species is no more than a straggler to theLesser Sunda Islands.
Measurements:

wing

tail

tarsus6

218-227(222.2)242-255(250)

110-115½(113,3)123-126(124.5)31.3-35(33.3)
34.5-38(36.7)

4

culmen fromforehead feathers
16.2-18(17.1)
18.5-19.6(19.0)

culmen fromcere
12.2-13.5(13.0)
15-15.3(15.1)

56

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Dendrocygna arcuata arcuata (Horsfield)
Anas arcuata Horsfield, 1824, Zool. Res. Java: pl. (65) and text – Java.
Collector.— Schmutz.
Material.— , 25.vi.1974, Pota (Schmutz, RMNH no. 85139).

Notes.— D. arcuata certainly breeds on Flores: a  shot by Schmutz near Djoneng,
on 1.ii.1977, contained a near-developed egg of 3 cm across.
In view of the fact that Watling (1982: 69) knew from Fiji only: “Two specimens, oneof which was a juvenile, collected before 1870 from Nadi Bay (a location which could beeither west Viti Levu or south-west Vanua Levu)”, it is worth recording the following
specimen:  ad., end of July 1877, Nandi Bay, Viti Levu, leg. T. Kleinschmidt (RMNH
cat. no. 31, ex Mus. Godeffroy no. 5716, original label), wing 194, culmen 42.3 mm. I
further note, that Salvadori (1895: 155) reports the presence of three specimens in the
British Museum: two from Viti Levu, leg. J.D. Macdonald, voyage of H.M.S. ‘Alert’,
September (the voyage of the ‘Alert’ took place in 1878/1882, and Fiji was probably
visited in 1879 or 1880) and one from Kandi, leg. E.L. Layard. These additional records
suggest that in the seventies of the 19th century, D. arcuata was locally not rare on Viti
Levu. Watling’s Nadi Bay is the same as Nandi Bay, Viti Levu: the letter d in Fijian is
pronounced as nd. Delacour (1954: 41) seems to be certain that “it has been exterminated by the introduced mongoose”. The wing-length of the RMNH specimen clearlyexceeds the measurements given for the subspecies D. a. pygmaeus, to which the presumably extinct Fiji population has been tentatively attached (cf. Mayr, 1945: 3).
There is also a specimen from New Caledonia in our collection (, no further particulars, from Verreaux in 1866, RMNH cat. no. 30). The specimen was published by
Schlegel (1866a: 89) under the name of Dendrocygna vagans, with the cat. no. 18. This is
the earliest published record of the occurrence in New Caledonia that I know of. Measurements: wing 199, culmen 46 mm. This does not help much in its subspecific identification, or whether there was a resident population in New Caledonia or birds foundthere were stragglers from Australia. Delacour (1966: 40) refers birds from New Caledonia to the nominate race. It becomes increasingly doubtful that a division into subspecies is meaningful.
Dendrocygna javanica (Horsfield)
Anas Javanica Horsfield, 1821, Trans. Linn. Soc. Lond. 13: 199 – Java.
Collectors.— Endih (2, cf. Somadikarta & Noerdjito, 1982), Verheijen (a wing, MCZ, cf. Paynter, 1963),
Schmutz.
Material.— , 20.vi.1974, Kulang Lake, East Flores (Schmutz, RMNH no. 85138).

Notes.— Like the preceding species, Schmutz found individuals in breeding condition. Rensch (1931a: 504) published the first record of this species from the Lesser
Sunda Islands on the basis of two females from Dompoe, Sumbawa, collected by Elbert on 21.xii.1909, preserved in the Senckenberg Museum. I should like to add that
there are in Leiden also two females, from the same date, locality and collector (RMNH
cat. nos. 30, 31).

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

57

Anas superciliosa rogersi Mathews
Anas superciliosa rogersi Mathews, 1912, Austral Av. Rec. 1: 33 – Augusta, West Australia.
Collectors.— Weber (1), Rensch (3), de Jong (2), Verheijen (1, MCZ), Verheijen/Schmutz (3, and eggs).
Material.— , xi.1888, Reo (Weber, RMNH cat. no. 39); , 27.x.1968, Ruteng (Verheijen, RMNH no.
65199); , 23.xi.1968, Nunang (Schmutz, RMNH no. 66075); , 2.vii.1969, Ruteng (Verheijen, RMNH no.
65200).
Eggs: c/10, 26.iii.1957, Mano (RMNH no. 70058); c/2, 28.ii.1961, Soa (RMNH no. 70059); c/1, 3.vi.1961,
Soa (RMNH no. 70060); c/6, 9.vi.1961, Soa (RMNH no. 70061).

Notes.— This is obviously the commonest duck of the Lesser Sunda Islands. In thisconnection it is perhaps worth recording that in our collection there are no less than six
specimens from Lombok, obtained by the “Sunda-Expedition” in May 1909 (RMNH
cat. nos. 44-49).
In assigning these birds to rogersi, I am following contemporary usage, as no adequate topotypical material of the nominate race (from New Zealand) is available forcomparison. Several authors have expressed doubt about the validity of rogersi (cf.
Amadon, 1943: 2; Frith, 1967: 166), although others seem to have accepted it (cf. Delacour, 1956: 63; Johnsgard, 1979: 471). On the other hand, Condon (1975: 70) definitely
rejected rogersi and included all Australian birds (and by inference also the populationsof the Lesser Sunda Islands) in the nominate race, and this has been followed, without
discussion, by Blakers et al. (1984: 76). A further study of the geographical variation ofthis duck is desirable.
In this connection it is worth drawing attention to the fact that in Australia this species is known to be a great wanderer, dependent as it is on the very unstable habitat
provided by temporary floodings, etc. This would work against geographical variation.
Mobile as they are, I cannot follow Van Marle & Voous (1988: 37, 68) in their suggestion,that birds from as far west as Sumatra (where they are supposed to be, or to have been,
periodically numerous), would be or even could be “irruptive” migrant visitors from
Australia.
Marchant & Higgins (1990: 1320, 1331) treat the species as monotypic, rejecting not
only rogersi, but also pelewensis, claiming that such geographical variation as may exist
is too minor for expression in nomenclature.
Measurements:




wing
260
[215]
245

tail
84
78
84

tarsus4142½
39

exposed culmen
53½
46½
48

Anas gibberifrons gibberifrons S. Müller
Anas (Mareca) gibberifrons S. Müller, 1842, Verh. Nat. Gesch. Ned. Overz. Bez., Land- en Volkenk.: 159 –Timor.
Collectors.— Allen (1), Weber (1, ZMA), Schmutz.
Material.— , 23/25.xi.1888 (Weber, ZMA no. 13253); , end vii.1969, Nanga-Lili (Schmutz, RMNH no.
81104).

58

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Eggs; c/1, 27.vi.1956, Réo (RMNH no. 70062); c/8, 22.vi.1959, Dampék (RMNH no. 70063). I have noidea on what the record by White & Bruce (1986: 139) of breeding on Flores in August is based, andpresume it to be an error.

Notes.— Previously, I have drawn renewed attention to the considerable differences
between A. g. gibberifrons and the Australian A. gibberifrons gracilis (cf. Mees, 1982a: 22),
as Ripley (1942a) did forty years earlier, but I still hesitate to follow Parker et al. (1985:10), who treat them as different species. Apart from the conspicuous skull-character, thetwo forms show a close resemblance.
It is unclear why Van Marle & Voous (1988: 37, 68) should consider Anas gibberifrons
in Sumatra as a “migrant from Australia”, referring it to “gibberifrons or gracilis”. Recordsof this species from Sumatra are unsatisfactory, and in my opinion do not qualify it forinclusion in the Sumatran list. If it does visit or colonize Sumatra, however, this would
certainly be by birds from Java (gibberifrons) and not from Australia (gracilis)!
There is some inconsistency in the citation of the type-locality of A. gibberifrons: inthe older literature and in a few more recent publications (Frith & Hitchcock, 1974: 123;
Mees, 1982a: 22) it is given as Timor, but in authoritative recent works like those of
Condon (1975: 70), Johnsgard (1979: 466) and White & Bruce (1986: 139), to mention justa few, this has been changed to Sulawesi. The facts are as follows: Müller (1842) described the species in a footnote, in an article on Timor. In the main text he records his
observations made on Timor, and the description in the footnote is taken from specimens of both sexes collected on Timor. Following the description, however, the next
paragraph of the footnote begins with: “Van deze eend heb ik ook een individu op Makassar verkregen, terwijl onlangs verscheidene exemplaren van dezelve door onzen
reiziger Dr Forsten, van Menado, aan het Rijks-Museum zijn toegezonden geworden” 4).
Therefore, material from Makassar and Menado, although not described, was definitely
included in the original publication, and has to be considered syntypical. If a restrictionof the type locality is required, however, this should obviously be to Timor.
All this material is still present in Leiden: , 1828/1829, Timor (RMNH cat. no. 1);
, iii.1829, Atapoepoe, Timor (RMNH cat. no. 2); , iii.1828, Makassar (RMNH cat. no.
3), and five Forsten specimens from various localities in northern Celebes (RMNH cat.
nos. 4-8). Perhaps not all of Forsten’s specimens are syntypes, some may have arrived
in Leiden after Müller’s description was written.
Megapodius reinwardt reinwardt Dumont
Megapodius Reinwardt Dumont, 1823, Dict. Sci. Nat. (éd. Levrault) 29: 416 – d’Amboine, dans les ÎlesMoluques (errore!); corrected to Lombok by Schlegel (1880: 57), and here further corrected to the
Banda Islands (see below).
Collectors.— Allen, Colfs, Everett, van der Sande (2), Rensch (1), Verheijen/Schmutz.
Material.— (?), 1880, Flores (Colfs, RMNH cat. no. 9); , 20.xii.1907, Wai Moké, south coast of middle
Flores, weight 0.55 kg (v.d. Sande, ZMA no. 25.831); , 1.ix.1969, Nanga-Lili (Schmutz, RMNH no. 66069);
, 30.iii.1976, Ceréng, 300 m (Schmutz, RMNH no. 81108). Iris brown, bill and legs orange-yellow.

4)

Of this duck I have also obtained an individual at Makassar, whilst recently our traveller Dr Forsten
forwarded several specimens from Menado to the Rijks-Museum.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

59

Eggs: 54 eggs (in 35 lots), taken in the months January (4), February (22), March (13), June (1), July (3),August (1), October (3), November (1), December (3), and three not dated (RMNH nos. 70112-70142, 73656,
73657, 75498, 78404). The very large eggs have a soft, chalky shell, varying from yellowish chamois to lightbrown, which comes off easily when the surface is scratched, to show the white chalk underneath.
Some measurements and weights:

RMNH no. 70122
RMNH no. 70126
RMNH no. 70131

80.5 × 49.2
82.0 × 53.1
83.4 × 51.0
87.0 × 54.0
87.9 × 54.0

7.28568.65658.23519.3206
9.311

Notes.— Megapodius reinwardt was originally believed to be from Ambon, but that
could not be correct, as Ambon is inhabited by a different, closely related species, M.
freycinet. Like Reinwardt’s other material, the specimen was insufficiently labelled(compare Nectarinia solaris). Discussing the holotype, Schlegel (1880: 57) noted: “individu rapporté par feu le Professeur Reinwardt. Ce savant voyageur ne nous ayant fait
parvenir qu’un seul individu du Mégapode aux pieds rouges et cet individu originairede l’Île de Lombock, comme je viens de le constater par l’etiquette originale...”. Since
then, Lombok has been generally accepted as the type locality.
Whereas from the systematist’s point of view there can be no objection to the type
locality Lombok, as that island is inhabited by the correct species and subspecies, historical evidence is against it. First, Schlegel’s claim that the bird had an original label,giving its provenance as Lombok, is amazing. I have examined the type specimen, andit does not have an original label. One might think of a label that has become lost since
Schlegel wrote, but that is most unlikely. The specimen is, and has been, mounted, for atleast a century and a half. Former practice in Leiden was, when a specimen was mounted, to throw away the label (assuming there was one), and to write any particulars thatwere thought worth keeping, underneath the socle. A mounted specimen in the old collection with an original label, would be exceptional in the Leiden collection, and if it ever
had one, it becomes completely incomprehensible why its provenance had earlier beenthought to be Ambon, and was only corrected almost sixty years after its description. It
may safely be assumed that Schlegel’s story was based on faulty memory: it is known
that Schlegel used to write from memory, and did not much checking up on facts, once
his manuscript was written (Brouwer, 1954: 57). Perhaps Schlegel had seen a Wallace
specimen from Lombok. This, however, is not the whole argument: Lombok might still
be considered a satisfactory designated type locality. But Reinwardt never visited Lombok, and could not have collected a bird there. Reinwardt did call at Bima, Sumbawa,where he spent a few days (20-23.iii.1821), probably too short to do any bird collecting.
However, Reinwardt spent five weeks (18.v-23.vi.1821) on the Banda Islands, within therange of M. reinwardt. Moreover, from Banda he sailed on to Ambon. The Banda Islands
are, therefore, the obvious, and almost the only possible place for him to have obtained
the type. Above I have corrected the type locality accordingly.
Measurements:



(?)

wing
231
240
235
236

tail
107
85
87
82

tarsus
63.5
66.5
66.2
67

entire culmen
32
30
29
32

60

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Coturnix ypsilophora raaltenii (S. Müller)
Perdix Raaltenii S. Müller, 1842, Verh. Nat. Gesch. Ned. Overz. Bez., Land- en Volkenk.: 158 – omstreken
van Babauw, Timor.
Synoicus ypsilophorus castaneus Mayr, 1944, Bull. Am. Mus. Nat. Hist. 83: 144 – Alor Island.
Collectors: Allen (few? – not in BM), Rensch (2), Verheijen (8, MCZ), Verheijen/Schmutz (many).
Material.— 26 specimens from Langkas in Rahong, 900 m, and Lumu (Verheijen, RMNH nos. 5978359797, 65212, 85285-85294).
Eggs: 65 clutches of c/1 (7 ×), c/2 (14 ×), c/3 (12 ×), c/4 (8 ×), c/5 (9 ×), c/6 (3 ×), c/7 (4 ×), c/8 (6 ×) andc/9 (2 ×), collected in the months February (1), March (9), April (12), May (21), June (5), July (1), August(4), September (2), October (1), and nine insufficiently dated (RMNH nos. 70151-70214, 70172a). The
eggs are white, almost without gloss, usually somewhat dirty.
Some measurements and weights:

RMNH no. 70175

RMNH no 70199

RMNH no. 70200

28.8 × 22.6
29.4 × 23.3
30.2 × 23.3
30.2 × 24.3
28.0 × 23.8
29.6 × 23.9
29.7 × 23.7
27.6 × 23.6
28.3 × 24.0

0.8122
1.0636
1.05470.92871.236
1.2715
1.15771.0824
1.0751

Notes.— In a previous paper I mentioned my inability to distinguish between these
specimens, and specimens from Timor and Roti (cf. Mees, 1975b: 123). In this connection: Mayr (1944a: 144) had seen no specimens from Flores when he described castaneus
from Alor, so that the inclusion of Flores into its range was only a guess. When Flores,
Timor and Roti are inhabited by one subspecies C. y. raaltenii, it is no longer likely that
Alor has a different one. White & Bruce (1986: 145) have given additional arguments forthe non-recognition of castaneus, and I follow them.
Hitherto, the range of this species along the chain of Lesser Sunda Islands has been
known to extend westwards only to Flores. In September 1984 I found it on the Tg.
Pioen peninsula, northern Sumbawa, where it appeared to be quite common.
Coturnix chinensis chinensis (Linnaeus)[Tetrao] chinensis Linnaeus, 1766, Syst. Nat. (ed. 12) 1: 277 – in China, Philippinis = China.
Excalfactoria chinensis palmeri Riley, 1919, Proc. Biol. Soc. Washington 32: 93 – Daroe, Java.
Excalfactoria chinensis lineatula Rensch, 1931, Mitt. Zool. Mus. Berlin 17: 473 – Badjawa, Flores.
Collectors.— Everett (no number given), Rensch (2), Verheijen (24, MCZ), Verheijen/Schmutz.
Material.— , 1957, Rahong (Verheijen, RMNH no. 66261); 2 , ix.1958, Rahong (Verheijen, RMNH nos.
66262, 66264); 2 , ix. 1958, Ruteng (Verheijen, RMNH nos. 66263, 66265); , ix.1959, Ruteng (Verheijen,
RMNH no. 66266); , 1960, Potjong (Verheijen, RMNH no. 66267); , 19.x.1968, Rahong (Verheijen,
RMNH no. 59800); , 5.xi.1968, Wangkung, Rahong (Verheijen, RMNH no. 59798); , 14.xi.1968, Rahong (Verheijen, RMNH no. 59799); , xi.1968, Rahong, 900 m (Verheijen, RMNH no. 59801); , 21.
ix.1971, Gurung Wangkung, 900 m (Verheijen, RMNH no. 85165); , 3.x.1971, Gurung, Langkas, 900 m(Verheijen, RMNH no. 85188); , 4.x.1971, Gurung, Langkas, 900 m (Verheijen, RMNH no. 85166); ,
8.x.1971, Gurung, Langkas, Rahong, 900 m. (Verheijen, RMNH no. 85167); , , 9.x.1971, Gurung, Lang-Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

61

kas (Verheijen, RMNH nos. 85168, 85189); , 11.x.1971, Suma, Langkas (Verheijen, RMNH no. 85169); ,
3.viii.1976, Lingko Cehet (Verheijen, RMNH no. 81301).
Eggs: 18 clutches of c/1 (1 ×), c/2 (5 ×), c/3 (2 ×), c/4 (7 ×), c/5 (1 ×) and c/6 (2 ×), collected in themonths March (1), April (1), May (4), June (8), July (2), and two not dated; from Tado, Potjong, Méngé,Rentung Lelak, Bénténg Djawa, Mano, Mataloko, Tjara, Léwé (RMNH nos. 70215-70232). The eggs varyfrom olive brown to coffee brown, and are either plain, or have a variable number of very small darkbrown spots.
Some measurements and weights:

RMNH no. 70229

RMNH no. 70230

RMNH no. 70231

23.0 × 18.4
23.4 × 17.7
23.8 × 19.0
24.2 × 18.8
25.0 × 18.6
25.0 × 18.8
25.6 × 19.4
25.7 × 18.9
24.2 × 19.5
24.2 × 20.00.24080.26900.28140.29890.41340.36720.43080.40090.36950.4114

Notes.— The geographical variation of C. chinensis requires further study. Of thetwo subspecies here synonymized, E. c. palmeri was diagnosed as being: “Similar to
Excalfactoria chinensis lineata from the Philippines, but the back and scapulars much
mixed with slate color, the wing with much rufous, and the rufous of the breast more
extensive”. Riley had for comparison 12 specimens from the Philippines, two from
Sulawesi (Celebes) and one from the Malay Peninsula (all ), but avoids mentioning
the size of his sample from Java. Significantly, no mention is made of nominate chinensis to which at least Malayan birds had previously been ascribed. Note that Robinson& Kloss (1927) included Sumatra and Borneo into the range of palmeri, listing E. c.
caerulescens Hachisuka, type locality Sarawak, as a synonym, but that Rensch (quoted
below) referred Bornean birds to lineata. The type-locality of palmeri, Daroe, is in thesouth-western part of the Residentie Batavia (as it was then), not in Bantam as claimed
by Deignan (1961: 71).
The diagnosis of E. c. lineatula reads: “Die 3 vorliegenden , wie auch ein  vonFlores aus dem Tring-Museum, unterschieden sich von der Java-Rasse palmeri Riley
durch stärkere braunschwarze Fleckung der Oberseite, wodurch sie weniger schiefergrau wirken. Die hellen Schaftstreifen der Oberseite sind viel feiner als bei Stücken
von Australien (australis Gd.), und von den Philippinen und Borneo (lineata Scop.).
Von der Rasse minima (Gd.) unterscheidet sich die neue Rasse durch bedeutendere
Grösse”.
Van Bemmel (in Van Bemmel & Voous, 1951: 104) called lineatula “a very weak race”,
but recognized palmeri. This was based on a comparison of specimens from Sulawesi(Celebes) (lineata) with specimens from Java (“palmeri”), which he considered different.
He had no material from the range ascribed to nominate chinensis, so that all he did, wasestablish that chinensis (including “palmeri”), differs from lineata.
The adequate series from Java available to me, made it clear that there is no difference between specimens from Java and Flores, the differences described by Rensch on
the basis of a very limited material being all within the range of individual variation.
The question remains whether, and how, the subspecies chinensis and lineata differ

62

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

from each other. Lack of adequate material prevents me from giving an opinion. A reviser will also have to explain the generally-accepted type-locality Nanking of nominate chinensis, for at present the species does not occur as a wild bird within 500 km of
Nanking (for a map of the distribution in China, cf. Cheng et al., 1978: 82).
The occurrence of this little quail on Sumbawa was predictable, but I know of noprevious records. On 22.ix.1984, I flushed a pair on the Tg. Pioen peninsula, northern
Sumbawa, hardly above sea level. White & Bruce (1986: 145) give it a vertical range of
400-1200 m in the Lesser Sunda Islands.
Measurements:
9
10 

wing
68-75
(71.8)
70-76(73.1)

tail28-34(29.8)
27-31(29.3)

tarsus19.3-22.5(20.5)
19-21(19.9)

entire culmen
11.2-14(12.9)
12-13.8(13.2)

exposed culmen
9-10.6(10.1)
9.4-11(10.2)

Gallus varius (Shaw)
Phasianus varius Shaw, 1798, Nat. Misc. 10: pl. 353 and text – “Probably an Indian bird” = Java.
Collectors.— Semmelink, Allen, Everett, Verheijen (2, MCZ), Verheijen (eggs), Schmutz.
Material.— ,  juv., 1862, Larantoeka (Semmelink, RMNH cat. nos. 5, 7); , not dated, Flores (Schmutz,
RMNH no. 81107).
Eggs: c/4, 9.vi.1955, Potjong (RMNH no. 70143); c/2, 7.vii.1955, Ruteng (RMNH no. 70144); c/2,
22.xi.1955, W. Rembong (RMNH no. 70145); c/4, 9.vi.1957, Tado (RMNH no. 70146); c/5, 17.vi.1959,
Potjong (RMNH no. 70147); c/4, 28.vi.1959, Montjok (RMNH no. 70148); c/1, 27.ii.1961, Soa (RMNH no.
70149); c/5, 28.v.1961, Mataloko (RMNH no. 70150). The eggs are creamish white, with pores madedistinct by dirt.
Some measurements and weights:

RMNH no. 70146

RMNH no. 70148

49.7 × 38.4
50.1 × 37.6
50.1 × 38.3
51.0 × 37.5
47.3 × 35.5
49.2 × 36.4
49.5 × 37.2
51.1 × 37.9

2.81232.51472.54022.673
2.69072.96653.1442
3.2480Turnix suscitator powelli Guillemard
Turnix powelli Guillemard, 1885, Proc. Zool. Soc. Lond.: 511 – Gunong Api Island, Sumbawa.
Collectors.— Everett (several), Verheijen (5, MCZ), Verheijen/Schmutz.
Material.— , 15.x.1968, Nunang (Schmutz, RMNH no. 66090). Iris white.
Eggs: 83 clutches of 1-4 eggs, of this species and T. maculosa combined (RMNH nos. 70233-70314,
73508).
Some measurements and weights:

RMNH no. 70272 (Montjok)
RMNH no. 70278 (Palué)

25.0 × 18.8
26.3 × 19.4
24.4 × 20.4
25.1 × 20.2
26.0 × 20.8

0.36560.38470.38420.36110.4188

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

63

RMNH no. 70279 (Palué)

RMNH no. 70280 (Palué)
RMNH no. 70281 (Palué)
RMNH no. 70287 (Palué)
RMNH no. 70290 (Palué)

RMNH no. 70291 (Palué)

23.6 × 19.6
24.7 × 20.0
26.1 × 19.5
24.2 × 19.5
25.6 × 19.9
24.7 × 20.1
25.4 × 20.1
25.0 × 21.0
23.8 × 19.9
24.4 × 20.6
24.6 × 20.2
25.0 × 20.7
24.9 × 19.9
24.9 × 20.0
25.0 × 20.5
25.0 × 20.5

0.37030.39570.37920.38980.41340.438
0.43150.44530.404
0.40620.43490.4229Verheijen collected a number of clutches of Turnix-eggs on Palué, all of which heidentified as belonging to T. powelli, the only Turnix he recorded from Palué. He further
wrote: “Boys told me about two kinds of mbu. Probably this difference is only a question of sexual dimorphism” (Verheijen, 1961: 184). In accordance with the above identification, Verheijen’s eggs were originally registered in Leiden as T. powelli. An examination of the Palué eggs revealed, however, that they can be divided into two size- andweight-classes: larger and heavier eggs which have been correctly identified as belonging to T. powelli, and smaller and lighter eggs which are obviously referable to T. maculosa, thus confirming the local boys’ claim that two species of Turnix occur on Palué.
Notes.— This form has sometimes been regarded as a separate species, but in myview it is well placed as a subspecies of T. suscitator. In this, I agree with White & Bruce(1986: 148), who support this by saying that T. s. baweanus bridges the difference between nominate suscitator and powelli. It is likely that this was not based on personalobservation, but uncritically copied from Hoogerwerf (1962: 197-200). However, theindividual variation in plumage of nominate suscitator has been underestimated, andbaweanus is a synonym (cf. Mees, 1986: 27). In their somewhat confused discussion of
the type locality of powelli, White & Bruce do not make very clear that Gunung Api andSangeang are two names for the same island.
Measurements:


wing
85

tail
44

tarsus23

entire culmen
16.2

exposed culmen
12Turnix maculosa floresiana Sutter
Turnix maculosa floresiana Sutter, 1955, Verh. Naturf. Ges. Basel 66: 121 – Süd-Flores.
Collectors.— Everett (number not recorded), Rensch (1), Sutter (1), Verheijen (12, MCZ), Verheijen/
Schmutz.
Material.— ?, IX-1968, Rahong (Verheijen, RMNH 59802); , 19-X-1968, Rahong (Verheijen, RMNH
59803); , 2-xii-1968, Rahong (Verheijen, RMNH 59804); , 2.xi.1970, Tjumbi, Rahong, (Verheijen,
RMNH no. 85170); , 15.ix.1971, Roka, Wangkung (Verheijen, RMNH no. 85171); , 20.ix.1971, Gurung,
Langkas, Rahong, 900 m (Verheijen, RMNH no. 85172); , 24.ix.1971, Gurung (Verheijen, RMNH no.

64

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

85173); , 25.ix.1971, Gurung (Verheijen, RMNH no. 85174); 3 , 26.ix.1971, Gurung (Verheijen, RMNH
nos. 85175-85177); , 2.x.1971, Gurung (Verheijen, RMNH no. 85178); , 3.x.1971, Gurung (Verheijen,
RMNH no. 85179); , 4.x.1971, Gurung (Verheijen, RMNH no. 85180); 2 , 8-X-1971, Gurung (Verheijen,
RMNH 85181, 85183); , 8-X-1971 Gurung (Verheijen, RMNH 85184). , 9-X-1971, Suma, Langkas (Verheijen, RMNH 85182); , 11-X-1973, Suma Langkas, (Verheijen, RMNH 85185); , 10-viii-1976, Langkas(Verheijen, RMNH 81427). , no date, Gn. Wangkung (Verheijen, RMNH 85186). Iris grey, bill black andlight brown, legs light brown. Several specimens collected in September and October show primary
moult.
Eggs: See under the preceding species. Verheijen (1964: 198) did not distinguish between the eggs of the
two Turnix-species, with the result that they have all been registered as T. s. powelli in the RMNH collection. I have sorted out some of these, of which measurements and weights are given below, but not all,as this would require measuring and weighing all eggs, several hundreds, and even then it is likely thatsome would be difficult to place. It did not seem worth the extra effort in the framework of this article.
The eggs are similar to those of T. suscitator powelli, but on average smaller and lighter.
Some measurements and weights:

RMNH no. 70249 (Potjong)

RMNH no. 70251 (Tjarang)
RMNH no. 70285 (Palué)

RMNH no. 70286 (Palué)

RMNH no. 70288 (Palué)

RMNH no. 70289 (Palué)

21.4 × 18.8
22.4 × 18.7
23.9 × 19.2
23.3 × 18.6
24.6 × 19.1
21.1 × 20.1
22.6 × 20.5
23.1 × 20.6
22.7 × 18.9
22.8 × 19.4
23.7 × 19.0
22.0 × 18.0
22.7 × 18.0
23.6 × 17.5
22.7 × 18.7
23.0 × 18.9
23.7 × 18.8

0.309
0.294
0.34130.2891
0.314
0.29610.32570.32090.33270.35300.32910.2636
0.27110.24880.2926
0.31370.3181

Notes.— Lack of adequate material from the surrounding islands prevented mefrom discussing the interesting geographical variation in the species in this region, but
a single specimen from Sumba (leg. Verheijen) differs from the series of floresiana in thecharacters described by Sutter (1955) and supports the validity of sumbana.
Measurements:
10
7

wing
68-75
(70.9)
76-81(78.4)

tail25-30½(27.2)
27-33(30.4)

tarsus18-20(19.1)
19.8-21.6(20.4)

entire culmen
13-14.2(13.7)
14.6-15.3(14.9)

exposed culmen
9.4-11.0(10.4)
11-12(11.5)

Rallus pectoralis exsul (Hartert)
Hypotaenidia brachypus exsul Hartert, 1898, Novit. Zool. 5: 50 – Mangarai district, South Flores.
Collectors.— Everett (1), Verheijen (3, MCZ), Verheijen/Schmutz.
Material.— (?), x.1958, Tjumbi, Rahong (Verheijen, RMNH no. 85156); (?) juv., 3.v.1959, Mano (Verheijen,
RMNH no. 85157);  juv., 1.ii.1974, Nunang, 650 m (Schmutz, RMNH no. 85154), all remiges growing,
noted as “a prey of Accipiter sp.”. Iris brown, bill and legs grey.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

65

Eggs (figs 13c, 13cc): c/3, 12.iii.1955, Potjong (RMNH no. 70315); c/3, 4.vi.1955, Potjong (RMNH no.
70316); c/2, xii.1955, Bénténg Djawa (RMNH no. 70317); c/1, 16.vi.1957, Mano (RMNH no. 70318); c/1,
23.xi.1957, Mano (RMNH no. 70319); c/4, 19.ii.1958, Mano (RMNH no. 70320); c/1, iv.1958, Potjong(RMNH no. 70321); c/5, 12.ix.1959, Montjok (RMNH no. 70322); c/4, 15.iii.1959, Potjong (RMNH no.
70323); c/1, iv.1959, Potjong (RMNH no. 70324).
Measurements and weights:

RMNH no. 70315

RMNH no. 70316

RMNH no. 70317
RMNH no. 70318
RMNH no. 70319
RMNH no. 70320

RMNH no. 70321
RMNH no. 70322

RMNH no. 70323

RMNH no. 70324

32.5 × 25.1
32.6 × 25.1
33.2 × 24.5
31.0 × 24.2
32.2 × 25.0
33.0 × 24.5
31.4 × 24.0
31.5 × 24.6
30.8 × 23.0
32.9 × 24.7
31.3 × 24.0
31.8 × 23.3
32.1 × 24.0
33.5 × 24.2
31.6 × 23.7
30.8 × 24.9
32.0 × 24.5
32.4 × 25.3
33.4 × 25.2
35.7 × 25.4
31.6 × 24.2
31.6 × 24.4
31.7 × 24.5
33.7 × 24.7
31.9 × 24.1

0.68290.69300.6677large holes
0.5596
0.57530.4564–0.61720.5986
0.6311–0.57310.56170.57530.55100.54470.6043

Notes.— The MCZ-specimens, one unsexed adult and two juveniles, were discussed
by Paynter (1963). This subspecies is known from very few specimens.
Measurements:(?) ad.

wing
106

tail
41

tarsus28

entire culmen
32

exposed culmen
29

Rallus philippensis wilkinsoni (Mathews)
Eulabeornis philippensis wilkinsoni Mathews, 1911, Birds Austr. I: 198 – South Flores.
Collectors.— Everett, Rensch (1), Verheijen (13, MCZ), Verheijen/Schmutz.
Material.— , 8.vii.1969, Rahong (Verheijen, RMNH no. 65222); , 19.i.1970, Montjok (Verheijen, RMNH
no. 65224); , 15.xi.1970, Ruteng (Verheijen, RMNH no. 66091); , 15.ii.1974, Nunang, 700 m (Schmutz,
RMNH no. 85144); , 20.x.1977, Wewak, 400 m (Schmutz, RMNH no. 81076). Iris coral red.
Eggs (fig. 13d): 68 clutches of from 1 to 7 eggs, not all of them with complete data. Months of collecting:January (3), February (1), March (9), April (17), May (15), June (10), July (1), August (1), September (1),October (1), November (2), December (1). (RMNH nos. 70330-70395, 70338a, 73507). The eggs are cream
colour to pale pinkish chamois, with moderate numbers of well-defined chocolate brown primary dots,
and light grey secondary dots.

66
Some measurements and weights:

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)
RMNH no. 70349

RMNH no. 70350

RMNH no. 70361

38.3 × 29.5
38.8 × 29.7
39.0 × 29.7
39.3 × 29.6
39.4 × 29.4
39.4 × 30.0
38.5 × 29.6
39.0 × 29.8
39.1 × 30.3
39.2 × 29.7
39.3 × 30.1
39.4 × 30.0
37.2 × 29.0
37.4 × 28.8
37.4 × 28.8
38.3 × 29.0
38.8 × 28.3

1.3504
1.34681.35891.39871.32261.4163
1.41981.45291.47601.41281.50571.46621.48001.35541.42461.42561.4349Notes.— Several authors have studied the geographical variation of Rallus philippensis, and the material at that time present in Leiden was thoroughly discussed byJunge (1953: 11-16). Mistrusting, from experience, any subspecies described by Mathews, I have compared these additional specimens from Flores with specimens fromadjacent islands. Birds from Ambon, Buru, New Guinea and Australia have a broadorange-brown pectoral band, which is absent in birds from Sulawesi (Celebes) and
Flores (immature birds from these two islands have a weakly-developed band). Birds
from Flores are large and have darker olive-brown upper-parts than specimens from
Celebes. On this basis, it seems possible that wilkinsoni is a valid subspecies. Mayr
(1944a: 145-146) regarded large size (, wing 158 mm) as the main subspecific characterof wilkinsoni, and an average larger size was the only difference between a bird from
Timor and birds from Celebes that Junge could find. But Mayr considered the larger
birds (wilkinsoni) to be confined to Flores, and included specimens from Timor, on thebasis of smaller size, in R. p. chandleri from Celebes, whereas Junge, without specimensfrom Flores and with only one, a rather large one, from Timor, concluded that birds
from Flores and Timor are similar, and differ by larger average size from Celebes birds.
Junge wisely refrained from applying trinomials. Stresemann (1941: 28) compared two
specimens from Alor (, wing 137 mm) and Flores (wing 146 mm) with a series from
Celebes (7 , wing 135-146 mm, 5 , wing 134-142 mm), and as he found no differences in plumage either, he placed wilkinsoni in the synonymy of chandleri. I agree with
Ripley (1977: 83), that: “The racial affinities of birds from the Moluccas and Lesser Sunda Islands are imperfectly understood”.
Subsequently, White & Bruce (1986: 150-151) also failed to arrive at definite conclusions. White discusses R. p. chandleri, described from Sulawesi (Celebes), and points
out characters by which it differs from nominate philippensis. Nevertheless, White &Bruce followed Mayr and Ripley in not recognising chandleri. In this connection it seems
inconsistent that they accepted wilkinsoni. I am unable to add much to the discussion,but for the moment, until its exact relations to chandleri are solved, I consider it cautious
to recognize wilkinsoni; the range of wilkinsoni, as here defined, probably includes Timorand Sumba.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

67

As few specimens are known from Sumba, I mention here: , 20.vii.1974, Wai Kabubak, Sumba (Verheijen, RMNH no. 85143).
Note that Ripley (1977: 85 map 3) has placed Goenoeng Api Island in the position of
Komodo, between Sumbawa and Flores, instead of in the Banda Sea, although in the
text he places it correctly.
Measurements:
Flores Sumba

wing
154
145
153
152
150
142

tail
68
60
63
66
58+
-

tarsus
49
40.8
43
41
42.5
45

entire culmen
40.5
33
35
33
39
33.3

exposed culmen
36
28
29
27.5
31
30.4

Rallina fasciata (Raffles)
Rallus fasciatus Raffles, 1822, Trans. Linn. Soc. Lond. 13: 328 – Sumatra.
Collectors.— Weber, Verheijen/Schmutz.
Material.— (?), xii.1888, Maumeri (Weber, RMNH cat. no. 15); , large gonads, 18.xii.1975, Nunang, 650
m (Schmutz, RMNH no. 97114), , 22.iii.1978, Nunang (Verheijen, RMNH no. 85155), with c/6 eggs. Billdark green, eyerim, legs and iris deep orange-red.
Eggs: c/2, 25.ii.1959, without locality (RMNH no. 70328); c/1, 22.ii.1961, Soa (RMNH no. 70329). Theegg from Soa was collected with an incubating parent, so that its identification is beyond question (cf.
Verheijen, 1964: 195, s. n. Rallina euryzonoides). The eggs are white, entirely without markings, with asmooth, glossy surface.
Measurements and weights:

RMNH no. 70328
RMNH no. 70329

32.4 × 23.9
33.0 × 23.7
37.0 × 23.5

0.56380.57830.6520The measurements and weights agree with those of eggs from Java, described byHellebrekers & Hoogerwerf (1967: 29) and with eggs from Burma described by Harrison & Parker (1967). The measurements of 39.9 × 24.3 mm given for one egg by thelast-mentioned authors are due to a misprint and should read 31.9 × 24.3 mm (Walters,in litt., 22.vii.1992).
Notes.— Ripley (1977: 160) notes that this species is highly migratory, and may be
only a winter visitor in the southern part of its range. The fact that several clutches of
eggs are known from both Java and Flores, the southern limit of its normal range, contradicts this suggestion.
In the description of the eggs, the only reference given by Ripley (l. c.) is to Harrison & Parker (1967). The last-mentioned authors laboured under the misconceptionthat authentic eggs of Rallina fasciata had not been described before they publishedtheir note. They seem to have overlooked Hoogerwerf’s (1949a: 49, pl. IV fig. 36) important publication, in which eggs from Java are described and (admittedly not verywell) illustrated, and Coomans de Ruiter’s (1951: 313, pl. VIII) description and excellent photograph of a clutch from Borneo. Our collection contains four clutches fromJava (Bartels, Hoogerwerf), one from Billiton (c/4, 29.i.1948, leg. A.H. de Bruijn, RMNH
no. 73331), one from Padjintan, Singkawang, Borneo (c/5, 9.vi.1934, leg. Coomans de

68

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Ruiter, RMNH no. 73552; this is the clutch recorded by Coomans de Ruiter, l. c.), andtwo from Flores as described above. Smythies (1981: 76) makes no mention of breedingin Borneo. Excluded from the above enumeration is a clutch from Java (leg. Sody), of
which the identification was questioned by Hellebrekers & Hoogerwerf (1967: 29). Besides being broader and a trifle heavier, as noted by these authors, the surface of theshells is a little less smooth, and without gloss. Sody’s label shows that no bird wasseen with the eggs, and that they were only subsequently identified as being of R. fasciata by M. Bartels Jr. Measurements and weights, in combination with the information, given on the label, that they were collected in a wet rice-field, suggest Ixobrychus
cinnamomeus.
Measurements:



wing
123
123

tail
48
46

tarsus44
44

entire culmen
23.8
23.4

exposed culmen
21.8
21Porzana pusilla pusilla (Pallas)
Rallus pusillus Pallas, 1776, Reise d. versch. Prov. Russ. Reichs 3: 700 – Dauria (reference not verified).
Porzana pusilla mira Riley, 1938, Proc. Biol. Soc. Washington 51: 95 – Tanggarong, Mahakkam River, East
Borneo.
Collector.— Verheijen (3, MCZ).
Material.— None.

Notes.— The dates of the Flores specimens suggest, but do not prove, that the species
is a resident of the island: late April and mid-May (cf. Paynter, 1963). In contradistinction,
all specimens from Java and Sumatra in our collection have been taken in the wintermonths:Java
Sumatra
Ceram

I
14
1
–

II
26
–
–

III
1
–
–

IV
–
–
–

V
1
–
–

VI
–
–
–

VII
–
–
–

VIII
–
–
–

IX
–
–
–

X
–
–
1

XI
–
–
–

XII
2
–
–

The last spring dates are 6.iii.1917 (Tjitamijang, Java, RMNH no. 52116) and 11.v.1918(Tjisaät, Java, RMNH no. 52117). Although there may be all kinds of bias in the samples,
there is at least a strong suggestion that the species is mainly a winter visitor (this has
never been denied), and there is no evidence for the presence of a resident population(cf. Kooiman, 1940b: 100; Junge, 1948: 313-314).
Van Marle & Voous (1988: 82) refer to eggs and a bird collected on Billiton, 5.ii.1949,
by A.H. de Bruijn, in RMNH. There is no such material in our collection, but correspondence between de Bruijn and A.C.V. van Bemmel, at that time curator of the MZB(archive RMNH) revealed that the record is correct. De Bruijn collected both eggs and abird which he had personally seen attending the nest. Van Bemmel identified the bird
as belonging to the nominate race, P. p. pusilla. However, skin and eggs are in the MZB,
not in Leiden; they provide proof that the species is not only a winter visitor but also a
resident in Malaysia.
The first Celebes specimen was collected on 17 June 1939: the date suggests that itMees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

69

belonged to a resident population and consequently it was referred, with some doubt,
to the subspecies mira (Van Marle, 1940b).
The existence of an endemic Sunda-Island race, P. pusilla mira, which has been
doubted before (cf. Junge, 1948: 313-314) becomes increasingly questionable; indeed I
see no reason at all to retain it, now that the one certain breeding bird has been identified as P. p. pusilla.
One might perhaps expect an as yet hypothetical resident population of the Lesser
Sunda Islands to be closer to Australian P. p. palustris than to nominate pusilla, but Paynter states expressly, that the specimens agree in size with pusilla. Paynter played with
the idea that the specimens from Flores represented an undescribed endemic race, but
their poor condition prevented a conclusion. Until definite evidence to the contrary
may be found, I consider that the species should be regarded as a winter visitor, and thespecimens as nominate pusilla.
Porzana fusca fusca (Linnaeus)
Rallus fuscus Linnaeus, 1766, Syst. Nat. (ed. 12) 1: 262 – in Philippinis.
Gallinula rubiginosa Temminck, 1825, Recueil d’Ois. 5 (livr. 60): pl. 357 – Java.
Collectors.— Everett (2), Verheijen (1, MZB, cf. Jany, 1955), Verheijen (7, MCZ), Verheijen/Schmutz.
Material.— (?), vii.1959, Wenus (?), Rahong (Verheijen, RMNH no. 85158); , 20.vii.1969, Paku, 350 m(Schmutz, RMNH no. 97156); , 10.ix.1971, Longko, Wangkung, Rahong, 900 m (Verheijen, RMNH no.
85136); (?) juv., 17.vi.1976, sawah R. Léma (Verheijen, RMNH no. 81322);  juv., 24.vi.1978, Natu (?)(Schmutz, RMNH no. 81307).
No eggs. The eggs listed with a question mark by Verheijen (1964: 198) must have been misidentified.

Notes.— Mayr (1944a: 146) drew renewed attention to the larger size of birds from
Flores (2 , wing 100 and 102 mm), compared with specimens from Java, Sumatra andthe Malay Peninsula. As one Luzon bird, topotypical of the nominate race, measured
equally large (wing 100 mm), he suggested that the Malaysian populations, if the size
difference was confirmed, might be known as P. f. rubiginosa. Voous (1948: 88), on thebasis of some specimens from Java and Sumatra, agreed with Mayr on their small size
and accepted the name rubiginosa for them. He was not clear, however, about the characters of P. f. erythrothorax, which is a very much larger subspecies, not known to migrate beyond Taiwan (Mees, 1970: 292-293). Ripley (1977: 254) included the resident
population of Taiwan in P. f. erythrothorax, but it belongs to P. f. phaeopyga.
White & Bruce (1986: 155) deny the smaller size of rubiginosa, but it is not clear thatthey had examined material, and specimens measured by me support it. Whether thedifference is enough to justify its formal recognition in nomenclature is another question; my personal opinion is, that it is not. Anyway, the Flores birds agree in size with
topotypical fusca and therefore must be included in the nominate race.
Measurements:
Flores  juv.
(?)(?) juv.

wing
97
100
97
94
102

tail
49
48
42
41
44

tarsus
33
33
34
31
36

entire culmen
24.2
24.3
23
23
23.9

exposed culmen
19.6
22.5
20
19
20

70

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)
Java

10 

Luzon 3

91-96
(93.0)
99
92-96(94.7)

34-42(38.8)
44
43-51
(46)

31.8-34.6(32.9)
31
32-33(32.7)

21.8-23.8(22.8)
20.6
21.7-22.9(22.1)

19.8-21.3(20.7)
18.8
19-20.6(19.6)

Porzana cinerea cinerea (Vieillot)
Porphyrio cinereus Vieillot, 1819, Nouv. Dict. d’Hist. Nat. 28: 29 – Le pays de cet oiseau m’est inconnu =Java (cf. Pucheran, 1851: 563).
Porzana leucophrys Gould, 1847, Proc. Zool. Soc. Lond. 15: 33 – Port Essington and Northern Australia.
Collectors.— Verheijen (3, MCZ, and eggs), Schmutz.
Material.—  in juvenile plumage, 27.vi.1978, sawah Lanar (Schmutz, RMNH no. 81314).
Eggs (fig. 13e): c/4, 18/25.viii.1955, Dampék (RMNH no. 70325); c/1, 18/25.viii.1955, Soa (RMNH no.
70326); c/1, 29.vi.1961, Soa (RMNH no. 70327).
Measurements and weights:

RMNH no. 70325

RMNH no. 70326
RMNH no. 70327
For comparison, clutches from Sumatra and Java:
RMNH no. 4295,
Deli, Sumatra

27.0 × 20.6
27.1 × 20.6
27.1 × 20.8
27.4 × 20.6
27.3 × 21.0
28.2 × 21.0

0.41190.45570.44070.39050.45560.478226.9 × 21.7
27.5 × 22.5
27.8 × 21.3
27.9 × 20.8
RMNH no. 52082,
28.7 × 22.8
Sitoe Palahlar, Java 29.0 × 22.0
29.0 × 22.5
29.4 × 23.0
30.3 × 22.8

A series of 18 eggs from New Guinea measured 27.2-30.1 × 20.7-22.3 mm (Mees,
1982a: 42), quite similar, and therefore I do not understand what Meise (in Schönwetter,
1988: 256) meant with the following words: “Daraus folgt, dass die kleinen MeraukeEier entgegen der Annahme von Mees....nicht zur Nominatform gehören können, die i.
D. viel grössere Eier legt”. The measurements of the New Guinea eggs are also quite
similar to those of eggs of the nominate race as provided by Schönwetter (1962: 348),
viz., 28.0-30.1 × 21.0-23.7 mm.
Notes.— Reasons for the rejection of the generic name Poliolimnas for this species
have been given elsewhere (Mees, 1982a: 42-49). In the paper mentioned, a partial
revision of the subspecies was given, but lack of material prevented me from giving adefinite judgement on the validity of Australian leucophrys, although I quoted Greenway’s (1973: 314) statement that birds from Malaya and Java (nominate cinerea)
seemed to differ from northern Australian birds (leucophrys), merely by having a
blacker head. I have since received on loan all three Australian specimens present in
the ANC (,14.vii.1975, Kapalga, Arnhemland, N.T., CSIRO no. 18072; , 5.xii.1975,

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

71

Humpty Doo, N.T., CSIRO no. 20234; , xi.1972, Darwin coast, CSIRO no. 16626).
These specimens, interestingly, have rather dark, blackish crowns, darker than mostJava birds, although not outside their range of variation. This is exactly the oppositeof the difference noted by Greenway. There are no other differences in colour, patternor measurements, hence I have no hesitation in synonymizing leucophrys with nominate cinerea.
Marchant & Higgins (1993: 567, 571) recognise no subspecies at all and confidently
list Porzana cinereus [sic] as monotypic. Surely the subspecies, brevipes, ocularis, meeki
and tannensis should have been discussed before such a step was taken.
Measurements:
wing
Flores  juv.
92
Northern Territory, Australia

90

90

93

tail
49

tarsus35

exposed culmen
18.5

middle toe
38.5

44
44.5
48.5

35.8
32
32.5

22.5
20
19.8

38.0
35.4
ca. 38Amaurornis phoenicurus leucomelanus (S. Müller)
Gallinula leucomelana S. Müller, 1842, Verh. Nat. Gesch. Ned. Overz. Bez., Land- en Volkenk.: 158 –Timor.
Collectors.— Everett (1), Rensch (1), de Jong (3), Verheijen (3, MCZ), Verheijen/Schmutz.
Material.— , 17.viii.1968, Nunang (Schmutz, RMNH no. 66092); , 23.xi.1968, Nunang (Schmutz, RMNH
no. 66093); , 8.ix.1969, Montjok, L.L. (Verheijen, RMNH no. 65225); , 20.ix. 1969, Look (Schmutz, RMNH
no. 66094); , 7.vii.1971, Waé-Ntjuang, Langkas (Verheijen, RMNH no. 66095).
Eggs: 16 clutches of c/1 (4 ×), c/2 (4 ×), c/3 (1 ×), c/4 (5 ×) and c/5 (2 ×), collected in the months January(4), February (2), March (3), April (1), July (1), December (1), and four insufficiently dated.
Some measurements and weights:

RMNH no. 70420

RMNH no. 70425

40.4 × 29.2
41.1 × 28.4
41.7 × 28.9
42.6 × 28.8
43.0 × 29.3
39.2 × 30.0
39.7 × 30.1
41.7 × 30.4
42.3 × 30.1

1.31261.293
1.3284
1.27981.3694
1.26871.30481.33881.3343

Notes.— The larger size of this subspecies, compared with the adjacent A. p. javanicus, was already noted by S. Müller when he described it. The eggs range also slightly
larger and heavier. White’s (in White & Bruce, 1986:155) remarks about A. p. javanicus,
and his grounds for its rejection are inappropriate.
In a previous publication (Mees, 1986: 27-30) I discussed the subspecies A. p. javanicus in relation to A. p. chinensis, pointing out its smaller size. In the Malay Peninsula thesituation is not clear (cf. Wells, 1999: 198-199), but such evidence as is available seems toindicate a steep gradient between javanicus and chinensis where they meet, suggesting
secondary rather than primary contact and strengthening-the case for keeping themapart.

72

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

There is every reason to discuss javanicus also in relation to its eastern neighbour,
leucomelanus, but first I want to comment on White’s treatment of javanicus. He presents
a list of measurements from various localities, apparently in support of his opinion that
there are no significant size differences between these populations. It struck me that formales from Java (number of specimens not given) he records a maximum wing-lengthof 158 mm, somewhat larger than the maximum I had found. I expected that White had
based his table on measurements personally taken, but discovered that actually they
have been raked together from various literature sources. The measurements from Java,of both sexes, have evidently been taken from Siebers (1930: 198). Siebers had 8 males
from Java, of which one in moult and disregarded by White. Among the Java birds there
is, however, one specimen from Noordwachter, an islet in the Java Sea, about 90 km
north of the western end of Java: this is the bird with a wing-length of 158 mm. In theothers the maximum is 153 mm (within the size range of 137-154 mm found by me in aseries of-29 males of javanicus). Its date, October 1889, and locality show that this is thebird previously recorded by Vorderman (1895a, s. n. Gallinula phoenicura), who unfortunately did not provide further details. It was unsexed, but because of its large size,
Siebers assumed it to be a male: it is a little too large even for a male of javanicus, andone would certainly not expect a resident population on Noordwachter, but the islet
would be an obvious landing place for a migrant of chinensis. Therefore I feel quite safe
identifying the specimen as a migrant of that race, and probably being a female rather
than a male. Although Noordwachter is about 90 km from Java and only ca. 65 km fromthe mainland of Sumatra, it is generally, faunistically and politically, included in Java.
As there are no previous records of the subspecies chinensis from Java, the specimen
constitutes an addition to its avifauna.
Now about the delineation between javanicus and leucomelanus. It is generally
known that there is a large area in which the populations are variably intermediate between these two subspecies. This area includes the whole of Sulawesi (Celebes), with
the island of Buton (van Bemmel & Voous, 1950: 89-91) and even Buru (Siebers, 1930).
Stresemann named the intermediate birds variabilis and although he believed them tobe of hybrid origin, he was not quite sure (cf. Stresemann, 1939: 346). In spite of theunusually large range these intermediate populations occupy, I also have little doubtthat they are hybrids: their extreme variability would be difficult to explain in any other way. Ripley (1977: 265) speaks of a “partially continuous phenotypic cline”. It should
be realized that a cline implies primary contact and hybridisation secondary contact, sothat Ripley’s and my views are diametrically opposed. Unlike Ripley, White consideredthese populations definitely hybrids. He rejected the name variabilis with the argumentthat a separate name “is not appropriate where unstable populations affected by introgression are involved”. The nomenclaturial solution both he and Ripley followed, wasto include all these hybrid populations in leucomelanus. This solution has the advantage
of being simple, but fails to do justice to the facts. The proper treatment would be to indicate them with the formula A. p. javanicus < A. p. leucomelanus. If this were considered
too awkward, another solution would be (dare I suggest it?) to retain for them the namevariabilis, keeping in mind that this is an unstable hybrid.
Dickinson et al. (1991: 147) come with the surprising statement: “Amaurornis isfeminine: phoenicurus is a Greek compound adjective and would end the same whether
masculine or feminine”. Although ornis may have either gender, it has expressly been

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

73

ruled that in nomenclature, words ending with ornis are to be treated as masculine (cf.
ICZN, 1985: art. 30). The name phoenicurus is a Latinized adjectivum, and as such,takes the gender of its genus; thus Phoenicurus phoenicurus, but Rhipidura phoenicura.
David & Gosselin (2002 34) treated the genus Amaurornis as feminine, with a reference
to the new edition off the Code (ICZN, 1999: art. 30.1.4.2), but they made no mentionof art. 30.1.2, in which the ruling that generic names ending in –ornis are to be treatedas masculine, is retained.
Measurements:
2
3

wing
165, 166
152, 157, 159

tail68, 7062, 63, 63

tarsus57, 6154, 55, 56

culmen39.3, 38
33, 34, 35

Gallicrex cinerea (Gmelin)[Fulica] cinerea Gmelin, 1789, Syst. Nat. (ed. 13), 1(2): 702 – Sina.
Collector.— Verheijen (2, of which 1, MCZ).
Material.— , 25.v.1969, Wangkung, Rahong (Verheijen, RMNH no. 85137). Plumage worn, but nomoult. Iris yellowish green, bill yellowish, tip of mandible yellowish white; legs olive green.

Notes.— Much about the status of this species in the southern part of its range remains to be elucidated. It is known as a breeding-bird from India and Sri Lanka (Ceylon), the northern part of the Malay Peninsula (south to Kedah), and throughout thePhilippines. In Java it is a fairly common winter visitor, not known to breed, as it is in
the southern part of the Malay Peninsula. The situation in Sumatra is less clear: it iscertainly mainly a winter visitor, but a nest is said to have been found, and there aresummer observations (Van Marle & Voous, 1988: 83-84). There is no evidence for breeding in Borneo, and from Sulawesi (Celebes) there are only a few records, all in the northern winter and therefore undoubtedly migrants. The birds from Flores may also be assumed to be migrants, although the dates are late.
Dated material from the southern part of the range in our collection is divided overthe months as follows:
I
2
34
-

Andamans
SumatraJava
FloresSri Laanka
Mindanao

II
15
1
-

III
1
3
-

IV
3
1

V
2
1
1

VI
-

VII
-

VIII
-

IX
1
-

X
-

XI
-

XII
1
2
19
-

The last dates from Java are 5 and 10 May, so that the Flores specimens, from 14(MCZ) and 25 May may certainly be regarded as late. A Mindanao bird is dated 15.
v.1889, Davao (leg. Platen, RMNH cat. no. 20), but the species is known to be a resident
there (cf. Dickinson et al., 1991: 147).
Measurements:


wing
160

tail
57

tarsus59

culmen with forehead plate36

74

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Gallinula tenebrosa frontata Wallace
Gallinula frontata Wallace, 1863, Proc. Zool. Soc. Lond.: 35 – Bouru.
Collector.— Allen.
Material.— None.

Notes.— The record of this species from Flores is based on Wallace’s (1864: 487) list,
which shows it as having been collected by Allen. The specimen (or specimens ?) has
not been traced (it is not in the BM, where so many of Wallace’s specimens have, over
the years, accumulated).
The status of the two species of moorhen in the Lesser Sunda Islands requires further investigation. G. chloropus occurs east to Flores, where it breeds, and Sumba. G.
tenebrosa is known from Timor (five specimens collected by Stein, cf. Mayr, 1944a: 133),
Sumba (Everett and Sutter each obtained one specimen), Flores as mentioned above,
and Sumbawa (leg. Elbert, cf. Rensch, 1931a: 469, presumably one specimen). AlthoughRensch (l. c.) concluded that: “Diese Art kann nicht in den Rassenkreis von G. chloropus
einbezogen werden, da sie in Celebes und Sumbawa neben diesem vorkommt”, thishas to be qualified for the Lesser Sunda Islands, where there is no proof yet of sympatricbreeding occurrence. Individuals of G. tenebrosa found west of Timor might conceivably
be migrants or vagrants.

Fig. 7. Gallinula tenebrosa, nominate race from Australia in full colour; note in particular the olive-grey
rings around the joints and toes; the toes are not as bright orange as the legs, usually more yellowish
orange, as in this bird (Busselton, Western Australia, 14.xii.1996).

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

75

Now that G. chloropus has been established as the breeding bird on Flores, Stresemann’s (1939: 318 and 1941: 39) hypothesis on the colonisation of Celebes by G. tenebrosa: “Einwanderung: von Flores her”, is no longer so obvious, unless there is some
substance to the suggestion that in Celebes and the Lesser Sunda Islands G. chloropus isgradually replacing G. tenebrosa (cf. White & Bruce, 1986: 160).
As White (1976b) pointed out, G. tenebrosa is in need of revision. The subspecies frontata is supposed to differ from nominate tenebrosa of southern and eastern Australia by
having bright red legs, with grey or olive green joints, whereas tenebrosa would have thelegs variable in colour, but not or only rarely bright red. Surprising as it may seem, atthat time it was apparently not generally known that in Australia G. t. tenebrosa shows
seasonal variation in the colours of the unfeathered parts, not only on the legs but also
bill and forehead-shield. This was studied in eastern Australia (Canberra) by Eskell &Garnett (1980). During the past few years I have noted a similar seasonal variation insouth-western Australia (Busselton, fig. 7). By far the most conspicuous are the changes
in colour of bill and frontal shield. In summer, shield and bill are bright vermilion, except
for a clear yellow tip. In this season, the legs are variable in colour, but mainly orange,
always with the joints (heels and toes) olive grey or greenish grey, and never as bright as
frontal shield and bill, orange rather than vermilion. These summer colours are universal in the last third of the year (September to December). Just as in the eastern states,
there is a great individual variation, although, once the change begins, it proceeds rapidly. In Busselton frontal shields may begin to shrivel and darken as early as the end of
December, but other birds retain their summer colours to March or even the first days ofApril. The legs, in the winter season, become generally dark grey, olive-grey or brownish
grey. Only at close range may it be seen that some scutes along the anterior margin oftibio-tarsus and tarsus are dark red, or that some irregular patches of reddish or yellow
remain. This does not influence the general dark and dull apearance of the legs. Thechange in colour of shield and bill begins with a general darkening, dusky concealing
the bright colours. There is a stage in which the bill looks dull dark red. Finally the bill
and the shriveled shield become blackish grey, except that the bill tip remains more orless yellow, dirty yellowish, at any rate pale, and the middle of the shield become lightbrownish, the colour of the shrivelled skin. From the beginning of June, summer colours
may begin to return. This is usually also a diffuse process, but in birds which I assume tobe immature, changing for the first time, bright vermilion appears first along the edge ofthe frontal shield, in a horsehoe shape, and in the succeeding days spreads inward anddownwards into the bill. Various stages may be seen in the following weeks, and by the
beginning of September, at the latest, all birds are in full colour.
What now is the relevance of all this for the subspecies frontata? Hitherto attention
has been focussed on leg-colour; seasonal variation in bill- and shield-colour was described for nominate tenebrosa by Eskell & Garrett, but there is no mention of these partsin the other subspecies, frontata and neumanni. If these two subspecies show no seasonal variation in leg-colour, it is at least very likely that they also lack seasonal variation in bill and frontal shield colour. Bill and shield are far easier to observe than legs (ifonly for the reason that a large proportion of observations will be of swimming birds),
and it would be easy to pay attention to this character.
There has been speculation about the relationship between G. tenebrosa and G. chloropus. Some authors have even suggested thet they are conspecific. Apart from other76

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

morphological differences, I am not aware that a seasonal variation in colours of theunfeathered parts is known from anywhere within the extensive range of G. chloropus,
and the raucous call of G. tenebrosa is also quite different from that of its congener.
Gallinula chloropus orientalis Horsfield
Gallinula orientalis Horsfield, 1821, Trans. Linn. Soc. Lond. 13: 195 – Java.
Collectors.— Verheijen (9, MCZ), Verheijen/Schmutz.
Material.— , 2.vi.1969, Tjantjar, Rahong (Verheijen, RMNH no. 85135); (?), 18.vi.1969, Montjok, L.L., ca.
700 m (Verheijen, RMNH no. 65230);  juv., 1.vii.1969, Tjumbi, Rahong (Verheijen, RMNH no. 65231); juv., 9.vii.1969, Wangkung, Rahong (Verheijen, RMNH no. 65232);  juv., 14.vii.1969, Wangkung (Verheijen, RMNH no. 65233); , 8.xii.1969, Wetok Ndekes (?), 900 m (Verheijen, RMNH no. 65234); ,
19.i.1970, Montjok, 800 m (Verheijen, RMNH no. 65235);  with large gonads, 10.ix.1971, Paku, 400 m(Schmutz, RMNH no. 85134).
Eggs: 15 clutches of from 1 to 7 eggs, collected in the months April (3), May (6), June (2), and four without data (RMNH nos. 70396-70404, 70406-70410, 70410A).
Some measurements and weights:

RMNH no. 70400
RMNH no. 70406

41.0 × 30.3
41.8 × 30.1
36.8 × 28.3
37.0 × 28.3
37.7 × 28.5
37.8 × 28.4
40.7 × 29.9
40.9 × 29.0
41.6 × 29.6

1.7092
1.7146
1.143
1.22381.1913
1.22911.58371.49351.5174

Notes.— For a discussion of this species, see the notes given under G. tenebrosa.
Measurements:2

wing
155, 160

tail68, 64tarsus49, 50culmen with shield
44, 45Porphyrio porphyrio indicus Horsfield
Porphyrio Indicus Horsfield, 1821, Trans. Linn. Soc. Lond. 21: 194 – Java.
Porphyrio porphyrio plessenorum Neumann, 1941 (November), Zool. Meded. 23: 109 – Bratan See, Nord
Bali in 1200 m.
Collectors.— Verheijen (1, MCZ), Schmutz.
Material.— , vii.1969, Sokrutung (Schmutz, RMNH no. 85160).
Eggs: c/3, 23.vi.1959, Dampék (RMNH no. 70411).
The eggs have a pale café-au-lait ground colour, with a moderate number of larger and smaller brownspots, and light violet-grey secondary spots. The ground colour is decidedly browner than that of eggs
from Celebes and Java in the RMNH collection, which have a greyish white to cream ground colour.
Measurements and weights:

RMNH no. 70411

46.8 × 34.0
47.3 × 35.9
47.6 × 34.5

2.19422.5156
2.1992

Notes.— The specimen examined obviously belongs to the subspecies indicus,
having the large azureous-blue breast-patch which separates this subspecies from theMees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

77

forms melanopterus and melanotus. I am not aware that, since its description, the name P.
p. plessenorum has again been recorded in the literature. The type-series of six specimens
is in Leiden and I am satisfied that they also are P. p. indicus. Ripley (1977: 299 map)
included the Lesser Sunda Islands, from Lombok eastwards, into the range of melanopterus, although the species is not yet known from Lombok and Sumbawa. A few pages
later, in his text, he records it in the Lesser Sunda Islands from Timor only. White (inWhite & Bruce, 1986: 158-159) calls birds from the Lesser Sunda Islands samoensis. His
judgement was based on literature only and he was not handicapped by personal study.
The species is clearly in need of a revision.
Measurements:


wing
210

tail
67

tarsus75

bill with shield
56.5

bill from nostril
15

Irediparra gallinacea gallinacea (Temminck)
Parra gallinacea Temminck, 1828, Recueil d’Ois. 5 (livr. 78): pl. 464 – Célèbes.... dans le district de Menado.
Collector.— Weber.
Material.— (?), skeleton, 26/28.xi.1888, Bari (Weber, RMNH cat. a).

Notes.— With much hesitation I retain a trinomial for this species, as I regard one of
the usually accepted subspecies (novaeguinae) as a synonym, and doubt the validity ofthe only other one (novaehollandiae, cf. Mees,1982a: 51).
Rostratula benghalensis benghalensis (Linnaeus)[Rallus] benghalensis Linnaeus, 1758, Syst. Nat. (ed. 10) 1: 153 – Asia.
Collector.— Verheijen (2, MCZ), Verheijen.
Material.— , 14.ix.1971, Flores (Verheijen, RMNH no. 85151).
Eggs: c/2, c/4, 29.vi.1961, Soa (RMNH nos. 70428, 70429); c/4, 23.v.1961, Soa (RMNH no. 70430); c/2,
31.v.1961, Soa (RMNH no. 70431).
Measurements and weights:

RMNH no. 70428
RMNH no. 70429

RMNH no. 70431

36.6 × 25.5
36.6 × 25.8
34.2 × 25.8
34.8 × 25.3
34.8 × 25.5
35.1 × 24.8
34.2 × 26.1
35.1 × 26.2

0.92080.93290.95740.88720.90950.8712
1.0272
1.0048Notes.— Birds from Flores belong to the nominate race (cf. also Paynter, 1963).
Measurements:


wing
125

tail
34

tarsus42

Pluvialis fulva (Gmelin)[Charadrius] fulvus Gmelin, 1789, Syst. Nat. (ed. 13) 1(2): 687 – Tahiti.

exposed culmen45

78

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Collectors.— Allen (1), Weber (1, ZMA), Everett, Rensch (1), Schmutz.
Material.— 2 , , 20.ix.1982, Nisar, 600 m (Schmutz, RMNH nos. 81029, 81030, 81032).

Notes.— Allen’s specimen was first published by Wallace (1864: 487, s. n. Charadrius
longipes) and later by Sharpe (1896: 205, s. n. Charadrius dominicus).
Of the material collected by Schmutz, the two males are still partially in summer
dress, with mottled black and yellowish white underparts.
There has long been discussion over the status of dominica and fulva in relation toeach other: subspecies of a single species or different species. Vaurie (1965: 385) dismissed earlier claims that they are not conspecific, but on the basis of differences inmoult and migration, Stresemann & Stresemann (1966: 53, 200-203) argued strongly infavour of considering fulva a separate species. Their views were not generally accepted;
for example Glutz (1975: 313-314) and Cramp & Simmons (1983: 196-200) chose to retain a subspecific relationship. The evidence provided by Connors (1983) for separating
P. fulva specifically from P. dominica is suggestive rather than conclusive, but the supporting particulars since published by Alström (1990) and Connors et al. (1993), haveconvinced me.
Charadrius leschenaultii leschenaultii Lesson
Charadrius Leschenaultii Lesson, 1826, Dict. Sci. Nat. (Levrault) 42: 36 – Pondichéry.
Collectors.— Allen, Rensch (1).

Notes.— As Wallace (1864: 487) listed this species from Flores, I assume that Allen
has collected material, but no actual specimen has been traced. Throughout the Archipelago, this is one of the commonest migrant waders.
I use a trinomial on the evidence provided by Spitzenberger (in Glutz, 1975: 257)
and Roselaar (in Cramp et al., 1983: 177-178), that there is a well-marked western subspecies C. l. columbinus. Roselaar recognizes in addition a subspecies C. l. crassirostris,
but the measurements supplied do not separate it convincingly from the nominate race,and the statement that in non-breeding plumage all races are similar, would make separation of C. l. leschenaultii and C. l. crassirostris in the winter quarters impossible.
Charadrius veredus Gould
Charadrius veredus Gould, 1848, Proc. Zool. Soc. Lond. 16: 38 – Northern Australia.
Collector.— Schmutz.
Material.— , 22.ii.1983, Nisar, 100 m (Schmutz, RMNH no. 81162). Iris brown.

Notes.— Considering that this is a fairly common winter visitor to north-western
Australia (cf. Blakers et al., 1984: 158; Lane, 1987: 70-72), it is surprising that the above
specimen constitutes only the second record for the Lesser Sunda Islands, the first one
being from Timor and being questionable. I have, without difficulty, traced the Timor
record from Hellmayr (1914: 106), through Mayr (1944a: 133) to White & Bruce (1986:165), but have failed to find its source. Hellmayr’s vague references to S. MüllerMees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

79(Koepang) and Everett (Atapupu) do not lead anywhere and I suspect an error. Likely
as it is that C. veredus visits Timor on migration, the evidence is unsatisfactory. In otherwords, Father Schmutz’s specimen constitutes not the second, but the first record fromany of the Lesser Sunda Islands.
Measurements:


wing
170

tail
63½

tarsus46½

entire culmen
28.3

exposed culmen
22.3

Numenius phaeopus variegatus (Scopoli)
Tantalus (variegatus) Scopoli, 1786, Del. Flor. Faun. Insubr. (2): 92 – no locality = Luzon.
Collectors.— Semmelink, Allen, Everett, Schmutz.
Material.— (?), 4.vi.1862, Larantoeka (Semmelink, RMNH cat. no. 50); , 22.x.1969, Nisar, beach (Schmutz, RMNH no. 85150); , 23.ii.1982, Joneng, beach (Schmutz, RMNH no. 81163); (?), Flores, without
data (Schmutz, received 1983, RMNH no. 81033).

Notes.— The Whimbrel is one of the commonest and most widely distributed winter-visitors in the Indo-Australian region, where it is present throughout the year (there
are specimens from all months in the RMNH).
Limosa lapponica baueri Naumann
Limosa Baueri Naumann, 1836, Naturgesch. Vögel Deutschl. 8: 429 – Neuholland = Norfolk Island (seeNotes).
Limosa lapponica var. novaesealandiae Gray, 1845, Voy. Erebus & Terror, Birds: 13 – New Zealand (reference
not verified).
Collector.— Weber (1).
Material.— (?), 23/25.xi.1888, Reo (Weber, ZMA no. 24087).

Notes.— Limosa Baueri was rejected as a nomen nudum by Hartert & Steinbacher(1938: 431) and later still by Cheng (1976: 202-203). Mayr (1941b: 31) was not convinced
and a few years later returned to the use of baueri (Mayr, 1944a: 140, footnote; Delacour& Mayr, 1946: 72). The name baueri was subsequently accepted as valid by Vaurie (1965:422), and by Condon (1975: 132), who expressly stated: “not a nomen nudum as claimed
by Hartert and Steinbacher”. Notwithstanding this definite statement, I consider that
there is still room for genuine doubt about the validity of L. baueri. The original description reads as follows: “Ein paar nähere Verwandten hat sie [meant is L. l. lapponica] an
Limosa adspersa (des Berliner Museums) aus Mexiko, und an Limosa Baueri (des Wiener
Naturalienkabinets) aus Neuholland, welche beide jedoch bedeutend grösser oderhochbeiniger sind, beide aber ebenfalls einen schmallgebänderten Schwanz haben”.
Note that, although some characters are given to distinguish L. adspersa and L. Baueri
from L. lapponica, there is nothing here to distinguish L. adspersa and L. Baueri from eachother. Therefore I feel obliged to agree with Hartert & Steinbacher that, strictly speaking,L. Baueri is a nomen nudum. Being reluctant to reject a name that has become almost
universally accepted in recent years, and because of the somewhat doubtful characterof the issue, I retain baueri for the moment.

80

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

The original description of L. Baueri is on page 429, not 29 as cited in several recent
lists (Condon, 1975: 132; Turbott, 1990: 151): one of those misprints that is carried onfrom list to list.
Even when the name L. baueri is considered invalid, the specimen on which it was
based has historical interest. The name suggests that F.L. Bauer was the collector, whichis interesting, as Whittell (1954: 43-44) describes Bauer as an artist only, and makes nomention of birds collected by him. The point intrigued me, and therefore I enquired of
Dr Schifter. Dr Schifter informed me that the original specimen from the collection of
Ferdinand Bauer is still present, and that in addition there is a sketch, agreeing with thespecimen, dated in Bauer’s own handwriting with: “Norfolk Island, 21. September
1804”. This further disposes of Mathews’s (1912: 220) arbitrary and unexplained restriction of the type locality to Victoria. Dr Schifter has since published a paper about Bauerand his collections, to which I refer for further particulars (Schifter, 1992).
Tringa hypoleucos Linnaeus[Tringa] Hypoleucos Linnaeus, 1758, Syst. Nat. (ed. 10) 1: 149 – Europa.
Collectors.— Semmelink, Allen, Verheijen.
Material.— (?), 1862, Larantoeka (Semmelink, RMNH cat. no. 72); , 22.ix.1971, Ruteng, 1150 m (Verheijen, RMNH no. 85152).

Tringa glareola Linnaeus[Tringa] Glareola Linnaeus, 1758, Syst. Nat. (ed. 10) 1: 149 – Europa.
Collectors.— De Jong (1), Verheijen (16, MCZ), Verheijen.
Material.— , 4.x.1968, Rahong (Verheijen, RMNH no. 65241); , 24.x.1968, Rahong (Verheijen, RMNH
no. 65242); , 25.x.1968, Rahong (Verheijen, RMNH no. 65243); , 28.x.1968, Rahong, 900 m (Verheijen,
RMNH no. 65244); , 31.x.1968, Wangkung, Rahong (Verheijen, RMNH no. 85148).

Tringa nebularia (Gunnerus)
Scolopax nebularia Gunnerus, 1767, in Leem, Beskr. Finm. Lapp.: 251 – near Trondheim, Norway (reference
not verified).
Collector.— Schmutz.
Material.— , 25.ix.1969, Nunang-Meer, 650 m (Schmutz, RMNH no. 65240); , 6.xii.1982, NunangMeer, 650 m (Schmutz, RMNH no. 81028).

Notes.— The Greenshank is a common winter visitor to the whole Indo-Australianregion (as well as to Africa).
Gallinago stenura (Bonaparte)
Scolopax stenura Bonaparte, 1830, Ann. Stor. Nat. Bologna 4: 335 – Abita nelle isole della Sonda, segnatamente in quella de Giava = Buitenzorg, West Java (cf. Mees, 1986: 33).
Scolopax stenoptera Schlegel, 1844, Krit. Übers. Europ. Vögel: 96 – Ostindien.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)81

Collector.— Verheijen (3, MCZ).
Material.— None.

Notes.— See the discussion under the following species.
Obviously, G. stenura must occur in Sulawesi (Celebes), but as White & Bruce (1986:
174) have pointed out, the documentation for its occurrence is rather unsatisfactory.
The best evidence comes from Coomans de Ruiter & Maurenbrecher (1948: 191) andCoomans de Ruiter (1954: 92-93); it is very likely correct, but I agree that it is too meagre as a basis for adding the species to the avifauna of Sulawesi. Sody (1954) claims apersonal observation from Makassar, and a second-hand one from Gorontalo, but hedemonstrates a complete lack of awareness that snipe seen anywhere in the Archipelago could be anything but G. stenura (he does not even mention G. megala), so that his
records must be dismissed. The same applies to an “unconfirmed record” by Escott &Holmes (1980). It would be better not to burden the literature with unconfirmed
records.
Gallinago megala Swinhoe
Gallinago megala Swinhoe, 1861, Ibis 3: 343 – by inference several localities in northern and eastern China(cf. Mees, 1982a: 63-64).
Collectors.— Verheijen (22, MCZ), Verheijen/Schmutz.
Material.— (?), xii.1958, Rahong (Verheijen, RMNH no. 66258); , 4.ii.1969, Rahong, 900 m (Verheijen,
RMNH no. 65237); , 10.ii.1969, Rahong (Verheijen, RMNH no. 65238); , 13.ii.1969, Rahong, ca. 1000m (Verheijen, RMNH no. 65239); (?), 26.xii.1982, Lembor, 250 m (Schmutz, RMNH no. 97173). Iris
brown or greyish brown, bill yellowish or brownish olive, towards the tip darker, legs greenish to yellowish olive.

Notes.— This is apparently the commoner of the two species of snipe wintering on
Flores, as Paynter (1963) received: “A long series” (22 specimens) from Father Verheijen,
against three G. stenura, whereas I received five G. megala and no G. stenura. Verhoeye &Holmes (1998: 21) misquoted Paynter, thus giving the impression that G. stenura is thecommon snipe of Flores and G. megala the scarce one. In contrast, G. stenura is the common snipe in Java, where G. megala is very scarce (cf. Van Bemmel, 1938). Delacour’s(1947b: 87) succinct statement that in Malaysia, G. stenura is rare, and G. megala common, is an error. Our collection contains from Java 97 specimens of G. stenura (among
these, several from East Java), and only two specimens of G. megala. There are no reliable records from Sumatra (Van Marle & Voous, 1988: 92). The inclusion of Sumatra andthe exclusion of Java from the winter range by Higgins & Davies (1996: 43) are both erroneous.
Calidris ruficollis (Pallas)
Trynga ruficollis Pallas, 1776, Reise versch. Prov. Russ. Reichs 3: 700 – Circa lacus salsos Dauriae campestris= Kulussutai, southern Transbaikalia (reference not verified).
Collector.— Weber.
Material.— 2 (?), 23/25.xi.1888, Reo (Weber, RMNH cat. no. 39, ZMA no. 8191).

82

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Notes.— These birds were recorded by Büttikofer (1894: 306) under the name of
Tringa minuta. Büttikofer listed a skin and a spirit-specimen; the latter has, evidently,
also been turned into a study-skin.
Calidris alba (Pallas)
Trynga (alba) Pallas, 1764, Vroeg’s Cat., Adumbr.: 7 – no locality, but refers to p. 32, no. 320 of the main
volume, where: “Valt aan de Noordsche Zeekusten” = North Sea coast of Holland.
Collector.— Schmutz.
Material.— (?), 27.ix.1971, Nanga-Lili, sandy beach near a river-mouth (Schmutz, RMNH no. 85153).
The small size suggests that this bird is a male.

Notes.— Bruce (in White & Bruce, 1986: 169) states: “Schmutz (1977) noted its inclusion in an unpublished list for Flores, but had no records”. The list referred to would bea list I sent to Father Schmutz, and the inclusion was based on the specimen listed here.
This species has rarely been recorded from the Lesser Sunda Islands, where otherwise it is only known from Sumba (2  collected by Stein, cf. Mayr, 1944a: 141). In Java,it has also been regarded as uncommon (cf. van Bemmel, 1940b), but in East Java, Kooiman (1940a; 1940b: 102) observed the species regularly, once even a flock of ca. 100
birds.
Measurements:
[]

wing
118

tail
45½

tarsus27½

entire culmen–

exposed culmen
25

Himantopus himantopus leucocephalus Gould
Himantopus leucocephalus Gould, 1837, Synops. Birds Austr. 2: pl. 37 – Australia generally and the islandsof Java, Sumatra, & c.
Collector.— Schmutz.
Material.— , 11.vii.1969, Kenari (Schmutz, RMNH no. 65245).

Notes.— Surprisingly, there are apparently no earlier records of this widely-distributed species.
Phalaropus lobatus (Linnaeus)[Tringa] tobata, misprint corrected to lobata in the Emendanda, Linnaeus, 1758, Syst. Nat. (ed. 10) 1: 148,
824 (Emend.) – America septentrionali, Lapponia.
Collectors.— None.

Notes.— Verheijen (1971) observed a Phalarope at Kedindi, on the north-west coastof Flores, on 1.xi.1970. The bird was associating with domestic ducks in a salt-water
creek in a coconut-plantation, ca. 200 m from the sea. At sea this species is a common
winter visitor throughout the region.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

83

Esacus magnirostris (Vieillot)
Oedicnemus magnirostris Vieillot, 1818, Nouv. Dict. Hist. Nat. (Nouv. éd.) 23: 321 – no locality = la N.elle
Hollande (see Pl. G. 39, fig. 1, in vol. 28, 1819).
Collectors.— Van der Sande (1), Rensch (1), Verheijen/Schmutz.
Material.— (?), i.1909, south coast (v.d. Sande, ZMA no. 24161).
Egg: c/1, 18.xi.1948, Papagaran (RMNH no. 70432).
Measurements and weight:

RMNH no. 70432

62.2 × 45.1

4.9815

Stiltia isabella (Vieillot)
Glareola isabella Vieillot, 1816, Analyse: 69 – l’Australasie.
Glareola grallaria Temminck, 1820, Manuel d’Orn. (2 éd.) 2: 503 – l’Austral-Asie.
Collectors.— Semmelink, Verheijen/Schmutz.
Material.— (?), 1862, Larantoeka (Semmelink, RMNH cat. no. 4);  im., 23.vi.1969, Ruteng (Verheijen,
RMNH no. 65249); , 2 , 25.vi.1969, Ruteng (Verheijen, RMNH nos. 65250, 65251, 65252); , 3 ,
18.vii.1969, Ruteng (Verheijen, RMNH nos. 65255, 65253, 65254, 85149).
Iris light brown to brown, bill dark brown, at gape reddish, legs purplish brown.

Notes.— On the label of RMNH no. 65249 is noted that this is one of five specimens
brought in, all (like this one) without remiges (these had presumably been pulled out
by the catchers). Evidently, this species is a periodically common visitor in the southern
winter.
Specimen RMNH cat. no. 4 was recorded by Schlegel (1865b: 17), but overlooked
by Rensch (1931a). Of the other old, historical specimens listed by Schlegel, RMNH cat.
no. 1 is of particular interest, being the type of Glareola grallaria, and very likely also a
syntype of Glareola isabella (having been received from the Paris Museum).
The other member of this family to be expected on Flores is Glareola maldivarum. I
mention it here to point out that on Timor, this species was collected as long ago as
November 1829 by S. Müller (Schlegel, 1865b: 17). The year 1820 given by Schlegel is, ofcourse, an error. I found that actually, the mounted specimen (RMNH cat. no. 19) has a
cardboard label with the year 1820 on it, but on the under surface of its socle, the correctyear 1829 is pencilled. White & Bruce (1986: 178) only mention a record by de Sousa(1883), but Müller’s specimen was obtained fifty years earlier.
One wonders whether, perhaps, the birds seen by Father Schmutz near Nisar on
23 November 1974 and between Djonéng and Lita early December 1969, might havebeen G. maldivarum rather than S. isabella. The dates of specimens of S. isabella in thewinter-quarters in our collection range from 6 June to 7 November, and one from May(without exact date). Dates of G. maldivarum in the winter-quarters range from 21 September to 16 April. The RMNH material from Java is dated September (2), October (46)and November (15).
Sterna albifrons sinensis Gmelin[Sterna] sinensis Gmelin, 1789, Syst. Nat. (ed. 13) 1 (2): 608 – Sina (ex Latham, 1785: 365: China).

84

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)[Sterna] Pusilla Temminck, 1840, Manuel d’Orn. (2e éd.) 4: 465 – des Îles de la Sonde et des Moluques ...
On la trouve jusqu’á la Nouvelle-Guinée = Java, cf. Junge (1948: 317).
Collector.— Rensch (4).
Material.— None.

Notes.— Rensch (1931a: 489) identified his material as S. a. sinensis, and this was
confirmed by Junge (1948: 315-318), who examined one of Rensch’s specimens. Junge
further concluded that the breeding birds of the region are sinensis, and that pusilla is asynonym. Nadler (1976), however, came with a different opinion: he considered that,
although the breeding birds of the Indian Archipelago are sinensis, the region is, witha great degree of probability, the winter-quarters of a smaller subspecies, which breeds
in northern India, and that the name pusilla was based mainly on such smaller birds, inwinter plumage, as is apparent from Temminck’s description and from the name hegave them (pusilla); for sinensis is larger, not smaller than nominate albifrons. These
smaller Indian birds were included in nominate albifrons by Junge (1948) and by Glutz(1982: 734). If this view is accepted, all that remains to be decided is the unimportantquestion, whether pusilla is a synonym of albifrons or a synonym of sinensis. Junge
made a mounted bird from Java, in breeding-dress, lectotype of S. pusilla (RMNH cat.
no. 1 of S. a. sinensis). In this bird I measured a wing-length of 191 mm. This makes S.
pusilla a synonym of S. a. sinensis, unless it can be proved that the specimen was notpart of the type series. The specimen was collected by Kuhl (1820/1821). Unfortunately, it has been newly mounted, so that there is no original hand-writing underneath the socle, the writing being in van Oort’s hand. Nevertheless, van Oort clearly
indicates that the bird was originally labelled as Sterna pusilla Kuhl, and there can be
no doubt that it was in Temminck’s hands. The fact that the name S. pusilla was given(as a manuscript name) by Kuhl, and that Kuhl collected exclusively in West Java,
where he died within nine months of his arrival from Europe, confirms that this bird
must certainly have been a syntype of S. pusilla Kuhl, and that Junge was completely
justified in making it the lectotype.
Storr (1973: 46, and several subsequent publications) elevated the weakly differentiated subspecies sinensis to species status, without explanation.
Sterna bergii cristata Stephens
Sterna cristata Stephens, 1826, in Shaw’s Gen. Zool. 13 (1): 146 – China; and many of the south-easternislands of Asia = China (restricted type locality).
Collector.— Semmelink.
Material.— (?), 1862, Larantoeka (Semmelink, RMNH cat. no. 39).

Notes.— The above specimen was recorded by Schlegel (1864a: 11, where it is cat.
no. 35).
Chlidonias hybridus subsp.
Collector: Verheijen (1, MCZ).
Material: (?), 4.vi.1961, Soa (Verheijen, MCZ no. 332909).

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)85

Notes.— This specimen was examined and identified for me by Dr Paynter (in litt.,
14.x.1993), according to whom it is an immature bird, which he has not attempted to
identify to subspecies. On geographical grounds, and month of collecting, the bird
would almost certainly belong to the Australian subspecies javanicus (cf. Mees, 1977b).
It is curious that this specimen seems to constitute the first definite record from theLesser Sunda Islands, where the species ought to be a regular migrant visitor. Recent
field observations from Timor confirm this (Andrew, 1986).
David & Gosselin (2002: 32) have taken me to task over treating hybridus, a, um, as
an adjectivum, “against the advice of a scholar”. The reason was that in a book of which
the said scholar is co-author, the name is expressly listed as a Latin adjectivum (Coomans
de Ruiter et al., 1948: 119). This book was the obvious source for a Dutch ornithologist
to consult and rely on, certainly 25 years ago when my paper was published (Mees,
1977b: 49). I had every reason to regard this book, a pioneer study in its field, as a reliable guide. I note its omission from the references given by David & Gosselin. The late
H. G. Deignan certainly was a good latinist. He provided the name Pteruthius hybrida
Harington with a “sic!” and changed it into hybridus (Deignan, 1964: 388). This issue isclearly controversial among classical scholars. Where the specialists disagree, who am
I, a complete ignoramus in this field, to take sides. The natural course to take is to retain
the spelling I have used in recent years, until the controversy has been completely
solved. See also below under Cisticola.
Chalcophaps indica indica (Linnaeus)[Columba] indica Linnaeus, 1758, Syst. Nat. (ed. 10) 1: 164 – in India orientali.
Collectors.— Weber, Everett, de Jong (2), Verheijen/Schmutz.
Material.— (?), xi.1888, Maumeri (Weber, RMNH cat. no. 136); , 5.xi.1968, Ruteng (Verheijen, RMNH
no. 65284); , 5.iii.1971, Rana Loba, Borong (Verheijen, RMNH no. 66097); , 11.iii.1971, Kisol, Borong(Verheijen, RMNH no. 66096); , 12.iii.1971, Kisol, Borong (Verheijen, RMNH no. 66098); , 1.v.1971,
Ngalor-Roga, Rahong (Verheijen, RMNH no. 66099); , 15.ix.1971, Naga, 250 m (Schmutz, RMNH no.
97112);, 10.x.1971, Nunang, 650 m (Schmutz, RMNH no. 85314); , 16.vi.1973, kuststrook Borong (Verheijen, RMNH no. 85315); , 16.ix.1976, Poco Lareng (Verheijen, RMNH no. 81095); , (Schmutz no.
1515, RMNH no. 81096).
Eggs: 24 clutches of 1 (19 ×) and 2 (5 ×) eggs, from various localities on Flores, and from Palué, collectedin the months March (2), April (7), May (5), June (2), July (3), October (1) and November (1), and three
undated (RMNH nos. 70525-70547, 73509). Unlike the eggs of most other species of pigeon, which arewhite, the eggs of Chalcophaps indica are easily identified because of their pale brownish cream colour.
Some measurements and weights:

RMNH no. 70536
RMNH no. 70538
RMNH no. 70541

27.7 × 20.9
28.6 × 21.3
27.2 × 19.7
26.8 × 20.9
26.9 × 21.1

0.37090.36140.28740.35440.3315

Notes.— Birds from Flores belong to the nominate subspecies, the males having awhite forehead and supercilium, and a blue-grey crown and nape.
Birds from Timor, C. i. timorensis Bonaparte, are abruptly different and closer to theAustralian group of subspecies, although perhaps to a certain extent intermediate

86

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)(Johnstone, 1984). The geographical variation of this pigeon, with the remarkably wide
range of the nominate race, evidently requires further study.
Treron floris Wallace
Treron floris Wallace, 1864, Proc. Zool. Soc. Lond. (1863): 496 – Flores and Solor Islands.
Collectors.— Semmelink, Allen, Martens, Weber, Everett, Rensch (2), Schmutz.
Material.— , 1862, Larantoeka (Semmelink, RMNH cat. no. 1); , (?) juv., xi.1888, Maumeri (Weber,
RMNH cat. nos. 2, 3); (?), xii.1888, Mbawa near Rokka (Weber no. 507, ZMA); , ?, (?), 18.vi.1974,
Orong, 550 m (Schmutz, RMNH nos. 85120, 85121, 85122).

Notes.— Although this species was described by Wallace, on the basis of specimens
collected in 1862 by Allen, twenty years earlier, a specimen from Bima, Sumbawa, had
found its way to Leiden (cf. Schlegel, 1866b: 212; now RMNH cat. no. 7).
Measurements:

?
(?)
(?)

wing
145
150
152
145

tail
78½
83½
88
76

tarsus23.5
24.5

entire culmen
24
22.5
23
22

exposed culmen
17
17
18.5
16.5

Ptilinopus cinctus albocinctus Wallace
Ptilonopus albocinctus Wallace, 1864, Proc. Zool. Soc. Lond. (1863); 496, pl. XXXIX – Flores (in the interior
only).
Ptilopus cinctus Florensis Schlegel, 1871, Ned. Tijdschr. Dierk. 4: 20 – Flores. Re-naming of P. albocinctus,
as it is not regarded as a species, but a “souche”.
Collectors.— Allen (1), Colfs, Rensch (6), Schmutz.
Material.— (?), 1880, Flores (Colfs, RMNH cat. no. 1); , 29.iii.1976, Poco Nernancang (Schmutz, RMNH
no. 97110); (?),5.vi.1976, Poco Nernancang (Schmutz, RMNH no. 81102); (?), 17.v.1976, Lingko Ncilor(Schmutz, RMNH no. 81098); (?), 22.v.1976, Lingko Ncilor (Schmutz, RMNH no. 81101); (?), 21.vii.1976,
Poco Nernancang (Schmutz, RMNH no. 81103); , (?), 1.v.1978, Golo Léhot (Schmutz, RMNH nos.
81100, 81099); (?), 4.v.1978, Golo Rucuk (Schmutz, RMNH no. 81097).

Ptilinopus regina flavicollis Bonaparte[Ptilopus] flavicollis Bonaparte, 1854, Consp. Gen. Av. II: 20 – Timor.
Collectors.— Semmelink, Colfs, Everett (1).
Material.— (?) ad., 1862, Larantoeka (Semmelink, RMNH cat. no. 5); (?) ad., vi.1880, Flores (Colfs,
RMNH cat. no. 6).

Notes.— As stated by Rensch (1931a: 488), this pigeon appears to be rare on Flores,
being known from the two specimens listed here, and a single one collected by Everett
at Maumeri. The Maumeri record, published by Hartert (1898a: 48), puzzled me, as I
could not find any other evidence that Everett had visited this locality. Mrs. LeCroy (inlitt., 12.iii.1991) has, however, confirmed the record: , viii.1897 (Everett, AMNH no.
609099).

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

87

According to Johnstone (1981), P. r. flavicollis is a synonym of the Australian P. r.
ewingii; the reason why I have not yet accepted this conclusion, is a reluctance to discard
a name that has been in use since 1854, without personal investigation (for which thematerial has not been available to me), and that a distribution of subspecies as indicatedby Johnstone is unusual. For much the same reasons, White & Bruce (1986: 201) haveretained flavicollis. Johnstone had not examined material from Flores. Higgins & Davies(1996: 993) also continue to recognize flavicollis.
Ptilinopus melanospilus melanauchen (Salvadori)J[otreron] melanauchen Salvadori, 1875, Ann. Mus. Genova 7: 671 – Flores.
Collectors.— Semmelink, Allen, Colfs, Weber, de Jong (1), Verheijen (2, MCZ), Schmutz.
Material.— , 13.vi.1862, Larantoeka (Semmelink, RMNH cat. no. 11, holotype of the subspecies); (?) =, 1880, Flores (Colfs, RMNH cat. no. 13); , xi.1888, Maumeri (Weber, RMNH cat. no. 14);  juv.,
xi.1888, Maumeri (Weber, RMNH cat. no. 15);  imm., 29.xi.1888, Rusa Radja (Weber, ZMA no. 2574); im., 29.xi.1888, Rusa Radja (Weber, ZMA no. 2574); , 15.vii.1969, Paku-Nggoang, 300 m (Schmutz,
RMNH coll. no. 0000); , 17.vii.1969, Paku (Schmutz, RMNH no. 85087); , large gonads, 20.i.1973, near
Nunag, 800 m (Schmutz, RMNH coll. no. 0000).
Eggs: c/2, 18.vi.1961, Soa (RMNH no. 70562); c/1, xi.1961, Soa (RMNH no. 70563); c/1, 1964, Soa (RMNH
no. 70564). Curiously, Verheijen (1964: 198) gave only a single record (with a query) and that one forMay. The eggs are plain white, only moderately glossy.
Measurements and weights:

RMNH no. 70562
RMNH no. 70563
RMNH no. 70564

29.5 × 22.3
29.6 × 22.4
28.1 × 22.7
29.0 × 22.6

0.49070.51550.47340.4745The eggs agree in size with eggs from Java recorded by Hellebrekers & Hoogerwerf(1967: 42), or they may be marginally broader, but they are distinctly heavier. On theother hand, they agree perfectly, in appearance, measurements and weight, with eggsof Streptopelia chinensis tigrina, so that a large dose of scepticism about their identification is justified.
Notes.— The name Jotreron melanauchen was entirely based on Schlegel (1866b: 207):“Individu de l’Île de Flores. Aile 4 pouces 2 lignes. Tache de la gorge d’un jaune de citron foncé. Tache noire de la nuque large”. This is RMNH cat. no. 11, the only specimen
from Flores that was available to Schlegel.
Ducula aenea polia (Oberholser)
Muscadivores aeneus polius Oberholser, 1917, U.S. Nat. Mus. Bull. 98: 18 – Pulo Siantan, Anamba Islands.
Collectors.— Semmelink, Allen, Weber, Everett, Rensch (1), Verheijen (1, MCZ), Verheijen.
Material.— (?), 1862, Larantoeka (Semmelink, RMNH cat. no. 3); (?), xii.1888, Maumeri (Weber nr. 44G,
ZMA); (?), xii.1888, Sikka (Weber, RMNH cat. no. 10); , well-feathered nestling, 15.vi.1970, Wesang, 700m (Verheijen, RMNH no. 65278); , not dated, Flores (Verheijen, RMNH no. 81092).
Eggs: c/1, 5.v.1953, Wesang (RMNH no. 70548); c/1, 15.i.1958, Wesang (RMNH no. 70549); c/1, 18.
ii.1958, Wesang (RMNH no. 70550); c/1, 24.v.1958, Flores (RMNH no. 70551); c/1, 24.v.1958, Flores

88Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)(RMNH no. 70552); c/1 ca. 29.vi.1958, Léwé (RMNH no. 70553); c/1, 16.vii.1958, Flores (RMNH no.
70554); c/1, 30.x.1958, Déngé, Todo (RMNH no. 70555); c/1, c/1, 1961, Lamba, Todo (RMNH nos. 70556,
70557); c/1, 23.i.1961, Kisol (RMNH no. 70558); c/1, 1.v.1961, Kisol (RMNH no. 70559); c/1, 13.v.1964,
Kisol (RMNH no. 70560); c/1, 17.v.1964, Kisol (RMMH no. 70561). The eggs are white, slightly glossy.
Some measurements and weights:

RMNH no. 70553
RMNH no. 70554
RMNH no. 70555
RMNH no. 70557
RMNH no. 70558
RMNH no. 70559
RMNH no. 70560
RMNH no. 70561

46.3 × 32.6
45.5 × 33.2
47.8 × 32.9
46.9 × 32.5
52.4 × 32.9
47.0 × 33.8
49.4 × 33.8
44.8 × 32.7

1.46481.61901.71301.61851.85701.86651.79601.7004

Verheijen (1964: 198) mentioned that the nine clutches he listed under this species,
might include eggs of D. l. sasakensis. The eggs of D. sasakensis remain undescribed, but
I would expect them to be a trifle smaller than those of D. aenea. Chances are, that theeggs are correctly identified. Note that some eggs are from Wesang, where also a skinof D. aenea was obtained (and no D. sasakensis), and from Kisol, which seems too low
for D. sasakensis. White & Bruce (1986: 204) record as breeding season in the Lesser
Sunda Islands the months April to July. Verheijen’s eggs, however, prove breeding inJanuary, February, May, June, July, and October; the nestling listed above would have
hatched from an egg laid in March. These data suggest that nesting may take placethroughout the year.
Notes.— In using the name polia rather than aenea for these birds, I follow Stresemann (1952: 520) whose arguments look convincing to me. Actually, the name polia
had been resurrected earlier, by Delacour (1946, 1947a), following Mayr’s (1944a: 147148) argument that birds from Malaysia are different from those of the Lesser Sunda
Islands (the latter were at that time regarded as the nominate race). Subsequently,
Hoogerwerf (1963b) found a lot of variation in birds from diverse localities within therange here ascribed to polia. Perhaps his most surprising conclusion was that four
specimens from Komodo and Rintja were “so different in certain aspects”, that it
seemed “quite impossible to unite them with polia”. Hoogerwerf compared these specimens with the type of D. problematica Rensch from Sumba, and found them to agree
well. He further speculated that problematica could be an endemic subspecies of Komodo and Rintja, and that the type from Sumba could have been a straggler from Komodo/Rintja, as: “These birds are extremely good fliers which certainly are able tocover a distance of 100 km or more within a rather short time, so that the presence on
Sumba of birds originating from Komodo or Rintja need not be rejected”. Especially inthe light of the fact that Hoogerwerf placed specimens from West Flores in polia, andthat Rintja is separated from western Flores by a sea strait of scarcely 1 km in width (cf.
p. 6), the concept of an endemic subspecies on Komodo and Rintja is quite unbelievable. Hoogerwerf’s specimens from Komodo and Rintja and probably also the typespecimen of D. problematica had been preserved in formalin before being skinned. In
spite of Hoogerwerf’s awareness of the effects such conservation fluids, as well as
grease and de-greasing fluids, could have on the plumage, it is likely that the differences he described have to be attributed to such factors.
In the unlikely case that differences between birds from the western part of theMees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)89

range and birds from the Lesser Sunda Islands are confirmed, the name problematica
will be available for the latter.
The authors of the chapter Columbidae in del Hoyo et al. (1997: 229) have becomeso confused by the several shifts of the type-locality of the species, that they include thewhole range of polia in the range of the nominate race, for which they (correctly) givethe type locality as Manila!
The specimen measured does nothing to support Mayr’s contention that birds from
Flores are smaller than birds from Sumba, but does not convincingly contradict it either.
Measurements:
(?)

wing
246

tail
138

tarsus36.6

entire culmen
31

exposed culmen
24.6

Ducula rosacea (Temminck)
Columba rosacea Temminck, 1836, Recueil d’Ois. 4 (livr. 98): pl. 578 – Timor.
Muscadivores rosaceus zamydrus Oberholser, 1917, Proc. U.S. Nat. Mus. 54: 179 – Solombo Besar Island,Java Sea.
Collectors.— Semmelink, Allen, Schmutz.
Material.— (?), 1862, Larantoeka (Semmelink, RMNH cat. no. 4); (?), vii.1974, Pota (Schmutz, RMNH no.
81093).

Notes.— The subspecies zamydrus was listed by Chasen (1935: 18) with an asterisk,
meaning that he had not examined material, and therefore had not been able to verify
its validity. Peters (1937: 49) accepted it without comment, as did Deignan (1961: 109),
who referred for this to Chasen. Hoogerwerf (1963c) accepted zamydra, with a very unlikely distribution. Goodwin (1970: 407) mentions zamydra, but with added the remark“I have not seen specimens of this form”.
In the mean time, Mayr (1944a: 133) came with the definite statement, that: “Thedescription of Oberholser’s amydrus [sic] (Solombo Besar) indicates that it is based on
the characters of grease-stained specimens”. Subsequently van Bemmel (1948: 376)
gave this pigeon a binominal, implying that he did not recognize subspecies, whereas
White & Bruce (1986: 207) also rejected zamydra. It seems highly unlikely that this“Kleininselbewohner” would have developed into a distinctive subspecies on Solombo Besar and Arends.
D. rosacea is a widely-distributed species inhabiting, mainly, smaller islands, westward through the Java Sea to P. Nangka off Billiton (cf. Van Bemmel, 1940a, overlookedby Van Marle & Voous, 1988), and eastwards to the Tanimbar and Kai Islands (Van Bemmel, 1948: 376). In addition, there is a record from Telofoso on Halmahera, some 900 km
away from the nearest localities in the main range. This is so remote, as to require careful verification. Fortunately, this is possible. The record is based on material collectedby Bernstein, and in the archives of the RMNH I found an original list from Bernstein,
dated Ternate, 21 March 1862, entitled: “Lijst van de naturaliën....grootendeels afkomstig van Noordelijk Halmahera en Morotai”.5) In the list: “Carpophaga formosa ? van Tolofoso op N. Halmahera; Iris donker kersrood, randen der oogleden iets ligter”. 6) Espe-

5)
6)

List of natural history objects . . . . mainly originating from northern Halmahera and Morotai.
Carpophaga formosa ? from Tolofoso on N. Halmahera; Iris dark cherry-red, eye-rims a trifle lighter.

90

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

cially as in the preceding period Bernstein’s activities were confined to the North Moluccas, and as he never visited the better-known part of the range of D. rosacea, the record
is above any suspicion. Schlegel (1866b: 201) already discussed the peculiar distribution; he also stated that Bernstein had collected five specimens at Tolofoko (there issome variation in the spelling of the name of this locality), evidently in error. Schlegel(1873: 88) correctly lists four specimens (Schlegel cat. nos. 20-23), but now there are only
two left (30.i.1862, RMNH cat. nos. 23 and 24). In the exchange book, I found that on 6.
vi.1899 a  was sent to D. S. Drew of the Public Museum, Wanganui, New Zealand, andabout the same time another one (sex not recorded) to D’Arcy W. Thompson, University College, Dundee, Scotland. With this, all four specimens have been accounted for.
Specimens from “Makassar”, collected by Teysmann, received in 1878 (RMNH cat.
nos. 21, 22, 31), may safely be assumed to have been mislabelled, and to originate fromthe islands to the south of Sulawesi (Celebes), even though Wallace is also supposed tohave collected two specimens at Makassar, which were discussed by Stresemann (1941:59). The case is very similar to that of Macropygia magna macassariensis (cf. Mees, 1972).
Therefore, the inclusion of Saleyer into the range of the species by White & Bruce (1986:
207) was based on inference only (the specimens labelled Makassar), although it is very
likely to occur there. On the other hand, Tual or Klein Kei (Little-Kei) can be added to
the list of islands given by these authors (cf. Van Bemmel, 1948; Hoogerwerf, 1963c).
Ducula (lacernulata) sasakensis (Hartert)
Carpophaga sasakensis Hartert, 1896, Novit. Zool. 3: 564 – Lombok.
Collectors.— Everett (“a series”), Rensch (1), Verheijen/Schmutz.
Material.— (?), 14.ix.1968, Ruteng (Verheijen, RMNH no. 65277); , 18.ix.1969, forest between Paku
and Sok-Rutung (Schmutz, RMNH no. 66108); , late ix.1969, forest between Paku and Sok-Rutung(Schmutz, RMNH no. 85077). Iris dark brown, eye-rim and legs red-cabbage violet, bill grey with ablackish tip (Schmutz).

Notes.— Interestingly, this well-marked subspecies shares the light grey crown withnominate lacernulata from West Java; it is abruptly different from adjacent williami of
East Java and Bali, which has a pinkish crown. In Java, there is still a huge gap in the
known distribution of lacernulata and williami (cf. Mees, 1996: 23-24).
Measurements:
2
(?)

wing
215, 222
215

tail162, 148

tarsus30, 32
148

entire culmen
26, 2832

exposed culmen
20.5, 22
28

Columba vitiensis metallica Temminck
Columba metallica Temminck, 1835, Recueil d’Ois. 4 (livr. 95): pl. 562 – Timor.
Collector.— Schmutz.
Material.— , 21.vi.1972, Léma, 900 m (Schmutz, RMNH no. 85076). Iris brownish yellow, basal twothirds of the bill, and bare skin around the eyes blood-red, bill-tip yellowish white, legs violet-red. This
bird was collected in primary forest.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)91

Notes.— This species is apparently uncommon on Flores, the above specimen providing, in fact, the first record from the island.
Measurements:


wing
236

tail
126

tarsus27.5

entire culmen
27.7

exposed culmen
20

Macropygia unchall unchall (Wagler)C[olumba] Unchall Wagler, 1827, Syst. Av., Columba: 38 – Java.
Collectors.— Everett (1), Verheijen/Schmutz.
Material.— , 18.vii.1969, Ruteng (Verheijen, RMNH no. 65289); , 18.viii.1969, Wangkung, Rahong(Verheijen, RMNH no. 85130); , fledgling, 30.vi.1970, Ruteng/Kumba (Verheijen, RMNH no. 65293); ,
4.vi.1976, Poco Nernancang (Schmutz, RMNH no. 97109); , 29.vi.1978, Golo Rucuk (Schmutz, RMNH
no. 81094); , 1.v.1978, Golo Léhot (Schmutz, RMNH no. 81054). No eggs (see below); the fledgling
points to laying in May.

Notes.— Previously, only a single specimen of Macropygia unchall was known from
Flores: , October 1896, collected by Everett (cf. Hartert, 1904: 182 footnote). I cannot
find any difference between specimens from Flores and specimens from Java, the typelocality of the nominate race.
When Verheijen (1964) published his list, he was not aware of the occurrence of thisspecies on Flores. It is possible that some of the eggs identified as from M. emiliana, actually are from M. unchall, but I know of no characters by which to separate the eggs ofthe two species, although those of the latter may average a trifle larger (cf. Hellebrekers& Hoogerwerf, 1967: 45).
Measurements:
5

wing
170-176(172.8)tail
158-177(168.8)tarsus
28.3-29(28.7)

entire culmen
22.2-24(23.0)

exposed culmen
15.8-17(17.0)

Macropygia ruficeps orientalis Hartert
Macropygia ruficeps orientalis Hartert, 1896, Novit. Zool. 3: 573 – Tambora, Sambawa [sic].
Collectors.— Everett (“a series”), de Jong (1), Verheijen/Schmutz.
Material.— , 12.v.1971, Langkas, Rahong (Verheijen, RMNH no. 85310); , 2.vi.1976, Poco Nernancang(Schmutz, RMNH no. 85311); , 23.vii.1976, Poco Nernancang (Schmutz, RMNH no. 81053); , , 24.
viii.1976, Poco Nernancang (Schmutz, RMNH nos. 85313, 85309); , 4.ix.1976, Poco Nernancang(Schmutz, RMNH no. 85312).

Notes.— There is a clear sexual dimorphism in plumage, the females differing fromthe males in having black spots on the breast and black streaks on the darker crown. A
similar sexual difference is found in the nominate race, M. r. ruficeps from Java, but inthe subspecies from Borneo (nana) and Sumatra (sumatrana), peculiarly, both sexes haveblack spots on the breast.
According to White & Bruce (1986: 188): “The supposed larger size is not a goodcharacter”. In specimens of the nominate race from western Java I took the following
wing-measurements: 12  141-153 (147.1) mm, 12  135-146 (142.1) mm. This suggests

92

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

to me that the size-difference alone is quite enough for the recognition of orientalis. The
measurements also show a sexual difference in size in the nominate race, that is not apparent in the (admittedly quite insufficient) material of orientalis.
Measurements:
4
2

wing
155-163(159.3)160, 162tail
164-186(172)
166, 171

tarsus19-20(19.3)
19, 20.5

entire culmen
19-20.2(19.6)
20, 20

exposed culmen
13-16
(15)
14.8, 15

Macropygia emiliana emiliana BonaparteM[acropygia] emiliana Bonaparte, 1854, Compt. rend. Acad. Sci. Paris 39: 1111 (27 in reprint) – Java.
Collectors.— Everett (2), de Jong (1), Verheijen (1, MCZ), Verheijen/Schmutz.
Material.— (?) nestling, 4.x.1968, Ruteng, 1100 m (Verheijen, RMNH no. 65288); , 13.vi.1969, Nunang,
650 m (Schmutz, RMNH no. 85131); , 12.iii.1978, Nggoang, 800 m (Schmutz, RMNH no. 81057); (?) (=), 25.vi.1978, Flores (Schmutz, RMNH no. 81056); (), without data, but undoubtedly 1978, Manggarai, Flores (Schmutz, RMNH no. 81055).
Eggs: c/1, 10.v.1947, Bénténg Djawa (RMNH no. 70505); c/1, 2.viii.1947, Flores, without locality (RMNH
no. 70506); c/1, 8.vi.1954, Wesang (RMNH no. 70507); c/1, 17.v.1956, Méngé (RMNH no. 70508); 3 c/1,
v.1957, Tado (RMNH nos. 70509-70511); c/1, 1.v.1957, Tado (RMNH no. 70512); c/1, 19.ii.1958, Wesang(RMNH no. 70513); c/1, vi.1958, Tulang (RMNH no. 70514); c/1, 1.vi.1958, Tado (RMNH no. 70515); c/1,
16.x.1958, Waso Bénténg Djawa (RMNH no. 70516); c/1, 21.vi.1959, Poéng (RMNH no. 70517); c/1, 15.
v.1959, Mano (RMNH no. 70518); c/1, 18.vi.1960, Lai, Palué (RMNH no. 70519); c/1, 22.vi.1960, Palué(RMNH no. 70520); c/1, 27.vi.1960, Palué (RMNH no. 70521); c/1, 29.vi.1960, Palué (RMNH no. 70522);
c/1, 1.v.1960, Palué (RMNH no. 70523); c/1, 20.vi.1961, Mataloko (RMNH no. 70524). The eggs arewhite, almost without gloss, I cannot confirm the statement by Hellebrekers & Hoogerwerf (1967) thatthey are never pure white, but always: “creamy or buffy white or ivory”. To me, they seem just white.
Measurements and weights:

RMNH no. 70505
RMNH no. 70506
RMNH no. 70507
RMNH no. 70508
RMNH no. 70509
RMNH no. 70510
RMNH no. 70511
RMNH no. 70512
RMNH no. 70513
RMNH no. 70514
RMNH no. 70515
RMNH no. 70516
RMNH no. 70517
RMNH no. 70518
RMNH no. 70519
RMNH no. 70520
RMNH no. 70521
RMNH no. 70522
RMNH no. 70523
RMNH no. 70524

32.5 × 23.3
33.7 × 22.8
- × 24.7
31.2 × 22.8
34.9 × 24.7
33.0 × 23.0
33.6 × 22.7
33.1 × 24.0
32.7 × 22.1
32.9 × 23.8
32.6 × 21.8
36.7 × 21.8
33.5 × 23.9
31.7 × 23.1
33.0 × 24.0
34.8 × 22.5
31.4 × 22.5
34.1 × 23.2
35.1 × 23.8
31.2 × 22.4

0.578
0.55750.624
0.505
0.59370.521
0.54630.56070.43020.576
0.51440.567
0.620
0.45670.612
0.5795
damaged
0.608
0.617
0.465

Verheijen (1964) listed these 20 clutches all under the name of Macropygia phasianella
emiliana, but at that time it was not generally known that, besides M. emiliana, M. unchallMees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

93

occurs on Flores. It is likely that the collectors of these eggs identified their material togenus only. In addition, M. ruficeps being of fairly common occurrence on Flores, one
might expect some of the eggs to belong to that species. In all three species, a single egg
seems to constitute the complete clutch. The eggs of M. emiliana and M. unchall are probably not separable, but the eggs of M. ruficeps should be distinguishable from the former
two on the basis of smaller size and, particularly, lower weight (cf. Hellebrekers &Hoogerwerf, 1967: 44-45). A snag is that the subspecies M. r. orientalis is distinctly larger
than nominate ruficeps from Java, and might have correspondingly larger and heavier
eggs. Anyway, there was ample reason to measure and weigh all the eggs.
A useful fact is that five of the eggs are from the island of Palué, where M. unchalland M. ruficeps may safely be assumed not to occur, so that the identification of theseeggs ought to be certain. The slight reservation I feel is not because I consider it likelythat Macropygia is represented on Palué by more than one species, but because of a veryslight possibility that this species is not M. emiliana but M. magna: Verheijen (1961) heard
and saw birds on Palué, but did not collect any. For comparison, here follow the weights
of eggs from Java as recorded by Hellebrekers & Hoogerwerf. M. r. ruficeps: 0.300-0.380;
M. e. emiliana: 0.45-0.57; M. u. unchall: 0.465-0.68. It is at once clear that, compared withweights from Flores, these figures are not of much help. The series from Flores suggeststhat eggs with a weight of less than 0.5 g could be M. r. orientalis, as they are well below
weights of the other eggs, but the minimum recorded in Java for M. emiliana is 0.45 g, sothat only one of the Flores eggs is below it (with 0.43). From the Java weights it mightlook as if eggs of over 0.6 g are M. unchall, but then three of the four eggs that could beweighed from Palué, definitely M. emiliana, are over 0.6 g. It looks as if the majority ofthe eggs, perhaps all of them, have been correctly identified as being of M. emiliana (assuming, of course, that the genus has been identified correctly).
Notes.— It is unexpected that none of the early collectors has obtained M. emiliana.
Hartert (1898a: 49) listed two specimens ( ad.,  im.) collected by Everett. A few years
later, he spoke of “several” specimens which had been sent by Everett (cf. Hartert, 1904:
182 footnote).
Whatever its merits on a morphological base, the division of the Macropygia amboinensis group into six subgroups, of which four are given species status, and two, theemiliana and the tenuirostris subgroups, inhabiting the Sunda region and the Philippines
respectively, are placed in subspecific relationship to M. phasianella from eastern Australia, has something artificial, as the three subgroups thus united under the specificname of phasianella, are geographically separated by the subgroups magna and amboinensis, both regarded as separate species.
The absurdity of this classification was first pointed out by Mayr (1944a: 148-149,
191), who, as a consequence, reduced the members of the subgroups magna and rufipennis to subspecies of phasianella, but hesitated to further unite this conglomerate withamboinensis, which he considered to belong: “at least to the same superspecies”. Mayr
concluded with the suggestion that: “This whole group of forms would make a favorable subject for a study in character geography and speciation”.
Now, over half a century later, Mayr’s suggestion has not yet been taken up, and in
spite of his very sensible comment, the illogical classification has been maintained, forexample by Goodwin (1970: 152-153) and Dickinson et al. (1991: 192-193). Recently,
however, White & Bruce (1986: 189) and Sibley & Monroe (1990: 198) have taken the94

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

obvious step of treating each of Mayr’s six subgroups as a separate species, and I follow
them, with in mind the additional argument that the co-existence of three rather similar
species of Macropygia on the comparatively small island of Flores, should be a warning
against oversimplification. The opposite view, to unite all subgroups, has been taken
mainly by Australian authors (Storr, 1973: 49 and 1984: 66; Condon, 1975: 166; Frith,
1976: 234; Blakers et al., 1984: 224) and is equally well defensible. It should be noted,
however, that in a later work, Frith (1982: 143-144) has changed his opinion and treated
the Australian M. phasianella as a separate species again, predicting that eventually theconglomerate may be found to consist of two species: M. amboinensis, M. phasianella andM. magna making one, and M. tenuirostris with M. emiliana and M. rufipennis the other.
Streptopelia bitorquata bitorquata (Temminck)
Columba Bitorquata Temminck, 1809, Hist. Nat. Pigeons, Les Colombes: 86, pl. XL – elle habite l’Inde,mais nous ignorons dans quelle île.
Collectors.— Allen (at least 5, cf. Salvadori, 1893: 422), Weber, Verheijen.
Material.— (?), xi.1888, Maumeri (Weber, RMNH cat. no. 8); , (?)juv., 23.ix.1970, Maro-Kama (Verheijen,
RMNH nos. 65280, 65279). Iris of adult golden yellow, of juvenile brownish cream colour; bill blackish
brown, legs dark purplish red.
No eggs. Verheijen (1964: 198) recorded with a query one case of breeding in May.

Notes.— Judging by the fact that only three collectors have obtained it, this doveseems to be rather local on Flores, where it is probably confined to the coastal regions,
as it is elsewhere. Schmutz (MS) saw it at Nunang (650 m), and more commonly near
the west coast, from sea-level to 400 m.
Streptopelia chinensis tigrina (Temminck)
Columba tigrina Temminck, 1809, Hist. Nat. Pigeons, Colombes: 94, pl. 43 – Timor; Batavia.
Collectors.— Semmelink, Allen (at least 5, cf. Salvadori, 1893: 444), Weber (1, ZMA), Everett, Rensch (1),de Jong (2), Verheijen (2, MCZ), Verheijen/Schmutz.
Material.— (?), 1862, Larantoeka (Semmelink, RMNH cat. no. 13); , 17.vi.1968, Ruteng (Verheijen,
RMNH no. 65256): , 30.i.1970, Ruteng (Verheijen, RMNH no. 65272); , 19.ix.1970, Kisol (Verheijen,
RMNH no. 65273); , 22.ix.1970, Maro-Kama (Verheijen, RMNH no. 65274); 2 , 23.ix.1970, Maro-Kama(Verheijen, RMNH nos. 65275, 65276); , 9.iii.1971, Maro-Kama (Verheijen, RMNH no. 66100); ,
30.vi.1971, Langkas, Rahong (Verheijen, RMNH no. 66101); , 2.v.1971, Langkas (Verheijen, RMNH no.
66102);  juv., 24.v.1971, Langkas (Verheijen, RMNH no. 66103); , 25.v.1971, Langkas (Verheijen,
RMNH no. 66104); 2 , 28.vi.1971, Langkas (Verheijen, RMNH nos. 66105, 66106); , 30.vi.1971, Langkas (Verheijen, RMNH no. 66107).
Eggs: 64 clutches of c/1 (39 ×) and c/2 (25 ×), collected in the months March (4), April (23), May (14),June (1), July (6), August (4), September (3), October (2), December (1), and six undated (RMNH nos.
70433-70496). The eggs are white, slightly glossy.
Some measurements and weights:

RMNH no. 70453
RMNH no. 70454
RMNH no. 70474
RMNH no. 70489

28.0 × 20.8
31.9 × 21.8
28.2 × 22.0
30.0 × 22.0
31.9 × 22.4

0.37350.47940.44470.468
0.4603

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)
RMNH no. 70490
RMNH no. 70491
RMNH no. 70492
RMNH no. 70493

95
29.7 × 22.6
29.1 × 21.6
28.9 × 21.8
30.7 × 21.1
31.0 × 21.3

0.51910.43830.41510.451
0.4511

Geopelia striata maugei (Temminck)
Columba Maugeus Temminck, 1809, Hist. Nat. Pigeons, Les Colombes: 115 – les îles de l’Australe-Asie.
Columba maugens Temminck, 1809, Hist. Nat. Pigeons, Les Colombes: pl. LII – no locality.
C[olumba] maugei Temminck, 1811, Hist. Nat. Pigeons, Index: xiv – correction of misprint (cf. Mees,
1975b: 126-127).
Collectors.— Semmelink, Allen, Martens, Everett, Rensch (2), de Jong (1), Verheijen/Schmutz.
Material.— (?), 1862, Larantoeka (Semmelink, RMNH cat. no. 2); 2 , 16.ix.1970, Maro-Kama (Verheijen,
RMNH nos. 65299, 65300);(?), 23.ix.1970, Maro-Kama (Verheijen, RMNH no. 65301); , 30.ii.1971, NisarBadjo, 50 m (Schmutz, RMNH no. 66110); , 9.iii.1971, Maro-Kama, Kisol (Verheijen, RMNH no.
66109).
Eggs: c/2, 23.vi.1947, Rekas (RMNH no. 70500); c/2, Dampék, 21.vi.1959 (RMNH no. 70501); c/2, 1.
v.1959, Dampék (RMNH no. 70502); c/1, 1960, Rekas (RMNH no. 70503); c/2, 3.v.1960, Téo (RMNH no.
70504), and three undated clutches (RMNH nos. 70497-70499). The eggs are white, smooth but without
gloss.
Some measurements and weights:

RMNH no. 70500
RMNH no. 70501
RMNH no. 70502
RMNH no. 70504

21.8 × 17.1
22.7 × 17.3
21.5 × 17.0
22.9 × 17.1
22.5 × 17.0
24.4 × 17.3
21.6 × 17.8
22.0 × 17.8

0.21350.20720.1916
0.18910.23
0.20660.21720.2142Trichoglossus (haematodus) weberi (Büttikofer)
Psitteuteles weberi Büttikofer, 1894, in Weber, Zool. Ergebnisse 3: 290, pl. XVII fig. 1 – Flores.
Collectors.— Weber, Everett (“a fine series”), Endih (1), Rensch (4), de Jong (4), Verheijen (1, MCZ),
Verheijen/Schmutz.
Material.— 2 (?), 23/25.xi.1888, Reo (Weber, RMNH cat. nos. 1, 2, syntypes); (?) juv., 26/28.xi.1888, Bari
(Weber, RMNH cat. no. 4, syntype); , i.1889, Endeh (Weber, RMNH cat. no. 3, syntype); 15 , 17 , 6(?), 1969/1976, from Rahong, Nunang, Ulu Ros, Larang, Langkas (Rahong), Nggolong-Tédé (Verheijen/
Schmutz, RMNH, cat. nos. 5-42). Iris red, bill red, legs brownish grey.
Eggs (fig. 13f): c/2, 25.viii.1960, Mataloko (RMNH no. 70577); c/2, 20.vi.1961, Mataloko (RMNH no.
70578); c/2, 25.v.1962, Mataloko (RMNH no. 70579). The eggs are white.
Measurements and weights:

RMNH no. 70577
RMNH no. 70578
RMNH no. 70579

26.3 × 21.6
26.7 × 21.0
28.1 × 22.1
27.0 × 21.7
26.4 × 22.5
26.5 × 22.0

0.45380.42470.45280.44560.41050.4202

96

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Notes.— The nomenclatural history of this distinctive endemic form is a chequered
one. It was introduced in ornithology by Büttikofer, as a geographical representative of
T. euteles (Büttikofer described it as a species, but he did not yet use ternary nomenclature). Mivart (1896: 129-130), the first to comment, was misguided by four specimensof T. euteles mislabelled as being from Flores (BM, leg. Allen), into believing that T.
weberi was a synonym of T. euteles. The matter was put right by Hartert (1898a), who
re-instated T. weberi as an “excellent species”. Mathews (1927: 296) listed it, under thegeneric name Eutelipsitta thought up by himself, as a species, E. weberi. The specificstatus was apparently given because he believed that weberi and euteles both occurred
on Flores. Three years later he corrected this: “Read Eutelipsitta euteles weberi (Büttikofer), and delete Flores from the distribution of E. e. euteles” (Mathews, 1930: 914).
The elimination of Flores from the range of T. e. euteles, enabled Mathews to treat weberi
as a subspecies.
Up to that time, the relationships, as a species or a subspecies, had always beensought with T. euteles. Rensch (1931a: 527) came with a radically different proposal: “Ichfasse “Psitteuteles” weberi als stark specialisierte Rasse von Tr. ornatus auf, da alle Zeichnungsmerkmale dieses Rassenkreises vorhanden sind (gelber Halsring, Blau am Kopfeetc.) und nur die Lipochromfärbung der Unterseite weniger intensiv ist”. Rensch’s T.
ornatus evidently included T. haematodus, but these were separated by Peters (1937: 148),
in whose classification T. ornatus became a monotypic species, and all other forms, including weberi, went with haematodus. A year earlier, Stresemann (1936: 363) had listed
T. ornatus as a separate species, and later he supported this opinion with the following
argument: “T. ornatus steht der weitverbreiteten Art T. haematodus ziemlich nahe, istaber so stark von dieser unterschieden, dass er als besondere Art geführt werden muss”(Stresemann, 1940: 435).
The same classification was accepted by subsequent revisers, such as Cain (1955)
and Forshaw (1973: 58-59). Cain (1955: 443) placed, within the species T. haematodus,
weberi in the “capistratus group”, stating: “weberi is a small greened flavotectus”. Cain’s
evaluation of weberi is difficult to contradict, but does hardly do justice to its aberrant
character in comparison with the surrounding subspecies. Considering the very close
relationship usually existing between birds from Flores and Sumbawa, the striking differences between weberi and T. h. forsteni need an explanation.
Rensch (1931b: 395) discusses a  from Wai Sano, collected by de Jong, but strangely, in the preceding enumeration of material no specimen from that locality is listed.
Measurements:
15 
16 

wing
124-132(127.8)123-131(127.1)tail
82-102(93.2)
82-109(93.5)

culmen from cere18-20.1(19.1)
17-19.2(18.4)

Cacatua sulphurea occidentalis Hartert
Cacatua parvula occidentalis Hartert, 1898, Novit. Zool. 5: 120 – Lombok.
Collectors.— Semmelink, Allen, Martens, Weber, Everett, Endih (2), Rensch (2), de Jong (2), Geeraeds,
Verheijen/Schmutz.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

97

Material.— (?), 1862, Larantoeka (Semmelink, RMNH cat. no. 4); 3 (?), 1862, Flores (Allen, RMNH cat.
nos. 1-3); (?), 23/25.xi.1888, Reo (Weber, RMNH cat. no. 5); (?), 17.ix.1968, Nunang (J. Geeraeds, RMNH
no. 66136); , 11.v.1969, Waé-Wako (Schmutz, RMNH no. 66137).
Eggs (fig. 13g, 13gg, 13ggg): c/1, 23.vi.1955, Flores (RMNH no. 70573); c/2, 28.vi.1955, W. Rembong(RMNH no. 70574); c/1, 20.xi.1960, Soa (RMNH no. 70575); c/1, 26.ii.1961, Kisol (RMNH no. 70576); c/1,
not dated, Soa (RMNH no. 70580). The eggs are elongate, dull white, without any gloss, their surface
slightly rough.
Measurements and weights:

RMNH no. 70573
RMNH no. 70574
RMNH no. 70575
RMNH no. 70576

35.8 × 27.2
43.5 × 29.0
44.5 × 29.2
42.0 × 27.1
39.9 × 28.6

1.31231.510
1.6332
1.3496
1.3719Notes.— Of the subspecies parvula, only the type (, 1829, Samao, leg. S. Müller,
RMNH cat. no. 1) was available to me for comparison. It certainly has a smaller (especially a more slender) bill than all specimens from Flores, and therefore I would tentatively retain occidentalis, which was synonymized with parvula by White & Bruce (1986:
211). Forshaw (1973: 122) also recognized it.
Measurements:

wing

tail

Flores (C. s. occidentalis)

207
107½
5 (?)
209-223
99-106
Samao off Timor (type of C. s. parvula)

216
114

tarsusculmenfrom cereculmendepthculmenwidth

18.6
19.4-20.5

30.4
31.2-38.3

20.4
19.8-24

19.5
19-21

19.3

30

19.6

16

Tanygnathus megalorynchos floris Hartert
Tanygnathus megalorhynchos floris Hartert, 1924, Novit. Zool. 31: 126 – South Flores (Mangarai).
Collectors.— Everett (5), Verheijen (1, MCZ), Verheijen/Schmutz.
Material.— (?), 1.i.1969, central Manggarai (Schmutz, RMNH no. 66133); , 11.xi.1970, Réo, near thecoast (Verheijen, RMNH no. 66134); , 17.vii.1971, Look-Mabacone, mangrove (Schmutz, RMNH no.
66135). Iris grey, bill red with a whitish tip, cere greyish, eye-rim above the eye grey-yellow, below it
grey, legs olive grey.

Notes.— Schmutz (MS) reports observations from the coastal region near Mburak(50 m), where at Sokrutung many hundreds of Tanygnathus and Cacatua were seen as
they came to sleep communally in some large trees along the river. Inland they wereseen at Kulan (430 m), also a swampy place, between Paku (350 m) and Kandang (600m), and Nggoang (900 m). According to locals, they came in that season (about the endof February) to Nggoang to harvest the nuts of Canarium asperum. The five specimens
collected by Everett were all females; so were two of the three birds collected by thefathers, the sex of the third one being uncertain. It seems that the male sex of T. m. floris
remains unknown. Note that Forshaw (1973: 188) examined 8 specimens, all females:
the total material of the subspecies known.
Forshaw (l.c.) recognized eight subspecies (nominate megalorynchos, affinis, subaffinis,

98

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

hellmayri, viridipennis, djampeae, floris and sumbensis). White & Bruce (1986: 227-228) reduced this to five, by placing viridipennis, djampeae and floris in the synonymy of nominate megalorynchos, on the basis of the argument that the characters of viridipennis anddjampeae were due to “an irregular small island effect unworthy of formal designation”,
and floris was based on: “a clinal effect leading to sumbensis”. Lacking adequate comparative material to form an independent opinion, I prefer to follow Forshaw’s morecautious approach.
Measurements:


(?)

wing

tail

tarsus

229
220
235

146
137
150

21.4
21
23

culmenfrom cere454246

depth ofculmen at base
23
23
27

width ofculmen at base
24.2
23.6
26.8

Loriculus flosculus Wallace
Loriculus flosculus Wallace, 1864, Proc. Zool. Soc. Lond. (1863): 488 – Flores.
Collector.— Allen.
Material.— (?), 1862, Flores (Allen, BM no. 73.5.12.1555, holotype).

Notes.— This species is known from a single unsexed specimen only, collected byAllen, hence without an exact date and locality. Finsch (1868: 728) regarded the bird
as immature, as the red throat patch appeared to be not fully developed. Reichenow(1882: 114) was apparently the first to add information that did not exist; he wrote“iride miniatis” and assumed that the type was a male. Sometimes Allen recorded thecolour of the eyes on the labels of specimens he collected, but there is no such information on the label of the holotype of L. flosculus. Enigmatically, Forshaw (1973: 320)
describes adult males as well as females; he also describes the iris colour: orange inmales, brown in females, with a reference to Wallace. I have been unable to find thisreference and as Wallace never had more than this one unsexed individual, it cannot
be correct.
During the last twenty years, there have been a number of sight records, as summarized by Butchart et al. (1996: 344-346); the cumulative evidence is sufficient to accept that the species still exists.
The eggs described by Forshaw, ex Schönwetter (1964: 526) cannot have been correctly identified. One might think of confusion with L. exilis of Sulawesi (Celebes),
which has been considered a subspecies of flosculus by Rensch (1931a: 525), but themeasurements seem too large for that dwarf of the genus, and moreover it inhabits
Sulawesi (Celebes), not Flores. Rensch has not been followed, and all later authors, likeStresemann (1936: 363; 1940a: 444, 445) and Peters (1937: 259) have treated L. exilis as aseparate species. Stresemann considered the presumed relationship between L. flosculus
and L. exilis at best remote.
Measurements:
(?)

wing
80

tail
34

tarsus
10.4

culmen from skull
16.4

culmen from cere
10.5

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

99

Geoffroyus geoffroyi floresianus Salvadori
Geoffroyus floresianus Salvadori, 1891, Cat. Birds Brit. Mus. 20: 406 – Flores.
Collectors.— Semmelink, Allen, Everett, v.d. Sande (1), Rensch (8), de Jong (2), Verheijen (5, MCZ), Verheijen/Schmutz.
Material.— , , (?) juv., 1862, Larantoeka (Semmelink, RMNH cat. nos. 5-7); , 1862, Flores (Allen,
RMNH cat. no. 1); , 9.xii.1968, Pongkor (Verheijen, RMNH no. 59809);  juv., 13.ix.1971, Tjeréng, 850
m (Schmutz, RMNH no. 85281); , 22.ix.1971, Kandang, 900 m (Schmutz, RMNH no. 85282); , 24.
ix.1971, Nisar (Schmutz, RMNH no. 97117); , 16.x.1971, Tjeréng, 750 m (Schmutz, RMNH no. 85283);
, 25.v.1974, Nunang, 1000 m (Schmutz, RMNH no. 81064); , 25.vi.1975, Paku, 300 m (Schmutz, RMNH
no. 81068); , 28.vi.1978, Golo Rucuk (Schmutz, RMNH no. 81067); , 9.x.1982, Cereng, 600 m (Schmutz,
RMNH no. 97118). Iris sulphur yellow.
Eggs: c/3, 14.vi.1954, Wesang (RMNH no. 70565); c/2, 14.v.1955, Mano (RMNH no. 70566); c/3, Mano(RMNH no. 70567); c/1, 10.vi.1957, Montjok (RMNH no. 70568); c/3, c/2, 7.v.1957, Méngé (RMNH nos.
70569, 70570); c/2, 3.vii.1957, Flores, without locality (RMNH no. 70571); c/2, 12.vii.1959, Léwé (RMNH
no. 70572).
Measurements and weights:

RMNH no. 70565

RMNH no. 70566
RMNH no. 70567

RMNH no. 70568
RMNH no. 70569

RMNH no. 70570
RMNH no. 70571
RMNH no. 70572

30.3 × 25.4
31.1 × 25.5
31.9 × 25.7
29.8 × 25.6
30.2 × 25.4
28.5 × 24.5
29.8 × 26.5
29.9 × 25.6
34.2 × 26.3
30.8 × 26.0
31.5 × 25.8
31.6 × 26.3
29.8 × 25.4
30.4 × 26.3
30.5 × 25.6
30.9 × 25.5
29.0 × 25.0
29.1 × 25.4

0.6595–0.65860.6591
0.6386
0.58530.72690.67170.737
0.66150.74370.73170.71650.688+
0.64550.694
0.59820.6525

damagedidentification correct?

very large blowhole

Notes.— These birds are rather smaller in wing and tail than the specimens measured by Forshaw (1973: 171).
Measurements:







 juv.

wing
156
154
153
160
163
149
152
152

tail
70
71
66
77
71
69
74
70

tarsus
20
17
16.8
19½
18
17½

culmen from skull
225
26.5
25
26
26
25
25.5
25

culmen from cere
21.7
22
22
22.5
21.8
20
21.3
20.5

100

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Cuculus saturatus horsfieldi Moore
Cuculus horsfieldi Moore, 1858, in Horsfield & Moore, Cat. Birds Mus. Hon. East-India Comp. II: 703 –Java.
Collectors.— Allen, Everett, de Jong (3), Schmutz.
Material.—  im., 29.i.1976, Joneng, dunes (Schmutz, RMNH no. 97107); , 12.xi.1977, Cereng, 400 m(Schmutz, RMNH no. 97106);  im., 12.ii.1982, Nunang, 650 m (Schmutz, RMNH no. 81059). Iris ochre,eye-rim yellow.

Notes. — Their measurements place these specimens definitely in the subspecies
horsfieldi (cf. Junge, 1937a). Payne (in del Hoye et al., 1997: 512, 555) has made the interesting suggestion that horsfieldi is not a subspecies of C. saturatus, but a separate species.
The evidence provided is somewhat sketchy and I look forward to a more comprehensive documentation.
Measurements:

 im.


wing
207
202
200

tail
147
144
127

entire culmen
28
27
28

exposed culmen
21.2
20
21.4

Cuculus saturatus lepidus S. Müller
Cuculus lepidus S. Müller, 1845, Verh. Nat. Gesch. Ned. Overz. Bez., Land- en Volkenk.: 236 – Timor.
Collectors.— Everett (6), Rensch (3).
Material.— None.

Notes.— Everett’s men collected half a dozen specimens towards the end of 1896,
and Rensch obtained three in June 1927. Although Rensch (1931a: 542) reported that hefound it “vereinzelt”, it is still strange that it was not obtained by the Fathers, neither
did Verheijen (1964) acquire its eggs.
C. s. lepidus is a mountain bird, not usually occurring below ca. 800-1000 m, although Rensch (1931a: 542) found it on Bali as low as 500-600 m. In the mountains of theLesser Sunda islands it is widely distributed. White & Bruce (1986: 231) list it from Lombok, Sumbawa, Flores, Pantar, Sumba and Timor. Also Bali, as just mentioned.
The Sumba record is, however, erroneous. The species was first listed by Rensch(1931b: 389), in an enumeration of the birds of Sumba, and casually mentioned by
Rensch (1931a: 542). I cannot find on what the Sumba record by Rensch, Peters (1940:20), and White & Bruce is based. The species was not recorded by A.B. Meyer, not mentioned by Hartert (collections of Doherty and Everett), not collected by Dammerman,
Stein or Sutter. The error probably originated from the confusion there has been overthe names of the Cuculus-species in the Indo-Australian region. Until recently, the name
C. poliocephalus lepidus was used for the form now called C. saturatus lepidus. For a while,
C. saturatus was known by the name C. intermedius. Note that Rensch (1931a: 541) gives
under C. p. lepidus, mistakenly, a reference to C. intermedius of Hartert (1898a: 45), whichis, however, C. saturatus horsfieldi. Rensch seems to have thought that C. intermedius ofHartert applied to C. s. lepidus, and has overlooked that C. saturatus lepidus has in thesame article the name C. poliocephalus.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

101

The fact that the record from Sumba is erroneous, does not necessarily mean that
therefore C. s. lepidus does not occur on the island. Pantar, from where it is reliably
known, is not much higher than Sumba (1365 m against 1225 m). Another factor influencing the distribution may be the availability of host species. In Java Phylloscopus trivirgatus and Seicercus grammiceps have been recorded as hosts. Related species (Phylloscopus presbytes, Seicercus montis) occur on Flores and Timor, but no resident Phylloscopi
are known from Sumba.
Cuculus pallidus (Latham)C[olumba] pallida Latham, 1801, Suppl. Ind. Orn.: lx – Nova Hollandia = New South Wales.
Cuculus poliogaster S. Müller, 1845, Verh. Nat. Gesch. Ned. Overz. Bez., Land- en Volkenk.: 236 noot 4 –Ternate.
Collector.— Schmutz.
Material.— , 26.x.1969, Nanga-Lili (Schmutz, RMNH no. 65302).

Notes.— Individuals of this species found outside Australia may be assumed to be
migrant visitors. Besides Flores, it has in the Lesser Sunda Islands been recorded from
Timor (McKean, Mason & O’Connor, 1975). I do not think that the occurrence on Timor
as late in spring as the end of October, justifies the assumption that it reproduces there(although I do not deny the possibility). Several other migrants from Australia may stayin their winter quarters until October and even November, for example Stiltia isabella,
Halcyon sancta, Myiagra r. rubecula (cf. Mees, 1982a). The Flores specimen is equally late.
C. pallidus is peculiarly scarce outside Australia: besides those from the Lesser Sunda
Islands, there is one record from Babar (cf. Hartert, 1906: 295, s.n. Cuculus variegatus),
one from Ternate (the type specimen of Cuculus poliogaster, collected by Forsten), andonly three from New Guinea. Evidently few birds migrate outside Australia.
Cacomantis variolosus sepulcralis (S. Müller)
C[uculus] sepulcralis S. Müller, 1843, Verh. Nat. Gesch. Ned. Overz. Bez., Land- en Volkenk.: 177 – Java en
Sumatra.
Collectors.— De Jong (1), Verheijen/Schmutz.
Material.— , 7/8.vi.1968, Ruteng (Verheijen, RMNH no. 59807); , 22.vi.1971, Langkas, Rahong (Verheijen, RMNH no. 85259); (?) pull., 17.ix.1976, Waé Lega (Schmutz, RMNH no. 81305); , 29.vi.1978,
Golo Léhot (Schmutz, RMNH no. 81319).
Eggs in the collection Ottow. Verheijen (1964) recorded eggs of this cuckoo in the months May (1), July(2), August (3), September (5) and October (2). According to Ottow & Verheijen (1969), Saxicola caprata is
the favourite host on Flores: eleven of the thirteen records concerned this species (the host-species of thetwo remaining nests could not be identified, it was probably a warbler).

Notes.— The species C. variolosus can conveniently be divided into two sub-groups:
the brighter, darker “sepulcralis” group, ranging from the mainland of south-eastern
Asia, to the Philippines, the Greater Sunda Islands, and the Lesser Sunda Islands east to
Flores and Sumba, and the “variolosus” group, duller in plumage, found in northernand eastern Australia, New Guinea, the Moluccas and Timor.

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On Timor, the species was discovered in 1829 by Salomon Müller, who recognized
its distinctiveness and described the one specimen obtained as C. tymbonomus. This remained generally recognized and with the introduction of ternary nomenclature, became Cacomantis variolosus tymbonomus.
Confusion was, unintentionally, started by Junge (1937b: 179-180), who comparedspecimens of C. variolosus from New Guinea (migrants from Australia, although this is
not stated clearly), with material from northern Australia (dumetorum) and from Timor(tymbonomus). He considered these all identical and as tymbonomus is an older namethan dumetorum, he concluded that northern Australia should be included into therange of tymbonomus. The specimens from northern Australia available to Junge were:one , 31.vii.1840, Pig Lagoon, Port Essington (RMNH cat. no. 1, name of collector notgiven, but almost certainly J. Gilbert), and a bird of unknown sex from Cape York, without date or collector’s name, purchased from Frank in 1870. Junge’s text: “the specimens of tymbonomus, which are in the Leiden museum” is due to his inclusion of specimens from Australia and New Guinea under that name, for there is not now, and there
has never been, more than the one specimen from Timor in the collection. That the
specimen is a holotype was already made clear by its describer: “C[uculus] tymbonomus,
n. sp. Het , hetwelk wij alleen van deze soort bezitten” 7) (Müller, 1843: 177). In judging Junge’s work, it should be kept in mind, how very little he had to go on. His material will be further discussed below.
Peters (1940: 25-26), informed by Mack that dumetorum cannot be separated fromnominate variolosus, and by Junge that tymbonomus is an earlier name for dumetorum,
naturally placed both names into the synomymy of nominate C. v. variolosus. Probably
through an oversight he omitted to mention Timor in the description of the range of thisexpanded subspecies, so that it remains unclear whether he would have regarded it asa resident or as a migrant visitor from Australia, although the latter is likely.
Mayr (1944a: 149-150), on the basis of Stein’s material from Timor, disagreed with
Junge: “It has been assumed by recent writers, for some curious reason, that no resident
race of this cuckoo occurs on Timor Island and that the name tymbonomus was a synonym of variolosus or at best an earlier name for dumetorum from western Australia.
Stein’s specimens prove that there is an endemic race of this species on Timor, as might
be expected from the general distribution of the species, and that the name tymbonomus
is, therefore, not applicable to western Australian birds”.
Mayr proceeds to describe the subspecific differences. His criticism of “recent writers”, just quoted, is somewhat overdone, for Junge never mentioned whether he regarded C. variolosus on Timor as a resident or as a migrant visitor: the fact that he wasunable to distinguish the type of tymbonomus from Australian birds does not automatically mean that he considered it a migrant. Peters did not give an opinion either.
This is as matters stood, when White got on the trail. He asked me to examine thetype specimen of tymbonomus, with the thought in mind that it might be a migrant from
Australia. From my reply (dated 18.xi.1976), I quote: “The type of tymbonomus is our
only specimen from Timor. It belongs to the pale Australian group, and allowing for 140
years difference in age, it does not look different from two specimens collected by me

7)

C[uculus] tymbonomus, n. sp. The single male we possess of this species

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

103

in northern Australia. The type of tymbonomus is plain above, without the rufous edges
to the feathers which Mayr claimed are characteristic of this race. I would suspect all
birds with rufous edges to be immature. I regret that we have no juvenile (‘immature’ ofMayr) birds, which according to Mayr would be ‘even more different’. One old specimen
from Timor does not make much of a series. In my opinion the type (merely labelled
1829, without date), might conceivably be a migrant”. White (1977) presented my reply
correctly, but added a concluding sentence for which I would certainly not have wanted
to share responsibility: “If there is a distinct resident form in Timor, it is evident thattymbonomus is not on present evidence available as its name”. Next we find: “Müller’s
tymbonomus has been confirmed as a migrant of the nominate form (G.F. Mees)” (White& Bruce, 1986: 236). Thus, my cautious words: “might conceivably be”, were, in two
steps, changed to “confirmed”, and on this basis, Bruce (l.c.) felt justified to provide thejuvenile birds from Timor, described by Mayr, with a new subspecific name, C. v. whitei.
It does not look as if Bruce had personally examined any of this material.
In the meantime, the type of tymbonomus had been sent to Dr Schodde, who informed me (in litt., 2.ii.1979) as follows: “We have now compared it with a selection of
Stein’s material from Timor and found they all belong to one and the same subspecies... tymbonomus may look like variolosus from northern Australia but retains juvenile barring on the under-tail coverts and a rufous wash on the rectrices longer into adulthoodand has a proportionally much longer tail”.
The two Australian birds examined by Junge are both in their first juvenile plumage: the Port Essington specimen is heavily barred, both above and below, the CapeYork one is also heavily barred below, but the upper surface is more spotted than barred.
It will be clear that these two specimens could not possibly form a basis for a meaningful comparison with one adult (“nearly adult” according to Junge) specimen from
Timor. The conclusion that the Timor bird did “fully agree” with specimens from North
Australia is therefore enigmatic, as is his remark about birds received from the Buitenzorg Museum, which in its context suggests that these were Australian birds. Actually,
they are birds from the Mamberamo expedition (leg. W.C. van Heurn), previously recorded by Hartert (1932: 453), which Hartert assumed (correctly, no doubt) were migrants from Australia. In addition, Junge had two specimens from southern New Guinea, one from Andai, and one from Misool. Mr White, in common with a majority of his
fellow-men, sometimes found it difficult to abandon a pet theory. He had another one
about C. variolosus: “I wonder whether C. variolosus and C. sepulcralis are in fact conspecific. Might not C. heinrichi, which I have not seen, but which was stated to have abarred juvenile plumage (unlike C. castaneiventris), possibly represent C. sepulcralis.
Stresemann’s description of C. heinrichi did not consider this possibility” (White, in litt.,
10.xii.1976). My reply (dated 21.xii.1976) read: “As C. heinrichi is not represented in ourcollection I am unable to speculate about its relationships. As regards your suggestion
it is relevant to state, however, that I know the songs of C. variolosus sepulcralis (Java), C.
v. chivae (Biak), C. v. variolosus (New South Wales) and C. v. dumetorum (Northern Australia), and they all sound pretty much the same to me, whereas according to Heinrich(in Stresemann, 1931) the song of C. heinrichi is ‘ganz anders’”. Later, without reference
to Heinrich’s observations, White (in White & Bruce, 1986: 235), still without having
examined material, repeated his opinion: “that C. heinrichi is derived from C. sepulcralis”. This opinion, of the co-occurrence on Halmahera and Batjan of C. variolosus with a

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Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

presumed derivative of C. sepulcralis, must have influenced White in his decision totreat these as different species. Against this, his remark about the close relationship between C. merulinus, C. sepulcralis and C. sonnerati, with the conclusion that: “It is, however, wrong to suppose that they are all representatives of a single species in view of thesituation in the Malay Peninsula, Greater Sundas and part of the Philippines, where 3
sympatric species occur”, seems odd, as nobody has ever claimed these three to be conspecific, so that he is attacking windmills here.
Sibley & Monroe (1990: 98-99) have accepted White’s ideas and claimed of C. sepulcralis: “this species is distinct and more closely related to C. merulinus with which it is
sympatric”.
The suggestion that C. variolosus and C. sepulcralis are not conspecific, although theirranges are complementary, deserves some consideration, but I find the assumption thatthe two are not even very closely related, and that C. sepulcralis is closer to C. merulinus(with which, indeed, it is sympatric over a large part of its extensive range), completely
unacceptable. The songs of C. merulinus and C. variolosus are clearly different, as undoubtedly every ornithologist with field-experience in south-east Asia knows, and as I
described from Java (Mees, 1954). In evaluating the relationship between the variolosus
and the sepulcralis groups, not only their similarity in voice will have to be considered,
but also the fact that the Timor form seems to be to a certain extent intermediate, resembling variolosus in plumage, but sepulcralis in relative length of the tail. Tail-lenght assuch cannot be considered a specific character in this case, for the longest-tailed subspecies, C. v. macrocercus, is found in the Bismarck Archipelago.
Measurements:
wing
tail
tarsusFlores (C. v. sepulcralis)

117
126
13.4
 juv.
111
112
14
Java (C. v. sepulcralis)
10 
115-122115-133
15-18(118.2)(122.5)(16.8)
Timor (C. v. tymbonomus)

123
119
17.5
Kimberley Division, Australia ( C. v. dumetorum)

127
95
16.5

128
105
17
Port Essington, Australia (C. v. dumetorum)
 juv.
122
99
16
Cape York, Australia (C. v. dumetorum)(?) juv.
122
94
17
Pionier bivak, New Guinea
 subad.
121
102
17.3
 subad.
125
98
16

entire culmen

exposed culmen

22
22

17.3
16

18-22.7(20.9)

16-17.8(17.2)

22.5

16.8

20.5
23

17
17.5

20.2

17.2

19.8

14.8

23

18

Chrysococcyx basalis (Horsfield)
Cuculus basalis Horsfield, 1821, Trans. Linn. Soc. Lond. 13: 179 – Java.
Collector.— Rensch (1).
Material.— None.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

105

Notes.— For a discussion of records of this species from Borneo, see the notes under
C. minutillus.
Chrysococcyx lucidus plagosus (Latham)C[uculus] plagosus Latham, 1801, Suppl. Ind. Orn.: xxxi – Nova Hollandia.
Collectors.— Ten Kate, Verheijen/Schmutz.
Material.— (?), iv.1891, Sika (ten Kate, RMNH cat. no. 8); , 4.vii.1971, Langkas, Rahong (Verheijen,
RMNH no. 69760).

Notes.— The identification of several species of this genus is notoriously difficult.
The specimen from Flores listed by Wallace (1864: 484) as Chrysococcyx chalcites (whichequals C. lucidus), was apparently referred to C. basalis by Sharpe (1877: 320, cf. Parker,
1981: 35; I can find no evidence that Allen ever collected more than a single specimen of
the genus Chrysococcyx on Flores), was re-identified as Chalcococcyx malayanus by Shelley (1891: 299) and as Chrysococcyx russatus jungei by Parker (1981: 37). The specimen
collected by Ten Kate was originally identified as Chalcococcyx malayanus (cf. Büttikofer,
1892: 194), and correctly re-identified as Chalcococcyx plagosus by Finsch (1900: 95).
It is with much hesitation that I have retained the subspecific name plagosus forthese birds, for in a well-argumented paper, Gill (1983) has shown that geographicaldifferentiation between populations of Australia and New Zealand is negligible. If two
subspecies are recognized, obviously birds wintering in the Lesser Sunda Islands must
belong to the Australian plagosus and not to nominate lucidus. Gill made in his revision
no mention of the subspecies aeneus, described from the New Hebrides by Warner(1951), and from the discussion he gives of the New Hebridean population, it is apparent that he would not expect an endemic subspecies there; like previous authors (e. g.:
Friedmann, 1968: 110), he includes the New Hebrides into the range of C. l. layardi. Chalcites lucidus aeneus Warner is a primary homonym of Chalcites malayanus aheneus Junge(cf. ICZN, 1985: art. 58 (6)). Many years ago I drew the attention of Dr Warner to thehomonymy, and the fact that he has never renamed C. l. aeneus, may mean that he also
did no longer consider it a recognizable subspecies. Greenway (1978: 108-109) has accepted the subspecies, presumably without new investigation. I know of no other mention of it in the ornithological literature.
The statement in Blakers et al. (1984: 300) that this species has resident populationsin the Philippines, with a reference to Condon (1975: 208-209), who made no such claim
and describes the range correctly, must be due to confusion with C. minutillus.
Chrysococcyx minutillus subsp.
Collectors.— Allen (1), Verheijen (eggs only).
Material.— Egg: c/1 + 2 of Gerygone sulphurea, 27.ix.1955, Flores (RMNH no. 76581),
Measurements and weight:
The egg is immaculate olive brown.

RMNH no. 76581

21.0 × 14.0

0.107

Notes.— Unfortunately, the Fathers never managed to obtain skins of this species,

106

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

although it is not rare (cf. Ottow & Verheijen, 1969), so that in Flores it remains known
from the single specimen collected by Allen.
Admirable as Parker’s (1981) revision is, his division between the species C. minutillus and C. russatus, as well as the ranges ascribed to them, has something artificial. Ford
(1981) has presented a strong case for russatus and minutillus in Australia being conspecific. Indeed, Parker himself referred to intermediate specimens from the Australiancontinent. Following the nomenclature used by Parker, the host species of C. russatus
jungei on Flores is Gerygone s. sulphurea, and the host species of C. minutillus albifrons inJava is also Gerygone s. sulphurea. This supports the case for uniting the populations
from Flores and Java to one species, which must bear the older name C. minutillus.
Without topotypical material of jungei, and without specimens from Flores, I cannot
give an opinion on the subspecific status of the Flores population, but it is unusual for
Sulawesi (Celebes) and Flores to share one subspecies (and Java to have another). Therefore, I prefer to leave the subspecific identity of the population from Flores open until
adequate material becomes available.
Re-uniting C. minutillus and C. russatus leaves the problem of the occurrence of
two species (C. minutillus cleis and C. russatus aheneus in Parker’s nomenclature) in
Borneo (not Sulawesi as claimed by Blakers et al., 1982: 301). The status of cleis as aseparate form could still do with supporting evidence. Thompson (1966: 397) brought
all the specimens then known from Borneo together and concluded that such variation as they showed in plumage was sexual, although he was not clear about variations in wing-length. Nothing is known of breeding habits (host species, eggs), andvoice of the birds from Borneo. If the co-occurrence of two forms as resident birds is
confirmed, one of them must be a distinct species: C. aheneus or C. cleis. Payne (in delHoyo et al., 1997: 563) came with a new concept: that both cleis and aheneus would besubspecies of C. minutillus, spatially separated as follows: C. m. cleis, N and E Borneo,
and C. m. aheneus, SE Borneo and Phillipines. This view is not supported by the localities listed for the two forms by Parker, which seem haphazardly mixed, with bothrecorded from Mt. Kinabalu. This only increases my doubts about the validity ofcleis.
Now that I am discussing Borneo, I should also like to mention the type specimenof Cuculus neglectus Schlegel (1864b: 35). This bird was made the type of a separate genus Heterococcyx by Salvadori (1874: 61), but subsequently relegated to the synonymy
of C. basalis, a migrant from Australia. The type, however, is a fledgeling, not or barely
able to fly. It cannot possibly be a migrant from Australia, and must belong to a resident
species. I forwarded the specimen to Mr Parker, for his opinion. He concluded that it
must be either C. russatus aheneus or C. minutillus cleis (in the nomenclature of Parker),
but was not able to decide which of these two. The name C. neglectus antedates by many
years both aheneus and cleis.
The type-specimen of Cuculus neglectus constituted the sole basis for the inclusion of
mainland Borneo into the winter range of C. basalis (cf. Smythies, 1957: 638, etc.).
At my request, Mr Parker also borrowed from the Singapore Museum the specimenof C. basalis from the North Natuna Islands, first recorded by Chasen (1935: 128). He
could confirm that it had been correctly identified. It constitutes the sole remaining
record of C. basalis from the “Bornean Province”.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

107

Eudynamys scolopacea malayana Cabanis & Heine
E[udynamis] malayana Cabanis & Heine, 1862, Mus. Heineanum 4(1): 52 – Sunda-Inseln; Sumatra.
Collectors.— Allen, Weber, Everett (“a small series”), Elbert (1), Endih (1), de Jong (1), Verheijen (eggs),
Schmutz.
Material.— , , xii.1888, Maumeri (Weber, RMNH cat. nos. 38, 39); , 4.ii.1910, Endeh (Elbert, RMNH
no. 2330); , 15.ix.1969, Nunang (Schmutz, RMNH no. 85129); , laying, 17.xi.1984, Rangga, 650 m(Schmutz, RMNH no. 97108). Iris blood-red.

Notes.— Verheijen (1964: 199) listed egg-finds in November and December (oneeach). Ottow & Verheijen (1969) recorded two eggs, both found in November, in nestsof Corvus macrorhynchus, the only host-species that could be established on Flores.
Scythrops novaehollandiae Latham
Scythrops novae Hollandiae Latham, 1790, Index Orn. 1: 141 – nova Hollandia.
Collectors.— Semmelink, Allen (1), Martens (1), Weber, Verheijen (egg, in coll. Ottow).
Material.— (?) skull, 1862, Larantoeka (Semmelink, RMNH cat. no. a); (?), 19/24.xii.1888, Kotting (Weber,
RMNH cat. no. 35).

Notes.— Van Oort (1907: 125) listed Semmelink’s specimen as a complete skeleton,
and in fact there is a complete skeleton, but only the skull actually belongs to Scythrops,
the remainder of the skeleton is of a gallinaceous bird. Not improbably, this was doneon purpose, at a time that complete skeletons went on display, and it was considereddesirable to mount the spectacular skull on something that looked likely.
Ottow & Verheijen (1969) describe an egg found on 22.xi (year and locality not
given) in a nest of Corvus macrorhynchos; besides this egg, the nest contained four eggsof the host and one egg of Eudynamys scolopacea. In addition they record that a Corvus
florensis was seen feeding a fledgeling Scythrops. In an earlier paper, Verheijen (1961)
referred to an observation made on 28.iii.1960 in Central Flores, of two full-grownyoung being fed by crows.
This species, previously regarded as monotypic, was divided into three subspecies
by Mason & Forrester (1996); their study included RMNH cat. no. 35, but they were unable to assign it to a subspecies. Therefore I have, for the time being, left it with a binomial name.
Centropus bengalensis sarasinorum Stresemann
Centropus bengalensis sarasinorum Stresemann, 1912, Novit. Zool. 19: 338 – Lombok, Sumbawa, Flores...
Celebes...
Collectors.— Allen, Colfs, Weber (2 and a skull, of which 1 and the skull returned to ZMA), Everett (3?),
Rensch (1), Verheijen (2, MCZ), Verheijen/Schmutz.
Material.— , 1880, Flores (Colfs, RMNH cat. no. 22); (?), xii.1888, Maumeri (Weber, RMNH cat. no. 24);(?) nestling, 29.vi.1969, Nunang, 650 m (Schmutz, RMNH no. 81110); , 17.xii.1969, Nunang, 650 m(Schmutz, RMNH no. 85128); , , 26.ii.1976, Nunang (Schmutz, RMNH nos. 81058, 81052).

108

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Eggs: 23 clutches, of which five are from Palué, of c/1 (6 ×), c/2 (9 ×), c/3 (6 ×), and c/4 (2 ×), collectedin the months January (1), February (2), March (6), April (4), May (5), June (2), July (1), August (1), andone insufficiently dated. The eggs are white, with smooth but not very glossy shells.
Some measurements and weights:

RMNH no. 70600

32.7 × 25.4
0.829333.9 × 25.9
0.893134.2 × 25.5
0.930635.1 × 26.0
0.922
RMNH no. 70602 31.4 × 24.7
0.758133.6 × 25.3
0.848533.7 × 24.8
0.803733.8 × 25.6
0.8397The eggs are a little larger and heavier than those of C. b. javanensis from Java (cf. Hellebrekers & Hoogerwerf, 1967: 56), as would be expected from the larger size of C. b. sarasinorum.

Notes.— Four of the birds are in adult plumage, one (cat. no. 24) is in change. As I
have pointed out before, the well-established notion that this species would have an“eclipse” plumage, or a “winter” plumage, is erroneous: once the adult plumage has
been attained, it does not change (cf. Mees, 1971a).
In measurements, this subspecies is intermediate between the smaller C. b. javanensis, and the larger C. b. medius. White & Bruce (1986: 244-245) have, because of its intermediate character and geographical distribution, chosen not to recognise it. The discussions given by these authors, though lengthy, leave one in doubt as to whether they
have personally examined material. In my opinion, the fairly large size-differences involved, and particularly the wide range of this intermediate subspecies, justify its recognition. White (l.c.) called even medius “a very poor form”, and united it also withjavanensis. This certainly went too far. Bruce (l.c.), in addition, proposed to suppress C.
b. philippinensis as a size-race, ignoring the fact that this is not a size-race, but was basedon plumage-characters.
Measurements:

3
(?) 1

wing
152
173, 174, 182
59

tail
156
226, 204, 235
203

tarsus36
45.3, 44, 46
42

bill
25
29, 27.8, 30
28

Tyto alba javanica (Gmelin)
Strix javanica Gmelin, 1788, Syst. Nat. (ed. 13), 1 (1): 295 – Java = Batavia, for the name is entirely basedon a paper by von Wurmb (1781), who lived in that town.
Collectors.— Semmelink, Verheijen (4, MCZ), Verheijen/Schmutz.
Material.— (?), 1862, Larantoeka (Semmelink, RMNH cat. no. 6); , 24.v.1969, Nunang (Schmutz,
RMNH no. 66139); , 9.x.1969, Ruteng, 1150 m (Verheijen, RMNH no. 65300); , 15.xi.1969, Ruteng(Verheijen, RMNH no. 65310).
Eggs: c/3, ca. 20.iii.1954, Mataloko (RMNH no. 70612); c/2, 26.vi.1958, Deno (RMNH no. 70613); c/1,
27.vi.1958, Deno (RMNH no. 70614); c/4, 15.vi.1961, Soa (RMNH no. 70615); c/1, 25.v., 1961, Soa (RMNH
no. 70616); c/3, 15.xi.1961, Soa (RMNH no. 70617); c/3, 5.v.1963, Soa (RMNH no. 70618); c/1, 17.v.1963,
Soa (RMNH no. 70619); c/1, 18.v.1963, Soa (RMNH no. 70621); c/2, 16.xi.1966, Soa (RMNH no. 70622);
c/1, without data, Flores (RMNH no. 70620). The eggs are immaculate white.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)
Some measurements and weights:

109

RMNH no. 70612

RMNH no. 70613

42.3 × 34.2
42.6 × 32.4
42.7 × 33.5
41.1 × 32.4
41.3 × 32.8

2.30681.78972.02271.90621.9504

Notes.— Rensch (1931a: 522) placed the Barn Owls from Java, Lombok, Flores andTimor in the subspecies javanica. Without comment he accepted the subspecies sumbaensis from Sumba, everetti from Sawu, and kuehni from Kisar, all three of which had been
described by Hartert.
Mayr (1944a: 150-151) studied material of two of these named populations (everetti
and kuehni), comparing them with specimens from Timor and from Australia, and concluded, that all should be united with T. a. delicatula from Australia. In a previous publication, I followed this, without own investigation (cf. Mees, 1964a: 37).
Assuming that both the cited authors were right, the implication would be that T. a.
delicatula does not differ from T. a. javanica, and that the colonisation of Australia by Tytoalba must be very recent indeed. Anyway, the subspecific status of the Lesser Sunda Island birds could not be evaluated properly, without a comparison with birds from Javaas well as birds from Australia.
Unfortunately, the material available from Australia was very inadequate, but there
is a clear suggestion that Australian birds are smaller than javanica, and this is supportedby the measurements of a large series, provided by Schodde & Mason (1980: 82).
A further comparison showed that birds from Australia are paler on the upper parts
than birds from Java, and that in this respect, the specimens from Flores agree with javanica and not with delicatula. Perhaps birds from Flores average smaller than birds from
Java, but larger series are required to confirm this, and even in size they may be closer totopotypical javanica than to delicatula. In conclusion: T. a. delicatula is a valid subspecies,differing from T. a. javanica by average smaller size and by paler coloration. Birds from
Flores are javanica, not delicatula. This leaves open the possibility that birds from Sawu,
Timor and Kisar are delicatula, as claimed by Mayr (l. c.) and White & Bruce (1986: 146).
There is only one specimen from Sumba in the collection, of somewhat problematical antecedents (sex not given, mounted, no date, no collector, received from theKoloniaal Instituut in 1875). One wing is clipped, from the other wing the longest primaries are missing. The reason why I mention this imperfect specimen is that it does
not have the very pale, almost whitish, tail presumed to be the main character of T. a.
sumbaensis. The tail does not differ significantly from the tails of T. a. javanica, or T. a.
delicatula for that matter. In view of the hazy antecedents of the specimen, the question
arises, whether its given provenance from Sumba is reliable. Few specimens from
Sumba are known (Stein collected only one, Sutter none), and I consider that the validity of T. a. sumbaensis requires confirmation.
Surprisingly, there seem to be no published records of the Barn Owl from Sumbawa.
On 22.ix.1984 I observed one on the Tg. Pioen peninsula, northern Sumbawa. There is
now a specimen record from Sumbawa Besar (Johnstone et al., 1996: 170).
Measurements:
Java (T. a. javanica)
15 

wing

tail

tarsusculmen from cere295-324(308.1)

114-132(121.5)68-77(71.6)

22-24(23.1)

110

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)17 286-321(305.4)

Flores (T. a. javanica)
2972
278-289(?)
297
Australia (T. a. delicatula)2
272, 277
277110-126(119.5)65-74(70.1)

22-24(23.5)

117
108-120
112

69
69-70
64

23
19-22.3
22.5

103, 103103

68½, 7270½

21½

20½, 22.3

Tyto longimembris longimembris (Jerdon)
Strix longimembris Jerdon, 1839, Madras J. Lit. Sci. 10: 86 – Neilgherries.
Collector.— Verheijen (1, MCZ).
Material.— None.

Notes.— From Flores, the MCZ-specimen, collected near Wangkung in March 1956,
remains the only one recorded (cf. Paynter, 1963), but Verheijen (in litt., 17.vi.1991) mentioned that about 1953 the late Father Jan Loeters showed him a place in an extensive
plain grown with alang-alang (Imperata cylindrica), where he had flushed a “Barn Owl”
from a nest with eggs on the ground. Father Verheijen found it strange that a Barn Owl
would nest on the ground, but forgot it until Tyto longimembris was discovered.
Stresemann (1939: 316, 325, 343 fig. 11) gave T. longimembris as an example of a species that has colonized Australia from mainland Asia through the “Graslandstrasse”
formed by southern China, Taiwan, the Philippines, Sulawesi (Celebes) and Flores. It israther interesting that he anticipated its occurrence on Flores, from where at that time it
was not yet known. Mayr (1944b: 119) accepted this reconstruction as: “clearly indicated”. Amadon & Jewett (1946) considered it “probable” that Stresemann was right, but
added that: “it occurs in southern Indo-China, so there seems still a possibility that it
will be found somewhere in Malaya”. In the revision of the subspecies presented bythese authors, the Australian walleri is stated to be separable “at a glance” from continental longimembris, whereas amauronota of the Philippines is considered intermediatein certain respects. Thus, the geographical variation described by these authors would
seem to support, or at least not to contradict, Stresemann. However, Amadon (1959),
following the examination of much additional material in the British Museum, again
reviewed the subspecies of T. longimembris, accepting as valid chinensis (southern China), pithecops (Taiwan), amauronota (Philippines), papuensis (New Guinea). The main difference from his earlier revision was in his evaluation of walleri, which he accepted, butwith the comment: “The race walleri is exceedingly similar to longimembris of India: infact, if their ranges were continuous there could be no thought of separating them...
Perhaps the species has spread comparatively recently from India to Australia and may
still be recorded from some of the intervening areas, for example Sumatra”. Although
Amadon’s hesitation is understandable, the logical next step was to synonymize walleri
with longimembris (cf. Mees, 1964a). The morphological evidence, as far as now available, no longer supports Stresemann’s reconstruction of its distributional history. Several
other grassland birds have at present an interrupted range: continental Asia and Java,Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

111

but not Malaya and Sumatra (e. g. Timalia pileata, Prinia polychroa, Prinia inornata blythi 8),
Dicrurus macrocercus), suggesting a grassland corridor in the not too remote past, probably during the last period of low sea level (Late Pleistocene).
Otus magicus albiventris (Sharpe)
Scops albiventris Sharpe, 1875, Cat. Birds Brit. Mus. 2: 78, pl. VIII fig. 1 – Flores.
Collectors.— Semmelink, Allen, Everett, Verheijen, Schmutz.
Material.— (?) juv., , 1862, Larantoeka (Semmelink, RMNH cat. nos. 1, 2); , 23.x.1968, Nunang, 650 m(Schmutz, RMNH no. 65313);  with large gonads, 3.x.1975, Cereng, 550 m (Schmutz, RMNH no. 85159),(?), xi.1982, Nunang (Schmutz, RMNH no. 81023). Stomach contents of no. 85159: Cicadas.
Eggs (fig. 14a): c/1, without data, Flores, received 1973 (Verheijen, RMNH no. 70604); c/1, without data,
received 1973 (Verheijen, RMNH no. 70605); c/2, 25.ix.1955, Bénténg Djawa (RMNH no. 70606); c/2,
1.vi.1961, Soa (RMNH no. 70607); c/2, 4.vi.1961, Soa (RMNH no. 70608).
Measurements and weights:

RMNH no. 70604
RMNH no. 70605
RMNH no. 70606
RMNH no. 70607

RMNH no. 70608

32.9 × 27.1
32.6 × 27.9
31.7 × 27.8
33.5 × 29.0
33.2 × 28.1
33.2 × 28.2
32.6 × 29.5
32.8 × 29.2

0.84490.9621
0.96381.1071very large blowholes,
no weights taken
0.875
0.959

Notes.— Without much confidence I follow current practice of listing this bird under the above trinomial. The state of flux in which the classification at the specific andsubspecific levels of the genus Otus is at present, makes any trinomial of no more than
temporary validity.
Measurements:


(?)

wing
156
164
164

tail
73
73½
69

tarsus27
28.4
27½

exposed culmen
21
21
19

Otus alfredi (Hartert)
Pisorhina alfredi Hartert, 1897, Novit. Zool. 4: 527 – Mount Repok and other hills at about 3500 feet in S.
Flores.
Collector.— Everett (3).
Material.— None.

8)
Watson et al. (1986a: 142) united P. inornata with the African P. subflava. The illogical sequence of subspecies: first the Asiatic one from west to east, followed by the African ones from north-west to south,
suggest that this was a last-minute decision. The interesting, isolated occurrence on Java (blythi) was
overlooked.

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Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Notes.— Within a year after its description, Finsch (1898b: 177, 183) concluded
that Pisorhina alfredi is not a valid species, but represents the red morph of Otus manadensis on Flores, hence of O. manadensis albiventris (Finsch did not recognize albiventris, but that does not affect his conclusion). Subsequent workers have ignored Finsch’s views, and Stresemann (1925: 193) gave as his opinion that Otus alfredi: “gehört
sehr wahrscheinlich zum Formenkreis O. spilocephalus”. It was kept in the vicinity ofO. spilocephalus by Rensch (1931a: 521), Peters (1940: 89) and Eck & Busse (1977: 72).
Certainly, nobody since Finsch appears to have doubted its status as a distinctive species.
Some years ago it struck me that, whereas the plumage differences between O. m.
albiventris and O. alfredi, as described and illustrated by Hartert (1898a: pl. 1) and Marshall & King (in Amadon & Bull, 1988: pl.), are considerable, the morphological differences are very slight. Hartert stated of alfredi: “Toes and one-fifth of tarsus bare, the remainder of the tarsus thickly feathered”, whereas albiventris would have “the tarsi...
feathered down to the toes... the beak larger”. The illustrations, however, show O. alfredi with a tarsus-feathering similar to that of albiventris, and with the measurements
one can do little, as Hartert omitted to give those of albiventris, but comparing the measurements provided for alfredi by Hartert, with those of albiventris taken from my material, I see little difference.
The above considerations led me to write to the American Museum of Natural History, with the request to re-examine the type material of O. alfredi with the thought inmind that it might represent a red morph of albiventris. In March 1991, Dr Marshallvisited New York to make the comparison, and concluded that the suggestion firstmade by Finsch is correct.
As Dr Marshall was the acknowledged authority on the genus Otus I had no reason
to doubt his conclusion, but recently the discussion about the status of O alfredi has been
re-opened. The case is put strongly in del Hoye et al. (1999: 154):“morphological study
clearly demonstrates its distinctness and validity as a species”. Unfortunately, comparative documentation of these clear morphological characters is withheld. Admittedly there is mention of recent records, but too succinct to be of much use. Where hasthat specimen collected in 1995 gone? The references given look unpromising; I checked
the only one available to me (Rasmussen, 1998) and found it unhelpful: “O. alfredi …..
is not the red morph of albiventris, but belongs to the O. spilocephalus group (Hartert,
1925, Widodo et al. unpublished)”. I have already given my opinion on references to
unpublished work, and those to Hartert and Stresemann have no much value in thisconnection, as the idea that alfredi might be the red morph of albiventris may not havebeen considered by these authors.
On the other hand, evidence indicates that O. alfredi is a mountain bird, with an altitudinal range different from that of albiventris. I am now quite prepared to accept thevalidity of O. alfredi. As regards its relationships, as quoted by Rasmussen, it deserves
mention that the Javanese mountain form angelinae has now been chopped off fromspilocephalus, so that the latter is no longer its nearest neighbour and its most obvious
relative.
Addendum.— The paper I had been hoping for appeared already in 1999 (Widodo
et al., 1999), but I became aware of this, and received a copy, only in 2003. It is comprehensive and completely convincing, and answers all my questions. The authors ascribe

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)113

the small size of the recently-collected male to its being a juvenile, but a part of the difference might be sexual, the Everett specimens being unreliably sexed and perhaps females.
Otus silvicola (Wallace)
Scops silvicola Wallace, 1864, Proc. Zool. Soc. Lond. (1863): 487 – Flores.
Collectors.— Allen (1), Everett (3), Verheijen (3, MCZ), Verheijen/Schmutz.
Material.—  juv., 22.xii.1968, Wangkung, Rahong, 900 m (Verheijen, RMNH no. 65311);  juv., 10.
ii.1969, Nunang, 650 m (Schmutz, RMNH no. 65312);  ad., 8.ix.1971, Nunang, 650 m (Schmutz, RMNH
no. 84864);  ad., 1.xii.1975, Nunang (Schmutz, RMNH no. 81038);  juv., 1.xii.1975, Nunang (Schmutz,
RMNH no. 81039). Iris sulphur yellow.
Eggs (fig. 14b): c/2, 12.ix.1949, Rekas (RMNH no. 70610); c/3, 10.x.1949, Rekas (RMNH no. 70611).
Measurements and weights:

RMNH no. 70610

36.8 × 32.0
38.3 × 32.4
RMNH no. 70611 40.5 × 32.3
41.6 × 32.1
45.1 × 32.4
One of the eggs of clutch 70611 is indeed conspicuously longer than the others.

1.685
1.629
1.828
1.954
1.856

Notes.— The material is of particular value as hitherto few specimens existed in collections: the type collected by Allen, three males from Flores, collected by Everett, twomales from Sumbawa, collected by Rensch, and three birds from Flores (fig. 8), collected
by Verheijen (MCZ, cf. Paynter, 1963).
Unfortunately, only two of the above
specimens are adult, one of each sex; theothers are fledgelings, with the remiges
and rectrices basally still in sheath.
Wallace did not mention the sex of
the type, for which he recorded a winglength of 8½ inches, tail 4½ inches. The
two males from Sumbawa had wing
215, 221, tail 101, 106 mm (Rensch, 1931a:
521). Hartert (1897b: 527) gave for males
a wing-length of 215-223 mm, tail 108
mm. According to him, the type was ayoung bird (sex doubtful) in down, with
only a few feathers, and: “the dimensions of the type are very much larger”,
which is not borne out by the figures of
the type provided by Wallace. Warren(1966: 271) claimed, without explanation, that the type was an “Immature
Fig. 8. Otus silvicola, Flores, 12.x.1975 (photo E.
Schmutz).

114

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

female”. The various contradictory statements about the type evidently required looking into. At my request, Mr Cowles has carefully examined the type specimen, about
which he informed me as follows (in litt., 22.xi.1988): “We do not have any other silvicola for comparison, but I have examined other Otus specimens and the young downy
ones. These are really very downy and fluffy, even when fairly well grown. The silvicola
type is rather different in that it is fully feathered and shows an intermediate stage, being a mixture of some adult plumage with dark streaked feathers on the back, two or
three on the upper breast and 4 or 5 on the abdomen. The head, neck, upper back andmost of the breast has finely barred plumage similar to the juvenile plumage and israther loose in construction, like the young birds. To my mind this specimen appears tobe just changing into first adult plumage and could be called immature or subadult,
certainly not downy young as Hartert suggested .... The original Wallace label is marked
 and that is no doubt why Miss Warren copied it into her type-specimens publication”. Measurements taken from the type by Mr Cowles are, wing 223 mm, tail 109 mm,
quite close to the measurements provided by Wallace in the original description. With
the evidence now available (that the specimen is fully grown), the sex given on the label
may be questioned, it is more likely to be a male. About the three MCZ specimens, I was
informed by Dr Paynter (in litt., 20.x.1988): , 11.ii.1959, Mano (MCZ no. 261890), wing
240 mm, tail 120 mm; (?) juv., no date, Mano (MCZ no. 261891); ?, 27.vii.1958, Mano
(MCZ no. 261892), very badly worn (captive bird?). The measurements of the two adultbirds in the RMNH collection are given below. The juvenile birds have wing-lengths of 204,  176 and 212 mm.
The surprising and interesting point is the large size of the female. In plumage, I can
see no obvious differences between the sexes. There has been much speculation on theaffinities of Otus silvicola. Paynter (1963) thought that: “This endemic owl is probably a
giant geographical representative of O. bakkamoena”. Hekstra (in Burton, 1973: 113)
claimed it to be closely related to Otus brookii, a very little-known mountain-bird ofBorneo, Sumatra and Java, but he gave no explanation for this view. Perhaps it was
merely based on O. brookii also being, for a member of the genus Otus, rather large; its
few published measurements, however, show it to be much smaller than O. silvicola(wing 5  average 162.4 mm, 1  183 mm, according to Marshall, 1978: 57). Eck & Busse(1977) have forced both silvicola and lempiji (Java, Bali) as subspecies in an enormously
expanded species Otus asio. Few authors will follow them in this extreme lumping, but
the notion that silvicola and lempiji are closely related persists (cf. White & Bruce, 1986:
253). Therefore, I want to stress not only the huge difference in size: lempiji, wing 20 
135-149 mm, 22  136-151 mm (cf. Mees, 1986: 52), but also the sexual difference in size,
which is marginal in lempiji (ca. 1½%), as opposed to ca. 12% in silvicola. Compare also
the difference in size of the eggs: average 33.5 × 28.8 mm, weight 0.97 (0.9-1.09) g in
lempiji (cf. Hellebrekers & Hoogerwerf, 1967: 58).
I should like to add that Otus asio is also a small species, with only a slight sexual
difference in size (cf. Ridgway, 1914, 689).
It is interesting to note that there is now another tendency, in the opposite direction, towards the recognition of more species. Roberts & King (1986) and Marshall &King (in Amadon & Bull, 1988) have separated O. lempiji from O. bakkamoena, withwhich it has been held to be conspecific for a long time, on the basis of differences invoice. I find their arguments convincing. This adds to the opinion, that the generalMees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

115

state of knowledge of the genus Otus does not yet allow theories on interspecific relations to be more than speculation.
Measurements:



wing
214
252

tail
103
120

tarsus32.6
35

entire culmen
30
35

Ninox scutulata florensis (Wallace)
Athene florensis Wallace, 1864, Proc. Zool. Soc. Lond. (1863): 488 – Flores.
Ninox macroptera Blasius, 1888, Braunschw. Anz., 11 Jan., Nr. 9: 86 – Manganitu, Gross-Sanghir (nonvidi).
Ninox sanghirensis Blasius, 1888, Ornis 4: 546 – lapsus or alternative name for N. macroptera.
Ninox scutulata ussuriensis Buturlin, 1910, Mess. Orn. 1: 187 – Ussuri and Korea (non vidi).
Collectors.— Allen (1), Schmutz.
Material.— (?), 1862, Flores (Allen, BM no. 73.5.12.1650, holotype of the subspecies); , 14.i.1983, Nunang, 650 m (Schmutz, RMNH no. 81164); , 19.iii.1983, Paku, 350 m (Schmutz, RMNH no. 97115).

Notes.— These are large birds (see measurements). Birds of this size surpass inmeasurements birds from Japan, and must originate from Ussuria. Therefore, florensis
replaces ussuriensis as the valid name for the large birds from eastern Siberia. Deignan(1951) has suggested that the name florensis could be applicable to “Chinese birds”, andthis is probably correct for birds from Manchuria. Unfortunately, Deignan does notprovide a single measurement, neither does he discuss Ninox macroptera Blasius, ofwhich the type-locality was restricted to Mindoro by Peters (1940: 141), although, as far
as I can see, all the type material was from the Sanghir Islands, and Mindoro is not evenmentioned by Blasius (1888: 545-555). Previously, Meise (1941: 355-356) recorded twolarge specimens (2 , wings 239, 241 mm), from Noesa Penida near Bali, for which heused the name macroptera, correctlty placing ussuriensis in its synonymy, but he seems
to have overlooked florensis, which is an older name than macroptera.
The name N. s. florensis has not been generally used in recent literature, but duPont(1971a), following Deignan’s suggestion, quoted above, that Chinese birds should becalled by this name, referred three winter birds (October and December) from Luzon toit, with the remark: “Not previously recorded from the Philippines”. As no measurements are provided, indeed, no description of any kind is given, it is impossible to judge
whether the identification was correct. A little later, duPont (1971b: 176) listed the Philippine subspecies, accompanied by very succinct diagnoses; the first one, randi, even withmeasurements (wing 240, tail 135, bill 28, tarsus 33), followed by palawanensis (“differsfrom randi by being smaller”, wing 195, tail 109; no characters are given to distinguish
this subspecies from borneensis), by japonica (“differs from randi by having the upperparts browner and the red-brown in the wings and tail much reduced; underparts not as
heavily streaked”; no measurements are given), and by florensis (“differs from japonica byhaving the upperparts much paler and the underparts not so heavily streaked”; nomeasurements given). This would suggest that both japonica and florensis have the measurements of randi, but japonica certainly never reaches a wing-length of 240 mm, which
also places the measurements of duPont’s florensis, hence the subspecific identity of thesebirds in doubt. Parkes (1971: 14-15) was not convinced of the applicability of the name

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Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

florensis in the sense in which it was used by Deignan and duPont, but nevertheless also
used it: “I believe it to be highly likely that many Philippine specimens that have been
identified in the past as japonica will prove, upon reexamination, to be referable to thepale, grayish race called florensis by Deignan”. As Parkes also failed to give measurements, the identity of the specimens he was discussing remains obscure.
On this basis, Dickinson et al. (1991: 229) included N. s. florensis in the avifauna of
the Philippines, with added a note “The type of florensis...has been compared withnorthern birds likely to be on their breeding grounds, and is a good match for BM
1934.1.1.1549 taken at Wei Hai Wei, NE Shantung, China on 12 May 1911; florensis...istherefore thought to have precedence over ussuriensis...as the correct name for thenorthern Chinese population”. The conclusion about the synonymy is correct, but abird collected on 12 May at Wei Hai Wei would almost certainly have been a migrantfrom further north, and provides no proof that the subspecies florensis breeds in China
proper (cf. LeFevre, 1962: 74). The region of Chefoo/Wei Hai Wei is a well-known staging post for migrants across the Yellow Sea. As still no measurements are given, it is notclear whether this comparison has any relevance to the identification of the Philippine
specimens. In spite of this unsatisfactory evidence, N. s. florensis ought to occur in thePhilippines as a passage migrant and, probably, a winter visitor. One would expect the
winter range to extend westward to at least eastern Java and Borneo, although there are
no records, but the occurrence on islands off the west coast of Peninsular Thailand(Deignan, 1963: 63) is unlikely and is presumably based on misidentification.
In reply to my request for the loan of the specimen of Ninox scutulata, which, naturally enough, I believed that Pfeffer (1958: 71) had obtained in Flores, I received one
collected by him in Borneo (, 25.iii.1957, Long-Kemuat, upper Bahau, MHNP no.
1960.762). This is the bird recorded by Pfeffer (1960: 201) under the name of N. s. borneensis, the resident subspecies. The measurements I took from this specimen (wing 212,
tail 109, tarsus 26, culmen from skull 25.8 mm) are, however, too large for borneensis (inwhich the wing-length does not exceed 200 mm), and show it to be an individual of the
migrant visitor N. s. japonica.
The wing-length of a carcass picked up on Ashmore Reef was recorded as only 226
mm (Schodde & van Tets, 1981), but is still within the range of variation of birds from
Ussuria (cf. Vaurie, 1965: 618, s. n. ussuriensis), hence of florensis. The  from Boemboelan, northern peninsula of Sulawesi (Celebes), 28.x.1939, reported by Ripley (1941a: 354,
s. n. N. s. scutulata), had a wing-length of 238 mm and therefore is definitely florensis.
That the subspecies N. s. japonica also occurs in northern Celebes, was pointed out by
Van Marle (1940b: 123, s. n. N. s. scutulata), but only N. s. florensis seems to reach theLesser Sunda Islands.
Measurements:


(?)

wing
235
235
242

tail
125
113
127

tarsusca. 3029.8
ca. 31.5

culmen from skull
25
25
25

culmen from cere
14
14.7
14.5

Caprimulgus macrurus schlegelii Meyer[Caprimulgus] Schlegelii Meyer, 1874, Sitzb. Akad. Wiss. Wien 69: 210 – no locality = Port Essington.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

117

Material.— , 15.ix.1975, Nisar, 250 m (Schmutz, RMNH no. 75297).
Egg: c/1, 11.x.1956, Djinggor (Verheijen, RMNH no. 70624). This and the other eggs of the species arecreamy to pale pinkish buff, clouded with poorly-defined light grey and brownish markings.

Notes.— The above material was recorded in a paper by Mees (1977a: 31, 36, 47), towhich I refer for further particulars. The subspecies schlegelii is only poorly differentiated from the nominate race, with type-locality Java.
Dickinson et al. (1991: 234) suggested that eggs of C. manillensis (previously known
as C. macrurus manillensis) from Cape Engano, Luzon, illustrated by Mees (1985: fig. 7
nos. 1 and 2), had been misidentified and were actually referable to C. affinis griseatus. I
asked Mr Walters, who selected these eggs for illustration, about this possibility, and his
reply is that he sees no reason to question their identification (as C. manillensis).
Caprimulgus affinis affinis Horsfield
Caprimulgus affinis Horsfield, 1821, Trans. Linn. Soc. Lond. 13: 142 – Java.
Caprimulgus affinis undulatus Mayr, 1944, Bull. Amer. Mus. Nat. Hist. 83: 152 – Flores.
Collectors.— Weber, Everett, Rensch (5), Verheijen/Schmutz.
Material.— (?), skeleton, 28.xi.1888, Reo (Weber, RMNH cat. b, cf. van Oort, 1907: 156); , x.1896, Desoe,
S. Flores, 3300 ft. (Everett, BM no. 98.5.4.36); , 25.v.1972, Nisar, 50 m (Schmutz, RMNH no. 85272); ,
29.vi.1974, Orong, 550 m (Schmutz, RMNH no. 85271); , 17.ix.1975, Joneng (Schmutz, RMNH no.
85270).
Eggs: c/2, 12.viii.1955, Mano (RMNH no. 70623); c/1, 16.x.1956, Djinggor (RMNH no. 70625); c/1,
23.ix.1957, Léwé (RMNH no. 70626); c/1, 2.xi.1957, Léwé (RMNH no. 70627); c/3, 4.xi.1957, Mano(RMNH no. 70628); c/2, 10.xi.1957, Léwé (RMNH no. 70629); c/1, 19.xi.1957, Mano (RMNH no. 70630);
c/2, 21.ix.1959, Léwé (RMNH no. 70631); c/2, 22.ix.1959, Léwé (RMNH no. 70632); c/1, 23.ix.1959, Léwé
(RMNH no. 70633). The eggs resemble those of C. macrurus in a general way, but average a trifle smaller;
the ground colour is a little colder, tending towards olive-grey rather than buffy; the markings aredarker and usually more sharply defined.
Measurements and weights:

RMNH no. 70623
RMNH no. 70625
RMNH no. 70626
RMNH no. 70627
RMNH no. 70628

RMNH no. 70629
RMNH no. 70630
RMNH no. 70631
RMNH no. 70632
RMNH no. 70633

29.6 × 20.2
29.8 × 19.5
28.2 × 20.3
28.9 × 20.6
28.1 × 21.5
26.3 × 20.2
26.5 × 19.8
26.9 × 20.3
27.0 × 19.7
27.8 × 20.3
27.7 × 20.2
29.5 × 20.8
29.7 × 21.6
26.5 × 19.9
28.4 × 20.7
29.7 × 20.5

0.43080.42350.46680.44200.4782
0.39800.3770 large hole
0.44600.42170.44270.4216
0.48690.50760.35090.44610.3642

Notes.— The material on which Mayr (1944a: 152) based the subspecies undulatus
was not rich: 2  and 3  from Sumbawa and Flores. As type, he chose the nesting from Flores, listed by Hartert (1898a: 42). The subspecies undulatus was described as

118

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

being: “Similar to affinis in the coloration of the upper parts, but tending to be lighter,more evenly colored, and more finely vermiculated; breast and belly are more closely
barred and the barring extends farther down along the flanks”. My examination of alarger material from Java and the Lesser Sunda Islands (RMNH, BM), failed to support
any of the characters claimed by Mayr, who evidently underestimated the amount of
normal individual variation. Differences in method of preparation also affect somecharacters, such as the visible extent of barring of the underparts. Hence it is with confidence that I unite undulatus with nominate affinis.
Mayr (l. c.) recognized two additional subspecies from the Lesser Sunda Islands: C.
a. kasuidari Hachisuka from Sawu (type locality) and Sumba, and C. a. timorensis, newly
described from Timor. Neither subspecies is satisfactory: the three specimens from
Sawu examined by me (viii.1896, leg. Everett, BM nos. 97.11.1.76, 125, 126) fit into thenominate race; from Timor, only a single old specimen could be examined (, 1861, East
Timor, leg. Wallace, BM no. 88.10.3.135), so that I prefer not to give a definite opinion on
the validity of timorensis, although I must admit to having little confidence in it. Two
specimens from Kisar, to the East of Timor (8 and 25.xii.1897, leg. Schädler, RMNH cat.
nos. 17 and 18) are characterized by rather pale, less brownish-tinged, underparts (compared with the average affinis). A bird from Alor (v.1897, ex Rolle, RMNH cat. no. 16), on
the other hand, is over average brown, with even the wing specula tinged very pale
brown. Junge (1954: 318) was unable to name the collector of Rolle’s small Alor collection. Although there is no collector’s name on the labels, the skins show the unmistakable, rather thin shape characteristic of H. Kühn’s work. In the Lombok series, one individual (, vi.1896, leg. Everett, BM no. 97.11.1.44) is very pale, almost like griseatus!
The Lombok series illustrates well the amount of individual variation existing even ona smallish island: there is not only variation in the depth of the cinnamon colour, butalso in the amount of black on the pileum. All this confirms me in the opinion that inbirds with this kind of complicated colour pattern, it is just not possible to base subspecies on samples of one or two specimens, as has been done in the past.
This brings me to C. a. propinquus, a subspecies described on the basis of a single
male from Parigi, Sulawesi (Celebes). Compared with four specimens of the same sexof nominate affinis (Java 3, Borneo 1), it was stated to differ by being: “much lighter,both above and below, and the buffy spots on the chest and wing-coverts are more numerous and pronounced and much lighter; the vermiculations on the back finer and thetail-bars above narrower” (Riley, 1918). I doubt the validity of propinquus, and only thefact that Mayr examined two additional specimens from the same area (Tawaya), whichshowed the same characters as the type, has made me retain it for the moment.
Three of the nine birds from Borneo ( 171,  169, 170) surpass in wing-length
specimens from all other localities. The large size of Borneo birds was first noted by
Mayr (1944a: 152) according to whom: “The large size of two of the three specimens isprobably due to altitudinal variation. They were collected at Tanggaroeng, Mahakkam
River (H. Raven)”. There is a misconception here, for Tanggaroeng (recte: Tenggarong)is in the delta region of the Mahakkam, only some 25 km west of Samarinda, and hardly above sea-level. The bulge inland to the headwaters of the Mahakam, shown inMayr’s (1944a: fig. 2) distribution map, has to be corrected. For notes on Raven’s itinerary, see Deignan (1960).
Specimens from Sumatra average smaller than specimens from Borneo, but larger
than birds from Java. The series from Java and Sumatra are long enough to make itMees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

119

likely that the difference, slight as it is, is real and not due to inadequate sampling.
The opinion that C. monticolus of the Asiatic mainland is conspecific with C. affinis,
was apparently first expressed by Stresemann in a letter to Rothschild (1927), who did notagree. Later by Rensch (1931a: 537) and Mayr (1944a: 153). The latter supported this withthe statement that: “The two ‘species’ affinis and monticola [sic] are perfectly connected by
the subspecies griseatus (Luzon) and stictomus (Taiwan)”. It is my feeling that Mayr has
put the case too strongly: Philippine griseatus and mindanensis (the validity of mindanensis
requires confirmation; it is at best only slightly differentiated from griseatus) differ conspicuously from typical affinis by being much greyer, less brown: the upper parts, including the head are mottly grey; the underparts are much paler brown; in measurements,
however, they agree with affinis (see table). The Taiwanese subspecies, stictomus, clearly
belongs to the monticolus group; it has similar tail-markings; admittedly it is greyer thanthe very richly-coloured mainland birds, and seems to average slightly smaller, but I
doubt that these characters prove a close relationship with griseatus. The size-gap between stictomus and griseatus is considerable. I much doubt Mayr’s (1944a: fig. 2) reconstruction of its history of expansion, from the Asiatic mainland, through Taiwan and thePhilippines to Celebes, Java and the Lesser Sunda Islands; certainly the morphologicalevidence provided by the Taiwanese and the Philippine forms, does not strongly support
it. For one thing, it would have required, in the course of this colonisation, first to havechanged from brown to grey, and subsequently from grey to brown again.
Peters (1940: 212-213) and Etchécopar & Hüe (1978: 498) preferred to keep the two
as separate species. The reason why, in spite of the unconvincing morphological evidence, I have nevertheless reluctantly accepted that C. monticolus and C. affinis are conspecific, is that their calls are said to be identical (cf. Sibley & Monroe, 1990: 191). 9)
Measurements:
Sumatra (C. a. affinis)

Borneo

Java

Bali

9)



n
11



10



3



6



21



14



5



3

wing
157-168(163.5)158-166(161.0)164-171(166.3)160-170(165.0)154-167(160.9)155-163(159.0)156-166(161.2)
156-161(159.0)tail
91-97(94.1)
86-94(90.8)
91, 94, 93
84-95(91.5)
86-98(92.7)
86-95(91.2)
91-98(94.5)
93-96
94.5

As there is now general agreement that names ending in –cola, and perforce also –colus, are substantiva, not adjectiva, monticolus and monticola are different words. Therefore, Peters (1940: 201) did not
create secondary homonymy whnen he transferred Stenopsis cayensis monticola to the genus Caprimulgus,
and there was no need for him to rename that form. Caprimulgus cayensis apertus Peters, 1940, nomen
novum, is a synonym of Caprimulgus cayensis monticola (Chapman, 1915).

120

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

158-160
90-96½(159.3)(93.8)
5
152-165
88-94(158.2)(91.0)
Sumbawa1
163
92
Flores
2162, 16895, 96
2160, 165
92, 96
Alor

1
156
87
Sawu

1
164
101

2
163, 16590, 94
Timor

1
157
91
Kisar

2
152, 156
89, 85
Northern Philippines: Luzon, Mindoro, Catanderanes (C. a. griseatus)5160-168
87-94(163.6)(91.2)

1
167
91
Mindanao (C. a. mindanensis)

1
166
89

1
157
81
Taiwan (C. a. stictomus) 11184-197105-121(191.5)(113.6)
9
175-192
99-119(184.1)(107.8)Macao (C. a. amoyensis) 
1
199
123
Southern China (C. a. amoyensis)2191, 198
118, 116
Lombok6

Collocalia fuciphaga fuciphaga (Thunberg)
Hirundo Fuciphaga Thunberg, 1812, Kongl. Vet.-Akad. nya Handl. 33: 153 – Java (reference not verified).
Collocalia francica dammermani Rensch, 1931 (September), Mitt. Zool. Mus. Berlin 17: 541 – Mboera, WestFlores.
Collocalia francica dammermani Rensch, 1931 (December), Treubia 13: 396 – Mboera, West-Flores.
Collectors.— de Jong (1), Verheijen (1, MCZ), Verheijen.
Material.— , 9.iii.1971, Maro-Kama, Borong (Verheijen, RMNH no. 85325). Primary moult.
Eggs: c/2 with nest, 24.xi.1981, south coast Todo (RMNH no. 80906). The eggs are long-oval, dull white.
Measurements and weights:

RMNH no. 80906

19.5 × 13.4
0.098820.6 × 13.2
0.0990The nest is of the best edible quality, with little or no extraneous matter, thus supporting the identification.

Notes.— In his description of C. f. dammermani, Rensch claims: “Von diesem Rassenkreise war bisher noch kein Vertreter von den Kleinen Sunda-Inseln bekannt”; yet, a
few pages earlier in the same publication, he (Rensch, 1931b: 377) records C. f. micans
from Sumba! The type-locality of micans is Sawu, another of the Lesser Sunda Islands(cf. Stresemann, 1914b: 6). Both dammermani and micans are regarded as synonyms of
nominate C. f. fuciphaga by White & Bruce (1986: 266), and although their opinion ispoorly documented, I follow them for the moment.
About few genera of birds there has been so much confusion as about Collocalia, andeven the most recent classification can by no means be called definitive. Rensch (1931a:

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

121539) refers specimens from Lombok, with a query, to micans, but according to White &Bruce (1986: 266) this is an error. Rensch (1930: 49-51; 1931a: 540) has twice described his
visit to the cave where these specimens were found. White & Bruce (1986: 265) are confused about the specimens discussed by Rensch, and speculate that he had examinedmaterial collected by Everett, but it is quite clear that Rensch only discussed the three
specimens from his own collection. As the nests were not “edible”, the birds can not be
C. fuciphaga, which is according to current opinion the only producer of edible nests inthis region. Obviously, there must be on Lombok a hitherto unidentified species of Collocalia, which is not C. fuciphaga; most likely it is C. salangana, a species common in Java(I follow Salomonsen, 1983: 86, in not regarding C. salangana as conspecific with C. vanikorensis).
Measurements:


wing
114

tail
48

tarsus9

entire culmen
7.7

exposed culmen
4

Collocalia esculenta sumbawae Stresemann
Collocalia esculenta sumbawae Stresemann, 1925, Mitt. Zool. Mus. Berlin 12: 189 – Tambora 3000’, Sumbawa.
Collectors.— Rensch (4), Verheijen (5, MCZ), Verheijen/Schmutz.
Material.— , 9.xii.1968, Ruteng (Verheijen, RMNH no. 59905); (?) nestling, 15.vi.1969, Nunang, 600 m(Schmutz, RMNH no. 81458). Iris light brown, bill black, tarsus blackish flesh colour, toes and nails
blackish.
Eggs: 15 clutches of c/1 (6 ×) and c/2 (9 ×), collected in the months March (2), April (2), May (2), June(2), July (1), August (2), September (1), November (1), and three insufficiently dated; also a sample ofthree eggs without data, and a sample of six eggs without data, from Liang Bitu, Dano, Méngé, Madjung, Léwé, Ntéwéng (Mano), Puntu, and several clutches without locality (RMNH nos. 70657-70662,
70662a). The eggs are long-oval, white without markings.
Some measurements and weights:

RMNH no. 70668
RMNH no. 70669

16.5 × 11.1
17.2 × 10.9
16.5 × 10.5
16.8 × 10.8

0.05450.0516
0.05140.0522

Notes.— RMNH no. 59905 has noted on its label, that it was captured on a nest witheggs. This is interesting, as the bird is in the last stage of primary-moult, the 1st (outer)primary on both sides being short, basally in sheath. December is not mentioned as abreeding-month by Verheijen (1964).
Stresemann (1925) diagnosed the subspecies sumbawae in the following words:“durch den lebhaft violettblauen statt grünen Glanz der Oberseite und der Flügel stark
abweichend von C. e. neglecta und linchi; Innenfahne der Steuerfedern an der Basisweiss, nicht dunkel wie bei linchi. Flügel 94 mm”. Rensch (1931a: 538) commented: “Ichkann diese unlängst beschriebene Rasse durchaus bestätigen: der blaue und violette
Schiller der Oberseite und der Schwingen weicht stark von dem grünen Schiller deröstlich und westlich anschliessenden Rassen ab und ist nur bei jungen Exemplaren
weniger deutlich (bläulichgrün)”. More recently, Salomonsen (1983: 29) described C. e.
sumbawae as being: “Like nominate esculenta, but gloss of upper-parts duller and not122Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

greenish-blue, but darker blue...”. Note that violet has been eliminated from the spectrum, although Salomonsen examined at least partly the same specimens previously
studied by Stresemann and Rensch.
Gloss as a subspecific character must be treated with the utmost caution, for not
only may its colour depend on the angle under which it is viewed, but it is also strongly dependent on the state of the plumage. With only one adult bird from Flores, andnone from other parts of the range ascribed to sumbawae, I am obviously not in a good
position to discuss the characters and variation of the subspecies. Just the same, it isworth observing that the specimen is mainly glossy green, perhaps with a slight admixture of bluish on the primaries and rectrices, but still essentially glossy green. Frombirds of Java and Bali (C. linchi) it differs by the latter being a duller oil-green, and lacking white basal spots to the rectrices. C. e. neglecta seems to be a duller subspecies (cf.
Mayr, 1944a: 153); the few specimens available to me conform to this.
Although it is not directly relevant, I have, for the identification of these specimens,
consulted Somadikarta’s (1986) paper. The large amount of material studied by him,
gives his paper a great value. Still, there remain some unexplained facts. He has specifically separated C. esculenta and C. linchi, the latter inhabiting Sumatra, Java, Bali and
Lombok. C. linchi is diagnosed as being glossy green, and having a naked hind-toe; thewestern populations of C. esculenta, on the other hand, would be glossy blue, and havea feather tuft on the hind-toe. I find it illogical that the more eastern populations of C.
esculenta, like C. e. sumbawae, immediately adjacent to C. linchi, are glossy green, andhave a naked hind-toe. There is also a large distributional gap between the eastern
populations of C. esculenta, and the western populations included by Somadikarta into
this species. It might be more logical to treat these western forms as constituting a distinct species, the name of which would apparently be Collocalia cyanoptila, or C. marginata, if the relationship of the Philippine forms currently included in C. esculenta lies
with these western forms (cf. Dickinson, 1989). If this is accepted, it becomes once more
possible to unite C. linchi and C. esculenta to one species.
In several publications, Somadikarta has hinted that C. linchi is not conspecific with
C. esculenta, but I cannot find that he has published arguments, until 1986, when hedescribed sympatric occurrence of C. linchi and C. esculenta in Sumatra. I note that Somadikarta himself said of this: “I have not yet decided upon the relationship between
the esculenta taxa without tail spots to the west and north of Stresemann’s line and thosewith tail spots to the east and south”.
In his discussion of nominate C. e. esculenta, Salomonsen (1983: 34-35), refers to apaper by me (Mees, 1965: 172) of which he finds some of the conclusions “quite incomprehensible”. When a man of the calibre of Salomonsen reads in my paper the oppositeof what I intended it to mean, only one conclusion is possible, and that is that I havecompletely failed in writing comprehensible prose. For this I accept the full blame. What
I was trying to point out was that, within the large range ascribed to nominate C. esculenta before Ripley separated C. e. nubila (Celebes, the Moluccas, Aru Islands, the wholeof New Guinea), North Moluccan birds actually are darker on the underparts than specimens from New Guinea and the Aru Islands, thus conforming to Ripley’s diagnosis, butthat the material available to me from Ambon in the South Moluccas, the type-locality of
nominate esculenta, was also dark, like birds from the North Moluccas, so that if C. e. esculenta had to be split by this character, birds from the North and the South Moluccas

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

123

would remain together as nominate esculenta, whereas the birds with whiter underparts
from the Aru Islands and New Guinea, if the difference was considered sufficient for
expression in nomenclature, would have to be known as C. e. hypoleuca.
There are other aspects of swift classification in which Salomonsen and I do not see
eye to eye, for example in the recognition of Hemiprocne mystacea confirmata. It is mainly
a matter of subjective judgement, whether one wants to recognize in nomenclature an
average difference, but I regret that Salomonsen (1983: 11) ignored the specimen fromsouthern New Guinea with a wing-length of only 212 mm, which was the starting pointof the discussion (Mees, 1964b: 11-12). This gave in the small series measured by me forthe smaller subspecies confirmata a wing-length of 208-236 (220.3) mm, for the larger
subspecies mystacea a wing-length of 212-239 (226.9) mm. It remains my opinion, contrary to Salomonsen, that the considerable individual variation (ca. 13%), with an almost complete overlap in measurements, constitutes a more important argument
against recognition of confirmata, than the average difference of 6.6 mm (ca. 3%) constitutes in its support.
In the case of Collocalia vanikorensis, Salomonsen (l.c.: 89) cited me correctly, as far as
it goes, but he did not heed my conclusions. My comment on strange contradictions inMayr’s (1937) “thorough study” and “excellent review” (Salomonsen, l.c.: 84, 87) appears to have been too subtle for him. When consulting the distributional map presented by Salomonsen (l.c.: fig. 26), it should be kept in mind that he did not examine asingle specimen from either the North or the South Moluccas, so that his inclusion of
the North Moluccas into the range of the subspecies waigeuenisis is no more than aguess. The distribution shown in fig. 26 for waigeuensis (North Moluccas, Waigeu, Misool, Salawati), with on the New Guinea mainland opposite Salawati a different subspecies, granti, is zoogeographically impossible. It is perhaps well to point out that thename moluccarum antedates the names aenigma, heinrichi, waigeuensis, steini and granti; itis therefore unintentionally misleading when Salomonsen describes moluccarum as a“very ill-defined form”, for it is the only name of the several just listed that may survive
future revision.
Apus pacificus pacificus (Latham)[Hirundo] pacifica Latham, 1801, Suppl. Ind. Orn.: lviii – Nova Hollandia.
Collector.— de Jong (1).
Material.— None.

Notes.— The fact that a single bird from Flores (, 18.x.1929, Mboera) remains the
only specimen record of this species from the Lesser Sunda Islands, should not be interpreted as meaning that it is of rare occurrence. It should pass over the islands twice ayear, on its migration from north-eastern Asia to Australia and back, and may also overwinter in small numbers, as it does in Sumatra and Java (cf. Holmes, 1977; Mees, 1982a:
90). Observations from Timor, in March and September 1985, confirm the above picture(Andrew, 1986).
Verheijen (1961) saw a bird which he identified as “Apus pacificus (or A. affinis)” on
his visit to Palué. It is perhaps worth recording that on 24.ix.1984, when on board of the124

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

RV Tyro, I observed two individuals of A. pacificus rather high over the ship in the position 7°15’903S, 120°38’563E, which is a little south of Djampea, and about 110 km northof western Flores.
The number of records from Sulawesi (Celebes) is also limited. In this connection I
want to discuss a record by Maurenbrecher (in Coomans de Ruiter & Maurenbrecher,1948: 176) of several Apus affinis at Makassar. White (1976, and in White & Bruce, 1986:268) suggested that these observations concerned A. pacificus, but apparently he overlooked, at least he ignored, the statement in the same place, that Dr J.R. van Blom had
found A. affinis nesting in the fort of Balangnippa, Sindjai (Bonthain). This seems definite enough, and proves breeding in southern Celebes at least some forty years earlier
than the record listed by White & Bruce, that was based on observations made in 1978by Escott & Holmes (1980). In early October 1984, when staying in Makassar for a few
days, I had, from my hotel room, a view of several busily-attended nests of A. affinis,
placed under the edge of the roof of a nearby tall house. Evidently, the species is a wellestablished resident.
Alcedo atthis floresiana Sharpe
Alcedo floresiana Sharpe, 1892, Cat. Birds Brit. Mus. 17: 140, 151 – Flores.
Collectors.— Allen (at least 4), Weber (1, specimen not traced), Everett (2), Rensch (3), de Jong (1),
Schmutz.
Material.— , 20.ix.1969, Look (Schmutz, RMNH no. 81403); , 12.x.1969, Nggoer, west coast (Schmutz,
RMNH no. 97138); , 14.ix.1971, Look river, coast (Schmutz, RMNH no. 97124).

Notes.— This is a good subspecies, deeper blue than the migrant subspecies fromthe northern hemisphere, but, like them, with brown cheeks. It is confined to the Lesser
Sunda Islands. A. a. ispidoides, the resident subspecies of Sulawesi (Celebes), the Moluccas, etc., is deep blue, like floresiana, but has the sides of the head blue (sometimes, there
are traces of brown).
In a previous publication I pointed out that there is no proof of the occurrence of
this species (the subspecies ispidoides) on the western Papuan islands Misool and Salawati (cf. Mees, 1965: 198). As not all subsequent authors are aware of this (cf. Rand &Gilliard, 1967: 281; Forshaw, 1983: 70; Beehler, Pratt & Zimmerman, 1986: 144), I repeat
it here.
Forshaw (1983: 69, map) has in addition given the species a much too generous
range in mainland New Guinea, where it is known from the eastern part only. An interesting question is: why this should be so.
Measurements:




wing

tail

tarsus

72
72
71

31
30½
30

10½
11
9.2

entireculmen44.8
4943exposedculmen
37.8
37.3
38

Ceyx rufidorsa Strickland
Ceyx rufidorsa Strickland, 1847, Proc. Zool. Soc. Lond. 14 (1846): 99 – Malacca.

culmenfrom nostril
33.9
32
34

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

125

Collectors.— Allen (at least 2, cf. Sharpe, 1892: 180); Colfs (1); Everett (“a fine series from the lower
country”, cf. Hartert, 1898a: 42).
Material.— (?), 1880, Flores (Colfs, RMNH cat. no. 26).

Notes.— It is interesting that this species has penetrated as far east as Sumba, where
evidently it is not rare (there are seven specimens in Sutter’s collection, cf. White &Bruce, 1986: 486). The distributional maps published by Forshaw (1983: 128) and Ripley& Beehler (1987) must be corrected in this respect.
The question as to whether C. rufidorsa and C. erithaca are to be regarded as conspecific has, after more than forty years of investigation and discussion, become a matter ofpurely subjective personal preference. The latest revisers (Ripley & Beehler, 1987) treat
C. rufidorsa as a separate species; they say also that the species, thus defined, shows nogeographical variation worthy of nomenclatural separation, so that, evidently they nolonger recognize C. rufidorsa jungei Ripley from Simalur, still listed by Van Marle &Voous (1988: 129).
Pelargopsis capensis floresiana Sharpe
Pelargopsis floresiana Sharpe, 1870 (1 April), Monogr. Alcedinidae (pt. viii): pl. 36 and text – Flores.
Pelargopsis floresiana Sharpe, 1870 (June), Proc. Zool. Soc. Lond.: 62, 68 – Flores.
Collectors.— Allen (at least 5, cf. Sharpe, 1892: 104), Everett, de Jong (2), Verheijen/Schmutz.
Material.— , 12.viii.1968, Waé Mulu, 700 m (Verheijen, RMNH no. 65314); , 7.vii.1972, Ruteng, 1150m (Verheijen, RMNH no. 85277); , , 1.viii.1972, Bara-Latji, 200 m (Schmutz, RMNH nos. 80855, 81069).
Iris chocolate brown.

Notes.— Specimen no. 85277 crashed through a window pane at the, for this species, surprising altitude of 1150 m.
The description of Pelargopsis floresiana in the Monograph of the Alcedinidae waspublished at least two months earlier than that in the “Proceedings”, which is usuallycited as the original description (for the date of publication of Part viii of the Monograph, see Zoological Record for 1870: 41).
The statement by Warren (1966: 99), on a syntype of Pelargopsis floresiana: “Flores,
1869. Collected by A. R. Wallace”, cannot be correct. The bird in question, like the othersyntypes referred to, must have been collected by Allen in 1862.
The status of the subspecies floresiana requires discussion. According to Sharpe (1892),
the main character by which this subspecies could be distinguished from P. gurial wasthat it shows “the crown of the head strongly washed with green”. Sharpe’s P. gurial is
the subspecies from continental India, now known as P. c. capensis, the form the mostremote from Flores of the whole conglomerate, but the only subspecies with which
Sharpe compared floresiana. Subsequently Vorderman (1895b) distinguished his P. sasakfrom floresiana: “door het gemis aan groen, waarmede de kopvederen van floresiana
sterk bewassen zijn”910).

10)

By the absence of green, with which the feathers of the pileum of floresiana are strongly washed.

126

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

The character was dismissed by Hartert (1898a: 43) in the following words: “A fineseries of this interesting form from South Flores. Specimens from Lombok cannot be
separated from typical floresiana. Vorderman’s sasak is only an immature specimen, as
the characters given by him are found in immature birds before me”. Note that Hartert calls floresiana an interesting form, but provides no evidence that he has compared
it with any other subspecies. Oberholser (1909) had no material, either from Java or
Flores, and therefore his description of floresiana was based on Sharpe (1870, 1892),
although it is not clear where he got the statement about paler ochraceous lower parts,
as this is not a character mentioned by Sharpe. Stresemann (1913b: 338) had a specimen from Bali which he referred without explanation to floresiana. Forshaw (1983:
145) gave as only character to separate floresiana from javana, that the former would
have “the brownish cap and sides of the head washed with dull green”. This looks
much like a repetition of a character originally given by Sharpe, and I cannot confirm
it. White (in White & Bruce, 1986: 274) repeated the greenish wash and resurrected thecharacter of the pale shade of the ochraceous underside, which had become lost in
recent years. None of the four specimens from Flores examined by me has a greenish
wash on the head, neither are the underparts conspicuously pale, compared withjavana. A specimen from Bali (RMNH no. 10122) shows no trace of green on the cap
either.
Only Hoogerwerf (1963d: 150), perhaps the only author to have personally compared specimens from Flores (2) with specimens from Java (4), expressed doubt about
the validity of floresiana, although, owing to his limited material, he did not venture adefinite conclusion. With a much larger material than Hoogerwerf had, I still feel dubious about the validity of floresiana. This is mainly due to the large individual variation.
Tentatively, I would diagnose floresiana (as differentiated from javana) as follows: cap on
average darker; mantle and tail, and also the rump-patch somewhat deeper and purer
blue, less greenish blue.
Measurements:

wing

tail

Flores 3

147155, 156
160
146

94
90+, 97

Bali



89

entireculmen80
81, 83, 78
104
81

exposedculmen70
74, 75, 71½

culmenfrom nostril
63
65, 67, 63½

74

66

Halcyon chloris chloris (Boddaert)
Alcedo Chloris Boddaert, 1783, Table Pl. Enlum.: 49 – no locality, but = Bouro (Buru), from the references
to Buffon and Latham.
Collectors.— Semmelink, Allen, Martens, Colfs, Weber, ten Kate, Rensch (1), de Jong (2), Verheijen (4,MCZ), Pfeffer (2), Verheijen/Schmutz.
Material.— (?), 1862, Larantoeka (Semmelink, RMNH cat. no. 69); (?), 1862, Flores (Allen, ex coll. Sody,
RMNH no. 26948); 2 (?), 1880, Flores (Colfs, RMNH cat. nos. 115, 116); , xii.1888, Sikka (Weber, RMNH
cat. no. 124); (?), 1/3.v.1891, Groot Bastaard (ten Kate, RMNH cat. no. 126); (?), 27.ix.1955, BénténgDjawa (Verheijen, RMNH no. 61512); (?), x.1958, Ruteng (Verheijen, RMNH no. 61513); , 5.ix.1968,
Ruteng (Verheijen, RMNH no. 59808); , 14.x.1970, Ru’a, Ruteng (Verheijen, RMNH no. 65325); ,
1.ix.1971, Nunang, 650 m (Schmutz, RMNH no. 85266); , 2.x.1971, Tendo, Ruteng (Verheijen, RMNH

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127

no. 85267); , 14.i.1983, Nunang, 650 m (Schmutz, RMNH no. 81160); (?) [= ], not dated, West Flores(Verheijen, RMNH no. 65326).
Eggs (fig. 14c): c/4, 27.ix.1955, Bénténg Djawa, Kentjo (RMNH no. 70635); c/4, 10.x.1955, Mano (RMNH
no. 70636); c/4, 2.xi.1956, Poéng (RMNH no. 70637); c/4, x.1957, Tulang, Potjong (RMNH no. 70638);
c/2, 11.x.1957, Mano (RMNH no. 70639); c/3, 19.x.1958, Potjong (RMNH no. 70640); c/4, 16.ix.1959,
Puntu (RMNH no. 70641); c/1, 12.x.1959, Montjok (RMNH no. 70642). The eggs are white, only moderately glossy.
Some measurements and weights:

RMNH no. 70635

RMNH no. 70637

RMNH no. 70641

28.2 × 23.8
29.1 × 23.7
29.1 × 25.0
29.1 × 25.3
30.6 × 25.4
30.9 × 25.3
31.4 × 26.0
31.7 × 26.3
29.5 × 24.6
30.5 × 24.5
30.7 × 23.9
30.8 × 24.9

0.536
0.49830.56950.59170.60570.59610.65290.64920.53150.52690.48980.5221

Notes.— It remains to be seen whether birds from Java and Sumatra, variously
named cyanescens, laubmanniana and palmeri, are really different from the nominate
form.
Measurements:
2


wing
106, 116
111

tail
63½, 7066

tarsus16.2, 16
15.8

culmen from skull
47.3, 53
48.7

culmen from nostril
38.4, 42
39

Halcyon sancta sancta Vigors & Horsfield[Halcyon] Sanctus Vigors & Horsfield, 1827, Trans. Linn. Soc. Lond. 15: 206 – New Holland.
Collectors.— Everett (3), Rensch (7), de Jong (2), Verheijen/Schmutz.
Material.— , 5.v.1956, Ruteng (RMNH no. 61482); , 27.vi.1971, Langkas, 750 m (Verheijen, RMNH
no. 85269); , 3.xi.1971, Nisar, beach (Schmutz, RMNH no. 85268); , 2 , 15.ix.1982, Nunang, 650 m(Schmutz, RMNH nos. 81161, 81073, 81075); , 20.ix.1982, Nisar (Schmutz, RMNH no. 81074).

Notes.— Rensch (1931a: 533) observed of this species: “Ein australischer Zugvogel,
der sich während der Trockenzeit in grosser Anzahl an den Küsten – besonders in der
Mangrovezone – vereinzelt auch im Kulturlande und im Buschwalde (bis 200 m Höhe)findet”. The material listed here increases the vertical range considerably, to over 1000m (Ruteng).
I take this opportunity to correct an error in a previous paper where a specimen of
H. sancta is listed, collected on Ambon in January (cf. Mees, 1982a: 93). The specimen
was misidentified and is H. chloris! The removal of this record underlines that “oversummering” of H. sancta in its winter quarters is exceptional. There is no evidence tosupport White’s speculation that the species might occasionally remain and breed in
the Lesser Sunda Islands (White & Bruce, 1986: 278).

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Halcyon fulgida Gould
Halcyon fulgidus Gould, 1857, Proc. Zool. Soc. Lond. 25: 65 – Lombock.
Monachalcyon fulgidus gracilirostris Rensch, 1928, Orn. Mber. 36: 48 – Sita (700 m), Westflores.
Collectors.— Allen, Colfs, Weber, Everett, Rensch (2), de Jong (3), Verheijen (4, MCZ), Verheijen/
Schmutz.
Material.— (?), 1862, Flores (“collected for Mr Wallace” = Allen, RMNH cat. no. 1); (?), iv.1880, Flores(Colfs, RMNH cat. no. 2); (?), v.1880, Flores (Colfs, RMNH cat. no. 3); (?), 26.xi.1888, Bari (Weber, RMNH
cat. no. 5); skeleton, xi.1888, Bari (Weber, RMNH cat. a); (?), xi/xii.1888, Flores (Weber, RMNH cat. no.
4); , 30.x.1958, Ruteng (Verheijen, RMNH no. 61543); (?) ad., 1.xii.1958, Potjong (Verheijen, RMNH no.
85274); , 17.ii.1959, Ruteng (Verheijen, RMNH no. 61544); , 10.i.1969, Todo, 700 m (Verheijen, RMNH
no. 65315); , 12.ix.1971, Tjeréng, 850 m (Schmutz, RMNH no. 81071); , 21.x.1971, Rana-Kulan (Verheijen, RMNH no. 85276);  im., 22.x.1971, Rana-Kulan (Verheijen, RMNH no. 85273); (?), ix.1972, Mano
(Verheijen, RMNH no. 85275); , 15.ix.1976, Laring (Schmutz, RMNH no. 81027); (?), 1.v.1978, Golo Lehot (Verheijen, RMNH no. 81070).
Eggs (fig. 14d): c/2, Flores, without further data (RMNH no. 70643); c/2, 7.xii.1952, Potjong (RMNH no.
70644); c/1, 22.xii.1953, Potjong (RMNH no. 70645); c/1, 9.i.1954, Potjong (RMNH no. 70646); c/1,
29.x.1955, Bénténg Djawa (RMNH no. 70647); c/2, xii.1955, Bénténg Djawa (RMNH no. 70648); c/2,
ii.1958, Potjong (RMNH no. 70649); c/2, 1.xi.1958, Lalang, Todo (RMNH no. 70650); c/2, 14.ii.1959, Todo
(RMNH no. 70651); c/1, 28.ii.1959, Lamé (RMNH no. 70652); c/1, 29.xii.1959, Poéng (RMNH no. 70653);
c/1, x.1961, Soa (RMNH no. 70654); c/1, 12.x.1961, Soa (RMNH no. 70655); c/2, 3.iii.1962, Mataloko(RMNH no. 70656). The eggs are white, more glossy than those of H. chloris.
Some measurements and weights:

RMNH no. 70643
RMNH no. 70644
RMNH no. 70645
RMNH no. 70646
RMNH no. 70647
RMNH no. 70648
RMNH no. 70649
RMNH no. 70656

35.5 × 30.0
36.8 × 30.0
37.5 × 30.0
35.4 × 30.0
36.3 × 30.7
34.7 × 29.7
36.4 × 29.7
36.9 × 29.8
34.9 × 30.9
36.6 × 30.1
35.1 × 30.0
35.5 × 30.0

– (damaged)
0.98470.93840.905
0.940– (very large hole)
1.00230.98650.96720.92760.90340.9341

Notes.— Young birds differ from the adults by having the breast cream or pale
brownish, with sparse, very narrow grey bars, not pure white as in the adults; back andthe larger part of the wings dull black, without purple-blue gloss.
Forshaw (1985: 408) considered the subspecies gracilirostris to be: “probably notseparable”, and White (White & Bruce, 1986: 273) thought it unnecessary to recognise
this subspecies formally.
For comparison with the series from Flores, only one specimen each from Lombok
and Sumbawa were available to me (see table of measurements below). They show that
Rensch’s statement that Flores birds have the bill: “erheblich schmaler und länger” is
not correct, but the second part of the diagnosis: “sein First kantiger als bei Exemplarenvon Lombok” has some substance. The difference is subtle and variable, and I agreewith White, that it is altogether too slight for recognition in nomenclature.
Inter alia it may be recorded that the RMNH male from Lombok is a Wallace

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129

specimen, 1856; it is the bird figured by Sharpe (1868: pl. (99)). Warren (1966: 104)
listed a single BM specimen from Lombok, ex coll. Gould, as holotype, probably following Sharpe (1892: 296), who lists this bird as type. In the original description,
Gould makes no mention of the number of specimens he examined, although his reference to the trader Frank suggests that more specimens were then in the hands ofthat gentleman. In his next publication (Gould, 1860, pl. 46), however, it is clearlystated: “I published a description of this remarkably fine species of Halcyon in 1857,
taken from specimens [note plural! - GM] received direct from Mr Wallace ...”; further
down: “two beautiful specimens grace my collection”. On this basis I would assume
that all specimens collected by Wallace were seen by Gould, and therefore are syntypes: four in the British Museum, enumerated by Sharpe (1892), the RMNH specimen, and presumably also the male from Lombok in the Museum Heineanum (cf.
Cabanis & Heine, 1860: 163).
The RMNH specimen was purchased from Frank on 1 May 1857; considering that
Gould expressly states having consulted with Frank, and that he offered the description
to the Zoological Society in its session of 24 March 1857 (it was published on 14 July
1857), it is obvious that the specimen sold by Frank (as Alcedo n. sp.; for ƒ 25.-, a highprice!) is one of those examined by Gould.
Incomprehensibly, Schlegel (1874b: 16) claimed that this specimen was acquired in
1866, although he had listed it in the previous catalogue, published in 1863 (Schlegel,
1863b: 24-25)! Again, Schlegel (1864d: 21, 54, pl. 9 fig. 1) described and illustrated thissame specimen, adding that it was the only one in the collection.
Measurements:
Flores
4
6 (?)

Lombok
Sumbawa (?)

wing

tail

tarsus134130-140(135)
126-139(133.7)
132
128

106
110-116(112)
97½-112½(105.3)
107
108

22.5
21.5-22.5(22.1)
21-22.2(21.6)
21
22

culmen from
skull
51
49-53
(50.5)
48-53(50.5)
52.5
51culmen from
nostril
37
35-38(36.4)
34.7-38(36.3)
37.6
37Merops ornatus LathamM[erops] ornatus Latham, 1801, Suppl. Ind. Orn.: xxxv – Nova Hollandia = New South Wales (cf. Latham,Gen. Synops. Birds Suppl. 2: 155-156).
Collectors.— Allen, Colfs, Weber (1, ZMA), Rensch (2), Verheijen/Schmutz.
Material.— (?), iv.1880, Flores (Colfs, RMNH cat. no. 102); (?), v.1880, Flores (Colfs, RMNH cat. no. 99);
2 (?), 17.vi.1968, Ruteng (Verheijen, RMNH nos. 59805, 59806); 3 , 1.ix.1971, Nunang, 650 m (Schmutz,
RMNH nos. 85261, 85262, 85263); , 13.vi.1976, Poco Nernancang (Schmutz, RMNH no. 81303); ,
17.v.1976, Lingko Ncilor (Schmutz, RMNH no. 81315); , 19.v.1976, Lingko Ncilor (Schmutz, RMNH no.
81308).

Notes.— On Flores, this is a winter visitor from Australia; for a discussion see Mees
(1982a: 102-104).

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Merops philippinus Linnaeus[Merops] philippinus Linnaeus, 1767, Syst. Nat. (ed. 12) 1 (2): errata at end of volume, referring back to (1):183 – in Philippinis.
Collectors.— Allen, Weber (1), Everett (1), de Jong (2), Verheijen/Schmutz.
Material.— (?), xii.1888, Maumeri (Weber, RMNH cat. no. 41); , 2.iii.1969, Lita, 50 m (Verheijen, RMNH
no. 85265); , , 27.x.1969, Djonéng (Schmutz, RMNH nos. 65327, 65328); , 5.ix.1971, Nunang, 650 m(Schmutz, RMNH no. 85264). Stomach contents Hymenoptera (wasps) and Odonata.
Eggs: c/4, c/4, and 4 loose eggs, 24.x.1974, Maro-Kama, 50 m (RMNH nos. 85431, 85432, 85433). The
eggs are glossy white; they are the most nearly round eggs I know: in the bluntest one (21.5 × 20.3 mm),the width is 94.4% of the length.
Measurements and weights:

RMNH no. 85431

RMNH no. 85432

21.5 × 20.3
22.0 × 19.9
22.1 × 20.2
22.9 × 20.1
22.5 × 20.4
22.7 × 20.1
22.7 × 20.2
22.8 × 20.2

0.31690.3166
0.33520.33540.30980.27360.30750.2856

Notes.— The  from Djonéng is marked as having been collected at the nestinghole. The eggs from Flores were previously recorded by Mees (1982a: 105); a discussionof geographical variation and distribution of the species is given in the same place, andis supplemented by Mees (1986: 63-64). An additional record of nesting in Celebes isfrom near Bintoehan, 26.viii.1945 (Coomans de Ruiter, 1947).
Eurystomus orientalis orientalis (Linnaeus)[Coracias] orientalis Linnaeus, 1766, Syst. Nat. (ed. 12) 1: 159 – India orientali, arbitrarily restricted to Javaby Stresemann (1913a: 298).
Eurystomus orientalis connectens Stresemann, 1913, Novit. Zool. 20: 302 – Moa.
Collectors.— Semmelink, Allen, Everett (4), Rensch (1), de Jong (2), Verheijen (1, MCZ), Verheijen/
Schmutz.
Material. –(?), 1862, Larantoeka (Semmelink, RMNH cat. no. 68); , 23.i.1969, Naga, 100 m (Schmutz,
RMNH no. 97176); , 22.vi.1969, Nisar (Schmutz, RMNH no. 85218); , 1.ix.1969, Rowang, Ruteng,
1200 m (Verheijen, RMNH no. 65355);  juv., 26.i.1976, Nisar, 100 m (Verheijen, RMNH no. 81091); ,
27.i.1976, Nisar (Schmutz, RMNH no. 81086); , 30.i.1976, Waé-Wako, 200 m (Schmutz, RMNH no.
81087); 2 , 7.ii.1976, Nunang, 650 m (Schmutz, RMNH nos. 81088, 81090);  im., 12.ii.1976, Nunang(Schmutz, RMNH no. 81089).
Eggs: c/2, 5.xi.1960, Soa (RMNH no. 70634). The eggs are white, moderately glossy. This set was previously described by Forshaw (1993: 101, s. n. E. o. pacificus).
Measurements and weights:

RMNH no.70634

36.9 × 27.0
37.2 × 27.7

1.06731.2072

Notes.— White & Bruce (1986: 285-287), the latter also referring to a manuscript
by H. Scholtes, placed the subspecific name connectens, which had been used for theMees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

131

Lesser Sunda Island populations since its introduction by Stresemann (1913a: 302), in
the synonymy of E. o. pacificus. This may possibly be correct for some of the moreeasterly populations traditionally referred to connectens, but certainly not for birds
from the Lesser Sunda Islands: specimens from Flores are clearly deeper coloured
than pacificus and resemble birds of the nominate subspecies from Java, from which,in fact, I am unable to separate them. The Flores material shows no evidence of being
part of a “continuous cline”, as first suggested by Ripley (1942b), and repeated by
White & Bruce (1986: 285-287). A specimen from Sumba (leg. ten Kate) also belongs to
the nominate subspecies. Specimens from Timor are not available. As far as I canjudge, E. o. pacificus has not yet been recorded from the Lesser Sunda Islands, although at least on the eastern islands of the chain it is to be expected as a winter visitor from Australia.
The type specimen of E. o. connectens is from the island of Moa, east of Timor (seealso Greenway, 1978: 218). The specimen was examined by Mrs LeCroy. It is an adultbird with a blue throat and a red bill with a small black tip (this is noted on Kühn’s fieldlabel: “vermilion with black tip”). Mrs LeCroy (in litt., 18.iii.1996) concluded that thetype can be matched to specimens of nominate orientalis from Java, and not to the duller Australian pacificus. The specimen proves that nominate orientalis ranges at least as
far east as Moa, and also, of course, that E. o. connectens is a synonym of E. o. orientalis,and not of E. o. pacificus as suggested by White & Bruce.
Mayr’s (1944b: 119) reconstruction of the colonization of Australia by E. orientalis is
not necessarily wrong, but it was mainly based on recognition of “connectens” from theLesser Sunda Islands as “an exact intermediate between orientalis and pacificus”, henceon (in my opinion) faulty systematics.
Rensch (1931a: 528) commented on the large size of some of his specimens. The
specimens examined by me are not exceptionally large, but are in the upper half of therange of variation ascribed to their subspecies.
Of this small series, only one (RMNH no. 65355) had the bill entirely red; all others
show it at least partly black or dusky, which is a sign of immaturity. The juvenile bird
lacks the blue throat patch, and its measurements suggest that it is not yet fully grown.
In one or two of the others the blue throat patch is not yet fully developed, but theirmeasurements indicate that they are fully grown. The reason why Father Schmutz collected several immature specimens, is that I asked him to pay special attention to birds
in dull plumage, hoping that this would lead to the discovery of E. o. pacificus.
Hoogerwerf (1969/1971: 466) believed bill-colour to be a subspecific character. Discussing his observations in Udjung Kulon, the westernmost peninsula of Java, hewrote: “Probably most records relate to the dark-billed migratory race calonyx, because
in only two cases was the breeding subspecies orientalis identified by its ivory yellowish bill”. I have already mentioned that a dark bill is a character of immaturity, but
Hoogerwerf’s statement that nominate orientalis would have an ivory yellowish bill
also requires some comment. Adults of all subspecies have red bills (based on personal observations and on literature records). My observations include nominate orientalis in Java with red bills. Actually, in my diary notes from Java, I found a single reference to a bird which seemed to have an orange-yellow bill; whether this was just due
to a trick played by the light or actually an intermediate stage, I cannot say. Even orange-yellow is far from ivory yellowish. Perhaps Hoogerwerf examined old museum

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Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

specimens, in which the red bills tend to bleach to a condition that may well be calledivory yellowish, but that does not explain his field-observations.
Stresemann (1939: 346; 1940a: 422) listed Eurystomus orientalis as an example (actually his best example) of ‘Doppel-Einwandering’ in Celebes: nominate orientalis fromthe Philippines, and connectens from the Lesser Sunda Islands. This case falls with there-identification of the Lesser Sunda birds as nominate orientalis. Actually, as waspointed out by White & Bruce (1986: 286), there is no proof yet that E. orientalis breeds
in Sulawesi (Celebes), although I think that it does. Whether the specimens from southern Sulawesi (Celebes) and Muna ascribed by previous authors to ‘connectens’ are migrant visitors of pacificus or dull, immature individuals of orientalis, remains to be seen.
Measurements:
4
3
(?)
 juv.

wing
189-200(194.3)
190, 196, 197
199
186

tail
91½-101(95.5)
95, 99, 95
101
87

entire culmen
34-35(34.4)
36, 36, 34
33
31

exposed culmen
25-27(26.2)
30, 27, 26
24
23

Dendrocopos moluccensis grandis (Hargitt)
Iyngipicus grandis Hargitt, 1882, Ibis (4) 6: 45 – In insulis Malayanis “Lombock” et “Flores”, restricted toLombok by Rensch (1931a: 546).
Collectors.— Allen (several), Weber, Everett, Rensch (5), de Jong (3), Verheijen (1, MCZ), Verheijen/
Schmutz.
Material.— , 23/25.xi.1888, Reo (Weber, RMNH cat. no. 2); , xii.1888, Maumeri (Weber, RMNH cat.
no. 1); , 7.vii.1969, Nunang, 650 m (Schmutz, RMNH no. 81442); , 5.vi.1978, Poco Nernancang (Verheijen, RMNH no. 81406); , 26.vi.1978, Waé Rukus (Verheijen, RMNH no. 81433); , 23.v.1978, Golo
Léhot (Verheijen, RMNH no. 81441); , 6.vi.1978, Waé Hiam (Verheijen, RMNH no. 81451).
Eggs: c/2, 13.v.1956, Tado (RMNH no. 70673); c/2, 24.vi.1960, Potjong (RMNH no. 70674). The eggs are
glossy white.
Measurements and weights:

RMNH no. 70673

RMNH no. 70674(the second egg is badly damaged, not measurable)

18.0 × 13.9
18.2 × 13.9
19.3 × 14.5

0.12400.12270.1462

Notes.— The species is apparently not known from Bali. There is a conspicuous
difference in size and proportions between D. m. grandis and nominate D. m. moluccensis from Java. Note that grandis has a proportionally much longer bill and tail thanmoluccensis. The large size and long bill of this subspecies may be at the root of theerroneous records of D. macei analis from the islands between Sumbawa and Flores by
Hoogerwerf (1956).
Measurements:
Flores (D. m. grandis)

West Java (D. m. moluccensis)

2
5
10 

wing
81, 8684, 84½, 85,
87, 8773-77(74.5)

tail42½, 45
45, 44, 46, 46,
48½
29-32½(30.9)

culmen from skull
17.9, 20.0
20.4, 19.8, 20.3,
20.3, 21.3
15.0-17.0(15.6)

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133

East Java (D. m. moluccensis)
7630½
The smaller of the two males from Flores is probably immature.

15.5

Pitta elegans concinna Gould
Pitta concinna Gould, 1857, Proc. Zool. Soc. Lond. 25, 1857 (14 July): 65 – Lombock.
Pitta concinna everetti Hartert, 1898, Novit. Zool. 5: 459 – Alor.
Collectors.— Semmelink, Allen, Everett (“a fine series”), de Jong (3), Verheijen (1, MCZ), Verheijen/
Schmutz.
Material.— (?), 1862, Larantoeka (Semmelink, RMNH cat. no. 3); (?), 13.vi.1973, Golo-Karot, Borong(RMNH no. 85295).
Eggs (fig. 14e): c/2, 27.i.1953, Nempong (RMNH no. 70675); c/1, 28.vi.1955, W. Rembong (RMNH no.
70676); c/1, ca. 20.iii.1957, Djinggor (RMNH no. 70677); c/3, iv.1957, Nunuk (RMNH no. 70678); c/3,
iv.1959, Bénténg Djawa, Djinggor (RMNH no. 70679); c/2, 20.iv,1959, Waé Tua (RMNH no. 70680); c/1,
1960, Djinggor (RMNH no. 70681); c/3, 21.ii.1961, Kisol (RMNH no. 70682); c/3, 1.vi.1962, Soa (RMNH
no. 70683).
There is some variation in the eggs. The ground colour is white, and they are either densely mottled withnot very dark pinkish brown markings, or with sparser, more sharply defined, dark grey or blackish
spots.
Measurements and weights:

RMNH no. 70677
RMNH no. 70678

RMNH no. 70679

RMNH no. 70680
RMNH no. 70681
RMNH no. 70682

RMNH no. 70683

25.5 × 21.0
0.251024.9 × 20.8
0.263025.4 × 21.2
0.2875the third egg is badly damaged
27.1 × 21.7
0.328427.4 × 21.8
0.352827.6 × 21.7
0.338426.4 × 20.7
0.239326.8 × 21.4
0.293227.1 × 22.0
0.335926.6 × 20.8
0.327527.1 × 21.0
0.3259the third egg is badly damaged
28.4 × 22.0
0.346129.0 × 21.5
0.337329.0 × 22.2
0.3521

Notes.— As in so many instances, the first specimen of P. e. concinna was in the Forsten collection from Bima, ca. 1842. This subspecies was described almost simultaneously by Gould and by Verreaux. Mayr (1979: 328) records as publication date of Pitta
concinna Gould, April 1857, for Pitta Mathilda Verreaux, July 1857, but according to Duncan (1937: 82), the actual date of publication of P. concinna is 14 July 1857, so that its
presumed priority over P. Mathilda requires further investigation.
Meagre as the available material is, it supports White & Bruce (1986: 295) in theirrejection of everetti from Alor, exclusively based on having a supposedly longer bill thanconcinna.
Measurements:
Flores2 (?)
Adonara (?)
Alor
2 (?)

wing
99½, 104
100
105, 105tail34½, 37
36
33, 37tarsus38½, 3737
37½, 38½

entire culmen
25, 27.3
27.8
25.6, 27.8

exposed culmen
20.4, 21.1
2120.3, 24

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Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Pitta elegans maria Hartert
Pitta maria Hartert, 1896, Bull. Brit. Orn. Cl. 5: xlvii – Mountains of Sumba.
Collectors.— Everett (1), Verheijen.
Material.— , 2.xii.1968, Ruteng, 1150 m (Verheijen, RMNH no. 85296). Iris grey-brown, bill brownish
black, legs pinkish pearl colour.

Notes.— A male of this, the Sumba subspecies, was taken by one of Everett’s nativecollectors in South Flores, in November 1896 (cf. Hartert, 1897b: 526-527 and 1898d: 470471). In spite of Hartert having obtained Everett’s personal assurance that the specimen
was correctly labelled, an element of doubt seems to have remained whether it was
actually from Flores and the record was completely ignored by later authors.
Now a second specimen of P. e. maria from Flores is available. The identification restson the following characters: the superciliaries are pale over their whole length, very palebuff in their anterior part, almost white posteriorly (in P. e. concinna, the anterior superciliaries are brown); only the fifth primary, on each side, has a small white speculum, allother primaries are completely black; there is only a very small patch of greyish black on
the lower breast, above the light red of the belly. In the colour of its superciliaries, thespecimen resembles P. e. elegans, from which, however, it differs by smaller size, thebroad terminal tail-band (much narrower in elegans), the reduction of the wing-speculum, and the slightly more extensive black throat-patch. Of particular interest is a noteby Father Verheijen on the label: This bird has never before been found at such a highaltitude and is unknown to the local population; normally from 0-500 m.
Whether P. e. maria is actually a migrant, or only shows some irregular dispersal, is
not yet clear from the limited evidence available, but very likely it is as least a partial
migrant (‘Teilzieher’). Note that the geographically adjacent subspecies P. e. elegans, a
breeding-bird of Timor and probably Kisar (, 15.xii.1897, leg. Schädler, RMNH cat. 7),
is certainly migratory.
As the migration of P. e. elegans has been little studied, I have brought together its
records in a map (fig. 9). On present evidence, the Sula Islands constitute its main nonbreeding quarters; specimens were obtained there by Allen (at least 3), Hoedt (5), Doherty (1), Menden (1) and an anonymous collector (1). The remaining records, fromBuru (2), Boano (1), Ternate (1) and Tahulandang or Tagoelandang (1, cf. Meyer &Wiglesworth, 1898: 355, specimen MTD no. C 13526), are all due North of Timor, and all
indicate migration in the same direction.
Distribution of Pitta e. elegans by month:
s.d.
I
II
III
Timor
22
2
Kisar
Buru
Boano
1
Sula Isl.
4
Sula Besi
Sula Mangoli
Taliaboe
TernateTahulandang
-

IV
2
-

V
1
-

VI
1
-

VII
-

VIII
1

IX
-

X
1
1
-

XI
2
4
1
-

XII
1
1
-

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Fig. 9. Pitta elegans, breeding area on Timor and Kisar and records from non-breeding quarters.

135

136

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Documentation for the map and the table of P. e. elegans:: Timor:  and , xi.1828,
juv., vi.1828 (S. Müller, RMNH 88875, 88876, 88877, types of the species); , 24.xii.1931,
Koepang (Stein, ZMB); 11 , 11, six of these in juv. plumage, 9/28.i.1932, Tjamplong(Stein, AMNH and 3 ZMB); , 4-11.1932, Noilama (Stein, AMNH). Kisar: , 15.xii.1897(Schädler, RMNH). Buru: , , 24.vi.1922, Mada Range, 5000’ (Pratt Bros., AMNH, cf.
Hartert, 1924). Boano: , received in 1863 (Hoedt, RMNH 88878). Sula Islands, Sula
Besi (Sanana): 3 , 1, 18, 18, 19 and 21.xi.1864 (Hoedt, RMNH). Sula Islands, Sula
Mangoli: , 26.xi.1864 (Hoedt, RMNH); , x.1897 (Doherty, AMNH). Sula Islands, Taliaboe: , 6.x.1938 (Menden, MTD, cf. Eck, 1977 and in litt.). Ternate: , 5.v.1861 (Bernstein, RMNH). Tahulandang (Tagoelandang): (?), 6.viii.1894 (MTD, cf. Meyer & Wiglesworth, 1898: 355).
The above enumeration shows how important it is not to rely on published records
alone. Hitherto I believed Mayr’s (1944a) to be a complete record of the collections
made by Stein, but he listed only five specimens, when Stein actually collected 24. Allen
spent two months on the Sula Islands in 1858, but as his specimens are only dated to theyear of collecting and the period of his stay is not known, his material cannot contributeto the table, although it does provide supporting evidence for the periodically common
occurrence of P. elegans on the islands. In this stage, negative information is also valuable: the CSIRO collections from Timor, obtained in de months April, May, August andOctober, do not contain any P. elegans. Note, however, that these collections are from
Portuguese Timor, where there would be less suitable habitat than in the western halfof the island.
As White & Bruce suggest that P. e. elegans is absent from Timor in the months JulyAugust, it is worth recording that a nestling from Timor is dated June 1829 (S. Müller,
RMNH no. 88877). Of course, with this old material, lacking original labels, there is always a possibility of error. It seems to me likely, that P. e. elegans is a ‘Teilzieher’, andthat at least part of the population is sedentary.
Rensch (1931a: 548) still believed that the resemblance between birds from Timorand birds from the Sula Islands, etc. was due to: “eine parallele Entwicklung ... und daßspäterhin wohl auch noch Kennzeichen gefunden werden, welche die nördliche elegansGruppe von den Timor-Stücken trennen”. Stresemann (1939: 408, Abb. 18) gave elegans
as an interesting example of a subspecies that had expanded across the range of a different one (vigorsi). Mayr (1944a: 154) compared specimens from Sula-Mangoli (1) andBuru (2) with a series from Timor, and failed to find characters by which to separate
them. He says nothing about migration. Again, Mayr (1979: 328) lists the islands fromwhich elegans is known, without any suggestion of migration.
Van Bemmel (1948: 356) included Ceram in the list of islands from where P. e. elegans
has been recorded. White & Bruce (1986: 295) mentioned in their text that they had been
unable to find on what the Ceram record was based, but nevertheless included Ceram
without a query in their enumeration of islands. In Dr van Bemmel’s card-index (which
formed the basis for his paper), I read that the record of Ceram was based on Salvadori(1881: 391). Looking up the reference, I only found Ceram mentioned in the following
short sentence: “Boano, ad occasum Ceram (Hoedt)”, i.e.: Boano, to the west of Ceram.
It is a safe assumption that, due to superficial reading, van Bemmel’s record of Ceram
is based on this short sentence. Therefore, Ceram has to be deleted from the list of islands whence P. e. elegans is known.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)
Measurements:


wing
105

tail
32½

tarsus38

137
entire culmen
24

exposed culmen
21

Mirafra javanica parva Swinhoe
Mirafra parva Swinhoe, 1871, Ann. Mag. Nat. Hist. (4) 7: 257 – Flores.
Collectors.— Allen (at least 3), Everett (2), Rensch (2), de Jong (1), Verheijen.
Material.— 5 , 9.iii.1971, Maro-Kama, Borong, 50 m (Verheijen, RMNH nos. 66160, 66161, 85030, 85327,
85328).
No eggs.

Notes.— Since I discussed these specimens in connection with M. j. aliena of eastern
New Guinea (Mees, 1982a: 110), five additional specimens of M. j. parva have become
available: four from Sumba and one from Sumbawa (leg. Mees, ix.1984). These specimens are in good plumage, and therefore suitable for a meaningful comparison withother subspecies.
First, however, differences between this fresh material, and the specimens from
Flores as well as two old specimens from Sumba (leg. Ten Kate, vi-vii.1891), have to be
described and evaluated. The fresh birds (there is no difference between specimensfrom Sumba and Sumbawa) have the upper parts with very little brown, the feathers
have broad greyish, whitish, or just a trifle brownish-grey margins; the worn birds from
Flores, on the other hand, have the pale margins narrower, and the aspect of the uppersurface is a little more brownish. I have no difficulty in ascribing the difference to thestate of the plumage. The underparts show little or no difference. More different are Ten
Kate’s specimens from Sumba. On the upper parts they are close to the specimens from
Flores, but, although they are less worn, the feathers look blacker. The puzzle is, however, that one of these birds especially has the outer margins of the primaries, as well as
the wing-coverts and the alula, darker brown than in all other birds. Ten Kate’s specimens have been prepared from alcohol (as indicated on their labels; see also Büttikofer,
1892: 197), but I do not believe that that is the cause of the difference, as alcohol tends tobleach out, not to darken and intensify colours. There remain two possibilities: either
individual variation is greater than the modest material available to me suggests, orthere is, within Sumba, some geographical variation.
Nominate M. j. javanica of Java (also southern Borneo and Bali, from where I havenot seen material) is well-differentiated from M. j. parva: the feather edgings of the upper parts, including the wings, are much browner, not greyish, and the whole undersurface is buffy. In M. j. parva, the throat and the lower abdomen are creamy white, anda pale buffy colour is confined to the breast and flanks. There is also a difference inmeasurements, javanica being a little larger, especially the bill, which is not so much
longer as thicker, the mandible deeper.
Measurements:
Flores (M. j. parva)
5
Sumba (M. j. parva)


wing

tail

tarsusentire culmen

exposed culmen

70-72(70.6)

42-47(44.6)

19.2-21.3(20.5)

12-14(13.2)

11-11.5(11.1)

72

45

21

14.6

11.9

138

69
2 (?)
72, 72
Sumbawa (M. j. parva)
69
Western Java (M. j. javanica)
10 
73-77
(75.0)

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)
43
43, 45½

21.2
21, 21.6

14
13.8, 15

11.5
10.9, 12.2

44

19.8

13

10.3

42-49
(44.6)

22-24(23.2)

13.5-16.2(15.2)

11-13.8(12.3)White & Bruce (1986: 297) have included Lombok into the range of nominate M. j.
javanica. This was obviously based on Mayr (1944a: 154), who had a single specimenthat he considered to agree better with javanica than with parva. As no measurements ofeither wing or bill are provided, this identification required confirmation. This was provided by Mrs LeCroy (in litt., 18.iii.1996): “The Lombok specimen of Mirafra javanica is
identified in our collection as parva and seems to agree with that race. The measurements I made are as follows: wing 73, tail, 46, tarsus 22, exposed bill 12, entire culmen
14.5, bill depth at front of nares 6”. Although the measurements by themselves are notquite conclusive, Mrs LeCroy’s opinion that the specimen is parva should be accepted.
No material of M. j. timorensis from Timor (Dilli) and Savu has been available, sothat for its characters I must refer to Mayr (1944a: 154). If the measurements taken by
Mayr are comparable, timorensis, with wing 8  72-77 (73.9) mm, 4  69-73 (71.4) mm,
is also larger than parva. Further, it is of interest to mention weights here. Sumba:  17.3,
 17.8, 2 (?) 18.7, 18.8 g. Sumbawa:  16.5 g. In contrast, in a large series (27 , 26 , 3(?)) of M. javanica subsp. from northern and north-western Australia (Western Australia
and Northern Territory), I found weights of 19.2-26.6 g, the minimum being above themaximum of parva. This confirms that parva is not close to Australian birds.
Hirundo rustica gutturalis Scopoli
Hirundo (gutturalis) Scopoli, 1786, Del. Flor. Faun. Insubr. 2: 96 - nova Guiana (error!) = Antigua, Panay.
Collector.— Weber (2).
Material.— (?), skeleton, xii.1888, Maumeri (Weber, RMNH cat. no. c, cf. van Oort, 1907: 207).

Hirundo tahitica javanica Sparrman
Hirundo javanica Sparrman, 1789, Mus. Carlsonianum, fasc. 4, no. 3, pl. 100 – Java (reference not verified).
Collectors.— Weber (3, two of which in alcohol), Rensch (4), Verheijen (eggs only).
Material.—  ad., xii.1888, Sikka (Weber, RMNH cat. no. 42).
Eggs: c/4, 8.xii.1957, Nanga pandu (RMNH no. 70684). The eggs are creamy white with dark purplishbrown spots, concentrated around the blunt end.
Measurements and weights:

RMNH no. 70684

15.4 × 11.9
16.6 × 12.5
18.0 × 13.3
18.7 × 12.9

0.0616
0.07130.084-

Notes.— Traditionally, the Lesser Sunda Islands have been included into the rangeof H. t. frontalis, but Hoogerwerf (1965: 251) concluded that birds from these islands

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

139

belong to H. t. javanica. My own observations support Hoogerwerf (cf. Mees, 1975b:
129). Hoogerwerf also reviewed the question of the validity of H. t. frontalis, a matterthat remains to be cleared up. Hoogerwerf cautiously suggested that frontalis might bedarker on the underparts. Curiously, this is the opposite of what Mayr (in Stresemann,
1940a: 131-132) found: “New Guinea birds have very pale and uniform colored bellies”.
I cannot confirm either of these opinions, to me the two subspecies seem identical inplumage, but frontalis averages a little larger.
An explanation for the contradictory statements of Hoogerwerf and Mayr is provided by Schodde & Mason (1999: 667-668), who note the occurrence of pale-bellied
populations in southern New Guinea, sufficiantly differentiated from frontalis to be designated by name: H. t. albescens. It is likely that the specimens compared by Mayr belonged mainly to this subspecies. No material from southern New Guinea has been
available to me and it is unlikely that any was seen by Hoogerwerf.
Cecropis striolata striolata (Schlegel)
Hirundo striolata Schlegel, 1844, Krit. Übers. Europ. Vögel: 42 – Java.
Hirundo daurica rothschildiana Rensch, 1931, Mitt. Zool. Mus. Berlin 17: 550 – Mborong, Flores.
Collectors.— Allen, Everett (1), Rensch (1), Sutter (1), Verheijen (3, MCZ), Verheijen/Schmutz.
Material.— (?) juv., 8.iii.1971, Maro-Kama, Kisol, 50 m (Verheijen, RMNH no. 66154);  juv., 9.iii.1971,
Maro-Kama, Kisol, 50 m (Verheijen, RMNH no. 66155); , 3.xi.1971, Waé Radja, 20 m (Schmutz, RMNH
no. 85323).
Eggs: 33 clutches of 1 (11 ×), 2 (7 ×), 3 (14 ×) and 4 (1 ×) eggs each, collected in the months November (22)and December (10), and one undated, from Mataloko, Méngé and Potjong (RMNH nos. 70685-70717).
The eggs are elongated, plain white, without gloss.
Some measurements and weights:

RMNH no. 70712

RMNH no. 70713

22.0 × 14.6
22.4 × 14.6
22.6 × 14.5
22.3 × 15.2
22.6 × 15.4
22.6 × 15.5
23.0 × 15.1

0.12120.11980.11980.14170.14010.13740.1322

Verheijen (1964) gave as an explanation for the very short breeding season apparent
from the above data, that this species, “which likes to collect nesting material from rainpools, obviously prefers the beginning of the rainy season”. It is not so obvious to me,
why it should not be able to collect nesting material and breed at the end of the rainyseason, in line with most other breeding birds, when certainly there would be no shortage of drying mud.
Notes.— For rejection of the subspecies rothschildiana, cf. Vaurie (1951). With only
one adult bird from Flores available, I cannot add much, but the comparative measurements, given below, show that it falls entirely within the range of variation of ten birdsof the same sex from Java. The subspecies rothschildiana was separated by Rensch exclusively on the basis of supposed small size. In all birds, the tail-measurements areminimum measurements, as the slender tips of the outer rectrices are usually worn or
damaged.

140

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Usually, Temminck & Schlegel (1847: 33) are cited as authors of Hirundo striolata, but
fortunately Schlegel’s description, cited above, has priority (cf. Hartert, 1922; Mees,
1971b: 238). The reason why I call this fortunate, is that Rüppell (1845: 18, Taf. 6) described a Cecropis striolata from Africa, which according to a footnote added by himself,
is identical with C. a. abyssinica (Guérin-Méneville). In spite of the footnote, I believe
Cecropis striolata Rüppell to have been validly described (not merely cited in the synonymy). Boie (1844: col. 174, 176) cites both names, Cecropis striolata Boie and Cecropis
striolata Rüppell, but fortunately, both are nomina nuda here.
Not a nomen nudum is Acanthylis coracina Boie (1844: col. 167). It is the nomenclaturally correct name for the species currently known as Chaetura leucopygialis (Blyth, 1849),
with the junior synonym Acanthylis coracina Bonaparte, 1850.
Measurements:
Flores  juv.
(?) juv.
Java
10 

wing
125
115
116
123-129(126.6)

tail
90
67
66
90-101(95.5)

tarsus14½
13
15
14.3-16(15.2)

entire culmen
12
11
10½
11-13
(12.4)

exposed culmen
8.7
77
7.8-8.9(8.4)

Petrochelidon nigricans timoriensis Sharpe
Petrochelidon timoriensis Sharpe, 1885, Cat. Birds Brit. Mus. 10: 192 – Timor.
Collector.— Allen (1).
Material.— (?), 1862, Flores (Allen, BM no. 88.7.12.352).

Notes.— The first to record this species from Flores was Wallace (1864: 485), in abare list. Sharpe’s (1885) inclusion of Flores into the range of this subspecies was aguess, as he had not seen material. Later he wrote: “Mr Wallace... records it as occurringin Flores; but we have never seen a specimen from the last-mentioned island, nor was
there one in Mr Wallace’s collection when it passed into the hands of the Museum”(Sharpe & Wyatt, 1887: (529)). This was followed by Hellmayr’s (1914: 64) remark:“Wallace gibt diese Schwalbe auch für Flores an, doch liegen keine Belegstücke vondieser Insel vor”. Rensch (1931a) did not include the species in the avifauna of Flores,
but White (1936) discovered the above specimen in the BM, where it was received with
the Tweeddale collection in 1888, the year after Sharpe wrote, and this record has been
accepted by Peters (1960: 119). Later, White (in White & Bruce, 1986) thought that the
specimen might be mislabelled. I have examined the specimen, that has an original
Wallace label, with pre-printed the first three digits of the year: 186, to which in Allen’s
hand is added a 2, as well as the locality Flores. My reasons for acceptance of this record
are given in the introduction.
Measurements:
Flores (?)
Timor (?)
(?)

wing
92
93
94

tail
42
38
41

moult
no
heavy, wings and tailno

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

141

Motacilla flava simillima Hartert
Motacilla flava simillima Hartert, 1910, Vögel paläarkt. Fauna I: 289 – Kamtschatka.
Collectors.— Allen (at least 1, cf. Sharpe, 1885: 520; listed as Motacilla flavescens by Wallace, 1864: 485),
Weber (1), de Jong (1).
Material.— None.

Notes.— I have not examined material from Flores, but the bird collected by deJong was identified by Rensch (1931b: 399) and Voous (1950: 651) as M. f. simillima,
which is the commonest of the several subspecies wintering in the Indo-Australianregion.
Motacilla caspica subsp.
Collectors.— Colfs, Everett, Verheijen.
Material.— (?), iv.1880, West Flores (Colfs, RMNH without number); (?), 4.xi.1969, Ruteng (Verheijen,
RMNH no. 65399).

Notes.— In its huge range across Eurasia, from the British Islands to Japan, this species shows only minor geographic variation, mainly a clinal decrease in tail-length from
West to East. The recognition and delineation of subspecies is difficult and often somewhat arbitrary.
Earlier in last century, birds inhabiting the eastern part of the range went under thename melanope, and individuals wintering in the Indo-Australian region were usuallyassigned to this subspecies. Then Vaurie (1957) revised the species; he suppressedmelanope as a synonym of nominate ”cinerea” but accepted a subspecies M. c. robusta
from north-eastern Asia and Japan. This is the subspecies one would expect to be thecommonest, if not the sole, winter visitor to the Lesser Sunda Islands. Van Marle &Voous (1988: 193) and Dickinson et al. (1991: 364) placed all records from, respectively,
Sumatra and the Philippines, in robusta. As far as I can make out, this was done without renewed investigation. It might be argued that the Flores specimens are robusta,
and their measurements (rather short tails) do not contradict this. However, disagreement over the subspecific classification continues. Some authors would not recognize
any subspecies on the Eurasian mainland and Japan, others would suppress robusta
and re-instate melanope for birds from eastern Asia, and finally, some would recognize
both melanope and robusta as valid subspecies (Roselaar in Cramp, 1988: 453). One canagree with Mauersberger (1983: 61) about ”die Revisionsbedürftigkeit der bisherigen
Gliederungen”.
It is generally known that for this species the name Motacilla Cinerea, published in
an obscure anonymous pamphlet ascribed to Tunstall, has been validated (ICZN Opinion 882). Peculiarly, neither in this Opinion, nor in the discussion preceding it, has it
been pointed out that M. Cinerea Tunstall is a nomen nudum, not validated by being
provided with English and French vernacular names [cf. ICZN, 1985: art. 12 (c)]. The
remark above the list of species: “Nomina....Gallica verd ex ornithologia Brissonii
plerumque decerpta sunt”, is too vague to be construed as an indication.

142

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Measurements:
(?)
(?)

wing

tail

tarsus

80
80

79+
89

20
18.8

entireculmen15+
16.2

exposedculmen
12.8+
12.9

hind
claw
8.1
6.7

Anthus novaeseelandiae albidus Stresemann
Anthus richardi albidus Stresemann, 1912, Novit. Zool. 19: 316 – Süd-Flores.
Collectors.— Everett (3), Rensch (4), Verheijen (5, MCZ), Verheijen/Schmutz.
Material.— , 1.vii.1969, Tjumbi, Rahong (Verheijen, RMNH no. 81418); , 8.viii.1969, Ruteng (Verheijen, RMNH no. 65392); , 8.iii.1971, Maro-Kama, south coast (Verheijen, RMNH no. 85086); ,
12.x.1971, Langkas (Verheijen, RMNH no. 85085); , 28.vi.1977, Lumu (Verheijen, RMNH no. 81419).
Eggs: 37 clutches of 1 (8 ×), 2 (23 ×) and 3 (6 ×) eggs, collected in the months January (1), February (2),
May (1), June (4), July (3), August (4), September (7), October (10), and November (4), and one undated(RMNH nos. 70718-70752, 73511, 73512). The eggs are greyish white, with irregularly-shaped dullbrown primary spots and light grey secondary spots.
Some measurements and weights:

RMNH no. 70719

21.6 × 15.9
0.153621.8 × 15.7
0.1595RMNH no. 70720 20.6 × 15.6
0.160721.5 × 15.5
0.151
RMNH no. 70723 21.6 × 15.9
0.143322.3 × 16.0
0.1521RMNH no. 70724 21.0 × 15.0
0.1496
21.6 × 14.7
0.137122.0 × 15.2
0.1564RMNH no. 70746 21.3 × 16.2
0.156922.0 × 15.9
0.1566RMNH no. 70748 19.9 × 14.9
0.137820.0 × 15.1
0.1365Eggs of A. n. medius from Ndao near Roti, for comparison(see notes):
RMNH no. 76720 23.0 × 15.8
0.153
23.2 × 15.6
0.153323.6 × 16.0
RMNH no. 76721 23.6 × 15.6
0.1486RMNH no. 76722 22.8 × 16.0
0.162523.2 × 16.0
0.161
23.3 × 15.7
0.16

Notes.— Compared with a small series of A. n. medius from the islet of Dao or Ndao
near Roti (previously recorded by Mees, 1975b: 129), the present specimens are conspicuously white below and can be separated at a glance.
Verheijen (1976: 14) observed that eggs of the subspecies A. n. medius from Ndao
were larger than eggs of A. n. albidus from Flores and of A. n. malayensis from Java. It
seemed an interesting point, and therefore I re-measured the eggs from Ndao, andfound their large size confirmed, although the weights hardly differ. The most obvious
reason for the differences would be that A. n. medius is a larger subspecies than A. n.
albidus, but the measurements I took from the five Ndao females, show them to be practically identical in size (as expressed by wing-length) with specimens of the same sex

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

143

from Flores. Therefore, the difference remains unexplained, and intriguing.
It was about time for somebody to take the axe to the overexpanded species A. novaeseelandiae, and Haffer (in Glutz, 1985: 523-525) has done this with gusto, with thecreation of five species out of one. In this concept, birds from the Greater and Lesser
Sunda Islands are included in the species A. rufulus, ranging westward to India. A. novaeseelandiae is confined to New Zealand and its outlying islands, and Australia is inhabited by a separate species A. australis. Haffer’s discussion is centred on the Africanand Palaearctic populations and he provides hardly any documentation for the proposed split rufulus/australis/novaeseelandiae. To me, the forms medius and albidus lookvery close to australis. Admittedly, at the root of the problem of classification in Anthus,
is that so many perfectly good species resemble each other closely. Nevertheless, I feeljustified in retaining australis and the rufulus group as subspecies of A. novaeseelandiae,
until more concrete arguments for their specific separation may be brought forward.
Other authors have been reluctant to accept Haffer’s proposals.
Measurements:
Flores (A. n. albidus)

Sumba (A. n. albidus)


4

[]

Ndao near Roti (A. n. medius)
5

wing

tail
59
53-55
(53.8)
62
55½

tarsusculmen27
26-26.2(26.1)
26
25

entireculmen17.6
17-17.8(17.3)
18.3
17

exposedclaw
1413.6-15(14.0)
14
13

80
76-78
(77.0)
8377
77-78(77.4)

hind
9.8
9.6-11(10.1)
9.6
9.5

50-54
(51.9)

24-25(24.7)

16-16.7(16.5)

13-14(13.2)-

Anthus gustavi Swinhoe
Anthus gustavi Swinhoe, 1863, Proc. Zool. Soc. Lond.: 90 – the island of Amoy.
Collector.— Everett (2).
Material.— None.

Notes.— For a history of the discovery of this species, and a discussion of its synonymy and of specimens in the RMNH, see Mees (1990). It seems to be a not very common winter visitor to the Lesser Sunda Islands, where otherwise it has been recorded
from Timor (, 1861, near Dilli, collected by Wallace, cf. Sharpe, 1885: 614) and Sumba(one specimen, December 1896, collected by Everett, cf. Hartert, 1898d: 468).
Coracina novaehollandiae floris (Sharpe)
Artamides floris Sharpe, 1879, Cat. Birds Brit. Mus. 4: 14 – Flores.
Collectors.— Semmelink, Allen, Weber, Everett, Rensch (7), de Jong (1), Verheijen (1, MCZ), Schmutz.
Material.— (?), 1862, Larantoeka (Semmelink, RMNH cat. no. 2); , xi.1888, Reo (Weber, RMNH cat. no.
5); , 22.vi.1969, Nisar, 180 m (Schmutz, RMNH no. 81062); , ca. viii.1969, Nunang, 650 m (Schmutz,
RMNH no. 85117); , 25.v.1972, Nisar, 50 m, in Zizyphus-Savanna (Schmutz, RMNH no. 85116).
Eggs (figs 15a, 15aa): c/1, 2.v.1957, Poéng (RMNH no. 70753); c/2, 12.vi.1962, Mataloko (RMNH no.
70754); c/2, 16.v.1962, Mataloko (RMNH no. 70755); c/2, 1.vi.1962, Mataloko (RMNH no. 70756); c/1,
19.vi.1962, Mataloko (RMNH no. 70757).

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Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Measurements and weights:

RMNH no. 70753
RMNH no. 70754
RMNH no. 70755
RMNH no. 70756
RMNH no. 70757

31.7 × 23.2
31.3 × 22.4
33.4 × 23.4
34.6 × 22.4
34.9 × 22.2
34.1 × 21.9
34.5 × 22.2
33.0 × 23.4

0.5297
damaged
0.48320.436
0.4562
0.4656
0.5275

Notes.— The species Coracina novaehollandiae as defined in most recent literature,
consists of a mosaic pattern of well-differentiated forms, which replace each other geographically (on different islands), ranging from continental south–east Asia and Taiwan, to New Guinea, Australia and Tasmania.
Rensch (1931a: 564-565) drew attention, within this conglomerate, to the relatively
shorter tail, and other characters of C. javensis, and also of C. personata, compared with
Australian C. novaehollandiae, concluding “daß eine Vereinigung zu einem Rassenkreise
nicht möglich ist”. Mayr (1944a: 142) found exactly the opposite: that sumbensis (of thepersonata relationship) has a relatively longer tail than Australian C. novaehollandiae. The
same was recorded by Mason & McKean (1982) for personata (I do not understand thewing/tail ratios given by these authors) and the limited number of measurements taken by me indicates that this is also true for floris.
Rensch’s conclusion has of course, in a period that strong expansion of the species
concept was fashionable, not restrained later authors from doing exactly what Renschconsidered impossible (cf. Ripley, 1941), and this was generally, although sometimes
somewhat reluctantly, accepted for the following forty years (cf. Voous & van Marle,
1949). As a reaction against its over-expansion, the species concept is now going througha period of contraction, and this has not left the C. novaehollandiae agglomerate untouched. Mason & McKean (1982) have argued for the resurrection of C. personata from
Timor as a separate species. Well may they be right, but it is a pity that they discussed
just this one form, a segment in the middle of the agglomerate, and by their action left
the rest, in particular all forms to the west of Timor, dangling, without any suggestion as
to how these should be broken up into species. Even the difference between personata
and floris is considerable, and might justify specific status for the latter in the new concept. Feeling not competent, at present, to complete the dismemberment of the agglomerate, I prefer, for the moment, to keep all these forms under novaehollandiae (compare thediscussion of a similar problem in the Pachycephala pectoralis/fulvotincta complex).
Measurements:




(?)

wing
166
169
172
166
164

tail
119
121
118
123
120

tarsus
27
26.8
27.5
27
27.8

entire culmen
29
27
30.2
30
28

exposed culmen
24
23
26.2
24.7
24.5

Coracina novaehollandiae novaehollandiae (Gmelin)[Turdus] novae Hollandiae Gmelin, 1789, Syst. Nat. (ed. 13) 1 (2): 814 – terra van Diemen (ex Latham).
C[orvus] melanops Latham, 1801, Suppl. Ind. Orn.: xxiv – Nova Hollandia = Sydney.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

145

Collectors.— Rensch (1), Schmutz.
Material.— ? im.,  ad., 24.v.1972, Nisar, Zizyphus-Savanna, 50 m (Schmutz, RMNH nos. 85114, 85115).
Iris dark brown.

Notes.— Flores is the most north-westerly island whence this winter visitor from
Australia has been recorded. Stein collected three on Sumba (cf. Mayr, 1944a: 142). On
Timor it was found as early as 1828 by S. Müller (RMNH cat. no. 17), and by several
later collectors. Verheijen obtained a specimen on the island of Dao or Ndao off Roti (cf.
Mees, 1975b: 129; Verheijen, 1976: 14).1011)
The origin, within Australia, of these birds remains to be elucidated. One would
expect them to be from Western Australia, but in the extreme south-western part of thatstate the species remains common throughout the winter, and there is no clear indication that it would be a partial migrant (there is no proof that it is not, though). Moreover, the birds continue to live in pairs in winter, and I have never seen any evidence offlocking. In the mid-western and north-western parts of Western Australia, on the other
hand, there is considerable flocking in winter, and mixing of the subspecies “melanops”
and subpallida in the breeding range of the latter, indicating extensive movements (cf.
Mees, 1964c). In addition, birds from the South-West tend to be large in the wing andsmallish in the bill, unlike the specimens from Flores (cf. Keast, 1958; Mees, 1961b: 111).
The difference is only an average one and is of limited value; nevertheless, it gives some
support to my opinion that the birds visiting Flores originate from the more arid partsof north-western Australia. Formerly (l. c.), I believed that migrants to the tropics had
to come from the most southerly part of the range, but at the time I did not realise sufficiently that several species of large insectivorous birds leave northern Australia in thelong dry season (the Australian winter). The fact that the subspecies subpallida, originating from mid-western Australia, has now also been found as a winter visitor, supports
the assumption that migrants of novaehollandiae are from the same, or rather an adjacentpart of that continent.
The question of whether continental Australia and Tasmania are inhabited by different subspecies (melanops and nominate novaehollandiae, respectively), as claimed by
Keast (1958), has worried me ever since. Because of lack of adequate Tasmanian material, I hesitated to give a definite opinion (Mees, 1961b), and in subsequent publications
continued to use the name melanops for Australian birds and birds I believed to be migrants from continental Australia, even until quite recently (cf. Mees, 1994: 31). Although I have still not been able to compare good material, I consider that a final standin this matter is required now. The trivial average differences in measurements of wing
and bill, given by Keast in support of the recognition of a separate Tasmanian subspecies, do not appear to be outside the range of variation found in the several continental

11)

It might cause surprise that these two papers on the avifauna of Roti, both published in the “Zoologische Mededelingen”, were not combined and published under joint authorship. This is because Verheijen’s article was written first, and had been handed in for publication in “Ardea” before the collection
was received in Leiden. Later, examining the material, I found so much of interest, that I decided on a
supplementary paper, that duly appeared. After two years of deliberation, Verheijen’s manuscript was
returned by the editors as unsuitable for “Ardea”, after which it was published, without further delay,in the “Zoologische Mededelingen”.

146

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

populations, some of which were recognized by Keast as separate subspecies, but wereunited by me. In the thirty-five years that have passed, nobody else seems to have taken
up the matter, although I have repeatedly drawn attention to the unsatisfactory state ofaffairs. I think it may now safely be concluded that such average differences as possibly
exist between Tasmanian and mainland birds are insufficient for recognition in nomenclature. The consequence is that melanops, as hitherto recognised by me, becomes asynonym of nominate novaehollandiae. It means also that possible migrations of Tasmanian birds (they are even supposed to migrate as far as New Guinea), must be studied
mainly by observation and ringing.
Addendum.— After a period of some 35 years of doubting and vainly waiting fora revision I finally decided to place melanops in the synonymy of nominate novaehollandiae. Since then a revision was published by Schodde & Mason (1999: 578-580). At
first sight this seems to be a review, based on adequate material, for which I had been
waiting so long. The authors recognise the Tasmanian form as distinct, but the comparative notes given to support this show that the differences between Tasmanian novaehollandiae and mainland melanops are at best trivial. In general size melanops is said
to be “large to small” and novaehollandiae “medium”; the bill of the Tasmanian bird issaid to be marginally smaller, without clear separation. Especially as, within the large
mainland range, there is some geographical variation in bill size, Schodde & Mason’s
work actually confirms my opinion that the difference is too minor for expression innomenclature.
Verhoeye & Holmes (1998: 35) refer to C. novaehollandiae as a common austral migrant; they also mention an observation on Komodo, a slight extension westward of its
known winter range, suggesting that it also reaches eastern Sumbawa.
Measurements:
? im.
 ad.

wing
185
192

tail
126
125

tarsus28
27½

entire culmen
30
31

exposed culmen
24
25

Coracina novaehollandiae subpallida Mathews
Coracina novaehollandiae subpallida Mathews, 1912, Novit. Zool. 18: 326 – North-West Australia = Strelly[recte: Strelley] River (cf. Mathews, 1913: 193).
Collector.— Schmutz.
Material.—  with small gonads, 25.v.1972, Nisar, 50 m (Schmutz, RMNH no. 85118), plumage very
worn, especially the tail; no moult; lores and ear coverts black, but throat pale, hence an immaturebird.

Notes.— This specimen provides the first reliable record of the subspecies subpallida
from outside its breeding range, a point that requires some explanation. Although Mathews correctly described this subspecies as being characterized by its pale upper surface,
he did not really have a good idea of its characters and range, as he included the Northern Territory into its range. A year later, he (Mathews, 1913: 193) repeated this range, andtentatively included C. n. didima, which he had described from Melville Island, in thesynonymy of subpallida. The following year he confirmed this (Mathews, 1914a: 122). He
further claimed that the birds were migratory on Melville Island, leaving in November

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

147

and returning in May. This suggests winter visitors from the South, but surely C. novaehollandiae is a breeding-bird on Melville Island. Having concluded that the whole North
and North-West of Australia is inhabited by subpallida, it was logical that Mathews(1930: 534) assigned winter visitors to the Lesser Sunda Islands to this same subspecies.
Subsequently, Ripley (1941b) commented: “All the forms are representative. There is
one case of apparent overlapping from Timor to the Little Kei Islands, but it is now
known that the specimens of the Australian race subpallida, found on these islands are
winter visitors, not residents, as hinted by Hellmayr (1914)”. Correct, but Hellmayr
called these birds melanops, not subpallida. Mayr (1944a: 142) reverted to the name didima
for winter visitors on Sumba: “C. n. didima (= kühni Hartert) differs from melanops onlyin its slightly paler color”. He made no mention of subpallida. Peters & Mayr (1960: 171),presumably following Keast (1958), give subpallida its correct Australian breeding range(“Midwestern Australia from the Gascoyne to the DeGrey River”), but add: “winteringin the Lesser Sunda Islands and Kei Island”. Mees (1964c) commented: “It may be thatthe subspecies does partially migrate to these islands, but I have not seen specimens,
and the majority of individuals seems to stay in Australia”. Later, I was informed that
there is no extra-Australian material of subpallida in the AMNH (LeCroy, in litt., 17.
vi.1974). Thus, all previous records of subpallida from the islands are due to Mathews’s
misidentification and must be rejected.
Measurements:


wing
187

tail
129

tarsus29½

entire culmen
30½

exposed culmen
24

Coracina dohertyi (Hartert)
Edoliosoma dohertyi Hartert, 1896, Novit. Zool. 3: 584 – Sumba.
Collectors.— Everett (“a series”), Rensch (3), Verheijen (1, MCZ), Schmutz.
Material.—  ad., 31.vi.1971, Sésok, 800 m (Schmutz, RMNH no. 85029).
Eggs not certainly known.

Notes.— Coracina dohertyi was not previously represented in the RMNH collection.
Yet, it cannot be particularly scarce on the two islands to which it is confined. On Flores,
Everett obtained “a series” and Rensch three specimens. On Sumba, Stein collected no
fewer than 16 specimens, and Sutter another 7.
Rensch (1931a: 566) wrote: “E. dohertyi ist durch die scharfe schwarzgraue Zeichnung so sehr von den Angehörigen des Rassenkreises E. morio geschieden, daß ich sie
nicht zu einem einzigen Rassenkreise zusammenziehen möchte”. Mayr (1944a: 142)
agreed that C. dohertyi: “belongs undoubtedly to the morio assemblage”, but also left it
as a separate species. Unexpectedly, in view of the foregoing, Peters & Mayr (1960: 185)
have placed it between C. caerulescens (a Philippine species) and the widely-distributed
C. tenuirostris, and some distance away from C. morio.
With only a single specimen, and that a male, I am not in a position to speculate
fruitfully about the affinities of this species with its peculiarly restricted distribution.
Measurements:


wing
127

tail
92

tarsus25

entire culmen
25½

exposed culmen
18.3

148

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Lalage sueurii sueurii (Vieillot)
Turdus Suerii [sic] Vieillot, 1818, Nouv. Dict. Hist. Nat. 20: 270 – la Nouvelle-Hollande = Timor (cf.
Pucheran, 1855: 352).
Collectors.— de Jong (1), Verheijen (3, MCZ), Verheijen (eggs), Schmutz (?).
Material.— Apparently no skins; eggs only.
Eggs (fig. 15b): c/3, 20.vi.1960, Palué (RMNH no. 70758); c/3, 28.vi.1960, Wodja, Palué (RMNH no.
70759); c/2, 29.vi.1960, Lai, Palué (RMNH no. 70760); c/2, 9.v.1960, Palué (RMNH no. 70761). The eggs
are pale greenish, heavily spotted with more or less longitudinally-directed medium-brown primary
markings, and less numerous light grey secondary markings. They do not differ significantly fromeggs of L. nigra.
Measurements and weights of two clutches: RMNH no. 70759

RMNH no. 70760

20.0 × 15.9
20.3 × 15.8
21.2 × 16.2
21.2 × 16.3
21.7 × 16.3

0.14720.14530.159
0.15840.1563

Hellebrekers & Hoogerwerf (1967: 86) did not know eggs of L. sueurii from Java.
Kooiman collected eight clutches, labelled Lalage sp., of which the original data (localityand date) have become lost (RMNH nos. 23632-23639). Unfortunate as it is that the eggs
are without data, all Kooiman’s collecting was done in far eastern Java, where he obtained skins of L. sueurii at Klatakan and Dampar (cf. Mees, 1986: 86-87). It is a perfectly
safe deduction that the eggs are from the same localities, and belong to L. sueurii. Thereis also a clutch of eggs from Ndao near Roti in the collection (c/2, 5.vi.1969, leg. Verheijen, RMNH no. 76725, cf. Verheijen, 1976: 15).
Notes.— On Flores, this species appears to be rather local in distribution. When
Verheijen (1961) met with it on Palué, he remarked that he had never yet seen it inWest Flores. From mainland Flores, only the specimen collected by De Jong at Laboean
Badjo is known. As Rensch (1931a: 562) observed one near Endeh, and Schmutz (MS)has several records from the neighbourhood of Nanga-Lili, and between Nanga-Liliand Wae-Wako, it is likely to be more widely distributed around the coast.
Pericrocotus lansbergei Büttikofer
Pericrocotus Lansbergei Büttikofer, 1886, Notes Leyden Mus. 8: 155 – Bima (Sumbawa).
Collectors.— Everett (several), Rensch (3), de Jong (3), Schmutz (2).
Material.— , 17.ii.1969, Nunang, 650 m (Schmutz, RMNH no. 81409);  im., 8.ii.1972, Nggoang, 850 m(Schmutz, RMNH no. 85324).

Notes.— Presumably, P. lansbergei is a derivative of P. cinnamomeus.
Measurements:
 im.


wing
71
70

tail
87
88

tarsus16.5
17.7

entire culmen
15
14

exposed culmen
1111

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

149

Lanius cristatus superciliosus Latham[Lanius] superciliosus Latham, 1801, Suppl. Ind. Orn.: xx – Java.
Collectors.— Weber (1), Everett (4), de Jong (2), Schmutz (1).
Material.— , 20.x.1969, Nanga-Lili (Schmutz, RMNH no. 85216). Iris dark brown. Stomach contents
remains of grasshoppers.
According to a manuscript list by Büttikofer, Weber’s specimen, received in spirits (cf. Büttikofer, 1894:299), was made into a study skin for the RMNH collection. I have been unable to find it, so that either
the quality was insufficient and it has never been entered into the collection (most likely), or it has been
given out in exchange.

Notes.— The fact that this species has been recorded by four collectors, one ofwhom, Everett, obtained four specimens, proves that Flores belongs to the normal winter range of this subspecies. There are also several records from Sumba, but I do notknow of any from the islands to the east of Flores, or from Timor.
Brachypteryx montana floris Hartert
Brachypteryx floris Hartert, 1897, Novit. Zool. 4: 170 – At and above 3500 feet in South Flores.
Collectors.— Everett (4), Verheijen.
Material.— 2 , 12.xi.1969, Nggolong Tedé, Ruteng, 1900 m (Verheijen, RMNH nos. 65381, 65382); , ,
18.ix.1971, Potjo Gurung, Ruteng, 1700 m (Verheijen, RMNH nos. 85089, 85088); , 31.iii.1973, mountainlake Rana-Ka (Verheijen, RMNH no. 85090). Iris brown, bill black, legs grey.

Notes.— This interesting endemic form appears to be known from very few specimens. Hartert (1897b: 516) recorded that Everett obtained only two pairs. The fivespecimens listed here seem to be the only ones obtained since.
Measurements:
4


wing
70-74
(72.3)
71

tail
56-62
(59.3)
57

tarsus32.5-35(33.4)
32.4

entire culmen
17.5-18.5(18.1)
17.4

exposed culmen
13.8-14(14.0)
13.5

Saxicola caprata fruticola Horsfield
Saxicola fruticola Horsfield, 1821, Trans. Linn. Soc. Lond. 13: 157 – Java.
Collectors.— Semmelink, Allen, Colfs, Weber, Everett, Rensch (4), de Jong (5), Verheijen (11, MCZ), Verheijen/Schmutz.
Material.— , 1862, Larantoeka (Semmelink, RMNH cat. no. 10); 2 , iv.1880, Flores (Colfs, RMNH cat.
nos. 15, 16); , v.1880, Flores (Colfs, RMNH cat. no. 17); skeleton, xii.1888, Maumeri (Weber, RMNH cat.
c); 4 ,  im., 12 , 1 (?), 4 juv., 1968/1977, from various localities in Manggarai (Verheijen/Schmutz,
RMNH nos. 59906-59917, 65384-65386, 65391, 66159, 81359, 85402-85404, 85411).
Eggs: 103 clutches, of c/1 (19 ×), c/2 (32 ×), c/3 (41 ×) and c/4 (11 ×) eggs, collected in the months April(1), July (1), August (18), September (31), October (38) and November (14), (RMNH nos. 70770-70871,
73513). The full data of the April clutch are: c/2, 29.vi.1954, Poéng (RMNH no. 73513); the clutch has an
original label, with the month April written in full, so that there can be no confusion about the date. The
clutch came to Leiden with the collection Coomans de Ruiter, and that may be the reason why it was not150

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

listed by Verheijen (1964): when he wrote his article, this clutch was no longer in his possession. The
clutch is important as being the only one in a series of over a hundred, collected outside the period JulyNovember. The eggs are light greenish blue, lightly to moderately freckled with medium-brown spots,
which tend to be closest together around the blunt end.
Some measurements and weights:

RMNH no. 70822

17.8 × 14.5
0.096918.0 × 14.7
0.111718.6 × 14.7
0.1113
RMNH no. 70823 18.1 × 14.6
0.109918.3 × 14.7
0.11
18.6 × 14.6
0.1112
RMNH no. 70826 15.7 × 13.8
0.095416.8 × 14.4
0.096117.7 × 14.3
0.104517.9 × 14.7
0.1081RMNH no. 70827 20.2 × 15.4
0.117720.9 × 15.8
0.1271
Some 10% of the nests were found parasitized by Cacomantis variolosus (cf. Ottow & Verheijen, 1969).

Notes.— There is a ruling that names ending with -cola have to be treated as masculine substantiva, unless from the original description it is apparent that an author
regarded the name as feminine, in which case the feminine gender is retained (ICZN,
1985: art. 30); unfortunately, the issue is confused by the example given: “Compound
Latin nouns ending in -cola, such as Sylvicola, are treated as masculine”. This is a nonsequitur, and suggests erroneously that names ending in -cola are always to be treatedas masculine (cf. Mauersberger, 1983: 52). Naturally, I wrote to the Secretariat of the
ICZN, pointing out this contradiction in the Code, and although my letter remained
unacknowledged, I presume that it has been acted upon and the error has been corrected. Saxicola has usually been considered feminine, and that is correct. The genus
was based on three species, in the combinations S. Oenanthe, S. Rubetra and S. rubicola:
Oenanthe and rubicola are substantiva, but rubetra (“bramblebush-inhabiting” is a Latin adjectivum and decides the gender (Coomans de Ruiter et al., 1948: 41). As far as
the name Saxicola caprata fruticola is concerned: fruticola is a masculine substantivum,
so that it is not affected by the gender of its genus, but what about caprata? The namewas introduced by Linnaeus (1766: 335), in the combination Motacilla Caprata, andwas based on Brisson (1760: 442), who described the species from Luzon, and recorded as its local name there Maria-capra. Evidently, the name Caprata was derived
in some way from this vernacular, although it is not clear how. The fact that Linnaeus,
when introducing Caprata, gave it an initial capital, suggests that he regarded it as a
substantivum, which means that its termination does not change with the gender ofthe genus.
The genus Saxicola contains several other species, the most widely distributed ofwhich is at present universally known as S. torquata and its European subspecies as S.
torquata rubicola, but the masculine version has sometimes been used earlier in the lastcentury (for example by Stresemann, 1920: 171). Coomans de Ruiter et al. (1948: 42) regarded rubicola, and by inference fruticola, as an adjectivum, but Dr Kraak now supports
the opinion that it is a substantivum.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)151

Zoothera interpres (Temminck)
Turdus interpres Temminck, 1828, Recueil d’Ois. 2 (livr. 75): pl. 458 – à Java et à Sumatra.
Collectors.— Everett, de Jong (3), Verheijen/Schmutz (1).
Material.— , 21.vii.1976, Poco Nernancang (Verheijen, RMNH no. 81455).
Eggs: Verheijen (1964: 199) records breeding in May; there are no eggs in his collection.

Notes.— Everett obtained “A series from S. Flores” (cf. Hartert, 1897b: 515) and deJong collected three near Wai Sano (cf. Rensch, 1931b: 398). The facts that this species
was not found in Flores by Rensch, and that I received but a single specimen, indicate
that, although it is not rare locally, it is not generally distributed.
I agree with Junge (1938: 352) that: “The differences between interpres and leucolaema
are large enough to regard leucolaema as a separate species”. Therefore Z. interpres has
no subspecies. It is surprising that in its comparatively large, and mostly insular, range,
this species shows no geographical variation.
Measurements:


wing
104

tail
58

tarsus27

entire culmen
20.3

exposed culmen
16.5

Zoothera dohertyi (Hartert)
Geocichla dohertyi Hartert, 1896, Novit. Zool. 3: 555, pl. xi fig. 3 – Lombok.
Collectors.— Everett, Rensch (3), Verheijen (2, MCZ), Verheijen/Schmutz.
Material.— , 21.xi.1969, Nggolong Tedé, 1900 m (Verheijen, RMNH no. 65368); , 24.xii.1969, Todo
(Verheijen, RMNH no. 65369); ?  juv., 29.x.1970, Ruteng (Verheijen, RMNH no. 85035); , 29.iii.1971,
Lingko Laring Pongkor (Verheijen, RMNH no. 85033); , 15.iii.1973, Nggolong Tedé (Verheijen, RMNH
no. 85322); , 29.iii.1973, Rana-Ka (Verheijen, RMNH no. 85034); , 3.vi.1976, Ulu Ros (Verheijen,
RMNH no. 81435); , 23.vi.1976, Ulu Ros (Verheijen, RMNH no. 81436); , 30.iii.1978, Ulu Wae Wua(Verheijen, RMNH no. 81454); , 26.v.1978, Wae Rungget (Verheijen, RMNH no. 81320); , 8.vi.1978,
Wae Golo Lolo (Verheijen, RMNH no. 81437).
Eggs (fig. 15c): c/1, 27.vi.1954, Nilo (RMNH no. 70762); c/2, 19.v.1956, Mano (RMNH no. 70763). The
eggs are creamish, heavily freckled with light brown; they resemble closely eggs of Z. interpres from
Java, described by Hellebrekers & Hoogerwerf (1967), but are a little larger and heavier.
Measurements and weights:

RMNH no. 70762
RMNH no. 70763
The second egg is too damaged for measuring and weighing

25.9 × 19.2
26.8 × 19.1

0.24900.2299Notes.— This species is in measurements very similar to Z. interpres, but has a distinctly longer tail. Note the complete absence of a difference in size between the sexes.
On the basis of a combination of measurements of his own material, and measurements
previously published by Rensch, Mayr (1944a: 155) concluded that there is some geographical variation in size: “The population from Timor shows the largest measurements, those of Sumba and Sumbawa the smallest, while those of Lombok and Flores
are intermediate”. Of the postulated differences, I find only that of the Timor birds convincing. According to White & Bruce (1986: 333), the larger size of the Timor birds:

152

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)“may be due to their living at a high altitude: the species has only been found there on
Mt. Mutis”. The Timor birds were taken at 1600-2300 m. My series from Flores includes
specimens collected at 1900 and 2200 m.
Measurements:7
3
? Juv.

wing
103-108(105)
103-108
(105)
102

tail
65-73
(69)
67-70
(68.7)
64

tarsus30-32(30.9)
28.2-31(29.9)
31

entire culmen
20.5-22.7(21.5)
20.8-22(21.3)
21

exposed culmen
16.5-18.7(17.6)
16-18
(17)
17

Zoothera andromedae (Temminck)
Myiothera andromedae Temminck, 1826, Recueil d’Ois. 2 (livr. 66): pl. 392 – les îles de Java et de Sumatra(the figured type is from Java), restricted type locality W. Java (Kuroda).
Collectors.— Everett (4), Verheijen/Schmutz.
Material.— , 2.xii.1969, Nggolong Tedé, Ruteng, 1900 m (Verheijen, RMNH no. 65367); , 27.iii.1973,
Danau Rana-Ka, 2200 m (Verheijen, RMNH no. 85032); , 30.iii.1973, Danau Rana-Ka, 2200 m (Verheijen,
RMNH no. 85031); , 8.iii.1976, Ulu tuke’ nikit (Schmutz, RMNH no. 97120). Iris brown, bill dark brownto horn-black, legs brown or light reddish.
Eggs (fig. 15d): c/2, 17.v.1960, Potjong (RMNH no. 70768); c/2, without data (RMNH no. 70769).
Measurements and weights:

RMNH no. 70768
RMNH no. 70769

29.1 × 19.4
29.3 × 19.0
27.4 × 20.6
28.1 × 21.1

0.24540.2506
0.28800.2868Notes.— The only previous record of this species from Flores is by Everett, whose
specimens were obtained at a level of about 3500 feet (ca. 1050 m). On Flores, as on
Java, this seems to be definitely a mountain bird. White’s (in White & Bruce, 1986: 333)
statement that this is a lowland species, certainly requires qualification. On Sumbawa
Johnstone et al. (1996: 173) recorded Z. andromedae from altitudes of 450-850 m, lower
than I would have expected; this indicates that its ecological requirements are far fromwell known.
Salvadori’s (1892: 134) record of three specimens from the island of Engano isabove suspicion. Nevertheless it is puzzling, for if it were a resident, as this recordsuggests, it is difficult to understand how the species was missed by Abbott in 1904
and by J.K. de Jong in 1937. Admittedly it is known to be skulking and not easily observed.
On the basis of the three males and only one female collected by Everett, Hartert(1897b: 515) concluded that females of this species are smaller than males: “The males
have the wing 136 to 137 mm, the female only 125”. My four specimens do not support
this, but I cannot vouch for the reliability of their sexing. Therefore I have taken thewing-measurements of some specimens from other islands in the RMNH–collection,
as follows. Sumatra, 2  124, 128; Java 8  122, 123, 124, 126, 126, 126, 127, 130; 14 
111, 117, 118, 119, 120, 120, 120, 121, 121, 122 (type), 123, 123, 124, 126. Timor  133.
These figures confirm the existence of a moderate average difference between the sexes, which is worth recording in view of the absence of such a difference in Z. dohertyi.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

153

In combination with the measurements published by Hartert, they also suggest thatbirds from the eastern part of the range are larger than birds from Sumatra and Java,thus conforming to the rule, formulated by Rensch, that from Java eastwards, birds
tend to become larger in size.
It is tempting, but perhaps not rewarding, to speculate why this widely, but irregularly, distributed species should show so little indication of geographical variation.
Measurements:
2
2

wing
128, 133
124, 129

tail73, 8069, 64tarsus32, 39.5
30, 33entire culmen
31, 31.5
30, 31.8

exposed culmen
25, 2825.5, 27

Turdus obscurus Gmelin[Turdus] obscurus Gmelin, 1789, Syst. Nat. (ed. 13) 1 (2): 816 – in Sibiriae silvis, ultra lacum Baical.
Collectors.— Colfs, Schmutz.
Material.— (?), (1880), Flores (Colfs, RMNH cat. no. 11); , 2.vi.1976, Poco Nernancang, Ruteng, 1500 m(Schmutz, RMNH no. 85036); , 12.vi.1976, same locality (Schmutz, RMNH no. 81317).

Notes.— It is a pity that the specimen collected by Colfs cannot be dated. These
records suggest that the occurrence of T. obscurus on Flores is not exceptional.
Smythies (1968: 415, and still 1981: 309) states: “not yet recorded from Indonesian
Borneo”, but Nieuwenhuis collected a specimen on the upper Mahakam as early as
November 1899 (RMNH cat. no. 15): this bird was recorded by Finsch (1901a: 176, andagain 1905: 21, 126). In Java, the species had hitherto been recorded from the West only
(cf. Dammerman, 1929: 57; Kuroda, 1933: 293), but four specimens collected by Bartels
at Sikatok, Bagelen, Sindoro, 11-14.iii.1915, extend this to Central Java. In West Java,
Bartels collected three specimens on the Pangerango (1.vi.1905, 28.xi.1910, 12.xii.1914),
and a series of 50 at Tjibening, Djampang Tengah, all in the short period 13-19.ii.1909. I
cannot judge whether Bartels had any particular reason for collecting so many specimens from what evidently was one large flock.
Measurements:


(?)

wing
128
122
124

tail
83
77
82

tarsus30
30
31

entire culmen
22.8
21.5
23

exposed culmen
16.6
16
16

Pnoepyga pusilla everetti Rothschild
Pnoepyga everetti Rothschild, 1897, Novit. Zool. 4: 168 – South Flores... at elevations of about 3000 to 3500
feet.
Collectors.— Everett (“a series”), Rensch (5), Verheijen.
Material.— , 28.iii.1973, near Danau Rana-Ka, 2200 m (Verheijen, RMNH no. 85221).
No eggs; the species is included in Verheijen’s (1964: 199) list with a question mark.

Notes.— Rensch (1931a: 569) states: “Diese Rasse ist von der javanischen wohl nur
durch etwas bedeutendere Grösse unterschieden”. I cannot confirm this: a wing-length

154

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)of 52 mm is quite normal in females from Java, but the above specimen from Flores has
a slightly longer and more slender bill than P. p. rufa, and the pileum is darker, blackish
brown, not dark ferruginous-brown.
Measurements:


wing
52

tail
-

tarsus20.5

entire culmen exposed culmen
14.8
12

Tesia everetti everetti (Hartert)
Orthnocichla everetti Hartert, 1897, Novit. Zool. 4: 170 – South Flores.
Collectors.— Everett (“a series”), Rensch (5), de Jong (5), Verheijen (9, MCZ), Verheijen/Schmutz.
Material.— (?), x.1896, South Flores (Everett, RMNH cat. no. 1, syntype); , 18.xi.1967, Nggolong Tedénear Ruteng, 1900 m (Verheijen, RMNH no. 65436); , 22.xi.1968, Ruteng (Verheijen, RMNH no. 59881);
, 7.xii.1968, Ruteng (Verheijen, RMNH no. 59882); , 4.xi.1969, Ruteng (Verheijen, RMNH no. 65435);
, 14.iii.1973, Nggolong Tedé, 1900 m (Verheijen, RMNH no. 85213); (?) juv., 26.iii.1973, Danau Rana-Ka(Verheijen, RMNH no. 85214); , 28.iii.1973, Danau Rana-Ka (Verheijen, RMNH no. 85215); , 4.ii.1976,
Ero (Verheijen, RMNH no. 81375); , , 24.vi.1976, Ulu Ros (Verheijen, RMNH nos. 81357, 81367). Iris
brown or grey-brown, bill maxilla brownish black, mandible dirty white, inside of mouth orange-yellow, legs greyish flesh colour.
Eggs (fig. 15e): 22 clutches of one (6 ×) and two (16 ×) eggs collected in the months February (3), March(3), April (2), May (1), June (4), July (2), August (2), September (1), November (1), and December (2), andone not dated (RMNH nos. 70872-70893). The distribution of these records over the year, without any
suggestion of a peak, is remarkable. Evidently, two is the normal clutch-size.
Some measurements and weights:

RMNH no. 70872
RMNH no. 70873
RMNH no. 70874
RMNH no. 70875
RMNH no. 70876
RMNH no. 70878

RMNH no. 70879
RMNH no. 70886
RMNH no. 70887
RMNH no. 70889
RMNH no. 70890

18.6 × 14.4
18.4 × 13.6
18.6 × 13.3
19.1 × 13.8
20.0 × 13.9
20.4 × 13.8
18.6 × 13.9
19.3 × 13.8

20.0 × 14.1
20.4 × 14.1
19.5 × 13.9
20.0 × 14.2
19.4 × 13.8
19.6 × 14.0
20.0 × 13.5
20.0 × 13.7
20.2 × 13.9
21.0 × 14.0

0.09000.09260.0842
0.09500.09150.09260.08590.0922the second egg consists ofbroken pieces
0.09280.09470.09990.09630.08880.09800.09240.0901
0.09570.1025

Notes.— The species has a large vertical distribution. Rensch (1931a: 568) found it
from sea-level (Endeh) to 1400 m. This is now increased to 2200 m (Rana-Ka).
There has been a notable diversity of opinions about the systematic position of thisspecies. It was originally described as the second member of the previously monotypic
genus Orthnocichla. In neither of his two early publications did Hartert (1897a, 1897b)

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

155

discuss this placement, and the reason is obvious: at that time he did not yet have material of O. subulata, the type species of the genus, so that only its description and geographical probability (O. subulata was described from Timor, near Flores) were considered. A year later, Hartert (1898b: 114) did receive material, on which he commented:“This Orthnocichla differs considerably from O. everetti Hart. of Flores in being verymuch smaller, the beak being narrower and more pointed, the wing shorter, legs smaller and lighter in colour, in having a very distinct buffy white superciliary line, which is
not developed in O. everetti, and in its white breast, which is pale ashy grey in O. everetti.
O. whiteheadi from Mount Kina Balu in Borneo differs in having a much darker, almost
black crown, the back being much darker brown, the superciliary line rusty, sides ofbreast and flanks darker; under the superciliary stripe runs a blackish brown line fromthe eye to the neck, along the sides of the head. This line is not developed in O. everettiand O. subulata. While O. whiteheadi Sharpe is evidently a mountain bird, both O. everettiand O. subulata occur in the low country”.
Only a few years later, Finsch (1901b: 212-215), had received a specimen of O. everetti (RMNH cat. no. 1), that he could compare with the type material of O. subulata. It is
Finsch’s merit to have realised that not O. subulata, but the species described as Microura superciliaris, at that time known as Oligura superciliaris, from West Java is the closest relative of O. everetti. He expressed this relationship by proposing a new genus,
Pseudoxenicus, for these two species (Pseudoxenicus, because of their superficial resemblance to Xenicus of New Zealand).
Soon afterwards, it was observed that the Himalayan Tesia cyaniventer is close to
Microura superciliaris, although Chasen (1935: 231), who held a very broad species-concept, is no longer followed in treating the latter as a subspecies of the former. As Tesia is
the older generic name, it replaced Pseudoxenicus.
In this stage, Orthnocichla = Pseudoxenicus everetti was somehow forgotten, or left
behind. Rensch (1931a: 568) must have been unaware of Finsch’s work when he wrote
in the discussion of Orthnocichla everetti: “O. subulata Shpe. von Timor und Babar (hierRasse advena Hart.) ist so scharf differenziert, daß sie nicht mehr mit everetti zu einem
Rassenkreis vereinigt werden kann. Noch ferner steht C. [meant is O.] whiteheadi Shpe.
von N-Borneo”. Evidently, he accepted that O. subulata and O. everetti are congeneric.
Chasen (1935: 231 footnote 2) referred casually to “T. subulata of Timor and Flores”, but
gave no explanation.
Thus, it was left to Delacour (1942: 514) to give arguments for leading O. everetti
into Tesia. He is, however, rather confused about it, beginning with: “In examining theshort-tailed species (C. whiteheadi and C. subulata), from Borneo, Flores and Timor, usually referred to the genus Orthocichla [sic], it has occurred to me that they had been
wrongly placed in, or near, the genus Tesia,.... I have therefore put them in the genus
Cettia (subgenus Urosphena), of which they represent extremely short-tailed, long-tarsus adaptations”. This sentence only makes sense when it is assumed that Delacour,like Chasen before him, regarded O. everetti as a subspecies of O. subulata (an error that
may have inspired Watson to make the same error over forty years later). Yet, in thevery next paragraph, Delacour defines the genus Tesia, concluding: “In my opinion thefollowing species belong to the genus Tesia Hodgson, 1832: T. cyaniventer, T. olivea (India to Indo-China), T. superciliaris (Java) and T. everetti (Flores and Sumbawa)”. A step
forward was, that Delacour transferred Tesia to the Sylviidae, in the neighbourhood of156

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Cettia, from the Troglodytidae, where it was still placed by Chasen (1935: 231) andHoogerwerf (1948a: 131).
Presumably on the basis of Delacour’s revision, Mayr (1944a: 136) called the Timor
species Urosphena subulata; note that he treated Urosphena as a genus, not as a subgenus
of Cettia. Verheijen (1964: 199) followed Hartert and Rensch in keeping to the original
binomen Orthnocichla everetti; the nomenclature he used was based on a list sent to him
by Dr Mayr in 1957 (through a misprint, the author’s names of Pnoepyga pusilla everettiand Orthnocichla e. everetti have become transposed in his list).
Next came Watson (in Watson et al., 1986a: 6), who, probably misled by Delacour’s
enigmatic passage quoted above, reduced O. everetti to a subspecies of Urosphena subulata, which is obviously ridiculous and only understandable when it is assumed that hehad not personally examined and compared these species.
White (in White & Bruce, 1986: 338) had intended to follow Delacour in placing O.
everetti in the genus Tesia, and retaining O. subulata in Urosphena, but Bruce, in consultation with the editors of Watson, finally decided to place O. everetti in Urosphena, although as a distinct species, not as a subspecies.
The last word is by King (1989), who again separated O. everetti and O. subulata generically, in the genera Tesia and Urosphena respectively.
Summarizing, it may be said that there are two lines of thought about the positionof O. everetti. One is that it belongs to the same wave of colonisation as O. subulata, fromwhich it has only subsequently differentiated, and which therefore is its closest relative.
The other, that it represents a second, (later) wave of colonisation, derived from Tesia
superciliaris of Java.
I support without hesitation the second opinion: morphologically O. everetti agrees
with T. superciliaris in the basally broad, somewhat flattened bill, the very short tail, thelong tarsus. O. subulata, on the other hand, has a more slender bill, not notably wider
near its basis, a relatively longer tail and a shorter tarsus (see table of measurements).
In addition, the eggs of O. everetti are indistinguishable from those of T. superciliaris,in colour, as I found by direct comparison, and also in measurements and weights (com-Fig. 10. Tesia superciliaris. Records from the mountains of Java.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

157

pare the figures of O. everetti, given above, with those of T. superciliaris provided byHellebrekers & Hoogerwerf, 1967: 118). This argument is weakened by the fact that theeggs of O. subulata remain unknown.
Against this is a point that has been given much weight by some authors: the inconspicuous brown coloration that O. everetti and O. subulata share, as opposed to the distinctive dull blackish and grey head markings of T. superciliaris. In this connection, it
should be pointed out that in juvenile T. superciliaris the head markings are not yet developed, head and mantle are warm brown (without olive as in the adult) and are remarkably similar to the adult plumage of T. everetti. The underparts of both T. superciliaris and T. everetti are mostly light grey, whereas the underparts of O. subulata are whitish. Interesting is that a juvenile of T. everetti does hardly differ from the adults: its
plumage is a little looser, and the undersurface a little darker grey. Compare this withthe conspicuously distinctive juvenile plumage of T. superciliaris of which the upperparts have been described above. The underparts are also distinctive, brownish, in one
case yellowish brown, as opposed to grey in the adult.
For some 20 years, T. superciliaris was regarded as a subspecies of T. cyaniventer ofthe Asiatic mainland (Robinson & Kloss, 1924: 287; Bartels & Stresemann, 1929: 129;
Chasen, 1935: 231; Lonsain, 1941: 10; Hoogerwerf, 1948a: 131) but it was, so far as I amaware without explanation, restored to species status by Delacour (1942: 514; 1947: 271),
and this has been accepted by later authors.
T. superciliaris, then, is an endemic species of Java. As regards its distribution in Java,
Chasen (l. c.) and Delacour (l. c.) give it as Java, implying that it ranges throughout themountains of the island. Kuroda (1933: 327) gave its range as: “Confined to West and
Mid Java”, but all the localities he listed are from West Java. Watson (in Watson et al.,1986a: 5): “Mountains of western and central Java”.
Material in the RMNH collection is from the Pangerango (Bartels), Tankoeban Prahoe 1400-1600 m (F.C. van Heurn, v. Balgooy), summit Manglajan 1800 m (F.C. van
Heurn), Tjinjiroewan (v.d. Weele), all in the western part of West Java, and Kali Goea,
Slamat (Bartels) in the western part of Middle Java. It has further been recorded fromTjibodas-Gedeh (Hoogerwerf, 1949b: 2, 99); Papandajan (Stresemann, 1930) and Tjerimai (Kuroda, 1933). It ranges therefore throughout the mountains of West Java, and to
Mt. Slamat in Middle Java (fig. 10). This range is almost the same as that of anotherendemic Javanese mountain-bird, Garrulax rufifrons, except that the latter is known
from Mt. Karang in Bantam, whence T. superciliaris has not yet been recorded.
Measurements:
Flores (Tesia everetti)
4
5

wing

tail

tarsusentire culmen exposed culmen

53-55½(54.1)
49-53(51.4)
51

16½-20(18.0)
15½-17(16.2)
16½

24-25(24.7)
22.5-24.4(23.6)
25.4

16-17(16.4)
15.0-16.2(15.8)
17.2

13-13.7(13.2)
12.2-13(12.6)
13

13-17.5(15.3)
11-15(13.6)

24-25.5(25.1)
23-25(24.1)

13.7-16(15.0)
13.2-15(14.0)

11-12.8(11.9)
10-12(11.0)(?)
Java (Tesia superciliaris)
10 
49-52
(50.6)
10 
47-49.5(48.4)

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Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Timor (Orthnocichla subulata) *)

53
24
19
14.3
12

55
24½
18.2
14.5
12.2
*) Syntypes of O. subulata (RMNH cat. nos. 1, 2), collected in 1829. Sexing of this old material is unreliable.

Cisticola juncidis fuscicapilla Wallace
Cisticola fuscicapilla Wallace, 1864, Proc. Zool. Soc. Lond. (1863): 489 – Timor, Flores. Type locality restricted to “Delli” (= Dilli), E. Timor, by Lynes (1930: 633).
Collectors.— Allen, Everett (3), Rensch (1), de Jong (2), Verheijen (8, MCZ), Verheijen/Schmutz.
Material.— , 30.i.1976, Waé Rempo (Schmutz, RMNH no. 81421); (?), no data (Verheijen no. 1598,
RMNH no. 81417).
Eggs: 48 clutches of c/1 (8 ×), c/2 (17 ×), c/3 (19 ×) and c/4 (4 ×), collected in the months January (1),February (5), March (7), April (14), May (12), June (5), July (1), and August (1), and two not dated(RMNH nos. 70978-71025).
Some measurements and weights:

RMNH no. 71017

RMNH no. 71018

RMNH no. 71019

14.7 × 11.2
14.8 × 11.2
14.8 × 11.2
15.0 × 11.2
15.3 × 11.2
15.4 × 11.0
15.8 × 11.1
15.4 × 11.7
15.5 × 11.7

0.04320.04180.04350.0451
0.04840.04770.0476
0.05870.0580Notes.— The gender of names ending with -cola has been discussed on a previous
page (p. 150). There is no need to change the traditionally feminine gender of Cisticola
to masculine, as has been done in some recent works (e.g., Sibley & Monroe, 1990).
David & Gosselin (2002: 38) argued that names ending in -capilla/-capillus are substantiva and therefore do not change with the gender of their genus. One of the examples
given is Cisticola fulvicapilla, not to be changed to fulvicapillus, but they ignore the fact,
mentioned above, that Cisticola has consistently been treated as feminine and that itshould be possible to preserve this gender for it, which would make a discussion superfluous. I must add, however, that there are three species with ending -capilla/-capillus
on the Dutch list, and therefore included in the work by Coomans de Ruiter et al. (1948)
Sylvia atricapilla, Regulus ignicapillus, and Parus atricapillus, and that all three names areunequivocally stated to be Latin adjectiva. Surprisingly David & Gosselin (2000: 264),
in a paper published only 15 months before the one just quoted, state that atricapillus isa classical Latin adjectivum. They discuss one case, clearly regarded as exceptional, in
which atricapilla seems to have been used as a substantivum. Thus they agree here with
Coomans de Ruiter et al. In their later paper the sudden change of stand remained unexplained. It seems wise to be reticent in the alteration of customary endings until classical scholars agree among themselves.
Measurements:

(?)

wing
50
49

tail
36
36

tarsus20
19.8

entire culmen
12.6
11.8

exposed culmen
9.9
9.8

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

159

Cisticola exilis lineocapilla Gould
Cysticola lineocapilla Gould, 1847, Proc. Zool. Soc. Lond. 15: 1 – Port Essington.
Collectors.— Allen (cf. Wallace, 1864: 485, s.n. C. ruficeps), Weber, Everett (“several specimens”), de Jong(1), Verheijen (1, MCZ), Verheijen.
Material.— , 31.xii.1888, Kotting (Weber, Nr. 509a, RMNH); , 7.x.1971, Gurung, Langkas, Rahong,900 m (Verheijen, RMNH no. 85222).
Eggs (fig. 15f): 86 clutches, of c/1 (15 ×), c/2 (29 ×), c/3 (30 ×) and c/4 (12 ×), collected in the months
January (1), February (4), March (11), April (21), May (27), June (15), July (2), September (1), November(1) and December (1), and two without date (RMNH nos. 70894-70977, 73514, 76582). The eggs are light
blue, with rather coarse light to medium brown markings, moderately glossy.
Some measurements and weights:

RMNH no. 70957
RMNH no. 70970

RMNH no, 70971

RMNH no. 70972

16.1 × 11.3
16.1 × 11.8
16.5 × 11.5
16.8 × 11.7
16.9 × 11.3
17.0 × 11.7
15.0 × 11.1
16.1 × 12.0
14.4 × 12.0
14.5 × 11.6
15.0 × 11.8
15.2 × 11.9

0.049
0.05250.0536
0.0541
0.05450.05780.04430.0526one egg damaged
0.05580.05500.05730.0562

Notes.— In his classical monograph, Admiral Lynes (1930) included all populationsof C. exilis from Java, the Lesser Sunda Islands, and northern Australia, into one subspecies lineocapilla, but his material was very limited. Indeed, he was not even aware of theoccurrence of the species in West Java, where it is common, as I pointed out many years
ago (Mees, 1961b: 114). Mayr (1944a: 135) observed that birds from the Lesser Sunda
Islands have the colour of the underparts more deeply washed with ochre than specimens from northern Australia (Melville Island), and possibly worthy of subspecificseparation. My impression, gained from a superficial comparison of a series from WestJava with only a few specimens from tropical Australia, is that the former are brighter,
deeper in colour, than the latter. Note that the subspecies rustica, inhabiting the Moluccas and Sulawesi (Celebes), differs in a similar way from Australian lineocapilla.
The bird collected by Weber (31.xii) is in summer plumage, with a comparatively
short tail, the one collected by Verheijen (7.x) in winter plumage, with a long tail. Considering that the main breeding takes place in the months March to June, it is unexpected to have a bird in summer plumage in December. In retrospect, I much regret nothaving encouraged the Fathers to collect more material of the two Cisticola-species, bothof which are common on Flores. Material from all months of the year would be required, both for a study of geographical variation and for an understanding of seasonality in plumage variation.
Measurements:



wing
48
47

tail
33½
48

tarsus20
19.7

entire culmen
13.4
13.8

exposed culmen
10.8
10.5

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Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Orthotomus cucullatus everetti (Hartert)
Phyllergates everetti Hartert, 1897, Novit. Zool. 4: 517 – from 3000 and 4000 feet in S. Flores.
Collectors.— Everett (2), Rensch (1), Verheijen/Schmutz (12).
Material.— ?, 6.viii.1969, Ruteng (Verheijen, RMNH no. 65409); , 10.xi.1969, Ruteng (Verheijen, RMNH
no. 65410); , 27.x.1970, Ruteng (Verheijen, RMNH no. 66193); , 24.v.1971, Waé-Ntjuang, Langkas, 900
m (Verheijen, RMNH no. 85206); , 12.vi.1971, Nantal, Rahong (Verheijen, RMNH no. 85207); , 23.
vi.1973, Waé-Ntjuang (Verheijen, RMNH no. 85208); , 25.i.1976, Ulu Ros (Verheijen, RMNH no. 85209),
2 , 4.ii.1976, Ero (Verheijen, RMNH nos. 85210, 85211); , 16.ii.1976, Raé (Verheijen, RMNH no. 81410);
, 22.ii.1976, Ulu Ros (Verheijen, RMNH no. 85212); , 26.ii.1976, Rae’ (Schmutz, RMNH no. 97135).
Eggs (fig. 15g): c/3, 26.v.1958, Léwé (RMNH no. 375a); c/3, 29.iii.1960, Mataloko (RMNH no. 375b).
Measurements and weights:

RMNH no. 375a

RMNH no. 375b

16.1 × 11.9
16.4 × 12.2
16.9 × 12.0
14.4 × 12.3
15.2 × 12.4
15.5 × 12.5

0.0522
0.0536
0.05750.05570.05720.0577

Notes. –– As will be evident from the remarks given under the heading “Collectors”, this endemic subspecies was previously known from very few specimens.
Measurements:
8
4

wing
47½-50(48.5)
43-49(45.9)

tail43½-47½(46)41, 44(42.5)

tarsus19.5-21(20.1)
17.8-19.6(19.0)

entire culmen
17-18.6(17.8)
17-18(17.7)

exposed culmen
13-15(13.8)
12.8-14.7(13.6)

Phylloscopus borealis xanthodryas (Swinhoe)
Phyllopneuste xanthodryas Swinhoe, 1863, Proc. Zool. Soc. Lond.: 296 – Amoy.
Phylloscopus borealis examinandus Stresemann, 1913, Novit. Zool. 20: 353 – Bali.
Collectors.— Allen (1 or more), Weber (2), Everett (at least 4), de Jong (2), Verheijen/Schmutz.
Material.— (?), 19/24.xii.1888, Kotting (Weber, RMNH, skinned from alcohol); (?, 20.xii.1969, Todo (Verheijen, RMNH no. 65432); 2 , 22.xii.1969, Todo (Verheijen, RMNH nos. 65434, 65435); , , 14.vi.1973,
Golo-Karot, Borong (Verheijen, RMNH nos. 69763, 69764).

Notes.— There is still controversy over the number of subspecies to be recognizedin this species. When authorities cannot agree on the subspecies, their characters anddistribution, in the breeding quarters, speculation about the identity of birds in theirwinter quarters appears to be even less meaningful. I note that Dickinson et al. (1991:
324-325) have no such qualms, and record as winter visitors to the Philippines no lessthan six subspecies, one of which an undescribed one, large, with a spotted breast. In
the absence of any explanatory notes, one is left to wonder on what the specific andeven the generic allocation of this form was based.
The subspecies examinandus was based on material from the Lesser Sunda Islands,
including Flores. Later, this form was synonymized with xanthodryas, which, therefore,
would be the correct subspecific name for the birds from this region. P. b. xanthodryas isMees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

161

distinguishable from the nominate race by larger size, according to the literature. On
the basis of published measurements (Stresemann, 1913b: 353, s.n. examinandus; Vaurie,
1959: 289), the specimens from Flores are, sex for sex, too large for nominate borealis, andbelong to xanthodryas.
Measurements:

3
2 (?)

wing
71
67, 67,
67
66, 68tail49
44, 45,45
44, 44tarsus19.4
18.6, 19.3,
19.5
19.8, 19.8

entire culmen
14.0
13.9, 14.6,
14.7
14.8, –

exposed culmen
11.0
10.0, 10.2,
10.2
10.0, –

Phylloscopus presbytes floris (Hartert)
Acanthopneuste floris Hartert, 1898, Novit. Zool. 5: 114 – Flores.
Collectors.— Everett (“a series”), Rensch (5), Verheijen/Schmutz.
Material.— , 2 , 30.ix.1971, Hotju, Ruteng, 1500 m (Verheijen, RMNH nos. 85192, 85191, 85193); ,
21.xi.1971, Laréng Pongkor, 1100 m (Verheijen, RMNH no. 85194); [], 15.iii.1972, Nggolong-Tedé (Verheijen, RMNH no. 85195); , 14.iii.1973, Nggolong-Tedé, 1900 m (RMNH no. 85197); , 8.iii.1976, Ulu
Tukenikit (RMNH no. 85199); [], 22.ii.1976, Ulu Ros (RMNH no. 81377); , 22.iii.1976, Poco Nernancang (RMNH no. 85200); , 24.iii.1976, Poco Nernancang (RMNH no. 85201); [], 29.iii.1976, Poco Nernancang (RMNH no. 85202); , 5.vi.1976, Ulu Ros (RMNH no. 81412); , 23.ix.1976, Poco Nernancang(RMNH no. 81415).
No eggs.

Notes.— On Flores, this species was taken by Everett at “elevations from 3000 and3500 feet”, which is 900-1050 m (cf. Hartert, 1897b: 525), by Rensch between 1200 and
1400 m, and by the fathers at 1100-1900 m, giving it a total vertical range of 900-1900 m.
In contrast, nominate P. p. presbytes from Timor seems to have a greater vertical range:
Everett obtained it at Atapupu, on the coast, Haniel at Lelogama (845 m) and Bonleo(1100 m), Stein at Noilmina (up to 300 m), Mutis (1800-2300 m) and Ramelan (20002300 m).
Measurements:
7
6

wing
57-60
(58.7)
53-55(54.2)

tail40-43(41.9)
38-40(38.9)

tarsus20-21.5(20.8)
19.4-20.8(20.2)

entire culmen
12-14(13.0)
12.8-13.2(13.0)

exposed culmen
9.8-10.7(10.1)
9.9-10.3(10.1)

Seicercus montis floris (Hartert)
Cryptolopha montis floris Hartert, 1897, Novit. Zool. 4: 171 – the hills of South Flores.
Collectors.— Everett (“a series”), Rensch (9), Verheijen/Schmutz.
Material.—  with large gonads, 18.ix.1971, Potjo Gurung, Ruteng (RMNH no. 85190); , 14.iii.1973,
Nggolong-Tedé, ca. 1900 m (RMNH no. 85196); , 15.iii.1973, Nggolong-Tedé, 1950 m (RMNH no.
85198); , 8.iii.1976, Ulu Tuké Nikit (RMNH no. 81363); , 18.iii.1976, Ulu Tuké Nikit (RMNH no.
81414); , 27.vi.1976, Puar Lui (?) (RMNH no. 85203); , , 23.ix.1976, Poco Nernancang (RMNH nos.
85205, 85204).
No eggs.

162

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Notes.— The re-appearance of this species on Flores and Timor, with the nearest
known populations in Borneo (S. m. montis) and Sumatra (S. m. inornata) is surprising.
For its distribution to make sense zoogeographically, the species ought to occur in Java.
It would be tempting to look upon S. grammiceps, from the mountains of Java and Bali,
as its geographical representative, but in Sumatra both species occur.
The Timor subspecies paulinae was partly based on larger size, compared with floris,
viz., wing of 2  53.5, 54.5, 3  51, 51, 52 mm, against wing  50-52,  47-49 mm, in
specimens of floris (Mayr, 1944a: 159). The present material of floris averages a little
larger than that studied by Mayr, and shows that the size-difference between floris andpaulinae is very slight.
Measurements:
3
5

wing
51-53
(52.2)
48-50(48.9)

tail39-40(39.7)
37-38½(37.9)

tarsus17-17.6(17.3)
16-17(16.7)

entire culmen
11-11.8(11.3)
10-10.9(10.5)

exposed culmen
8-8.4
(8.2)
7-8.1
(7.8)

Gerygone sulphurea sulphurea Wallace
Gerygone sulphurea Wallace, 1864, Proc. Zool. Soc. Lond. (1863): 490 – Solor Island.
Acanthiza tenkatei Büttikofer, 1892, Notes Leyden Mus. 14: 195 – Flores.
Collectors.— Allen, ten Kate, Everett (1), Rensch (9), de Jong (3), Verheijen (2, MCZ), Verheijen/Schmutz.
Material. –(?), iv/v.1891, Flores (ten Kate, RMNH cat. no. 1, skinned from alcohol, holotype of Acanthiza
tenkatei);
Eggs: 75 clutches of c/1 (30 ×), c/2 (33 ×) and c/3 (12 ×), collected in the months March (2), April (3),
May (25), June (15), July (3), August (5), September (8), October (9) and five not dated (RMNH nos.
71026-71097, 73515, 76583). See also under Chrysococcyx minutillus. The eggs are glossless white, withbrown spots concentrated around the blunt end.
Some measurements and weights:

RMNH no. 73515
RMNH no. 71085
RMNH no. 71086

16.7 × 12.2
17.0 × 12.1
16.6 × 11.4
16.8 × 11.2
16.7 × 11.5
16.7 × 11.6
16.8 × 11.7

0.05930.0631
0.05230.04830.05620.05730.0549Notes.— For a discussion of geographical variation of this species, I refer to a previous publication (Mees, 1986: 130-133).
Ficedula westermanni hasselti (Finsch)
Muscicapa Hasselti Finsch (ex Temminck), 1898, Notes Leyden Mus. 20: 94 – Java.
Collectors.— Everett, Rensch (4), Verheijen/Schmutz.
Material.— (?)juv., 20.viii.1969, Ruteng (Verheijen, RMNH no. 65424); , 19.ix.1971, Hotju, Ruteng, 1500m (Verheijen, RMNH no. 85092); (? )juv., 12.xi.1971, Langkas (Verheijen, RMNH no. 85091); , received
January 1974, Lingko-Laréng Pingkor (Verheijen, RMNH no. 85093); , 4.v.1976, Poco Nernancang (Verheijen, RMNH no. 81411); , 3.vi.1976, Ulu Ros (Verheijen, RMNH no. 81413); , 24.viii.1976, Poco Nernancang (Verheijen, RMNH no. 81311); , 28.viii.1976, Poco Nernancang (Verheijen, RMNH no. 85094).

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

163

Notes.— In spite of revisional work by, amongst others, Mayr (1944a: 161-162) andRipley (1952), such geographical variation as this bird shows is not much better understood now, than when Finsch (1898a) wrote his note, almost a century ago. The temptation to follow White & Bruce (1986: 358) in not admitting any subspecies has beenstrong, but in deference to the authors just mentioned I have retained the name hasselti
for birds from Flores. Ficedula westermanni is a mountain bird, and a forest bird, moreover, very widely distributed in both mainland and islands, all ingredients for strong
geographical variation, one would think.
Measurements:
3

wing
tail
56½-6039-40½(57.7)(39.8)

55
36*) Tarsus and culmen from one specimen only.

tarsus16

entire culmen
14.2

exposed culmen
9.5 *)–

12.8

9

Ficedula hyperythra vulcani (Robinson)
Dendrobiastes hyperythra vulcani Robinson, 1918, J. Fed. Malay St. Mus. 7: 235 – Tjibodas, slopes of theGedeh Volcano, 4-6,000 feet, Western Java.
Collectors.— Everett, Rensch (2), Verheijen.
Material.— , 12.xi.1969, Nggolong-Tedé, Ruteng, 1900 m (Verheijen, RMNH no. 65425); , , (?)juv.,
8.iii.1973, Mt. Mbépé, Ruteng, ca. 1700 m (Verheijen, RMNH nos. 85111, 85112, 85113); , , 13.iii.1973,
Mt. Nggolong-Tedé, 1950 m (Verheijen, RMNH nos. 85107, 85110); , 28.iii.1973, Danau Rana-Ka (Verheijen, RMNH no. 85108); (?) im., 3.viii.1973, Mt. Tado-Walok, Ruteng, 1700 m (Verheijen, RMNH no.
85109); , 8.iii.1976, Ulu Tuké Nikit (Verheijen, RMNH no. 81426). Iris brown; bill black, its basis below
slate, legs greyish.
No eggs.

Notes.— In accord with all previous authors, I have been unable to find any difference between specimens from Flores and topotypical F. h. vulcani, of which a large series
was available for comparison.
Measurements:
3
5

wing
59½-60½(60)
58-59(58.6)

tail
37-41
(39)
36-38
(37)

tarsus18.3-19(18.8)
18-19.1(18.7)

entire culmen
12.5-13.2(12.9)
12-13.2(12.7)

exposed culmen
8.6-9.8(9.1)
8.3-9.5(9)

Ficedula dumetoria dumetoria (Wallace)
Saxicola (?) dumetoria Wallace, 1864, Proc. Zool. Soc. Lond. (1863): 490 – Lombock.
Collectors.— Allen, Everett, de Jong (1), Verheijen (1, MCZ), Verheijen/Schmutz.
Material.— , 12.ix.1971, Tjeréng, 850 m (Schmutz, RMNH no. 85219); , 21.x.1971, Rana Kulan, Biting,
400 m (Verheijen, RMNH no. 85220). Iris brown, bill black, legs bluish purple.

Notes.— This species “was met with not infrequently in the lowlands of South
Flores by Everett” (Hartert, 1897b: 524). The specimen from Tjeréng was collected inprimary forest.

164
Measurements:2

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)
wing
62, 64tail42½, 44

tarsus19.0, 19.2

entire culmen exposed culmen
15, 15.9
11, 11.8

Culicicapa ceylonensis sejuncta Hartert
Culicicapa ceylonensis sejuncta Hartert, 1897, Novit. Zool. 4: 526 – South Flores.
Collectors.— Everett (number not given; apparently a series); de Jong (2), Schmutz (1).
Material.— , 17.vii.1969, Paku, Mbelawang creek, 300 m (Schmutz, RMNH no. 81407).
Eggs (fig. 15h): c/1, 24.xi.1958, Montjok (RMNH no. 71098), 15.5 × 12.0 mm, weight 0.0495 g.

Notes.— The egg was identified with a query, and was also listed with a query inVerheijen’s (1964: 199) table. I have compared it with eggs of C. c. ceylonensis from Java,
with which it agrees so well, that there remains hardly any doubt.
The geographical variation of C. ceylonensis is interesting, in that over its huge continental and Sunda Shelf range, from Pakistan, Sri Lanka (Ceylon) and India, to Sumatra, Borneo, Palawan, Java and Bali, it shows hardly any geographical variation,
whereas Flores and Sumba each have a reasonably well-differentiated subspecies. Incidentally, the type of the Sumba race, C. c. connectens is in Leiden (RMNH no. 14070), notin Berlin as Rensch (1931b: 378) wrote.
Measurements:


wing
56

tail
42

tarsus12

entire culmen exposed culmen
13.6
10.0

Rhinomyias oscillans (Hartert)
Microeca oscillans Hartert, 1897, Novit. Zool. 4: 170 – At elevations from 3000 to 3500 feet in South
Flores.
Collectors.— Everett, Rensch (7), Verheijen/Schmutz.
Material.— , 29.iii.1971, Linko Lareng Pongkor, ca. 700 m (Verheijen, RMNH no. 85103); , 21.xi.1971,
Laréng (Verheijen, RMNH no. 85102); , 20.ii.1976, Lingko Ros (Verheijen, RMNH no. 81440); ,
19.vi.1976, Ruteng, 1500 m (Schmutz, RMNH no. 85101); , , 30.vi.1976, Poco Nernancang (Verheijen,
RMNH nos. 81452, 81445); , 4.v.1976, Poco Nernancang (Verheijen, RMNH no. 81462); , 16.vi.1976,
Ulu Ros (Verheijen, RMNH no. 81405); , 19.vii.1976, Potjo Nernancang (Verheijen, RMNH no. 85106);
, 2.ix.1976, Poco Nernancang (Verheijen, RMNH no. 85105); , 25.ix.1976, Poco Nernancang (Verheijen,
RMNH no. 85104); , 29.vi.1978, Golo Léhot (Verheijen, RMNH no. 81447); (?), without label, received
in 1983 (RMNH no. 81444). Iris brown, bill and legs black.

Notes.–– This interesting endemic form, placed in the genus Microeca until Rensch(1931a: 559 ) transferred it to Rhinomyias, has been regarded as conspecific with R. stresemanni, an endemic form of Sumba. The relationship has been discussed repeatedly, e. g.
Mayr (1944a: 143), who considered the matter a borderline case, and the judgement
whether to treat the two as conspecific or as different species, as purely subjective. Vaurie (1952: 15-16) enumerated the not inconsiderable differences in plumage and measurements between the two forms. Besides, R. oscillans is an inhabitant of mountain forest at ca. 900-1500 m, whereas R. stresemanni is a lowland bird, of light forest and more
open habitat. From this, one would almost expect the conclusion that they are different
species, but surprisingly, Vaurie does not further evaluate the differences, and without

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

165

explanation lists R. stresemanni as a subspecies of R. oscillans. This has been followed bylater authors, such as White & Bruce (1986: 353). My own conclusion, influenced by thecurrent, narrower, species concept, is the opposite: although there is an evident relationship between them, I would regard R. stresemanni and R. oscillans as different species. Whether these species have originally reached the Lesser Sunda Islands from Javaand Bali, or from Sulawesi (Celebes), is a question that cannot be answered. Vaurie(1952: 6) placed R. oscillans (with R. stresemanni) remote from all its congeners, with thesole comment: “This species consists of two well-marked races”.
Measurements:
9
3
(?)

wing
74½-80(76.8)
76-78
(77)
77

tail
52-59
(55.6)
53-54(53.7)
56

tarsus16-18(16.9)
17.7-18(17.8)
16

entire culmen
14.8-16.2(15.2)
15-16(15.3)
15

exposed culmen
10-12(11.2)
12(12.0)
11.7

Rhipidura diluta diluta Wallace
Rhipidura diluta Wallace, 1864, Proc. Zool. Soc. Lond. (1863): 491 – Flores.
Collectors.— Allen (several specimens, cf. Warren & Harrison, 1971: 149-150), Colfs (1), Weber (2), Everett(“a fine series”), Rensch (7), de Jong (1), Verheijen/Schmutz.
Material.— (?), 1880 (Colfs, RMNH cat. no. 1); 2. xii.1888, Maumeri (Weber, RMNH cat. nos. 2, 3); ,
20.v.1969, Wae Laci, 50 m (Verheijen, RMNH no. 81428); , 20.xi.1971, Laréng, Pongkor (Verheijen,
RMNH no. 85097); , , 21.xi.1971, Laréng, Pongkor (Verheijen, RMNH nos. 85096, 85099); , 14.
iii.1973, Mt. Nggolong-Tedé, Ruteng, 1900-1950 m (Verheijen, RMNH no. 85217); , , 5.i.1976, Puar
Léwé (Schmutz, RMNH nos. 85098, 81425); (?), 20.iii.1976, Ulu Tuké Nikit (Verheijen/Schmutz, RMNH
no. 81423);  juv., 31.iii.1976, Puar Lui (Schmutz, RMNH no. 85095). Iris brown, bill and legs black.
Eggs: c/2, 6.vi.1962, Mataloko (RMNH no. 71114). This clutch was identified as R. rufifrons, and an oblique reference to it, under that name, was made by Verheijen (1964: 194). The combination of the unusual locality (1000 m), the fact that neither of the Fathers has seen R. rufifrons on mainland Flores, theslightly larger size and greater weights of these eggs, compared with authentic eggs of R. rufifrons semicollaris from Palué, makes me confident of the re-identification. In colour and pattern, there is no difference.
The four clutches recorded by Verheijen (1964: 199) under R. diluta, were obviously misidentified. They
will be discussed below.
Measurements and weights:

RMNH no. 71114

17.0 × 13.3
17.7 × 13.5

0.08280.0871

Notes.— Four clutches identified as R. diluta were received with Verheijen’s collection, to wit: c/1, 21.v.1958, Wesang (RMNH no. 71100); c/2, 18.vi.1958, Léwé (RMNH
no. 71101); c/2, 6.v.1972, Léong (RMNH no. 71102); c/1, 19.ii.1959, Léong (RMNH no.
71099).
The months of collecting agree with those given by Verheijen, and confirm thatthese are the same clutches on which he based his record. The egg dated February(RMNH no. 71099) measures 26.4 × 18.5 mm; it is much too large to belong to any kindof flycatcher, and must belong to a thrush-sized bird. The other eggs measure 20.0-22.3
× 14.7-15.7 mm. Schönwetter (1976: 762-763) noted that the eggs of members of the genus Rhipidura are remarkably uniform. From the measurements provided by him, it166

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

appears that only R. leucophrys (of which the  has a wing-length of ca. 100 mm), has
eggs of about the size of the eggs ascribed to R. diluta (wing-length of the  ca. 81 mm).
The pinkish colour of the eggs is also different from assorted Rhipidura-eggs in our collection, and from Schönwetter’s description.
In this connection it must be remembered that R. diluta is a member of the R. rufiventris group of forms, and is often regarded as a subspecies of R. rufiventris (cf. Watson
et al., 1986b: 538). It is quite inconceivable that its eggs would be larger, and differently
coloured, than those of the other members of its group. Verheijen’s (1961: 185) remarkthat the nests of R. rufifrons: “resemble those of the Monarcha flycatchers rather thanthose of the Rhipidura diluta”, supports the suggestion that the nests he ascribed to R.
diluta had been misidentified, for actually the nests of R. rufifrons and R. rufiventris arequite similar (cf. Frith, 1976: 390-392). Before I left Leiden, it did not occur to me to compare these eggs with eggs of Terpsiphone paradisi, to which I now think that they may
belong; this would also explain the Monarcha-like nests.
Measurements:
4
2

wing
80-82
(81.0)
78, 79tail
83-85(84.5)
81½, 79

tarsus
17.5-19(18.3)
17.8, 18.5

entire culmen
15-17(16.3)
15, 16.5

exposed culmen
12-13(12.8)
11.5, 11

A few remarks about Rhipidura from farther away. The senior author of White &Bruce (1986: 374-375) had correctly given R. rufiventris hoedti priority over buettikoferi, but
this was reversed by Bruce, who, unlike White, had apparently not read my discussion
on the subject (Mees, 1975b). The date of publication is correctly recorded by Watson &Mayr (1986: 538). The last-mentioned authors err, however, in including Java in the rangeof R. perlata, and the specimen of R. euryura in Leiden, supposedly from Sumatra, towhich they refer, is obviously mislabelled. Authors using a very wide species concept(Chasen), have sometimes treated R. perlata and R. euryura as conspecific, using the combination R. perlata euryura for birds from Java, and that may have contributed to the
misconception that R. perlata occurs in Java. I note that Watson & Mayr (l. c.) placed thetwo species not even close together, but separated by R. aureola and R. javanica.
Rhipidura rufifrons semicollaris S. Müller
Rhipidura semicollaris S. Müller, 1843, Verh. Nat. Gesch. Ned. Overz. Bez., Land- en Volkenk.: 184 – Fatoe
Leeoe, Timor.
Collectors.— Everett (several skins), Verheijen (2, MCZ; eggs only in RMNH).
Material.— No skins in RMNH.
Eggs: Eleven clutches from Palué, c/1 (2 ×) and c/2 (9 ×), collected in the period from 14.iv-5.v.1960(RMNH nos. 71103-71113). The eggs are cream colour, with larger and smaller dull-brown dots and usually also some darker, blackish brown spots, the former ill- defined, concentrated in a wreath around theblunt end.
Some measurements and weights:

RMNH no. 71104
RMNH no. 71105

17.0 × 12.7
17.1 × 12.7
16.9 × 11.9
16.9 × 12.3

0.0711
0.06730.0653-

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

167

RMNH no. 71108
RMNH no. 71111
RMNH no. 71113

16.1 × 12.8
16.6 × 12.9
16.4 × 12.4
16.6 × 12.6
16.8 × 13.0
16.9 × 12.9

0.07290.06950.07180.07330.07210.0726

Notes.— Verheijen (1961: 185) stated that he had never with certainty seen this species on the mainland of Flores (where it was collected by Everett), but he found it very
common on Palué. A year later, however, he obtained a clutch from Mataloko, which
he ascribed to this species, but in my opinion it is referable to R. diluta (see under thatspecies).
Considering that on Timor this same subspecies ranges from sea-level to at least
2000 m, and is evidently widely distributed, as is the very similar subspecies R. r. sumbensis on Sumba (cf. Mayr, 1944a: 136, 142), it is peculiar that on Flores it is so local andapparently confined to Palué and the coastal lowlands near Nanga Ramau (Everett).
Monarcha trivirgatus trivirgatus (Temminck)
Drymophila trivirgata Temminck, 1826, Recueil d’Ois. 3 (livr. 70): pl. 418 fig. 1 – Timor.
Collector.— Everett (at least 5).
Material.— None in RMNH, but: (?), x.1896, South Flores (Everett, AMNH no. 654672);  ad. (A.E.),
xi.1896, South Flores (Everett, AMNH no. 654670); (?), xi.1896, South Flores, shot below 1000 feet (Everett, AMNH no. 654671);  (A.E.), xi.1896, South Flores, shot below 1000 feet (Everett, BM no. 98.5.4.100);
 “nat. coll.” (=  ?), xi.1896, South Flores (Everett, BM no. 98.5.4.99). The initials A.E. mean that the
sexing was done by Everett personally.

Notes.— M. trivirgatus is a somewhat enigmatic species on Flores, as only A.H.
Everett and his native collectors ever obtained it, and the specimens are merely labelled“South Flores”. In view of possible interaction with the closely related M. sacerdotum,
more exact knowledge of the occurrence of M. trivirgatus is much desired. From Everett’s itinerary it is known that he made his headquarters at Nanga Ramau and as one of
the specimens is marked as having been shot below 1000 feet, and two were sexed byEverett personally (evidently, Everett stayed close to the village, whereas his hunters
went farther afield), it may be assumed that the specimens originate from the cultivatedlowlands and wastelands near Nanga Ramau. From the number of specimens collected(there may have been more than the five I examined), it is apparent, that at that time thespecies was not particularly uncommon there.
Measurements of the BM specimens:
wing
tail

67
65
[]
68
65

tarsus18.5
18.1

entire culmen
15.5
16.0

Monarcha trivirgatus wellsi (Ogilvie-Grant)
Piezorhynchus wellsi Ogilvie-Grant, 1911, Bull. Brit. Orn. Cl. 27: 105 – Goram Laut.

exposed culmen
11.0

168

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Notes.— Discussing M. t. nigrimentum, Stresemann (1914a: 128) wrote: “Von dieser
Form unterscheidet sich die ihr nächststehende M. t. wellsi ... lediglich durch bedeutendere Ausdehnung der weissen Spitzen der drei äusseren Steuerfederpaare”. Mayr (inWatson et al., 1986b: 509, footnote) says of wellsi, that it is “very close to nigrimentum”.
White (in White & Bruce, 1986: 366) states that: “nigrimentum is very like nominate trivirgatus but has less white on the tail (includes.... wellsi)”.
Peculiarly, all RMNH specimens show another, and conspicuous, difference: nigrimentum has only a small triangle on the chin black, whereas wellsi has a black bib, extending over chin and throat; in this character, wellsi would seem to agree with nominate trivirgatus and not with nigrimentum. Unfortunately, BM material examined by me,
does not support the difference, but it is worth recording, as further study is evidently
required.
Differences from M. t. trivirgatus: the bill is longer and more slender; the belly looks
whiter (the rust colour of the flanks reaching less far backwards), the third pair of rectrices has only a little white (sometimes none at all).
The sex of our eight specimens is indicated by Finsch as “”. Finsch took the adultplumage (that is similar in both sexes), for the  plumage, and the juvenile plumage(of both sexes), for the  plumage. As of the eight specimens, seven are adult and one
subadult, he provided them with a sex indication, corresponding with his erroneous
notion.
Monarcha trivirgatus nigrimentum G.R. Gray
Monarcha nigrimentum G.R. Gray, 1860 (1861?), Proc. Zool. Soc. Lond.: 352 – Amboyna.
Monarcha bernsteinii Salvadori, 1878, Ann. Mus. Civ. Genova 12: 322 – Salvatti (errore!) = Ambon, seediscussion below.

Notes.— The occurrence of a distinctive subspecies of M. trivirgatus on Salawatti,
which has been generally accepted since the description of M. bernsteinii (cf. Mayr,
1941b: 135; Rand & Gilliard, 1967: 396; Mayr in Watson et al, 1986b: 509), did not seemto me obvious from the zoogeographical point of view. Salawatti is separated frommainland New Guinea by a narrow and shallow strait, and until only a few thousandsof years ago it was still connected with the main island. One would not expect endemism and as far as I am aware not a single endemic bird species or subspecies has been
described from Salawatti, except for M. bernsteinii.
Note that in the original description the spelling is bernsteinii, but that all later authors, beginning with Salvadori (1879: 493) himself, use the spelling bernsteini.
In the original description, bernsteinii was only compared with M. bimaculatus: “Iltipo esistente nel Museo di Leida è una femmina, differente da quella del M. bimaculatus
per le dimensioni maggiori, e per avere questa soltanto il mento nero”.
A comparison of the type of M. bernsteinii ( ad.) with material of diverse species
and subspecies of Monarcha inhabiting the Moluccas, showed that it agrees almost
completely with a  ad. of M. t. nigrimentum from Ambon, both in measurements andin plumage, the only difference being that it has more black on the forehead. Therefore
I do not hesitate to place M. bernsteinii in the synonymy of M. t. nigrimentum. Although
it would be difficult to prove that the type of M. bernsteinii is not from Salawatti, itMees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

169

seems most unlikely that two islands so far apart would be inhabited by the same subspecies. It is an obvious guess, on the basis of the arguments presented above, that theprovenance “Salawatti” ascribed to the type is erroneous, due to an inadvertent exchange of labels, and that actually it was collected on Ambon. There is no mention of
the specimen in Bernstein’s diary (v. Musschenbroek, 1883), but that is inconclusive
evidence, as he did not take notes on every specimen brought in by his hunters. Thismeans that probably also the date of collecting and perhaps the name of the collector
are erroneous. All that can be said is that the bird must have been collected about 1865
(on Ambon).
It is peculiar that (excluding the type of bernsteinii) our collection holds only one
adult specimen of M. t. nigrimentum from Ambon, as opposed to 7 in immature plumage. A bird found nesting on Ambon by Lieftinck (1950) was also in immature plumage,as mentioned by van Bemmel in his postscript to Lieftinck’s note. Of the five birds from
Ceram, on the other hand, four are adult. Birds from Ambon (type-locality of nigrimentum) and Ceram differ somewhat: the latter are smaller, the breast is of a deeper ferruginous, the abdomen whiter. The differences are slight and do not require expression innomenclature.
Measurements:
wing
tail
“Sailolo, Salawati” (type of M. bernsteinii)

81
70
Ambon (M. t. nigrimentum)
80
70
Ceram (M. t. nigrimentum)
4
75-76
64-68

72½
63

tarsusentire culmen exposed culmen

20

16+

13+

19

17.6

13.7

17.2-18
18

16-17
17

13-13.3
13.4

Monarcha (trivirgatus) boanensis van Bemmel
Monarcha trivirgata boanensis van Bemmel, 1939, Orn. Mber. 47: 152 – Boano.
Material.— , 13.v.1918, Boano (L.M.R. Rutten, RMNH no. 14055, holotype).

Notes.— In the diagnosis, M. t. boanensis was stated to have: “einen weissen Bürzel”,
a character that would link it to M. everetti from Tanahdjampea, the only other Monarcha
species with a white rump (cf. van Bemmel, 1939). Examination of the type (RMNH no.
14055) revealed, however, that the rump and upper tail-coverts are black, and that thesuggestion of a white rump was caused by the long silky white feathers of the lower
flanks being brushed up, to cover the black rump. Hence, the supposed affinity to M.
everetti was based on incorrect observation.
Measurements:


wing
81

tail
71

tarsus19½

entire culmen exposed culmen
17¼
13¼

Van Bemmel gave wing 79, tail 78 mm; I am definitely unable to attain such a large
tail-length, there must be a fundamental difference in method of measuring (cf. Mees,
1982a: 110; 1986: 21). The other characters enumerated by van Bemmel to distinguishboanensis from loricatus are correct: a white cross-bar on the forehead, a smaller throat-170

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

patch, and the outer rectrices with the basal one-third black (in loricatus white overtheir whole length).
Recently a second specimen was collected: welcome evidence that this interesting
form, be it species or subspecies, still exists. The naive way in which the discoverers
announced their find, and the outright stupid, if not dishonest, reaction of some authorities, has unfortunately led to some negative publicity (van den Broek, 1991). The
specimen will be deposited in the RMNH, but I have not yet seen it.
This, therefore, is an excellent form, either species or subpecies, apparently closer insome respects to M. loricatus than to nigrimentum. The white (or pale) bar across theforehead is otherwise only found in diadematus (Obi), which, however, has much smaller white tail-markings, is a smaller bird, etc.
Since the preceding notes were written, M. boanensis has become the subject of further
field studies, as well as an interesting study of its affinities (Moeliker & Heij, 1995,
1996).
Monarcha sacerdotum Mees
Monarcha sacerdotum Mees, 1973, Zool. Meded. 46: 179 – Sesok, Flores, 1000 m.
Collector.— Schmutz.
Material.— , 25.ix.1971, Sésok, 1000 m (Schmutz, RMNH no. 68135, holotype); , 25.vi.1975, Paku, 800
m (Schmutz, RMNH no. 85260).

Notes.— For the original description of this species, a single male only was available, so that there remained some room for speculation about the appearance of the female (cf. Mees, 1973; Mayr & Vuilleumier, 1983: 219).
In plumage, the female agrees almost entirely with the male, thus confirming thespecific distinction of M. sacerdotum, but it is about 5% smaller in linear measurements.
The male has the outer pair of rectrices 60 mm in length, the outer vane white to its
base, the inner vane white over a distance of 38 mm. The female has the outer pair of
rectrices 56 mm in length, the outer vane white over a distance of 42 mm, the inner vane
over a distance of 34 mm.
Both specimens were collected in pockets of rainforest, such as on Flores are foundonly in the westernmost part. Observations by Butchart et al. (1996: 346-348) support
Schmutz’s description of the habitat and indicate a rather wider distribution than waspreviously known, with records from levels of 350-970 m. Father Schmutz has informedme that the altitude given for the type specimen, 1000 m, was probably overestimatedand might be no more than 800 m. It appears that M. sacerdotum is an inhabitant of the
middle levels, not a mountain bird.
I agree with Mayr & Vuilleumier (l.c.) that a revision of the genus Monarcha is desirable; it could also be most fascinating. I fail to understand, however, why these authors
consider it “unfortunate” that, in the original description, I did not compare this species
with Monarcha manadensis, as I cannot see any close resemblance at all: M. manadensis isblack and white, without grey, and its tail is entirely black. There is no doubt in mymind that the closest relatives of M. sacerdotum are M. trivirgatus and M. mundus, as
stated in the original description, and I am glad to see that it is between these two species that Mayr (in Watson et al., 1986b: 508) has now placed it.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)
Measurements:



wing
74
69

tail
73
66

tarsus17
17.5

171
entire culmen
15.25
14.75

exposed culmen
12
11.5

Monarcha cinerascens cinerascens (Temminck)
Drymophila cinerascens Temminck, 1827, Recueil d’Ois. 3 (livr. 72): pl. 430 fig. 2 – Timor.
Collector.— Verheijen (1, MCZ; and eggs).
Eggs (fig. 15i): c/1, 14.vi.1960, Léi, Palué (RMNH no. 71115); c/1, 20.vi.1960, Léi, Palué (RMNH no.
71116); c/2, badly damaged, 21.vi.1960, Léi, Palué (RMNH no. 71117); c/2, 24.vi.1960, Léi, Palué (RMNH
no. 71118); c/2, 3.v.1960, Léi, Palué (RMNH no. 71119).
Measurements and weights:

RMNH no. 71115
RMNH no. 71118
RMNH no. 71119

22.8 × 15.2
21.7 × 15.8
22.6 × 15.9
22.2 × 16.0
22.9 × 16.5

0.14290.15670.16310.16120.1700Notes.— This species is not known from the mainland of Flores, but only from theisland of Palué, where it was discovered by Verheijen (1961: 185), who noted: “eggs,
nest and bird collected”. The eggs, as listed above, and three nests, are in our collection,
but not the bird, so that I cannot say much about its subspecific relationship. Mayr (inWatson et al., 1986b: 502) included it in M. c. disjunctus, but in my opinion, expressed
elsewhere, this is a synonym of the nominate race (Mees, 1965: 184-186). Therefore I
have felt justified in referring the population inhabiting Palué to the nominate race, although I have not examined specimens.
Hypothymis azurea symmixta StresemannHypothymis azurea symmixta Stresemann, 1913, Novit. Zool. 20: 294 – Alor.
Hypothymis azurea javana Chasen & Kloss, 1929, Bull. Raffles Mus. 2: 22 – Badjoelmati, East Coast ofJava.
Hypothymis azurea penidae Meise, 1941, J. f. Orn. 89: 361 – Noesa Penida.
Collectors.— Allen, Colfs, Weber, Everett, Rensch (3), de Jong (2), Verheijen (1, MCZ), Verheijen/
Schmutz.
Material.— , 1862, Flores (Allen, RMNH); , v.1880, Flores (Colfs, RMNH); (?), xi/xii.1888, Maumeri(Weber, no. 32, ex alcohol, RMNH); , 3.vi.1969, Nunang, 650 m (Schmutz, RMNH no. 81374); , 24.
vi.1969, Sésok, 610 m (Verheijen, RMNH no. 81371); , 23.xii.1969, Todo (Verheijen, RMNH no. 65401);“” = , 23.xii.1969, Todo (Verheijen, RMNH no. 65402); , 19.ix.1970, Kisol (Verheijen, RMNH no.
65403); , 4.iii.1971, Rana Loba, Borong (Verheijen, RMNH no. 85406); , 12.vi.1973, Golo-Karot, Borong (Verheijen, RMNH no. 85407); , 14.vi.1973, Golo-Karot, Borong (Verheijen, RMNH no. 85408); 2, , 16.vi.1973, Borong coast, in mangrove (Verheijen, RMNH nos. 85405. 85409, 85410); , 5.v.1978,
Flores (Schmutz, RMNH no. 81373).
Eggs: 25 clutches of c/1 (7 ×), c/2 (16 ×) and c/3 (2 ×), collected in the months April (2), May (10), June(4), July (1), August (1), September (5), November (1), and one insufficiently dated (RMNH nos. 7112071144). The eggs are white with smallish brown spots, sometimes concentrated in a wreath around theblunt end.

172
Some measurements and weights:

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)
RMNH no. 71136
RMNH no. 71138
RMNH no. 71141
RMNH no. 71142
RMNH no. 71143

17.6 × 13.9
18.0 × 13.3
17.8 × 13.2
17.0 × 13.7
17.5 × 14.1
17.0 × 13.0
17.4 × 13.0
17.7 × 12.9
17.9 × 12.6

0.0901
0.09800.08570.0906
0.09690.07960.07820.08140.0736

Notes.— Hypothymis azurea is a species with a fascinating geographical variation:very little variation over its huge continental range, and the large continental islands,
but with on some small, peripheral islands conspicuously modified forms, deeper incolour and larger in size (cf. Rand, 1970). That in the great days of the study of intraspecific variation it should have attracted the attention of systematists is only natural, see
the studies by Oberholser (1911), Stresemann (1913a) and several other authors who,
after Stresemann’s revision, added another 10 subspecies. Many of these named subspecies date from a period that the subspecies-concept was different from the present
one, and that average differences (in very small series!) were considered adequate for
their recognition. For example, when Stresemann described symmixta, he did not havea single specimen from Java for comparison. Similarly, H. a. javana was subsequently
described without material from the Lesser Sunda Islands, and even without mentionof symmixta.
A comparison between series from Java and from the Lesser Sunda Islands failed toreveal any difference. Males from Borneo (prophata) are very slightly different: on theundersurface, the black pectoral band is usually wider, and the blue of throat and breastis brighter, more clearly blue, and also continued farther downwards than in birds from
Java, which have this colour duller and a trifle more violet-blue, less clear blue. Notethat this description is almost the opposite of that given by Stresemann, who claims thatsymmixta is brighter blue, less violet-blue than prophata!
Hoogerwerf’s (1964) notes clearly support my own conclusion, that when adequate
material is examined, individual variation covers the differences that have been claimed
to exist between the subspecies symmixta and javana. It was only the insufficiency of hismaterial, that kept Hoogerwerf (1964: 213-214) from uniting the two.
White & Bruce (1986: 364-365) referred populations from the Lesser Sunda Islands
to prophata, stating that it was: “doubtful if they are separable from Sundaland prophata”(in the nomenclature used, they have transformed this doubt into certainty!). In their
discussion, I find no evidence that these authors based their opinion on a personal examination of material.
The Philippine material available to me is inadequate, but if symmixta is not recognized as different from prophata, I do not believe that prophata could be maintained as
different from nominate azurea (at present considered to be confined to the Philippines).
Van Marle & Voous (1988: 191) state that the subspecies abbotti is: “endemic on Banyak Islands”, but that is not correct, for this striking form is only known from the islands of Lasia and Babi, south of Simalur (cf. Junge, 1936: 48). The Banjak Islands, nearer mainland Sumatra, are inhabited by H. a. prophata (cf. Ripley, 1944: 397-398).

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

173

The subspecies H. a. penidae was based on a comparison with but very few specimens from Java, Bali, Sumbawa, and Flores. Meise found the males indistinguishable
from Javanese birds of that sex, the females, on the other hand, only distinguishable
from females of symmixta by having the cap a trifle lighter blue, as in javana; the mantle
and back, however, agreeing with symmixta and differing from javana by being slightly
greyer, less brownish. As mentioned above, I am unable to distinguish between javana
and symmixta, and that removes automatically the basis for the recognition of penidae,
which was described as intermediate between these two. Note that previously Stresemann (1913b) and Rensch (1931a: 556-557) had made the claim that: “Die Stücke vonBali sind intermediär zwischen dieser [meant is symmixta] und der westlich anschliessenden Rasse prophata”.
Large size is a common phenomenon in birds inhabiting small islands, comparedwith their relatives on the nearest larger land masses, but the fact that the males of thesmall-island populations are more brilliantly coloured (abbotti, karimatensis), is remarkable, as the usual tendency is, in sexually dimorphic species, for males in these conditions to lose some of their male plumage characters, and to be closer in plumage to thefemales.
Measurements:
wing
Flores (H. a. symmixta)
7
68½-71(69.6)
2
65, 69Java (H. a. “javana” = symmixta)
6
68-72(69.8)
Taiwan (H. a. oberholseri)
10 
71-74
(73.1)

tail

tarsusentire culmen

exposed culmen

62½-70(66.0)
59, 66

14.5-16(15.4)
15, 16

14.0-14.9(14.5)
14.0, 14.3

10.0-10.5(10.1)
10.0, 10.5

64-70
(66.3)

16.16.5(16.1)

13-15(14.3)

9.7-11(10.2)

69-74(71.7)

15.3-16.8(16.0)

14.2-16(15.3)

9.5-11(10.2)

Terpsiphone paradisi floris Büttikofer
Terpsiphone floris Büttikofer, 1894, in Weber: Zool. Ergebnisse 3: 293, pl. XVIII fig. 1-3 – Flores.
Collectors.— Allen (3 or more), Colfs, Weber (7), Everett (a series), Moraux (1), Rensch (1), de Jong (5),Verheijen/Schmutz.
Material.— , iv.1880, Flores (Colfs, RMNH cat. no. 7, described and figured by Büttikofer); 5 , ,v.1880, Flores (Colfs, RMNH cat. nos. 2-6 and Doubl.); , vi.1880, Flores (Colfs, RMNH cat. no. 1); ,
23/25.xi.1888, Reo (Weber, RMNH cat. no. 8a, skinned from alcohol); , 26.xi.1888, Bari (Weber, RMNH
cat. no. 8, skinned from alcohol);  in change, 26/28.xi.1888, Bari (Weber, RMNH cat. no. 10, skinned
from alcohol);  in change, xii.1888, Maumeri (Weber no. 48a, RMNH cat. no. 9); , , skeletons,
xii.1888, Maumeri (Weber, RMNH cat. a, b, cf. van Oort, 1907: 213); , vi.1897, Labuan Badjo (P. Moraux,
RMNH cat. no. 11, flat skin); , 16.v.1969, Nunang (Schmutz, RMNH no. 81109); (?), 19.xii.1969, Todo
(Verheijen, RMNH no. 65415); , 18.vii.1971, Tjereng, 500 m (Schmutz, RMNH no.xxx); , 17.vi.1973,
Waé Reca, Borong (Verheijen, RMNH no. 85100).
Eggs (fig. 15j): c/2, 2.x.1948, Rekas (RMNH no. 71145); c/2, 23.xi.1954, Montjok (RMNH no. 71146); c/2,
8.xi.1955, Flores, 600 m (RMNH no. 76584); c/3, iii.1956, Rekas (RMNH no. 71147); c/2, xii.1958, Nunuk(RMNH no. 71148).

174

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Measurements and weights:

RMNH no. 71145
RMNH no. 71146
RMNH no. 76584
RMNH no. 71147

21.1 × 15.9
21.3 × 15.6
20.4 × 16.2
20.7 × 16.1
21.7 × 16.2
21.7 × 16.3
21.4 × 16.6
21.6 × 16.5
21.7 × 16.7

0.13880.14620.13380.14480.15470.15740.1636
0.159 (large hole)
0.1654

Notes.— Although Büttikofer’s paper in the “Zoologische Ergebnisse” is generally
accepted as having been published in 1894, I note that Meyer (1894), in a postscript dated
5 January 1894, states that he had already received it. Probably reprints have been mailed
before the end of 1893 (in Orn. Mber. 2, 1894: 63, the year of publication is actually given
as 1893), but certainly it has been published in the very first days of January 1894.
Watson et al. (1986b: 488) give as type-locality for T. p. floris, Reo. This is not quite
accurate: in the description of the subspecies, Büttikofer began with an enumeration of
the material collected by Weber: “Ein altes Männchen in Spiritus von Reo, ein zweites...
ebenfalls in Spiritus, von Sikka, Balg eines Männchens im Uebergangskleide und ein
Weibchen in Spiritus, beide von Maumeri, Balg eines alten Männchens und zwei Exemplare in Spiritus (Männchen im Uebergangskleide und 491a, altes Männchen), alle drei
von Bari”. On the following page, Büttikofer lists all the material he ascribed to T. floris:
Sumbawa (3), Flores (16), Sumba (3), Ombaai (4). Included in the number from Flores isevidently a fully adult male, labelled Adoenara, Flores, June 1880 (Colfs, RMNH,
mounted). I am not quite sure that at that time Büttikofer realised that Adonara is aseparate island, not a place on Flores. T. p. floris seems to be a (hardly surprising) addition to the very insufficiently known avifauna of Adonara.
Two of the specimens from Sumbawa studied by Büttikofer, are from the enigmatic
Forsten collection from Bima, 1842. For some reason Rensch (1930: 103; 1931a: 631) believed that this species was unknown from Sumbawa before he collected it in 1927.
In Java, T. paradisi is in my experience a rare, or at least very uncommon, inhabitant of
the lowland forest. Therefore it is interesting that on some of the Lesser Sunda Islands,
where one would expect lowland forest to be less well-developed (especially Sumba), it
appears to be much more common.
Pachycephala fulvotincta fulvotincta Wallace
Pachycephala fulvotincta Wallace, 1864, Proc. Zool. Soc. Lond. (1863): 492 – Flores.
Collectors.— Semmelink, Allen, Weber, Everett, Rensch (7), de Jong (6), Verheijen (3, MCZ), Pfeffer (1),Verheijen/Schmutz.
Material.— , 1862, Larantoeka (Semmelink, RMNH cat. no. 1, mounted); , skeleton, xii.1888, Maumeri(Weber, cf. van Oort, 1907: 154); , 1/8.i.1889, Endeh (Weber, RMNH cat. no. 6); 19 , 8 , 2  im., froma number of localities in West Flores, from sea-level to 1100 m (Verheijen/Schmutz, RMNH nos. 5991859921, 65419, 65420, 66156, 81306, 81309, 81310, 81312, 81372, 81376, 81424, 85038-85054, 85326). Iris
brown, bill black, legs slate.
Eggs (figs 16a, 16aa): 29 clutches of c/1 (8 ×) and c/2 (21 ×), collected in the months April (2), May (9),June (9), July (2), August (5), and two insufficiently dated (RMNH nos. 71149-71174, 73516, 73517,
76585).

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)
Measurements and weights ofsome clutches:

175

RMNH no. 71149
RMNH no. 71153
RMNH no. 71155
RMNH no. 71156
RMNH no. 71163
RMNH no. 71165
RMNH no. 71171

22.3 × 16.5
22.3 × 16.6
22.1 × 16.4
22.3 × 16.4
22.5 × 17.4
22.6 × 17.1
21.3 × 17.0
22.0 × 17.0
21.7 × 15.4
22.8 × 15.6
21.3 × 16.2
21.3 × 16.4
22.9 × 16.1
23.0 × 16.4

0.15780.16580.18180.187
0.18040.18460.14630.13490.15910.1525

Notes.— When Galbraith (1967) separated specifically Pachycephala melanura ofnorthern Australia and New Guinea, from Pachycephala pectoralis of southern Australia, the numerous well-marked forms of the Moluccas and the Lesser Sunda Islands,
previously included in the all-encompassing P. pectoralis, were left hanging in the air.
Galbraith (l.c.) promised: “a reconsideration of the superspecies in the light of thepresent discovery”, a promise he repeated in different words some years later (Galbraith, 1974: 248), but it was not brought to fruition. Even White & Bruce (1986: 379381) refrained from taking the next step, which had become inevitable, of removing allthese forms from P. pectoralis, and re-arrange them. The simplest solution is to give
Galbraith’s (1956) various subgroups species status. In the area of the Lesser Sunda
Islands, this dismemberment results in the recognition of three species: P. calliope, P.
teysmanni, and P. fulvotincta. The last-mentioned species has the subspecies P. f. fulviventris, P. f. javana, P. f. everetti and perhaps P. f. jubilarii (the validity of jubilarii has beenquestioned).
Measurements:
Flores
20 
9
Alor

1

wing
76-83
(78.9)
77-81(78.6)
85

tail55-63½(58.6)
56-62(58.8)
62

tarsus19-22½(20.7)
20-21.5(20.9)
21

entire culmen
15-18.2(17.2)
16.7-18.2(17.3)
18.9

exposed culmen
13-15.8(14.2)
13.3-15.5(13.8)
14.8

Arguably, the subspecies jubilarii from Alor, based on large size, is too weakly differentiated for recognition (White & Bruce, 1986: 380); nevertheless, the single specimen
from Alor examined by me (not one of those previously studied by Rensch), supports
the rather large size of birds from that island.
Pachycephala nudigula nudigula Hartert
Pachycephala nudigula Hartert, 1897, Novit. Zool. 4: 171 – Flores meridionalis.
Collectors.— Everett, Rensch (21), Verheijen/Schmutz.
Material.— , x.1896, South Flores, 3500’ (Everett, RMNH cat. no. 1, syntype); , 22.xi.1971, Lareng,

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Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Pongkor (Verheijen, RMNH no. 85056); , 9.iii.1973, Tado Walok, Ruteng, ca. 1700 m (Verheijen, RMNH
no. 69761); , 15.iii.1973, G. Nggolong-Tedé, Ruteng (Verheijen, RMNH no. 69762);  im., 22.iii.1976,
Poco Nernancang (Verheijen, RMNH no. 81408); , 7.vi.1976, Poco Mulu, 1500 m, Ruteng (Schmutz,
RMNH no. 85055); , 2.x.1976, Poco Nernancang (Verheijen, RMNH no. 81416); , 28.vi.1978, Ulu Waé
Rukus (Verheijen, RMNH no. 81420); , 4.v.1978, Waé Rukus (Verheijen, RMNH no. 81422); , no data,
Flores (Verheijen/Schmutz, RMNH no. 81401). Iris dark brown, bill black, legs grey.

Notes.— In the mountains of Flores, above ca. 1000 m, this species is evidently common. The large series obtained by Rensch speaks for itself. The Sumbawa subspecies P.
n. ilsa, on the other hand, is still known only from the original 2  collected by Renschin 1927. This does not necessarily mean that it is rare. Rensch (1930: 85) indicated thatin the mountains near Batoe Doelang it was not uncommon, but a year later called it“wesentlich seltener” than the nominate race. Recent publications (Butchart et al., 1996:
358; Johnstone et al., 1996: 173) describe P. n. ilsa as common and widely distributed on
Sumbawa, with a vertical range of 200 to 1700 m. This suggests another species of which
populations on adjacent islands differ in vertical distribution.
According to Schmutz (MS), the species is not generally distributed above 1000 m;
for example from the mountain massif between Nunang and Sésok, which reaches 1230m (Potjo Dédéng), it seems to be completely absent. The only explanation I can think of,is that these mountains are not high enough, or the high ground is not sufficiently extensive, even though they are to well above the lower limit of distribution of P. nudigula
where it occurs on higher mountains.
Measurements:3 juv.
6

wing
102-108(104.3)
103
93-97(95.2)

tail
79-82
(80)
79
70-75
(73.3)

tarsus24-25.6(24.9)
24
22½(23.4)

entire culmen exposed culmen
20.6-23.0
17-18.2(21.9)(17.7)
20
15
2419.6-21.8
15.5-17.5(20.5)(16.8)

Parus major cinereus Vieillot
Parus cinereus Vieillot, 1818, Nouv. Dict. Hist. Nat. 20: 316 – Batavia.
Collectors.— Semmelink, Allen, Colfs, Weber (2, ZMA), ten Kate, Everett (3), Rensch (2), de Jong (4),Verheijen (5, MCZ), Verheijen/Schmutz.
Material.— 2 (?), 1862, Larantoeka (Semmelink, RMNH cat. nos. 28, 29); (?), iv.1880, Flores (Colfs, RMNH
cat. no. 30); (?), iv.1891, Sika (ten Kate, RMNH cat. no. 31); 18 specimens (12 , 5 , (?) juv.), collected
between 3.xii.1968 and 26.ii.1976, at Nunang (650 m), Ruteng (up to 1450 m), Todo, Langkas, WaéNtjuang Langkas, and Laé (Verheijen/Schmutz, RMNH nos. 59895-59900, 81404, 65377-65380, 66157,
66158, 85058, 85059, 85057, 85061, 85060).
Eggs: c/3, 18.v.1956, Mano (RMNH no. 71175); c/1, 6.v.1957, Léong, Méngé (RMNH no. 71176); c/1,
20.viii.1957, Manus (RMNH no. 71177); c/1, iv.1958, Todo (RMNH no. 71178); c/4, 27.iii.1961, Mataloko(RMNH no. 71179); c/5, 31.v.1961, Mataloko (RMNH no. 71180); c/4, 7.vi.1961, Mataloko (RMNH no.
71181); c/5, 15.vi.1961, Mataloko (RMNH no. 71182); c/4, 22.vi.1962, Mataloko (RMNH no. 71183). The
egg collected in August (a month not listed by Verheijen, 1964), has an original label, and is correctly
identified.
The eggs are white, well-covered with fairly light chocolate-brown primary spots, which are distributed
over the whole shell, but usually more concentrated around the blunt end, and somewhat sparse lighterand more greyish secondary spots.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)
Some measurements and weights:

RMNH no. 71181

RMNH no. 71182

177
15.6 × 13.6
16.3 × 13.7
17.1 × 13.5
17.8 × 13.1
16.6 × 13.3
17.0 × 13.1
17.4 × 13.3
17.9 × 13.3
18.0 × 13.2

0.086
0.08520.08110.08050.08310.08180.08530.08540.0895

Notes.— P. m. cinereus ranges over the whole length of Java, and the chain of Lesser
Sunda Islands to Alor and Sumba. It has failed to reach Timor. The absence of geographical variation in Java and the Lesser Sunda Islands indicates perhaps a fairly recent eastward expansion of the species. Its eastern limit also marks the eastern limit of distributionof the family Paridae, which is absent from the Australian region. The adjacent subspecies
to the West is P. m. ambiguus, which is reasonably well-differentiated from P. m. cinereus,by having the sides of the underparts light grey instead of almost white. P. m. cinereus is
an inhabitant of semi-open, including cultivated, country, such as is found most commonly in the lower and middle levels, but where suitable habitat exists it may occur highup in the mountains. Contrary to Stresemann (1939: 383) it is certainly not exclusively oreven mainly a mountain bird, nor has it been reliably recorded from Timor.
Dicaeum annae (Büttikofer)
Acmonorhynchus annae Büttikofer, 1894, in Weber: Zool. Ergebnisse 3: 301, pl. XVIII fig. 4 – Kotting,
Flores.
Acmonorhynchus annae sumbavensis Rensch, 1931, Mitt. Zool. Mus. Berlin 17: 617 – Batoe Doelang, WSumbawa.
Collectors.— Weber, Everett, Rensch (8), de Jong (4), Verheijen (4, MCZ), Verheijen/Schmutz.
Material.— , xii.1888, Kotting (Weber, RMNH cat. no. 1, holotype); , x.1896, South Flores, 3000’ (Everett, RMNH cat. no. 2); 15 , 19 , 1968-1976, various localities in West Flores up to 1400 m (Verheijen/
Schmutz, RMNH).
Eggs (fig. 16b): 63 clutches, of which 22 with one egg and 41 with two eggs, collected in the months
January (1), March (5), April (8), May (24), June (11), July (4), August (1), September (3) and October(5).
Some measurements and weights:

RMNH no. 71342
RMNH no. 71210
RMNH no. 71211
RMNH no. 71213

16.9 × 12.0
17.2 × 12.0
17.7 × 12.0
18.1 × 12.2
16.6 × 12.6
17.6 × 12.2
19.2 × 12.9
19.2 × 13.3

0.05930.06500.06440.06350.0741
0.06960.08280.0835

Verheijen (1964) stated that the series of eggs he listed under this name, probably
included eggs from another species of Dicaeum. However, the eggs are too large to belong to D. sanguinolentum and presumably also too large for D. igniferum (the eggs of D.
igniferum are unknown, but it is a small species). Probably D. agile has eggs correspond-

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Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

ing in size with those of D. annae, but it is a rare species, not observed for almost a
century, so that it is unlikely that its nest has been found.
Notes.— Wing measurements of 17  54-61 mm (the smallest specimen is adult), 19
, 54-59 mm. Both Rensch (1931a) and Salomonsen (1960: 3-4) mentioned that the subspecies sumbavensis is very slightly differentiated. It was based on insignificant differences in tone, and on a difference in wing-length, sumbavensis being supposedly smaller.
The much larger series from Flores now available, encompasses in its range of variation
the recorded measurements of the few birds known from Sumbawa (3 , 55-57 mm, 2
, 54-56 mm, cf. Rensch, 1931a: 617). Although I have not examined specimens from
Sumbawa for a direct comparison, I agree with White & Bruce (1986: 407-408), that sumbavensis “is too poorly differentiated to merit recognition”.
In his revision of the Dicaeidae, Salomonsen (1960: 3) states: “The sexes in D. annae
are alike, just as in Prionochilus olivaceus and P. maculatus”, and this was one of his reasons for considering D. annae a primitive species, forming a link between Prionochilus
and the more typical species of Dicaeum. Salomonsen’s statement is surprising, for Hartert (1897b: 518), over sixty years before, had correctly described the difference between
the sexes: only males have the yellow uropygial patch; in females the rump is concolorous with the back. This was also correctly described by Mayr & Amadon (1947: 17). Interestingly, the difference is already found in juveniles. This conflicts with Hartert’s(1897b: 518, 1898c: 456) claim that yellow on the rump is wanting in young of both
sexes. Hartert’s erroneous claim was at least partly due to confusion with a different
species (the juvenile from Alor, where D. annae is not known to occur, which was subsequently re-identified as D. agile; cf. Mayr & Amadon, 1947: 17).
Measurements:
17 
19 

wing
54-61
(58.7)
54-59(56.3)

tail31-34½(32.1)
26-32(29.3)

tarsus13-15(14.0)
13-15(14.2)

entire culmen
11.0-12.2(11.8)
10.5-13(11.9)

exposed culmen
8.2-10
(9.0)
8-10
(9.0)

Dicaeum agile tinctum (Mayr)
Piprisoma obsoletum tinctum Mayr, 1944, Bull. Amer. Mus. Nat. Hist. 83: 167 – Waingapu, Sumba.
Collectors.— Semmelink, Allen (2), Everett (1).
Material.— (?), 1862, Larantoeka (Semmelink, RMNH); 2 (?), 1862, Flores (Allen, BM nos. 73.5.12.1031
and 95.5.1.2157).

Notes.— This species must be rare or very local on Flores, for only four specimens
seem to be known: the RMNH one (which was mentioned by Sharpe, 1885: 75), two
unsexed specimens without exact locality, collected by Allen, and one collected by Everett. As Larantoeka was in the 19th century the only place on Flores with a garrison, it
is quite likely that Allen also made it his headquarters, in 1862, so that three specimens
may have been collected at the same place and at about the same time. The Everett
specimen, however, would be from near Nanga Ramau, almost on the opposite end of
the island from Larantoeka, and suggests a wide distribution.
On Sumba, on the other hand, evidence is that D. agile is rather common. Hartert’sMees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

179(1896: 567) record from Sumbawa was, within two years of its publication, withdrawn
by Hartert (1898b: 117) himself, as having been based on misidentified, poorly preserved skins of Dicaeum annae. There are no reliable records from Sumbawa. It does,
however, occur on Alor, where a  juv. was collected in April 1897 (leg. Everett, nat.
coll., AMNH no. 698410); the specimen was misidentified and published as D. annae by
Hartert (1898c), see also the notes under D. annae.
Dicaeum agile has been revised by Salomonsen (1960) and recently again by Sheldon(1985). Salomonsen called it: “undoubtedly a near ally of D. annae”, but partly on the
mistaken assumption that D. annae shows no sexual dimorphism in plumage. I have nomajor contribution to make to the systematics of this species, but having examinedmaterial of two little-known forms that were not studied by the said authors, I can put
some of their speculations on a firmer footing.
It concerns the subspecies D. a. atjehense and D. a. finschi. About the latter, Salomonsen (1960: 12) has this to say: “This form, which must be very rare and local, is known
from the type locality. Apart from the type specimen, which I have not seen, only one
specimen is known; it was collected at Wynkoops Bay in 1920 by Kloss and is now in
the British Museum, where I have examined it”. The information that this subspecies isknown from only two specimens, is repeated by Sheldon (1985: 606).
Actually, there is a series of 13 specimens in the RMNH collection (all leg. Bartels).
In view of the importance of this series, I give here a full enumeration:  , ,  juv.,
20.vi. 1918, Tjisoedjen (RMNH nos. 63905, 63904, 63906); 3  ,  juv., 5.vi.1922, Goenoeng Massigit (RMNH nos. 63907-63910); ,  juv., 6.vi.1922, Goenoeng Massigit(RMNH nos. 63911, 63912);  ,  juv.,  juv., 13.vi.1922, Goenoeng Massigit (RMNH
nos. 63914, 63913, 63915);  med., 27.xii.1922, Radjamandala (RMNH no. 63916). Although Bartels (1923) has published his 1918 encounter with the species, he does notstate specifically that he had collected material, and that may be the reason why thisrecord was ignored by the authors just mentioned (but not by Kuroda, 1933: 119).
It will be noted that the type specimen of Dicaeum finschi (  ad., 31.xii.1913, Goenoeng Beser near Wijnkoopsbaai, coll. Bartels no. 9482) fails in the above enumeration.
The type specimen was forwarded to Finsch in Germany, and the description was published in July/August 1914 (cf. Bartels, 1914), at the outbreak of the First World War.
From a note I found in the archive belonging with the Bartels collection, I learned thatthe type was never returned. War conditions may have prevented its return to Bartels
in Java, or it may have become lost in the mail. Finsch died in 1917, during the war. The
type specimen must be assumed lost.
The above series requires the following comment. In the first place, there is a clear
difference between adult birds and juveniles. Juvenile birds have the striations of thebreast much less distinctive than the adults (indeed, they are almost absent). In bothstages, the sexes are identical in plumage. Neither Salomonsen nor Sheldon mention a
distinctive juvenile plumage. The measurements taken by me indicate clearly that adultfemales are a little smaller than adult males, and therefore I regret that Sheldon (1985:
table 1) has failed to separate the measurements of the sexes.
D. a. finschi was described as having no white tips to the lateral rectrices, and this has
been accepted as a subspecific character. It even led to some speculation about its affinities. Actually, most specimens from Java show some white in the tail: the outer rectrix,
and sometimes also the second rectrix, have a little whitish, or at least pale coloration at

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Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

the tip, best developed on the inside of the tip (fig. 11). In fact, the white is not less developed than in our two specimens of D. agile from Borneo (identified by Sheldon).
Sheldon speculated whether: “a race such as finschi, which lacks white tail spots, merits
specific status”. It does not and is no more than a slightly differentiated subspecies.
Sometimes one finds this form cited as Dicaeum Finschii, but the original binomen is
Dicaeum finschi.
Finally, it is worth observing that the 13 specimens have been collected at three localities only: 3 near Tjisoedjen, on 29.vi.1918; 9 on the Goenoeng Massigit, on 5/13.
vi.1922; and one at Radjamandala. This points to the species being not only rare anddifficult to locate, but at times being, locally and temporarily, common.
Neither Salomonsen nor Sheldon have examined D. a. atjehense, which they statecorrectly that is known from only one single specimen (, 21.ii.1937, Gajoe Loeös, Pending, ca. 500 m, leg. Hoogerwerf, RMNH no. 14069). The year 1939, given for this bird
by Chasen & Hoogerwerf (1941: 107, s.n. Piprisoma modestum sumatranum), followed by
Van Marle & Voous (1988: 202) is a misprint. This bird has more white in the tail thanany specimen of finschi (and also than the two specimens from Borneo) examined by
me. The bill is rather small.
It may be of historical interest to present full data on the RMNH specimen of D. a.
modestum, of which the measurements are given below: , 27.xi.1879, Maplay choung,Thoungyeen Valley, leg. C. T. Bingham. The measurements confirm that this is the specimen described by Bingham (1880: 171, s. n. Prionochilus modestus).
A word remains to be said about the nomenclature. As Sheldon could not find any
difference between specimens from Malaya (remotum) and specimens from Borneo, andas I am unable to find any difference between specimens from Java and two of thesesame specimens from Borneo, it would follow that remotum is a synonym of finschi, aconclusion reached seventy years ago by the authors of remotum themselves (Robinson& Kloss, 1921-1924: 393). The only reason why I still hesitate is that in this concept Sumatra should obviously also be included into the range of finschi, but the unique specimen from Sumatra is characterized by having large white tail-tips, and a rather small
bill. The most likely explanation for this is, that the type of atjehense just happens tobe an extreme variant in a population otherwise like finschi, but until more material

Fig. 11. Dicaeum agile, outer and second left rectrix, viewed from below, to show the extent of white attheir tips. From left to right: , Sumatra (RMNH no. 14069, type of D. a. atjehense); , Kalimantan(RMNH cat. No. 4); , Java (RMNH no. 63910, bird with the greatest amount of white in the Java-series).
Although generally referred to as “white”, this should rather be called “pale”, for it is neither brilliant
nor contrasting.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

181

becomes available, it is impossible to be certain, and I consider that for the moment D.
a. atjehense will have to be recognized.
My field-experience is limited to the observation of a few individuals of D. a. tinctum on Sumba. I did not notice the tail-wagging and tail-fanning described by Sheldon
as characteristic of the species. Also, the vague whitish markings on the tail seem to me
much too inconspicuous to have an important signal function, as he assumes.
Measurements:
wing
Tenasserim (D. a. modestum)
59
Sumatra (D. a. atjehense)
63½
Borneo6263Java (D. a. finschi) 6390563 6390763 6390862½
 6390962
6391461
 med 63916
59 6390458
 6391159
 juv. 63913
58 juv. 63906
56
 juv. 63910
55½
 juv. 63912
58 juv. 63915
57
Flores (D. a. tinctum)(?) RMNH
58(?) BM 73
59(?) BM 95
57
Timor (D. a. obsoletum) RMNH59
 RMNH59
 RMNH56

tailtarsusentire culmen

exposed culmen

26½

13

9.7

7

33½

12.9

9

6.2

29
29

13
13

10.1
10

7.7
7.7

28½
31
30
29
29½
28
28
29
27
26
25
26
25

13
13.6
12.5
13
13
13
12.9
12.6
12.5
12.4
12
12
13

11.3
11
11
10.8
11
10.8
10
11.2
9.8
9
9.3
10
9.8

8.4
8
8
7.3
8
7.9
7
7.7
7.2
7
7
7.4
7.8

27½
27½
25

15
12.8
13

10
10.2
9.4

7
7.2
6.5

27
26½
25

13.4
13.5
13

9.9
9
9.8

7.1
7
8

Dicaeum igniferum Wallace
Dicaeum igniferum Wallace, 1864, Proc. Zool. Soc. Lond. (1863): 494 – Flores.
Dicaeum igniferum cretum Rensch, 1929, Journ. f. Orn., Ergänzungsb. 2: 201 – Alor.
Collectors.— Allen, Colfs, Weber, Everett, v.d. Sande (2), Rensch (2), de Jong (10), Verheijen/Schmutz.
Material.— , iii.1880, West Flores (Colfs, RMNH cat. no. 1); (?), skeleton, xii.1888, Sikka (Weber, RMNH
cat. a); , 22.xii.1969, Todo (Verheijen, RMNH no. 65519); , 12.vi.1971, Nantal (Verheijen, RMNH no.
85075); , 12.vi.1973, Golo Karot, Borong, 50 m (Verheijen, RMNH no. 85074); , 13.vi.1973, Golo Karot,
Borong (Verheijen, RMNH no. 85073 ); , 1.vi.1978, ? Méléng (Verheijen, RMNH no. 81382).

Notes.— The measurements of this material agree well with those of specimensfrom Flores, provided by Salomonsen (1961: 4). The subspecies cretum (from Alor and182

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Pantar) was based on the single character of a slightly larger size. Salomonsen measured specimens from Alor (4) and Pantar (2), including the type material of cretum, andcould confirm the larger size, but he added: “The difference is trifling indeed, and I am
not very keen on accepting such weakly established forms. However, for the moment
I recognize cretum, pending more material from Alor and Pantar”. Considering how
cautious Salomonsen was in treating weakly-defined subspecies, the above comment
practically amounts to a rejection of cretum. White & Bruce (1986: 409) had no such
qualms, and stated without presenting further evidence: “Subspecies are not established and Rensch’s cretum is a synonym”. I follow White & Bruce in the rejection ofcretum for two reasons: the first is that the measurements indicate that a larger series
from Pantar and Alor would almost certainly show considerable overlap with measurements of nominate igniferum, the second that on the ornithologically poorly known
connecting islands of Adonara and Lomblen, D. igniferum is likely to occur and might
have intermediate measurements: of course, this reasoning would be unacceptable if
there was a convincing difference between igniferum and cretum.
Measurements:
4
2

wing
49-52
(50.5)
48, 49½

tail
24-26(24.7)
23, 24tarsus13-14(13.3)
13, 13.7

entire culmen
10-12(10.7)
11, 12

exposed culmen
7+-8
(7.8)
9, 10.3

Dicaeum sanguinolentum rhodopygiale RenschDicaeum sanguinolentum rhodopygiale Rensch, 1928, Orn. Mber. 36: 80 – Rana Mesé, West-Flores (1200 m).
Collectors.— Everett (1), Rensch (2).
Material.— None.

Notes.— Apparently only three specimens of this subspecies are known: one  collected by Everett (cf. Hartert, 1897b: 518), and two ( and ) obtained by Rensch.
Rensch’s (1931a: 615) remark: “Alle drei Exemplare erbeutete ich...” must be a slip, asfrom his own enumeration (Rensch, 1928c and 1931a: 614) it is clear that he collected
only two, and that the third bird was from Everett’s collection. These specimens arefrom a level of ca. 1200 m, just about the level of the greatest activity of the fathers Verheijen and Schmutz, and this suggests that the species is really uncommon and that thepaucity of records is not merely due to the vagaries of collecting. The few known specimens of D. sanguinolentum hanieli from Timor are also from ca. 1200 m. The related D.
wilhelminae, of Sumba, however, is a lowland bird (Hellmayr’s, 1914: 57: “Gebirge vonSumba” is in error), and is common. The highest point of Sumba is 1225 m.
Salomonsen’s (1961) historical-zoogeographical hypothesis in which Java, etc.,
would have been colonized from Sumba, seems to me extremely dubious. The apparent
rarity of the forms inhabiting Timor (hanieli) and Flores (rhodopygiale) already constitutes an objection: I find it difficult to envisage the Flores form “invading” (over Lombok and Sumbawa, where it is not known to occur) Bali and Java, as required by Salomonsen. A consequence is that I do not regard the thick-billed lowland bird Dicaeum
wilhelminae as ancestral to the slender-billed mountain bird D. sanguinolentum, and evenMees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

183

less as conspecific with it. I would return D. wilhelminae to the status of a separate,monotypic, species, leaving its exact relations to D. maugei, D. igniferum, D. sanguinolentum, etc., once more open to discussion.
Perhaps my doubts about the picture of evolution and speciation presented by
Salomonsen, arise from the feeling that in attemps to reconstruct it, the limits between
justified deduction, plausible hypothesis, speculation, and nonsense, are vague.
Anthreptes malacensis convergens Rensch
Anthreptes malaccensis convergens Rensch, 1929, Journ. f. Orn. Ergänzungsb. II: 200 – Sita, Flores.
Collectors.— Semmelink, Allen, Colfs, Weber, ten Kate, Everett (2), v.d. Sande (1), Rensch (4), de Jong (3),Verheijen (2, MCZ), Verheijen/Schmutz.
Material.—  subad., , 1862, Larantoeka (Semmelink, RMNH without numbers); , iv.1880, West
Flores (Colfs, RMNH without no.); , iv.1891, Sika (Ten Kate, RMNH without no.); , 18.ix.1969, Rekas,
350 m (Verheijen, RMNH no. 81379);  juv.,  juv., 20.xii.1969, Todo (Verheijen, RMNH nos. 65517,
65516);  im., 21.xii.1969, Todo (Verheijen, RMNH no. 65494); , 24.xii.1969, Todo (Verheijen, RMNH
no. 65495); , 21.ix.1970, Kisol (Verheijen, RMNH no. 65563); , 15.iii.1971, Kisol, 150 m (Verheijen,
RMNH no. 85369); , , 20.vi.1971, Waé-Ntjuang (Verheijen, RMNH nos. 85370, 85377); , 12.vi.1971,
Nantal, Rahong (Verheijen, RMNH no. 85371).
Eggs (fig. 16c): 15 clutches of c/1 (7 ×) and c/2 (8 ×), from the months April (4), May (3), June (2), July(2), and August (4), (RMNH nos. 71327-71341).
The eggs are white to pinkish white, well provided with light to medium grey secundary dots, andsparser black primary markings of varying shapes.
Some measurements and weights: RMNH no. 71328

17.2 × 13.0
0.086217.5 × 13.2
0.0864RMNH no. 713219
19.0 × 13.1
0.095219.7 × 13.6
0.1009RMNH no. 71339
17.8 × 12.6
0.082218.9 × 13.1
0.0923The eggs agree in measurements with those of nominate A. m. malacensis from Java, as given by Hellebrekers & Hoogerwerf (1967: 141), but average slightly heavier.

Notes.— It is well known that the subspecies A. m. convergens is closer to A. m. celebensis from Sulawesi (Celebes), than to the nominate race, which occurs east to Java,
Bali, and Borneo. Stresemann (1940a; 59) put as a tentative suggestion that the species
might have colonized Celebes from the Philippines, or from the South (Sumbawa,
Flores), or from both directions. This was taken up and modified by Mayr (1944a: 164)in the following words: “The island of Celebes seems to have been colonized both fromthe Lesser Sunda Islands (convergens) and from the Philippines (chlorigaster), both ofwhich are very similar to celebensis”, thus changing Stresemann’s casual remark to nearcertainty. As Celebes is inhabited by a homogeneous subspecies (it is generally agreed
that citrinus is a synonym of celebensis), such a double invasion seems to me rather unlikely. That Stresemann himself was not very serious about it, is apparent from the factthat in the special section “Doppel-Einwanderung auf Celebes”, he makes no mentionof it (cf. Stresemann, 1939: 346).

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Measurements:7
2  juv.
5

wing
65-69
(66.3)
64, 6660-64½(61.7)

tail37-41½(39.0)
38, 4033½-39(36.7)

tarsus
16.6-17.3(16.9)
16.8, 17.5
16.3-17.2(16.9)

entire culmen
20.6-21.2(21.0)
20, 20.6
19-21(20.1)

exposed culmen
17-18.9(17.9)
16.7, 17.3
15.8-17.1(16.6)

Nectarinia jugularis ornata (Lesson)
Cinnyris ornata Lesson, 1827, Dict. Sci. Nat. (éd. Levrault) 50: 15 – no locality, but based on Nectarinia
eximia Temminck, pl. 138 fig. 1 = Java.
Collectors.— Semmelink, Allen, ten Kate, Everett, v.d. Sande (1), Rensch (7), de Jong (1), Verheijen (8,MCZ), Verheijen/Schmutz.
Material.— , , 1862, Larantoeka (Semmelink, RMNH without numbers); , 1/3.v.1891, Groot
Bastaard (ten Kate, RMNH without number); , 28.x.1969, Nunang, 650 m (Schmutz, RMNH no. 81439);
, , 16.ix.1970, Maro-Kama (Verheijen, RMNH nos. 65497, 65496); 2 , 17.ix.1970, Maro-Kama (Verheijen, RMNH nos. 65498, 65499); , 2 , 22.ix.1970, Maro-Kama (Verheijen, RMNH nos. 65500, 65502,
65503); 3 , 23.ix.1970, Maro-Kama (Verheijen, RMNH nos. 65504, 65505, 65506); , 16.iii.1971, Kisol,
175 m (Verheijen, RMNH no. 85368); , 9.iii.1974, Maro-Kama (Verheijen, RMNH no. 66199); , 4.
ii.1976, Ero (Verheijen, RMNH no. 85384). Iris brown, bill and legs black.
Eggs (figs 16d, 16dd): 46 clutches of c/1 (22 ×) and c/2 (24 ×), collected in the months March (4), April(19), May (11), June (7), July (3), and two without date. There are 17 clutches from Palué, the others arefrom Dampék, Mataloko, Soa (RMNH nos. 71247-71273, 71275-71293).
The eggs are white with large, somewhat cloudy, medium olive-brown markings.
Some measurements and weights: RMNH no. 71289
RMNH no. 71291
Measurements:
10 
4

wing
51½-55(53.6)
49-52½(51.4)

tail30½-36(33.4)
28-30(29.1)

tarsus14-15(14.8)
13.7-14.6(14.1)

14.5 × 10.8
16.2 × 10.9
15.7 × 10.6
17.0 × 10.9
entire culmen
18-21.2(20.0)
18.3-19.9(19.1)

0.04220.0485...
0.0509exposed culmen
15.3-18.2(17.0)
16.4-17.4(16.9)

Nectarinia solaris solaris Temminck
Nectarinia solaris Temminck, 1825, Recueil d’Ois. 4 (livr. 58): pl. 347 fig. 3 – Amboine, l’une des Moluques(errore) = Timor.
Cinnyris solaris degener Hartert, 1904, Novit. Zool. 11: 214 – Endeh, S. Flores.
Collectors.— Semmelink, Allen (several), Martens, Colfs, Weber (4), Everett, Rensch (5), de Jong (4),Verheijen (4, MCZ), Verheijen/Schmutz.
Material.— , 1862, Larantoeka (Semmelink, RMNH); , 1880, Flores (Colfs, RMNH), (?), skeleton,xii.1888, Sika (Weber, RMNH cat. a); 22 ,  im., 8  from Kisol, 175 m, Lingko Lumu, Ruteng, GoloKarot, 50 m, Tinung Ndéhes, 700 m, Ruteng, Langkas, Waé-Ntjuang, Raé and Todo, from the coast to ca.
1100 m (Verheijen/Schmutz).
Eggs (figs 16e, 16ee): 33 clutches of c/1 (15 ×) and c/2 (18 ×), collected in the months March (1), April (5),May (10), June (7), July (6), and August (4), from Rekas, Bénténg Djawa, Tado, Pau, Potjong, Wesang,Mano, Léwé, Lamé, Waé Tua (RMNH nos. 71294-71326).

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185

The eggs are close to those of N. jugularis ornata, but their markings are stronger, darker, sharper defined, and more purplish-brown in colour.
Some measurements and weights: RMNH no. 71319
RMNH no. 71320
RMNH no. 71323
RMNH no. 71326

15.8 × 10.8
16.0 × 11.0
15.7 × 11.1
16.1 × 10.9
15.3 × 11.1
14.9 × 10.9
15.4 × 10.9

0.05230.047
0.05230.05
0.049 (damaged)
0.046
0.0478Notes.— I agree with White & Bruce (1986: 406) that the subspecies degener (Sumbawa, Flores, Lomblen and Alor) is too poorly differentiated from nominate solaris(Timor and Samao) for recognition; indeed, I am unable to see any of the characters bywhich it was claimed to be distinguished. I also agree with Bruce (l.c.) that N. s. exquisita (Wetar) merits recognition, on the basis of having a slightly heavier bill, and alittle yellower upper parts, than birds from Timor and Flores. As a matter of historicalinterest: this species was discovered by Reinwardt, on his visit to Koepang and its surroundings, Timor, from 3/18.vi.1821, but because of poor labelling was thought to havecome from Ambon when Temminck described it in 1825. Thus it was left to S. Müller,
during his stay on Timor in 1828/1829, to establish its true provenance.
Knowledge of interactions between the two closely related species N. solaris andN. jugularis, and of a possible ecological segregation, are of much theoretical interest.
Father Schmutz (MS) has the following notes on the subject. In the immediate neighbourhood of the coast, only N. jugularis is found (note that from the islands of Komodo, Rintja and Palué only this species is known), but already a slight distance inland, at levels of 10-200 m, N. solaris occurs also, for example in Nisar, Waewako (180m) and Look. At Waewako and Rekas (350 m), both species occur, and that is also thecase in Nunang (650 m), where N. jugularis seems to be still the commoner species. On
the other hand, at levels of ca. 1000 m, only N. solaris remains. When travelling from
Orong to Tjantjar, over a pass of 1200 m, Schmutz never saw N. jugularis. N. solaris
was the only species known to Father Verheijen from the surroundings of Ruteng(1000-1100 m) and the mountains of Manggarai. Therefore he was surprised and intrigued to find that on the cultivated high plain and in the coffee gardens near Mataloko (ca. 1000 m, the same altitude as Ruteng), N. jugularis was common, and N. solaris was absent. Evidently altitude is not the only factor that decides which species
occurs in a given locality.
Measurements:
24 
8

wing
51-56
(52.8)
48-51(49.8)

tail30-35(32.1)
27-31(28.8)

tarsus
13.9-15.2(14.5)
14-14.5(14.2)

entire culmen
18.2-20.6(19.4)
18-19.9(18.7)

exposed culmen
15.2-17.7(16.5)
15.1-17.1(16.1)

Zosterops palpebrosa unica Hartert
Zosterops unica Hartert, 1897, Novit. Zool. 4: 520 – Nanga Ramau, South Flores.
Collectors.— Everett (1), Verheijen (1, MCZ), Verheijen.

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Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Material.— 2 , (?), 27.x.1970, Ruteng (Verheijen, RMNH nos. 66187, 66188, 85388); , , 27.vi.1971,
Wai-Ntjuang, Langkas, Rahong (Verheijen, RMNH nos. 66190, 66189); , 29.vi.1971, Wai-Ntjuang, 900
m (Verheijen, RMNH no. 66191); , 12.vi.1971, Rahong, Manggarai (Verheijen, RMNH no. 66192); ,
21.vii.1976, Poco Nernancang (Verheijen, RMNH no. 81449). Unfeathered parts: Iris light brown; bill
black, the base of the mandible plumbeous; legs slate.
Eggs (fig. 16f): 47 clutches of c/1 (17 ×), c/2 (20 ×) and c/3 (10 ×), collected in the months March (4),April (9), May (16), June (12), July (3), August (2), and October (1), from Mano, (Puntu), Poéng (Méngé),Wésang, Léwé, Tjarang, Tado, B. Liang, Mbélar, Potjong, Léing, Lété, Soa and Mataloko (RMNH nos.
71427-71473). The eggs vary from very pale blue to white.
Some measurements and weights: RMNH no. 71428

RMNH no. 71455
RMNH no. 71466
RMNH no. 71467

14.6 × 10.9
15.2 × 10.9
15.7 × 11.2
15.4 × 11.9
15.8 × 12.0
15.0 × 11.9
15.1 × 11.8
17.1 × 11.6

0.05340.05600.05850.06000.0596
0.06250.05790.0620Notes.— These specimens were compared with an adequate series of Z. p. melanurafrom Java, and were found to differ only in having a slightly yellower rump and upper
tail-coverts, agreeing with the diagnosis given by Mees (1957: 87). Previously this species was known on Flores from the type-specimen of Z. p. unica only. From Sumbawa,
seven specimens are known, collected by Rensch (1931a: 623).
Unfortunately, I have to discuss the gender of Zosterops again, a question I thought
had been settled some 35 years ago.
The history is as follows: The authors of Zosterops did not give its gender, neither
can it be deduced from the species they placed in it (Vigors & Horsfield, 1827: 234).
The first author to associate Zosterops with a gender may have been Swainson (1831:
205, 222: Zosterops Javanica), feminine. About the middle of the 19th century, several
authors treated the genus as masculine (Blyth, Gould, Guerin-Méneville, Temminck &Schlegel), although others treated it right from the beginning as feminine (Bonaparte,
Swainson).
The leading ornithologists of the second half of the 19th and first years of last century treated the genus as feminine (Salvadori, Sharpe, Wallace, E.L. Layard, A.B. Meyer,
Hartlaub, Tristram), with only one important exception (Reichenow, who was inconsistent but with a strong bias to masculinity). Last century saw the advocates of feminine gender further strengthened (by Neumann, Stresemann, Salomonsen, Delacour,etc.), and there is no doubt that this would have become universal (in spite of a short
hiccup of masculinity in Japan), had not W.L. Sclater (1930: 673), in an influential workthat was to destine the nomenclature of African birds for many years, chosen for the
masculine gender.
The result was that in the Indo-Australian Region, Zosterops continued as feminine,and so was it treated by non-British authors on African birds, but most British andSouth African authors followed, uncritically one may assume, Sclater, and treated it asmasculine, a situation that was obviously undesirable.
It appeared that the controversy was finally and definitely decided in favour of thefeminine gender through a Recommendation in the Copenhagen Decisions (Hemming,

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

187

1953: 49) and the acceptance also by British authors (Snow & Moreau, 1955).
In the first edition of the Code, this Recommendation was changed to an Article, butwith some explanatory text (ICZN, 1961: 30-31). The matter is made more complicated,in that a distinction is made between words ending in -ops, derived from Greek οψ,which are feminine, whereas words ending in -ops derived from Greek ωψ, “of which
the usual classical gender is masculine, are to be treated as masculine unless the author
indicated otherwise or unless, failing such indication, zoologists have generally treatedthem as feminine”. This gives a lot of leeway, but if Vigors & Horsfield did not provide
the gender of Zosterops, they did give its derivation: “Ζωστηρ cingulum, and οψ oculus”, hence, definitely feminine. Note, however, that these authors expressly state theletters οψ to mean "eye", which seems to put in doubt the distinction between οψ andωψ made in the Code (in agreement with Amaral, 1964).
Although of course this did not immediately penetrate everywhere (especially inSouth Africa it was a slow process), in important works, like White (1963) and Moreau(1967), the gender of Zosterops was treated as feminine.
This was the position when the ICZN exercised their prerogative of changing theirmind: the original decision was reversed, and Zosterops was proclaimed masculine. The
new ruling was included in the third edition of the Code (ICZN, 1985: art. 30 (a) (ii)). The
renewed discussion on which this change of mind was based, began with a complaintthat the Article as it stood in 1961, was too complicated (Follett & Dempster, 1963). Therefore they suggested a feminine gender, but on the same page, Holthuis expressed a preference for the masculine gender. In the following year, several zoologists/linguists (amost valuable combination), discussed the problem and failed to agree (Amaral, 1964;
Griffin, 1964; Sabrosky, 1964), upon which the matter was decided by vote.
Since then, I have seen a few very definite statements in the literature, such as: “Zosterops is of masculine gender” (Dickinson et al., 1991: 402). Such a simple statement isnonsense, and should have read: “Zosterops has been decreed to be of masculine gender”. Having treated the large genus Zosterops as feminine for close on fifty years, I am
reluctant now to change its gender, a change diametrically opposed to the stated objectof the Code (“to promote stability”).
Addendum.— To illustrate the arbitrary, not to say haphazard way this decision
was taken, I provide the following anecdote. When the question came up in 1963, thetechnical assistant of a distinguished member of the ICZN, who is a carcinologist, had
just finished labelling a series of European lobsters with the name Nephrops norvegicus
in the beautiful copperplate writing for which he was known. Not wanting to havethese labels re-written, a time consuming work, and moreover finding that Nephrops
norvegica “did not sound right”, this member, not a classical linguist, strongly pushed
for the masculine gender. For the sake of one lobster, Zosterops, the largest bird-genus
known, as well as several related genera (Speirops, Lophozosterops) had to change gender.
Admittedly, the story goes that E. Mayr was consulted about Zosterops and that he did
not object to the change, but this seems strange as several years later, in the authoritative “Checklist of birds of the world” (1967), Mayr used the feminine gender throughout, as he had done in all his earlier publications. It had not occurred to the above-mentioned carcinologist, who was my colleague, to mention the problem to me.
Other bird genera ending in –ops. Treatment of Prionops (8 species) has been inconsistent, but several leading authors considered it feminine. Xenops (4 species): masculine.

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Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Half a dozen small genera, with one or two species, carry little weight and in some cases
the gender does not affect the specific name (Scythrops novaehollandiae in this paper).
Measurements:
4
3
(?)

wing
51-54½(52.1)
50-51(50.3)
50

tail
31-34
(32.5)
30-32(30.7)
29

tarsus14-15(14.4)
15-15.8(15.3)
14.5

entire culmen
12-13(12.44)12(12.0)
11

exposed culmen
83/4-10
(9.2)
9
(9.0)
8.6

Zosterops montana montana BonaparteZ[osterops] montana Bonaparte, 1850, Consp. Gen. Av. 1: 398 – Sumatra = Mt. Merapi, Padang Highlands(cf. Mees, 1957: 176, 184-185).
Zosterops palpebrosa florensis Rensch, 1928, Orn. Mber. 36: 9 – Geli Moetoe (1500 m), Flores.
Collectors.— Everett (2), Rensch (7), Verheijen (10, MCZ), Verheijen/Schmutz.
Material.— 38 specimens, all from near Langkas (Rahong) and Ruteng, from elevations of 900-1500 m(Verheijen, RMNH nos. 59811, 59812, 59814, 59817, 59818, 59820-59826, 59829, 59833-59835, 59840, 6617566180, 66182-66185, 81438, 82185).
Eggs: c/2, 4.vi.1955, Wangkung (RMNH no. 71357); c/3, 4.ix.1955, Tjolol (RMNH no. 71358); c/2,
12.viii.1958, Robo (RMNH no. 71359); c/2, 1.viii.1959, Wangkung (RMNH no. 71360). The eggs are unmarked, white with just a suggestion of pale blue, almost without gloss.
Measurements and weights:

RMNH no. 71357
RMNH no. 71358

RMNH no. 71359
RMNH no. 71360

16.5 × 12.9
16.7 × 12.7
16.0 × 12.3
16.4 × 12.7
16.7 × 12.7
17.3 × 11.1
18.0 × 12.1
16.0 × 12.1
16.2 × 11.9

0.07470.07480.06540.07080.07010.07710.07900.0631
0.0616

Notes.— For a review of the extensive synonymy of this species, see Mees (1957 and1969: 270-273). In the last-mentioned paper I speculated about the identity of birds from
Lake Lanao, Mindanao, recorded by de Schauensee & du Pont (1962) under the nameof Zosterops montana montana. These authors gave for specimens from Lake Lanao awing-length of 53-56 mm, and for specimens from 3250-5000 ft. on Mindanao a winglength of 54-58 mm, concluding that “it appears that highland birds are larger thanlowland specimens”. The surface of Lake Lanao is at 400 m a. s. l., which puzzled me,
as Z. montana ought not to occur so low. Only later did I discover that the name Lake
Lanao is now applied to the new university, high above the Lake, at 1800 m, and thatthat is the place where these specimens would have been taken. As 1800 m is 6000 ft., acomparison with specimens taken at 3250-5000 ft. elsewhere in Mindanao, should show
the former to be larger, not smaller than the latter, if there was an increase in size withan increase in altitude. As I mentioned before, the lowland specimens from Luzon listedby these authors under the name Z. m. montana, were undoubtedly Z. japonica meyeni(cf. Mees, 1969: 263).

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189

On the basis of the above discussion, I gladly accept du Pont’s (1971a) statementthat “there was no misidentification of species” of the specimens from Mindanao. He
does not comment on the specimens from Luzon, so that I suppose he has accepted
their re-identification. In the same paper, du Pont describes a new subspecies, Zosterops montana gilli, from the island of Marinduque, 1000-1500 ft. In the description ofthis subspecies, du Pont compares it only with Z. m. halconensis, from which he saysthat it differs by being more yellowish. There is no mention of Z. japonica meyeni, withwhich it was apparently not compared. Neither is the iris-colour recorded. Z. j. meyeni is a yellower bird than Z. montana. The occurrence of Z. montana on an island witha highest peak of only 652 m, and at levels of 1000-1500 ft. (300-450 m) seems unlikely.
Therefore I suggest that Z. montana gilli is a synonym of Z. japonica meyeni (cf. Mees,
1992).
Erritzoe (1995: 13, 14) lists two specimens from Laguna de Bai in the flat lowlandsof Luzon, as Z. montana. Although he refers to my work (Mees, 1957) for this identification, I cannot believe it correct. There is no mention of the eye-colour and identificationon the basis of a plumage description alone is tricky.
Measurements:
24 
14 

wing
54-59
(56.5)
54-57(55.6)

tail36-41(37.6)
34-38½(36.7)

tarsus
16-17.6(16.8)
16-17(16.3)

entire culmen
12.2-14(13.1)
12.3-14(13.1)

exposed culmen
9-11(10.2)
9-10(9.8)

Zosterops chloris intermedia Wallace
Zosterops intermedia Wallace, 1864, Proc. Zool. Soc. Lond. (1863): 493 – Macassar and Lombock, restricted
to Makassar by Walden (see notes).
Collectors.— Weber (2), Rensch (8), de Jong (1), Verheijen (9, MCZ), Verheijen/Schmutz.
Material.— (?), 29.xi.1888, island Rusa Radja = Palué (Weber, RMNH cat. no. 8).
Eggs: c/2, 27.vi.1956, Dampék (RMNH no. 71361), and 65 clutches of c/1 (11 ×), c/2 (21 ×), c/3 (30 ×)and c/4 (3 ×), all from Palué, collected between 14.iv. and 5.v.1960 (RMNH nos. 71362-71426). The eggs
vary from pale blue to white, and have little gloss.
Some measurements and weights: RMNH no. 71375

RMNH no. 71402

RMNH no. 71414

RMNH no. 71416
RMNH no. 71417

RMNH no. 71418

16.0 × 13.1
16.4 × 12.8
16.5 × 12.8
16.3 × 12.3
16.7 × 12.3
16.7 × 12.8
15.6 × 12.8
16.1 × 12.9
16.3 × 12.8
16.5 × 13.0
15.7 × 12.5
16.0 × 12.4
16.1 × 12.1
16.5 × 12.5
16.7 × 12.4
17.0 × 12.6

0.07890.07540.07520.07050.07570.07890.07650.07620.07360.08690.07330.07400.06900.06880.07090.0722

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Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Notes.— Although Rensch collected it near Badjawa at 1200 m, this is mainly an
inhabitant of the coastal regions, the mangroves and adjacent cultivated lowlands, andespecially of the small islands: note that Verheijen obtained only a single clutch of eggs
from mainland Flores (Dampék, on the north coast), but 65 clutches from the island of
Palué during a stay of less than a month. The nine skins he collected (MCZ) are also allfrom Palué.
Z. intermedia was originally described from “Macassar and Lombock”, but in thedescription, Wallace (1864: 493) explained the provenance of the name given in the following words: “Remark: Mr G.R. Gray attached the MS. name of intermedius to my
Macassar specimen”. This was repeated in slightly different words by Walden (1872:72): “The above specific title was attached to a Macassar example in the British Museum
by Mr G.R. Gray, and was adopted by Mr Wallace, who first described the species”.
Neither of these statements is a valid restriction, or a valid lectotype-selection, and thesame pertains to Sharpe’s (1884: 185) listing of the Makassar specimen as “Type of species” (cf. ICZN, 1985: Art. 72 (b) (vii)).
White (in White & Bruce, 1986: 412, 415), discussing my treatment of the very similar species Z. palpebrosa, Z. montana, Z. chloris and Z. citrinella, concluded: “there seems
no reason why they should not...be closely related”, and: “Double invasion and character displacement could account for this situation...It is difficult to see why these species
should not have a common history associated with successive invasions and reinvasions”. The way this is put certainly suggests that White thought that he was disagreeing with me, but the fact that I and all other authors have placed these species in thesame genus, implies a common origin, and nobody would deny their common historyof speciation, associated with expansions and contractions of range. White’s reference
to character-displacement is less obvious, because the essence of the problem with these
species is their morphological similarity, which is the opposite of character-displacement.
In the first half of last century, when authors were still trying to create very widelyranging polytypic species, the four above-mentioned species, to which should be added
Z. japonica, Z. everetti, Z. lutea, and several others, have all, at some time, been treated asconspecific. My own work has largely been devoted to showing that this went much too
far, and that these large conglomerates, based on geographical replacement and on superficial similarity, were artificial (cf. Mees, 1969: 312). In this, I was conscious of thedanger of going too far in the opposite direction, and indeed, I was reluctant to deviate
too much from the conclusions reached by previous students of the Zosteropidae. That
is why only in 1969 I separated Z. chloris from Z. citrinella.1112) I kept Z. meyeni as a subspecies of Z. japonica, mainly as I believed birds from Hainan (Z. j. hainana) to show intermediate characters. Parkes (1971: 54) has since argued that a specimen collected by
Whitehead, supposedly on Hainan, and very similar to meyeni, which was at the basis of
my opinion, was mislabelled, and actually was taken on Luzon. This removed the main
objection to granting to Z. meyeni specific status. Parkes’s speculation that Z. meyeni might
be a lowland representative of Z. montana remains just that; indeed, it has less substance

12)
Authors following the directive of treating Zosterops as masculine, have invariably changed the name Z. citrinella to Z. citrinellus, but apparently in error (cf. Barbagli & Violani, 1997).

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

191

than the link with Z. japonica which he had just condemned, as here there is not even theargument of geographical and ecological representation. As other authors have held Z.
montana to be conspecific with Z. palpebrosa, and Z. palpebrosa with Z. japonica, this would
bring us right back to square one, where the affinities of Z. meyeni are concerned.
White gives an impression of denying the developments of the past forty years, andof wanting to go back to the expanded species of half a century ago. His remarks, however, do not contribute to a serious discussion, but are little more than hollow rhetoric.
For the same reasons, I cannot agree with Ford’s (1983: 396) claim that Z. chloris is “undoubtedly” the closest relative of Z. lutea.
Zosterops wallacei Finsch
Z[osterops] wallacei Finsch, 1901, Tierreich 15: 23 – Kleine Sunda-Inseln Sumbawa, Flores, Sumba undLomblen; restricted to Bima, East Sumbawa by Mees (1961a: 59).
Collectors.— Semmelink, Allen, Everett, Rensch (3), de Jong (2), Verheijen (4, MCZ), Verheijen/Schmutz.
Material.— (?), 1862, Larantoeka (Semmelink, RMNH cat. no. 3); 11 , 5 , 3 (?) from Ling Ndela, 600
m; Todo, 700 m; Nunang, 650 m; Waé Rempo; Weleng (Ruteng); Golo-Karot (Borong).
Eggs (fig. 16g): 58 clutches of c/1 (20 ×), c/2 (35 ×) and c/3 (3 ×), taken in the months April (12), May(14), June (11), July (13), August (4), September (1), October (2), and one undated (RMNH nos. 52320,
71474-71529, 73518).
The eggs are pale blue, lightly and finely mottled with brown.
Some measurements and weights: RMNH no. 71500
RMNH no. 71501
RMNH no. 71526
RMNH no. 71528

17.0 × 13.2
17.2 × 13.3
17.8 × 12.3
18.6 × 12.7
16.5 × 12.0
17.4 × 11.7
17.0 × 12.7

0.08600.08550.07050.07080.06930.07340.0744

Notes.— The measurements confirm that birds from Flores average slightly smaller
than birds from Sumba (cf. Mees, 1961a: 61-62).
Measurements:
11 
5

wing
53-56
(55.0)
53-55(53.9)

tail38-42½(40.8)
37-42(39.7)

tarsus
15.5-17.7(16.7)
17-17.8(17.4)

entire culmen
13.3-14.6(14.0)
13-15(14.0)

exposed culmen
10-11(10.8)
10-10.8(10.4)

Lophozosterops superciliaris superciliaris (Hartert)
Zosterops superciliaris Hartert, 1897, Novit. Zool. 4: 172 – South Flores.
Collectors.— Everett (at least 10, cf. Mees, 1969: 197), Rensch (21), Verheijen (4, MCZ), Verheijen/
Schmutz.
Material.— , x.1896, South Flores (Everett, RMNH cat. no. 1, syntype); 42 specimens from various localities in West-Flores, from 1050-2200 m (Verheijen/Schmutz).
No eggs.

192

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Notes.— Verheijen (1964: 196) considered it possible that amongst his large series of
eggs of Lophozosterops dohertyi, there might be a few of the present species, “since eggs
and nests of both species resemble each other strongly, and I gathered them in places
where they occur together, or at least adjacent to each other”. This may be correct, but I
doubt it for the following reason: the eggs of L. dohertyi, with their brown spots, areconspicuously different from those of all other Zosteropidae of which the eggs are
known. Although L. superciliaris and L. dohertyi are congeneric, I believe that L. superciliaris is more closely related to L. javanica than to L. dohertyi, and therefore I would
expect its eggs to resemble the plain, pale greenish blue eggs of L. javanica. Verheijen’s
remark, quoted above, about the strong resemblance, cannot be taken literally, as by his
own admission he did not know nest and eggs of L. superciliaris with certainty. A simple
and obvious explanation is that, although the species ranges down to ca. 1000 m, it does
not, or only in small numbers, nest below ca. 1300 m, the upper limit of Father Verheijen’s nest-searching activities
Measurements:
19 
21 

wing
67-71
(68.8)
64-69(66.9)

tail
46-54(48.2)
45-51½(47.6)

tarsus19.8-23(20.8)
19.4-21(20.2)

entire culmen
15.2-17(16.1)
14.4-16.8(15.8)

exposed culmen
12-14(13.0)
11.6-13.2(12.6)

Lophozosterops dohertyi subcristata Hartert
Lophozosterops (ad potius Zosterops) subcristatus Hartert, 1897, Novit. Zool. 4: 171 – hills of South Flores,
3000-3500 feet.
Collectors.— Everett (at least 8, cf. Mees, 1969: 203), Rensch (4), Verheijen (7, MCZ), Verheijen/Schmutz.
Material.— 36 specimens from Ruteng, Todo, Poco Nernancang, L. Ngkiong, Nantal, Waé-Ncuang,
Ntjuang, Waé Laci, Lingko Lareng Pongkor, Golo Mbélar, Rekas, at elevations from 350-1050 m.
Eggs (fig. 16h): 114 clutches of c/1 (34 ×), c/2 (79 ×) and c/3 (1 ×) eggs, collected in the months February(2), March (5), April (16), May (21), June (32), July (14), August (9), September (8), October (5), and 2 notdated (RMNH nos 52319, 71530-71620, 71622-71640, 73519-73521). The eggs are light blue, sparsely tomoderately dotted with brown.
Some measurements and weights: RMNH no. 71594
RMNH no. 71595
RMNH no. 71596
RMNH no. 71608
RMNH no. 71622
RMNH no. 71624

18.2 × 13.9
18.4 × 13.7
17.4 × 13.7
17.8 × 13.3
17.2 × 13.3
17.6 × 13.7
17.4 × 13.4
18.3 × 13.8
16.9 × 13.5
17.0 × 13.4
19.3 × 13.7
19.6 × 13.8

0.09500.08340.09000.09940.08920.08730.0972
0.0952
0.09330.0924
0.09620.1052

Notes.— As was to be expected in this larger material, the present series shows agreater vertical range than that assumed for the species by Rensch (1931a: 625).

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

193

Measurements:
17 

wing
tail
tarsusentire culmen exposed culmen
60½-6443½-4918-19.6
13-15.9
11-12.7(62.3)(45.6)(18.7)(14.8)(11.8)
18 
58½-62½ 41-47
17.5-19.2
13.2-16
10.8-12(61.1)(44.4)(18.2)(14.3)(11.5)
Although the two smallest females (wings 58½ and 59 mm) are in extremely worn plumage, there areamong the males also individuals in very worn plumage. Therefore it seems safe to conclude that in thisspecies, as in other Zosteropidae, the females average a little smaller than the males. There is no difference in plumage between the sexes.

Heleia crassirostris crassirostris (Hartert)
Zosterops crassirostris Hartert, 1897, Novit. Zool. 4: 172 – South Flores.
Collectors.— Everett (at least 11, cf. Mees, 1969: 211), Rensch (1), Verheijen (3, MCZ), Verheijen/Schmutz.
Material.— , 20.v.1969, Waé Laci (Verheijen, RMNH no. 81393); , 4.xi.1969, Ruteng (Verheijen, RMNH
no. 65459); , , 23.xii.1969, Todo (Verheijen, RMNH nos. 65461, 65460); , , 20.vi.1971, Waé- Ntjuang,
Langkas (Verheijen, RMNH nos. 85354, 85353); (?) nestling, 5.v.1971, Lingko-Moak, Langkas (Verheijen,
RMNH no. 66194); , 9.i.1976, L. Ngkiong (Verheijen, RMNH no. 81387); , 15.i.1976, Wae Cuang(Schmutz, RMNH no. 97134); , 7.ii.1976, Rempo (Verheijen, RMNH no. 85356); , , 2.vi.1976, Ulu Ros(Verheijen, RMNH nos. 81394, 81389); , 25.ix.1976, Poco Nernancang (Verheijen, RMNH no. 85355); ,
30.v.1978, Waé Hiam (Verheijen, RMNH no. 81390); , 31.v.1978, ? Méléng (Verheijen, RMNH no.
81396).
Eggs (fig. 16i): 23 different sets, of which c/1 (5 ×), c/2 (12 ×), c/3 (5 ×) and c/4 (1 ×), taken in the months
April (4), May (4), June (6), July (3), August (3), September (2) and October (1), (RMNH nos. 52318,
71642-71663). Assuming the c/1 to be incomplete, the usual clutch-size would be 2 or 3. The eggs arewhite, only slightly glossy, without markings.
Some measurements and weights: RMNH no. 71642

RMNH no. 71645
RMNH no. 71646
RMNH no. 71657
RMNH no. 71658

RMNH no. 71659

19.2 × 13.7
0.108419.2 × 14.1
0.1052
19.3 × 14.1
0.097619.3 × 14.6
0.114519.7 × 13.9
0.1227
19.7 × 13.8 }large holes
19.8 × 13.8 }19.0 × 14.2
19.0 × 14.2
18.4 × 13.6
0.0941
18.6 × 13.9
0.0997
18.7 × 13.8
0.107518.3 × 13.6
0.085819.1 × 13.5
0.0992

Notes.— H. crassirostris has been divided into two subspecies, the nominate one
confined to Flores, and H. c. junior confined to Sumbawa. Whereas from Flores now
good series are known, only the original two specimens (collected by Rensch) havebeen recorded from Sumbawa, and as I have pointed out (Mees, 1969: 212), the validityof junior requires confirmation, or, rather, its validity is dubious. White & Bruce (1986:419) stated with confidence that junior is a synonym, but how they could be so certain
is not clear, for as far as I can judge, they had not examined any specimens.

194

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The present material confirms that there is no difference in measurements between
birds from Flores and birds from Sumbawa, but until more material from Sumbawa
may become available, I would keep to my former opinion, that junior merits the benefit of the doubt.
Measurements:
6

wing
tail
tarsusentire culmen exposed culmen
66½-7349-56
19.8-20.7
18-19.2
14-16(69.7)(53.0)(20.2)(18.6)(15.2)
8
65-69
49-51
19-20
16.2-17.8
12.6-14.8(66.6)(50.0)(19.6)(17.0)(14.0)The smallest male has the tips of the primaries extremely worn, but even so, there is a clear average difference in size between the sexes. Previously, I was unable to establish this because of lack of females(only three having been traced, all in very worn plumage) (cf. Mees, 1969: 210).

Lichmera indistincta limbata (S. Müller)
Meliphaga (Ptilotis) limbata S. Müller, 1843, Verh. Nat. Gesch. Ned. Overz. Bez., Land- en Volkenk.: 162 –Timor.
Collector.— Allen (1).
Material.— (?) juv., 1862, Flores (Allen, BM no. 1873.5.12.994).

Notes.— Lichmera indistincta limbata is very common on most of the Lesser Sunda
Islands, from Bali to Timor. This makes it even more peculiar that on Flores the specimen without exact date and locality, collected by Allen in 1862, remains the only record.
Rensch (1931a: 608) wrote: “auf Flores merkwürdigerweise nicht beobachtet”. Verheijen
did not find the species on Palué, and Schmutz (MS) states expressly that on Flores he
has never heard the conspicuous song. I have observed (seen and heard) it on Gili Lawa
Laoet, an islet off the north-east coast of Komodo, about mid-way between Sumbawa
and Flores, and only ca. 30 km from the nearest point of mainland Flores (September
1984). Hoogerwerf (1956) does not mention L. indistincta from Komodo.
In this connection, Allen’s specimen assumes an increased importance. Dr Cowles
has examined it for me; he confirmed that it has been identified correctly and that it has
what seems to be an original label in the handwriting of Allen, with the words “Flores”,“eyes dark” and a figure “2”, to complete a pre-printed year 186”2”. Therefore, therecord is as reliable as can be expected from a specimen collected so long ago.
Since the introduction of ternary nomenclature, L. i. limbata had always been considered a, not particularly strongly differentiated, subspecies of L. indistincta. Sibley &Monroe (1990: 429) have promoted it to an “allospecies”. In this connection I should liketo mention that I know Australian L. indistincta quite well. My first field experience withlimbata is from the island of Sumba, where on my arrival in Melolo, I heard song fromthe mangroves, which I did not hesitate a moment recognising as being of L. indistincta:
in addition to their similarity in morphology and plumage, the two forms evidently
agree also in song. I believe that the relationship between indistincta and limbata is correctly expressed by treating them as subspecies.
The eggs support the close relationship. Father Verheijen (1976: 18) collected someclutches on Ndao near Roti. They are white, with small and sparse brown spots (one ispractically spotless).

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)
Measurements and weights of the eggs from Ndao are:
RMNH no. 76731

RMNH no. 76732
RMNH no. 76733

19517.4 × 14.0
17.5 × 14.1
17.8 × 14.0
16.7 × 14.3
16.9 × 13.5
17.2 × 13.4

0.09050.08940.08490.0691
0.072
0.0669Both in markings and in measurements and weights, they agree with eggs of nominate L. indistincta. White & Bruce’s (1986: 399) statement that this species is “commonest
800-1200 m”, is not correct for most of its range. My own observations, given above,show it to inhabit small islands and, on the large islands, lowlands, including coastalmangroves. This is supported by Verheijen for Roti and its adjacent small islands, andalso by the material collected by Stein on Timor and Sumba (cf. Mayr, 1944a: 137, 143).
Lichmera lombokia fumidigula (Rensch)
Meliphaga virescens fumidigula Rensch, 1928, Orn. Mber. 36: 9 – Geli Moetoe (1500 m), Flores.
Collectors.— Everett (1), Rensch (9), Verheijen/Schmutz.
Material.— 61 specimens from several localities in Manggarai, to 2200 m (mountain lake Rana-Ka).
(RMNH nos. 59886-59894, 65526-65541, 81358, 81361, 85225-85244). Iris dark brown, bill black, legs slate.

Notes.— The large series obtained shows that this is one of the most common birdspecies at the higher levels. Therefore it is surprising that not a single nest seems to havebeen found. Perhaps, the explanation is the same as that suggested for Lophozosterops
superciliaris on a previous page.
White & Bruce (1986: 398-399) have placed fumidigula in the synonymy of thenominate race, but a comparison of the series from Flores with our four specimensfrom Lombok (leg. Vorderman, 1895b: 15) showed me that all the characters claimed
by Rensch (1928a) for fumidigula, hold good. Especially the smoke-coloured throat
from which fumidigula takes its name, is a valid, albeit not particularly conspicuous,
character. The average difference in size between the two subspecies is fully confirmed.
It is interesting that on Lombok, L. l. lombokia is a lowland form and an inhabitant ofcultivated country, recorded from 0-1200 m, whereas L. l. fumidigula of Sumbawa andFlores inhabits mountain forest, with an altitudinal range of ca. 1000-2200 m. Although
the subspecies concept is primarily and necessarily a morphological one, I consider thatin judging the validity of fumidigula, such ecological differences may certainly be considered.
Note that the geographical variation of L. lombokia goes in two respects against
Rensch’s theories: it is the eastern subspecies which is smaller than the western subspecies, and it is the mountain subspecies which is smaller than the lowland subspecies.
Naturally, it is tempting to relate the variation in habitat and vertical distribution on
the different islands, with the presence of the closely related L. indistincta, but on Flores
L. indistincta is so scarce (if present at all), that it can hardly influence the distribution of
L. lombokia.

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Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Measurements:
wing
Lombok (L. l. lombokia)
4
67-70(68.5)
Flores (L. l. fumidigula)25 
61½-68(65.5)
36 
58-63(60.7)

tail

tarsusentire culmen

exposed culmen

50-52(51.25)20-21.6(20.7)

20+-23+–

16.3+-20+–

48-55(51.6)
44½-50(47.9)

18-21.2(19.6)
17.0-20(18.6)

20.4-25(23.4)
20-24.9(21.7)

17.9-21.8(19.8)
17.4-21.5(18.4)

Philemon buceroides neglectus (Büttikofer)
Tropidorhynchus neglectus Büttikofer, 1891, Notes Leyden Mus. 13: 213 – Flores, Sumbawa.
Philemon timoriensis plesseni Rensch, 1929, J. f. Orn. Ergänzungsb. II: 198 – Lomblen.
Collectors.— Semmelink, Allen, Martens, Weber, ten Kate, Everett (2), Endih (1), Rensch (4), de Jong (2),Verheijen (2, MCZ), Pfeffer (2), Verheijen/Schmutz.
Material.— (?), 1862, Larantoeka (Semmelink, RMNH cat. no. 3, syntype); 2 (?), xii.1888, Maumeri(Weber, RMNH cat. no. 7, and Doubl., syntypes); (?), 28/30.i.1891, Endeh (ten Kate, RMNH cat. no. 8,
syntype); (?) fledgeling, 13/24.vi.1891, Sika (ten Kate, RMNH cat. no. 9, syntype); , 28.x.1969, Nunang, 650 m (Schmutz, RMNH no. 97148); , 18.ix.1970, Kisol, 150 m (Verheijen, RMNH no. 65365);
, 10.iii.1971, Kisol, 175 m (Verheijen, RMNH no. 66147); , 9.iii.1971, Kisol, 175 m (Verheijen, RMNH
no. 66146); 4 , 2 , 11.iii.1971, Kisol, 175 m (Verheijen, RMNH nos. 66148-66153); , iii.1971, Kisol,
175 m (Verheijen, RMNH no. 66145); , 20.vii.1971, Nunang, 600 m (Schmutz, RMNH no. 85297); ,
27.vii.1971, Nunang, 650 m (Schmutz, RMNH no. 85298); , 12.x.1971, Nunang, 650 m (Schmutz,
RMNH no. 85299); 2 , 13.vi.1973, Golo-Karot, Borong, 50 m (Verheijen, RMNH nos. 85300, 85301); ,
14.vi.1973, Golo-Karot, 50 m (Verheijen, RMNH no. 85302); 4  , , 23.iii.1976, Nunang, 650 m (Schmutz,
RMNH nos. 81031, 81034, 81036, 81035).
Eggs (fig. 17a): 53 sets of c/1 (16 ×), c/2 (14 ×) and c/3 (23 ×), taken in the months January (1), February(2), March (2), April (11), May (12), June (5), July (1), October (8), November (8) and December (1), andtwo not dated (RMNH nos. 71664-71715, 73522). The eggs vary from white to a deep salmon colour, withbrown primary and light grey secondary dots, variable in size and density.
Some measurements and weights: RMNH no. 71678
RMNH no. 71690

RMNHH no. 71691

RMNH no. 71692

RMNH no. 71703

RMNH no. 71704
RMNH no. 71705

33.8 × 22.3
30.4 × 21.8
30.9 × 22.1
34.6 × 21.8
29.6 × 22.7
30.4 × 23.0
33.1 × 22.4
30.7 × 21.7
30.7 × 22.0
32.0 × 22.3
31.0 × 21.8
32.7 × 21.6
32.9 × 21.4
29.2 × 23.1
31.3 × 21.8
32.2 × 20.7
32.3 × 21.6

0.55270.37230.4041
0.48710.4116
0.4263
0.48140.38030.39460.43860.44300.49690.49070.51780.48350.43240.4511

Notes.— White (in White & Bruce, 1986: 398) has dismissed both the subspecies
plesseni and sumbanus, on the basis of what appear to be mostly Rensch’s own figures,

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

197

adding: “this minor size variation is probably not worth formal designation”. Whatever
the studied material was, it must have been small and therefore I considered it worth
measuring the above specimens from Flores and other specimens from the Lesser Sunda Islands. The results show that the increase in size eastwards is questionable: evenbirds from Lombok have a wing-length of up to 153 mm, birds from Flores to 154 mm.
The individual variation is apparently much greater than Rensch thought, and therefore I agree with White that P. t. plesseni is a synonym of P. b. neglectus. With only a single
unsexed specimen from Sumba at hand, I am unable to give a definite opinion on thevalidity of sumbanus, but note that the measurements of wing and tail of the Sumba
specimen exceed those of birds from all other islands, and therefore support Rensch’scontention that: “Sumba-Vögel sind langflügeliger und langschwänziger...”. The figures presented by Mayr (1944a: 165) also confirm the size difference. Whether this difference is enough for recognition of the subspecies sumbanus is a different, and perhaps
largely subjective, question. The case for retention of sumbanus is weakened by Hoogerwerf’s (1966) claim that birds from Komodo are equally large.
Measurements:
Flores
14 

wing
tail
tarsusculmen from posterior edge of knob
140-154
111-127
34-40.5
40.4-43.4(147.7)(119.7)(36.5)(42.0)
6
134-143
111-120
34-36
38-41(139.8)(114.8)(35.2)(39.6)
3 (?)
149-153
125-128
39-41
41-44.5
Judging by these figures, the three unsexed birds (collected by Semmelink and Weber), should be males.
Lombok 3 145,151,153 121-131
36-38
43.2-46
3 (?)
142-146
119-123
36-37½
41.8-45
Sumbawa 
147
118
40
44
Alor

153
123
38
43
Sumba
(?) (= )
155
135
40
46
The Sumbawa specimen is from the Forsten collection (1842); it is a syntype of P. b. neglectus (RMNH cat.
no. 1); all other syntypes are from Flores, as listed above.

Amandava amandava flavidiventris (Wallace)
Estrelda flavidiventris Wallace, 1864, Proc. Zool. Soc. Lond. (1863): 495 – Timor and Flores.
Collectors.— Semmelink, Allen (at least 8, cf. Sharpe, 1890: 323-324), Everett, Rensch (2), de Jong (2),Verheijen (8, MCZ), Verheijen/Schmutz.
Material.— , 1862, Larantoeka (Semmelink, RMNH cat. no. 1); , 26.v.1969, Wangkung, Rahong (Verheijen, RMNH no. 65577); , 23.vi.1971, Wae-Ntjuang, Langkas (Verheijen, RMNH no. 66207); ,
24.vi.1970, Ruteng (Verheijen, RMNH no. 65583); , 25.vi.1971, Lingko Watu Mésé, Langkas (Verheijen,
RMNH no. 85317); , 27.vi.1971, Wae-Ntjuang, Langkas (Verheijen, RMNH no. 66208); , 29.vi.1971,
Langkas, Rahong (Verheijen, RMNH no. 66209); , 5.vii.1971, Lingko-Moak, Langkas (Verheijen, RMNH
no. 85318);  im., 16.x.1971, Ruteng (Verheijen, RMNH no. 85321); ,12.vi.1976, Longko (Schmutz,
RMNH no. 81370). Iris wax-red, bill dark red, legs pale brown.
Eggs: c/1, 4.vi.1954, Potjong (RMNH no. 71747); c/4, 17.vi.1954, Puntu, Mano (RMNH no. 71748); c/4,
31.v.1954, Montjok (RMNH no. 71749); c/1, 7.vi.1954, Wesang (RMNH no. 71750); c/3, 14.v.1955, Mano(RMNH no. 71751); c/2, 23.v.1955, Potjong (RMNH no. 71752); c/2, 6.viii.1955, Mengé (RMNH no.
71753); c/4, 3.vi.1956, Wangkung (RMNH no. 71754); c/6, 5.vi.1957 (RMNH no. 71755); c/4, Mano,
10.vi.1958 (RMNH no. 71756); c/6, v.1959, Wangkung (RMNH no. 71757); c/4, 2.v.1959, Potjong (RMNH
no. 71758). The eggs are white, a little glossy.

198

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Some measurements and weights: RMNH no. 71749

RMNH no. 71757

14.0 × 11.0
14.1 × 10.6
14.1 × 10.7
14.2 × 10.8
13.8 × 9.9
13.8 × 10.1
14.2 × 10.5
14.3 × 10.8
14.5 × 10.7
14.6 × 10.8

0.05330.0496
0.05220.05140.035
0.04270.0476
0.05140.05030.0507

Notes.— This species has the distinction of being the smallest bird of Flores, both inwing-length and in size and weights of its eggs.
Measurements:
6
 im.
2

wing
44½-46(45.4)
46
46, 46tail30½-32½(31.3)
32
30½, –

tarsus
12.5-13(12.8)
14
13, 13

culmen9.0-10.0(9.5)
10.0
10.0, 10.2

Poephila guttata guttata (Vieillot)
Fringilla guttata Vieillot, 1817, Nouv. Dict. d’Hist. Nat. 12: 233 – Îles Moluques = Timor.
Amadina insularis Wallace, 1864, Proc. Zool. Soc. Lond. (1863): 495 – Timor and Flores.
Collectors.— Allen, Weber (3), Rensch (6), de Jong (8), Schmutz (1).
Material.— , not dated, acquired in 1878, Flores (no collector, RMNH cat. no. 13); , xii.1888, Maumeri(Weber, prepared from alcohol, RMNH cat. no. 20); , 9.v.1969, Nisar/Sésok, south coast (Schmutz,
RMNH no. 81304).
No eggs (but see notes).

Notes.— Verheijen (1961) recorded that, both on Flores and on Palué, Zebra finches
were only seen in the immediate vicinity of the coast. Although Father Verheijen did notcollect eggs of this species on Flores, he obtained several clutches on Roti and the adjacent islet of Dao or Ndao (Verheijen, 1976: 18-19).
Erythrura hyperythra obscura (Rensch)
Chlorura hyperythra obscura Rensch, 1928, Orn. Mber. 36: 6 – Sita (700 m), West-Flores.
Collectors.— Everett (1), Rensch (2), Verheijen (2, MCZ), Verheijen/Schmutz.
Material.— , 14.v.1971, Langkas, Rahong (Verheijen, RMNH no. 85316); , 17.v.1976, Lingko Ncilor(Schmutz, RMNH no. 81313).
Eggs: Verheijen (1964: 200) records nests found in the months May and June. There is one clutch in hiscollection, but its identification is very dubious.

Notes.— The subspecies obscura was placed in the synonymy of E. h. intermedia by
White & Bruce (1986: 420-421), but without explanation, unless the remark: “No further
subdivision seems necessary in this species with a range of broken, isolated popula-Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

199

tions”, be taken as such. Lacking specimens from Lombok (intermedia), I have been
unable to make a comparison with Rensch’s diagnosis of obscura, and until a criticalcomparison may prove otherwise, I accept it.
The sexes differ in that the male has more black on the forehead, and the blue of theforecrown deeper in colour and extending farter backwards, than the female; the brown
colour of the sides of the face and the underparts is darker in the male than in the female.
In this connection I note that in the original description of obscura, Rensch compared 5 
ad. from Sumbawa and Flores, with 5 ad., of which he did not record the sex, of intermedia from Lombok. One of these (the one he collected, cf. Rensch, 1931a: 602) was certainly a female; this may explain in part why Rensch found the cheeks and underpartsof obscura darker brown than those if intermedia. The differences in size as well as in colour were confirmed by Ziswiler, Güttinger & Bregulla (1972: 19, 23), but note that theirseries of intermedia consisted of 4 , 7 . Of obscura, they examined 5  and one  (from
Sita, Flores, hence undoubtedley the bird listed as  juv. by Rensch, 1931a: 603).
Measurements:



wing
56
58

tail
29
29

tarsus
15.7
16.2

culmen10.0
9.8

Lonchura molucca propinqua (Sharpe)
Uroloncha propinqua Sharpe, 1890, Cat. Birds Brit. Mus. 13: 368 – Flores.
Uroloncha kangeangensis Vorderman, 1893, Nat. Tijdschr. Ned. Ind. 52: 199 – Kangean.
Munia molucca vagans Meise, 1929, J. f. Orn. 77: 440 – Binongka (Tukang besi-Inseln).
Collectors.— Semmelink, Allen, Weber, ten Kate, Everett, Rensch (1), de Jong (2), Verheijen/Schmutz.
Material.— (?) ad., (?) juv., 1862, Larantoeka (Semmelink, RMNH cat. nos. 4, 5); (?), iv.1891, Sika (tenKate, RMNH cat. no. 6); (?), 5.vi.1969, Nunang, 650 m (Schmutz, RMNH no. 81368); , 19.ix.1970, Kisol(Verheijen, RMNH no. 65576); , , 28.vii.1971, Langkas, Rahong (Verheijen, RMNH nos. 85320, 66226);, , 20.ii.1978, Waé-Wako, 180 m (Schmutz, RMNH nos. 81364, 81365).
Eggs: c/3, 27.vi.1956, Dampék (RMNH no. 71740); c/4, 22.iii.1960, Mataloko (RMNH no. 71741); 2 c/1,
1.v.1960, Palué (RMNH nos. 71742, 71743); c/3, 1.v.1960, Palué (RMNH no. 71744); c/2, 20.vi.1960, Palué(RMNH no. 71746); c/4, 3.v.1961, Mataloko (RMNH no. 71745). The eggs are white.
Some measurements and weights: RMNH no. 71741

RMNH no. 71744

RMNH no. 71745

15.8 × 10.7
16.1 × 10.9
16.3 × 10.8
16.6 × 10.7
14.4 × 11.0
15.0 × 10.9
15.2 × 10.7
16.4 × 11.0
16.9 × 11.3
17.0 × 11.3
17.3 × 11.5

Notes.—
Measurements:
2
3

wing
52, 5251, 51, 51

tail
-, 36
36-40

tarsus15.5, 15.7
15-16

culmen11.4, 12
11, 11, 12

0.05640.05840.05740.05380.051
0.05150.05190.05640.05940.06230.0635

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Lonchura punctulata blasii (Stresemann)
Munia punctulata blasii Stresemann, 1912, Novit. Zool. 19: 317 – Timor-Deli.
Collectors.— Semmelink, Allen, Everett, Rensch (6), Verheijen (2, MCZ), Verheijen/Schmutz.
Material.— (?), 1862, Larantoeka (Semmelink, RMNH cat. no. 11, mounted); 16 , 12 , 1(?), months i,
iv, v, vi, vii, viii, xii, years 1968, 1969, 1971 (Verheijen/Schmutz).
Eggs: 14 clutches of 1 (2 ×), 2 (2 ×), 3 (3 ×), 4 (2 ×), 5 (2 ×) and 6 (3 ×) eggs, from the months March (5),April (7), May (1) and July (1), (RMNH nos. 71726-71639). The eggs are white.
Some measurements and weights: RMNH no. 71729

RMNH no. 71731

RMNH no. 71735

14.6 × 11.0
14.8 × 10.9
15.0 × 11.0
15.1 × 11.1
15.2 × 10.7
15.4 × 11.0
14.0 × 11.1
14.2 × 11.0
14.5 × 11.1
14.6 × 10.9
14.8 × 11.2
13.9 × 10.3
14.1 × 10.6
15.5 × 10.4
15.7 × 10.8

0.06030.06070.06070.05920.062
0.063
0.05340.05550.0536
0.05350.054
0.05120.05490.0551
0.0532

Notes.— Lonchura punctulata is a widely-distributed species, which shows a considerable amount of geographical variation. The most important variation is found in threecharacters: general size; colour-pattern of the breast and the flanks, which look either
scaly, or, more finely, vermiculated; and colour of the upper tail-coverts, and edges tothe rectrices. Minor characters which have been used for subspecific discrimination are
the depth of the brown colour, and the presence of pale shaft-streaks to feathers of thehead and mantle. Mayr et al. (1968: 375-377) list 12 subspecies (one of which with aquestion-mark). A superficial comparison of material from Flores with specimens fromneighbouring islands, gave me the impression, that the validity of some of the subspecies required confirmation. This led to the following review:
Lonchura punctulata blasii (Stresemann)
Characters.— Small; upper tail coverts and central rectrices tinged strongly withstraw-yellow; scaly markings om breast and flanks dark, almost blackish brown. Billaveraging larger than in sumbae.
Distribution.— The Lesser Sunda Islands from Flores eastwards, including Savu
but not Sumba.
Lonchura punctulata sumbae Mayr
Lonchura punctulata sumbae Mayr, 1944, Bull. Amer. Mus. Nat. Hist. 83: 169 – Sumba Island.

Characters.— Small; very similar to L. p. blasii, but the scaly pattern on (especially)
the breast, also the flanks, averages a little less dark, dark brown as in L. p. nisoria andL. p. particeps, not almost blackish brown. The bill averages smaller than in all otherMees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

201

subspecies, although the difference from blasii and particeps is slight.
Mayr’s (1944a) diagnosis of L. p. sumbae reads: “Similar to blasii (Timor), but Ushaped bars of underparts rufous brown, not blackish brown as in blasii, or rufous as inparticeps or fortior; white area on each feather of underparts narrower, brown vermiculated more irregular; under tail-coverts more heavily barred; differs from fortior (Lombok, Sumbawa) by smaller size and darker underparts.... Savu and Flores birds have theblackish brown bars of blasii”.
I cannot support Mayr’s statement that particeps and fortior (a synonym of nisoria)
would be “rufous” on the underparts, for as stated above, I fail to see any difference atall between the underparts of sumbae and “fortior”. Nor can I agree, however, withWhite & Bruce (1986: 423), who ascribe the similarity in underparts between sumbae andnisoria to “intergradation”, and by no stretch of the imagination could I call the underparts of these subspecies “reddish”, as they do.
Distribution.— Sumba.
Lonchura punctulata particeps (Riley)
Munia punctulata particeps Riley, 1920, Proc. Biol. Soc. Washington 33: 57 – Rano Lindoe, Celebes.

Characters.— Small; upper tail coverts with a little less yellow than in blasii (or is thedifference due to wear?); the scaly pattern of breast and flanks averages slightly lessdark than in blasii, agreeing with nisoria and sumbae. Bill as in blasii.
Distribution.— South-western and Central Sulawesi (Celebes). Records from thenorthern peninsula of Celebes are erroneous.
Lonchura punctulata nisoria (Temminck)
Fringilla nisoria Temminck, 1830, Recueil d’Ois. 3 (livr. 84): pl. 500 fig. 2 – Java.
Munia punctulata fortior Rensch, 1928, Orn. Mber. 36: 7 – Swela (400 m), Lombok.
Munia punctulata fretensis Kloss, 1931, Treubia 13: 363 – Kuala Lumpur, Selangor, Federated Malay
States.
Lonchura punctulata baweana Hoogerwerf, 1963, Bull. Brit. Orn. Cl. 83: 38 – Bawean.
L[onchura] p[unctulata] holmesi Restall, 1992, Avicult. Mag. 98: 115 – based on cage-birds, presumed tooriginate from near Pontianak and Banjermasin, Borneo.

Characters.— Small; upper tail coverts and central rectrices light pearl-grey, without yellow (occasionally there may be a trace of yellow), scaly markings on breast andflanks varying from dark brown to blackish brown (not clearly different from blasii, but
in series perhaps a trifle lighter). Bill large. Riley states correctly: “rump barred withwhite” (although this is variable and sometimes almost absent), whereas this is muchless in blasii, sumbae and particeps, and what there is, is yellowish rather than greywhite.
Distribution.— The Malay Peninsula, northwards to Trang and Phattalung (cf.
Medway & Wells, 1976: 394), Sumatra, Java, Bali, Lombok, Sumbawa, Bawean, and thenorthern peninsula of Celebes. In the last-mentioned locality, and perhaps in others, it
has presumably been introduced. Birds recently found in south-eastern Borneo (Harvey & Holmes, 1976), would probably also belong to this subspecies.
Notes.— Medway & Wells (1976: 394) suggest that L. punctulata may have been introduced in Malaya; they mention that Kelham, “working in Singapore and the western
Malayan states in the 1870s”, did not collect it. However, Horsfield & Moore (1858: 506)
list a specimen from Penang, received from Dr Cantor in 1854. This bird was later transferred to the British Museum (BM no. 60.4.16.340), which also holds two specimens

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from Malacca, collected by Wallace in 1854 (BM nos. 73.5.12.1370 and 81.5.1.4480), anda series of 36 skins from various parts of Malaya, and Singapore, collected by W. Davison in the years 1875-1879. These records do not disprove the suggestion that the species was originally introduced, but they put back in time the date of the presumed introduction.
From Bawean, our collection contains two adult birds: a paratype of L. p. baweana(leg. Franck, 1928), and one leg. Vorderman, 1896. Also two paratypes in juvenile plumage. The adult paratype shows the rather light markings of the underparts which Hoogerwerf made an important subspecific character; the markings are not lighter, however,than in the lightest Java birds. The other adult bird has the markings darker, agreeing inthis respect with average Java birds. The smaller brown throat-patch ascribed to baweana is entirely due to the make-up of the skins. The two paratypes of baweana in juvenile
plumage differ, contrary to Hoogerwerf’s claim, in nothing from Javanese birds in juvenile plumage, and are not paler. I conclude that baweana is a synonym of nisoria.
Specimens from the range ascribed to L. p. fortior are poorly represented in the collections studied by me. It is a size-race, that was differentiated merely on the basis ofwing-length: Java and Sumatra 50-53 mm, Lombok and Sumbawa 53-55 mm, according
to Rensch (1928a). The variation found by me in a not particularly large series of nisoria
from Java and Sumatra is 50-54 mm. Two birds from Lombok have wing-lengths of 52
and 53 mm. One specimen examined by Hoogerwerf, had a wing-length of 52 mm. Inadequate as my material is, I cannot see any argument for retention of fortior. The type
locality of fortior is Lombok, as cited above, not Sumbawa as claimed by White & Bruce(1986: 423).
Later, Rensch (1931a: 598) added, that fortior would have the brown of chin andthroat less extensive than nisoria, and the white parts of the feathers of breast and belly
more extensive. My comment on this is the same as that given in the discussion of
baweana. White & Bruce (1986: 423) have already dismissed fortior with the remark: “thedifferences distinguishing... fortior... seem insignificant”.
No material from Sumbawa has been available. Zoogeographically, I find it surprising that this island is inhabited by L. p. nisoria, and the adjacent island of Flores by L. p.
blasii. Perhaps, these are fairly recent range-extensions, helped by Man.
The validity of fretensis was questioned by Hoogerwerf (1963a), but it was still accepted by Mayr et al. (1968: 376) and Van Marle & Voous (1988: 208). I have been unable
to find any difference in plumage or in measurements (see table), which might justify
its retention.
Five specimens from Tondano, collected by Coomans de Ruiter in 1939 (RMNH),
and recorded by Van Marle (1940a) and Stresemann (1940a: 37) as L. p. particeps, arequite unequivocally L. p. nisoria. Undoubtedly, the population owes its origin to introductions from Java.
I have also examined two specimens from Bangkok, 8.viii and 8.xii.1917 (leg. Williamson, BM nos. 1955.1.4236 and 1955.1.4231), which are clearly nisoria. They musthave been imported, for all other specimens from Bangkok in the Williamson collection
are topela.
The recently-described holmesi is considered another synonym of nisoria, based on
birds introduced to Borneo from Java. For a further evaluation of this form, see the discussion of L. p. insulicola.

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Lonchura punctulata punctulata (Linnaeus)[Loxia] punctulata Linnaeus, 1758, Syst. Nat. (ed. 10), 1: 173 – Asia.
Munia lineoventer Hodgson, 1836, Asiatic Res. 19: 154 – Nepal (reference not verified).

Characters.— Large; upper tail-coverts brownish-orange; markings of the breast of
the “punctulata”-type.
Distribution.— Practically the whole of India; the southern and eastern parts of
Nepal; Sikkim; southern Bhutan; Sri Lanka (Ceylon); south-western Burma.
Lonchura punctulata subundulata (Godwin-Austen)
Munia subundulata Godwin-Austen 1874 (June), Proc. Zool. Soc. Lond. (1874): 48 – Manipúr valley bothon Lake Logtak and the head of the Barak river.
M[unia] superstriata Hume, 1874 (October), Stray Feathers 2: 481 footnote – Tavoy.
Lonchura punctulata catervaria Koelz, 1954, Contrib. Inst. Reg. Expl. 1: 19 – Mawphlang, Khasi Hills.

Characters.— Large, upper tail-coverts mostly as in topela, but sometimes orange asin punctulata, markings of the breast variably intermediate between the “punctulata”
and the “topela” types.
Distribution.— Eastern Assam, Manipur, western and southern Burma.
Notes.— Birds from Manipur and much of Burma are very variable, intermediate
between the well-differentiated punctulata and topela. It is a moot question, whether
such populations should be indicated by a separate subspecific name, but in view of thewide range of these intermediate populations, I consider it convenient to do so. This isalso in agreement with current classifications.
Smythies (1986: 396 and pl. xi fig. 6) gives in the text the names subundulata (as thecommon subspecies) and topela (status uncertain), but the illustrated bird seems to belong to the nominate race, at least is not topela-like, and is captioned L. p. lineoventer. As
lineoventer is a synonym of nominate punctulata, this is correct, but there is no mention,
under either name, of this subspecies in the text.
Lonchura punctulata topela (Swinhoe)
Munia topela Swinhoe, 1863, Ibis 5: 380 – In China it is abundant from Canton to Shanghai, and in Taiwan
all throughout the plains.
Lonchura punctulata yunnanensis Parkes, 1958, Proc. U.S. Nat. Mus. 108: 285 – “hills around Tengyueh” [=Tengchung], western Yunnan, alt. 6,000 feet.

Characters.— Large; upper tail coverts tinged greenish yellow (not straw yellow asin blasii); markings of the breast of the “topela”-type, quite different from those found in
the preceding subspecies.
Distribution.— Eastern Burma, Thailand, Indo-China, southern China, Hainan andTaiwan. The birds introduced into eastern Australia, where the species is now widelydistributed (Blakers et al., 1984: 604, 609), have been assigned to this subspecies (Boles,
1988), as have those introduced in Hawaii (Violani, 1979).
Notes.— The original description of Munia topela appeared in a paper on the birdsof Taiwan, and its distribution was given as quoted above. Later authors, including
Parkes (1958: 285) and Mayr et al. (1968: 376) have listed Amoy as the type locality.
About Amoy as the type locality, see Deignan (1963: 217), who says: “type locality (inferentially) restricted to Amoy... by Sharpe... 1890, p. 352”. The fact that, in the place
indicated, Sharpe refers to seven specimens from Amoy, ex Seebohm coll., as “Types ofthe species”, does in my opinion not constitute a valid restriction of the type locality,

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and even less a lectotype-designation (ICZN, 1985: art. 72(b) (vii)). Warren & Harrison(1971: 564) list a specimen from Amoy as a syntype, and comment correctly: “This specimen is labelled as the type but the author refers to a large series”.
In view of the title and contents of the paper in which the original description appeared, it would be more logical to have ‘Formosa’ as type locality, and this is also theopinion of Chinese authors (Cheng, 1987: 953), which I share. As long as the populations inhabiting Taiwan and the east coast of China are considered consubspecific, thepoint is, of course, a purely academic one.
The four specimens from Yunnan examined by me, one of which is from the typelocality of L. p. yunnanensis, are not perceptibly different from topotypical L. p. topela,
under which name I bring them.
Lonchura punctulata cabanisi (Sharpe)
Munia cabanisi Sharpe, 1890, Cat. Birds Brit. Mus. 13: 353 – nomen novum for Oxycerca (Uroloncha) Jagori
Cabanis, 1872, nec Munia (Dermophrys) Jagori Martens, 1866.
Oxycerca (Uroloncha) Jagori Cabanis, 1872, J. f. Orn. 20: 317 – Luzon.

Characters.— Small; similar to topela from Taiwan, but smaller, with also an on average smaller bill; markings of the breast even a little finer and lighter (but variable!);upper tail-coverts and edges of the rectrices greenish-yellow as in topela.
Distribution.— Luzon, Mindoro and Panay in the Philippines; also found on theMicronesian islands of Yap and Babelthuap (Palau group), where probably introduced(Baker, 1951: 340).
Lonchura punctulata insulicola subspecies nova
Type: ,13.iii.1865, Réunion (leg. Pollen & van Dam, RMNH cat. no. 36).

Characters.— Large; upper tail coverts straw yellow (as in blasii), markings of thebreast coarse, of the “punctulata”-type.
Distribution.— Mauritius, Réunion and Mahé (Seychelles).
Notes.— The characters of specimens from Réunion have first been discussed by
Büttikofer (1892): “Our specimens from Bourbon agree entirely with M. nisoria from
Java, with the exception of the upper tail-coverts and centre tail-feathers, which are notashy gray, but sensibly tinged with pale olive-green, in which character they agree with
M. topela. They are, however, undoubtedly to be united with M. nisoria”.
With the introduction of ternary nomenclature, the names M. p. nisoria (cf. Meinertzhagen, 1912: 91) and M. p. punctulata have been used for birds from the Mascarenes.
To this, Berlioz (1946: 66) commented: “Les spécimens de La Réunion que possède leMuséum de Paris, tout en étant certainement moins chaudement colorés que ceux del’Inde, présentent tout aussi bien des caractères ambigus référables tant aux formes indochinoises qu’aux formes de Malaisie”. Presumably it is on the basis of this evaluationthat Mayr, Paynter & Traylor (1968: 376) have referred birds from Mauritius, Réunion,
and the Seychelles to L. p. topela.
I have examined the same six specimens from Réunion that were available to Büttikofer (RMNH cat. nos. 33-38). I have also examined one specimen from Mahé (BM no.
1906.12.21.431, cf. Nicoll, 1906: 706), but none from Mauritius. I cannot agree with Büttikofer that they are nisoria, for not only do they have straw yellow upper tail coverts,
but in addition they are clearly larger, too large for nisoria. On the other hand, they are
not topela either, for they have the markings of the breast of the “punctulata/nisoria”

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type, not of the much finer “topela” type. The most obvious places of provenance for
introduced birds on Réunion would be coastal India or Sri Lanka (Ceylon), but the deepbrown-orange upper tail-coverts of nominate punctulata are completely absent.
In conclusion, birds from Réunion are neither nominate punctulata, nor nisoria, nor
topela. Their characters do not suggest a hybrid population between any of these forms
either. The possibility that through some sort of deficiency, they are punctulata whichhave lost the bright orange colours, must be envisaged, but does not seem very likely.
It has been generally assumed that Lonchura punctulata was introduced to the Mascarenes, but when exactly, and from where, is lost in the mist of history, see the review
by Cheke (in Diamond, 1987: 80). Meinertzhagen’s claim that on Mauritius it was: “Introduced from Java about 1800”, is given without a reference, and therefore cannot be
verified. Anyway, the characters of the population clearly disprove such a provenance.
It remains to find an explanation for the existence of this population, and the question of how to deal with it nomenclaturally. Its characters are sufficiently distinctive, toregard it as a moderately differentiated subspecies, less distinctive than some, but more
distinctive than others (sumbae, particeps). I can think of three possible explanations,each of which will be further discussed below: (1) The species was introduced from amainland locality where an as yet undescribed subspecies occurs. (2) The species wasintroduced from the mainland, but has undergone such rapid change since its introduction, that within a century or even less it has differentiated so much from its ancestralform, that it is now a separate subspecies. (3) The occurrence on at least one of the islands is not due to recent introduction by man, but is natural.
Explanation (1) seems unlikely, as the distribution and geographical variation of L.
punctulata throughout its range is now well-known; moreover, it is reasonable to assume
that an 18th century introduction would originate from an old-established coastal trading town, not from some isolated inland area, where an endemic subspecies with a limited range might have been overlooked. Explanation (2) is not one I like, but it has to beconsidered seriously as, significantly in this very same genus Lonchura, Diamond (1972:411), discussing L. spectabilis gajduseki, a subpecies with cinnamon underparts, apparently confined to the Karimui Basin in New Guinea, came with the following conclusion:“Either this race must have slowly evolved in some part of the basin not yet discovered– an unattractive possibility because of the circumscribed size of the basin and the large
areas of grassland necessary for L. spectabilis – or else it must have evolved in a very shorttime, perhaps as little as 15 years”. In support of this last hypothesis, Diamond mentioned studies of introduced Passer domesticus populations in North America, whichwere supposed to have undergone morphological changes in a very short time. Since
Diamond wrote, however, outside the Karimui Basin, a population of L. spectabilis has
been discovered which is even deeper cinnamon on the underparts than gajduseki; it wasnamed L. s. sepikensis, and is found in the Sepik Plains (cf. Jonkers & Roersma, 1990). This
makes it likely that when L. s. gajduseki colonized the Karimui Basin, it already had thecharacters which Diamond believed had been developed in such an extremely shorttime; therefore it is a case of explanation (1), not explanation (2) as Diamond thought.
There remains to be discussed explanation (3); one could call it an explanation by elimination. Whether it is correct I cannot say, but whichever explanation is the right one, by
current standards the Mascarene birds are sufficiently distinct to be named as a separate
subspecies. It is a conclusion I have arrived at only after long hesitation.

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The paper by Restall (1992) with the description of a new subspecies, L. p. holmesi,from Borneo (Pontianak and/or Banjarmasin) became available only some time after
the preceding notes were completed. At first sight it might seem to contradict my conclusion that the discovery of new subspecies from well-known coastal regions is unlikely. But in spite of the table of characters of the various subspecies presented by Restall, I consider it likely that the species is a recent introduction in Borneo (certainly near
Pontianak, where Coomans de Ruiter would not have missed it, if it had been present
in the 1930s). The description of holmesi does not separate it convincingly from nisoria.
The colour descriptions given by Restall are not directly comparable with mine. For
example, he describes the upper tail-coverts of blasii as “Yellow straw”, similar to mydescription of those parts, but then has them “Warm olive” in sumbae, when in my opinion the upper tail-coverts of blasii and sumbae are identical. Interesting, but difficult tointerpret, is Restall’s statement that there is a clear sexual difference in tail-length, fornone of the subspecies examined by me show such a difference. More directly relevant:the tail-lengths given, 30 and 35 mm, indicate that holmesi is a small subspecies (likenisoria), and not close to insulicola.
Without mentioning his description published three years previously, Restall (1995)
again described holmesi as a new subspecies, this time diagnosing it as more or less intermediate between nisoria and baweana. Since I regard baweana as a synonym of nisoria,
this supports my opinion that holmesi does not differ from nisoria. No type specimen
was indicated in the 1992 description, which was based on live cage birds. In the 1995
description, three specimens in the American Museum of Natural History are indicatedas types (syntypes), collected at Semitau in western Borneo. There is no indication thatthese specimens were skinned from captivity and no dates are given. Therefore it is
impossible to ascertain from the evidence presented, whether they had been seen by
Restall before he published his 1992 decription.
Measurements:
Flores (L. p. blasii)

16 
12 

Timor (L. p. blasii)

Sumba (L. p. sumbae)

2
2
(?)
5
6

Southern Sulawesi (L. p. particeps)

3

Sumatra (L. p. nisoria)

2
8
6

Java (L. p. nisoria)

29 

wing
47.5-54(50.4)
49-52.5(50.2)
52, 5250, 5052
49-51(50.2)
49-51(50.0)
51-54(52.7)
50, 5250-54(52.8)
50-53(51.6)
50-54(52.0)

tail32-36(33.5)
30-37(33.4)
32, 3632, 3233.5
34-38(35.8)
29-39(34.8)
33-40(36.7)
30.5, 38
33.5-39(36.6)
33-37(34.9)
30-40(34.9)

tarsus13-14(13.6)
13-14.3(13.7)
13.5, 14
13.6, 1414.4
12.8-14(13.6)
13-14(13.5)
13.8-15(14.5)
13.4, 1413.2-15(14.4)
13.7-14.7(14.1)
13-15(14.0)

culmen10.2-11.2(10.8)
10-11(10.7)
10.8, 11.2
10.6, 10.9
11.0
10-10.8(10.2)
9.6-11(10.2)
10.8-12(11.3)
11.2, 11.0
11.0-12.2(11.7)
11-12.8(11.9)
10.2-12(11.2)

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)22 
Bali (L. p. nisoria)
Bawean (L. p. nisoria)
Lombok (L. p. nisoria)
Northern Sulawesi (L. p. nisoria)
Sri Lanka (L. p. punctulata)

Taiwan (L. p. topela)




(?)
 im.
(?)
5
2
3
9
3
7 (?)

Hainan (L. p. topela)
Yunnan (L. p. topela)

Annam (L. p. topela)

Cochinchina (L. p. topela)
Thailand (L. p. topela)

2 (?)
2

(?)

5

12 
7

Burma, Moulmein (L. p. topela)
Burma, Naga Hills (L. p. topela)
Luzon (L. p. cabanisi)



3
3

Réunion (L. p. insulicola)(?)
5

Seychelles, Mahé (L. p. insulicola)


(?)

50-55
(52.7)
52
53
51
53
53
52
49-51
(50.1)
56, 5654-56(55.0)
53-57(54.5)
54-55(54.7)
53-55(54.4)
55, 5555, 55
55
53
56
54-56
(54.7)
55
53-56(55.0)
54-56(54.7)
55
55
49-50(49.7)
49-50(49.5)
50
54-56(54.9)
53.5
54207
30.5-40(34.8)
35
35.5
35
31
33
32.5
34-37
(35.7)

13.5-15.5(14.3)
14.2
14.2
14.2
14.2
15
14
13.3-14.7(14.0)

10.4-11.8(11.1)
11.0
11.8
12
12.0
11.6
11.0
10.9-11.1(11.0)

32-39

14.9-1536-40(38.0)
36-38(37.0)
31-39(34.1)
33, 3639, 39
39.5
38
37.5
36-42
(38.1)
32
36-43.5(38.6)
35-40(37.9)
3731+
34-37(35.2)
30-36(33.3)

13.5-15(14.3)
14.2-14.8(14.6)
13.7-14.8(14.1)
14.2, 14––14.4

12.0-13.0(12.4)
11.6-12.8(12.2)
12.0-12.1(12.0)
11.5-12(11.8)
12.0, 12.0
11.8, 12
11.9
12.1

14.2
14-15.2(14.7)
14.2-15(14.6)
14.5
–
–
–
–
–

11.2-12.0(11.6)
10.4
11.9-13.1(12.2)
11.5-12.8(12.2)
11.0
12.0
11.3-12.3(11.6)
11.0-11.7(11.3)

35-40(37.5)
33.5
38

13.6-15(14.4)
14
14

11.6-12(11.9)
12.0
11.0

–

Lonchura quinticolor (Vieillot)
Loxia quinticolor Vieillot, 1807, Ois. Chant.: 85, pl. 54 – les Îles Moluques = Timor, designated by Hellmayr (1914: 59) (reference not verified).
Lonchura quinticolor sumbae Restall, 1995, Bull. Brit. Orn. Cl. 115: 143 – Waingapo, Sumba.
Collectors.— Allen, Everett, Rensch (4), de Jong (2), Verheijen (3, MCZ), Verheijen/Schmutz.
Material.— 14 , 11 , 4 (?) from diverse localities in Manggarai (Verheijen/Schmutz, RMNH nos.
65574, 65575, 66227-66245, 81356, 81369, 81429, 81432, 85378-85380, 85387).

208

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Eggs: c/6, 19.v.1951, Bénténg Djawa (RMNH no. 71716); c/6, iii.1952, Bénténg Djawa (RMNH no. 71717);
c/5, 23.ii.1953, Mano (RMNH no. 71718); c/2, 4.vi.1954, Potjong (RMNH no. 71719); c/5, 12.vi.1956,
Wangkung, Ruteng (RMNH no. 71720); c/2, iv.1958, Mukun (RMNH no. 71721); c/3, 8.iii.1962, Mataloko (RMNH no. 71722); c/5, 18.vi.1962, Mataloko (RMNH no. 71723). The eggs are white, without
gloss.
Some measurements and weights: RMNH no. 71716

RMNH no. 71718

RMNH no. 71719

16.1 × 12.3
16.3 × 12.1
16.9 × 11.8
17.2 × 11.8
17.3 × 12.0
17.9 × 11.6
16.3 × 11.1
16.6 × 11.4
16.8 × 11.8
17.2 × 11.6
17.5 × 11.8
15.7 × 11.3
16.6 × 11.2

0.07030.07040.06910.071
0.07160.071
0.06720.06890.07250.07440.06980.0666
0.0649Notes.— This is one of the larger members of its genus. There does not seem to be
any difference, either in plumage or in measurements, between the sexes.The name L.
quinticolor sumbae, recently proposed by Restall (1995), is invalid, being a primary homonym of L. punctulata sumbae. I do not consider, however, that a replacement name isrequired, for two reasons. The first is that no other author has found geographical variation in this species, and that the existence of geographical variation worthy of expression in nomenclature requires confirmation. The second, more fundamental reason isthat if a division into three subspecies, with ranges as outlined by Restall, is accepted,
the type locality of the species should be established first. The species was describedfrom a French collection, and its designated type locality is Timor. This makes it overwhelmingly likely that the type locality of the species should be restricted to Koepang,western Timor, so the subspecies called ”sumbae” by Restall, actually is the nominate
race. Until these points have been cleared up, the application of trinomials in L. quinticolor can only cause confusion.
Measurements:
10 
10 

wing
54-58
(55.8)
54-58(56.0)

tail34-37½(36.4)
34-38(35.8)

tarsus16-17(16.4)
15.6-16.7(16.1)

culmen12.8-14(13.3)
12.8-13.8(13.2)

Lonchura pallida pallida (Wallace)
Munia pallida Wallace, 1864, Proc. Zool. Soc. Lond. (1863): 486, 495 – Lombock and Flores.
Collectors.— Allen (1?), Verheijen (1, MCZ).
Material.— , 1862, Flores (Allen, BM no. 73.5.12.1356), “bill and feet light blue, eyes dark” (original
notes by Allen).
Eggs: c/1, 4.vi.1954, Potjong (RMNH no. 71724); c/6, 29.v.1959, Léong (RMNH no. 71725). The eggs arewhite.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)
Measurements and weights:

RMNH no. 71725

209
16.0 × 11.0
16.1 × 11.1
16.2 × 10.8
16.2 × 11.1
16.3 × 11.0
16.3 × 11.1

0.0605
0.06180.05810.05970.06040.0608Notes.— This is apparently an uncommon species on Flores, as only Allen and Verheijen have obtained it. Rensch (1931a: 597) claims field-observations at Aimere (0-200m) and Geli Moetoe (1000 m), and Schmutz (MS) observed it at Kenari and Orong. In
the Catalogue of Birds (Sharpe, 1890: 346), 5 Wallace-specimens from Lombok are listed,but only one (f.  ad.) from Flores, the specimen examined by me.
Hitherto, Wallace was believed to have been the discoverer of L. pallida (on Lombok,
in 1856), but Forsten’s Bima collection contains a mounted specimen, labelled Loxia auriventer Temm. n. sp.  (RMNH cat. no. 1): as in so many cases, the bird arrived in Leiden, was recognized as new and provided with a manuscript-name, but remained undescribed.
Stresemann (1940a: 38) mentioned as first collector of this species in Sulawesi(Celebes), C. Platen, 1878. In our collection are two mounted specimens (RMNH cat.
nos. 2 and 3), labelled “Macassar”, collected by Teysmann in 1877.
Measurements:


wing
55

tail
32½

tarsus16

culmen13

Passer montanus malaccensis Dubois
Passer montanus Var. Malaccensis Dubois, 1885, Faun. Ill. Vert. Belg., Ois. 1: 572 – la presqu’île de Malacca
et... Java.
Collectors.— Verheijen, Schmutz.
Material.— , 29.x.1969, Ruteng (Verheijen, RMNH no. 65565);  without data, West Flores (Schmutz,
RMNH no. 81430).
Eggs: c/5, c/5, c/5, 18.vi.1963, Réo (Verheijen, RMNH nos. 71759, 71760, 71761).

Notes.— According to Schmutz (MS), this species has been established on Flores
since about 1955. The subspecific identity has been assumed on the basis of geographical probability (cf. Keve, 1978); I have made no comparisons.
Measurements:



wing
70
69

tail
50
48.5

tarsus
16.7
16.3

entire culmen
13
13

exposed culmen
10.2
10Aplonis minor minor (Bonaparte)
L[amprotornis] minor Bonaparte, 1850, Consp. Gen. Av. 1: 417 – Timor.
Collectors.— Allen, Rensch (4), Verheijen/Schmutz. In the Catalogue of Birds (Sharpe, 1890: 143) only
one juvenile specimen (cat. g) from the Wallace Collection is listed.
Material.— 2 , (?) juv., 19.ix.1970, Kisol (Verheijen, RMNH nos. 65371, 65372, 65373); , 21.ix.1970,

210

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Kisol (Verheijen, RMNH no. 65375); , 1.v.1971; Ngalor-Rogo, Langkas, Rahong (Verheijen, RMNH no.
85303); , 25.ix.1972, Manggarai (Schmutz, RMNH no. 85304); ,  juv., 28.viii.1976, Poco Nernancang(Verheijen, RMNH nos. 85305, 81302); , 2.ix.1976, Poco Nernancang (Verheijen, RMNH no. 85306); ,, 4.ix.1976, Poco Nernancang (Verheijen, RMNH nos. 85307, 85308). Iris bright red, bill and legs black.
No moult.
Eggs (fig. 17b): c/3, 26.x.1960, Mataloko (RMNH no. 71763); c/1, 8.vi.1961, Mataloko (RMNH no. 71764);
c/3, 13.vii.1961, Mataloko (RMNH no. 71765); c/3, 29.ix.1961, Mataloko (RMNH no. 76586); c/2, 19.
v.1962, Mataloko (RMNH no. 71766); c/3, 2.vi.1962, Mataloko (RMNH no. 71767); c/2, c/2, 7.vi.1962,
Mataloko (RMNH nos. 71768, 71769); c/3, 13.vi.1962, Mataloko (RMNH no. 71770); c/4, 18.vi.1962,
Mataloko (RMNH no. 71771); c/2, c/2, 18.vi.1962, Mataloko (RMNH nos. 71772, 71773). The eggs are
pale blue, moderately mottled with light brown. They are indistinguishable, in colour as well as inmeasurements and weights, from the eggs of A. panayensis strigata.
Some measurements and weights: RMNH no. 71763

RMNH no. 71765

RMNH no. 71766
RMNH no. 71767

RMNH no. 71768

23.2 × 18.5
24.0 × 19.2
24.0 × 19.4
23.7 × 18.8
24.1 × 18.3
24.2 × 18.5
24.8 × 18.5
25.1 × 18.3
24.1 × 18.5
24.6 × 18.1
25.2 × 18.3
22.9 × 17.5
24.7 × 17.7

0.26040.24200.27000.26370.26020.26480.26600.26830.25780.25050.26740.22090.2221

Notes.— The peak of the breeding-season is apparently in June. This is worth recording, as in Java the species is believed to be a non-breeding visitor, an east-west
migrant from the Lesser Sunda Islands, and: “In Java the birds do appear in the end of
May and leave again in September” (Van Bemmel, in Van Bemmel & Voous, 1951: 36;
see also Hoogerwerf, 1965: 287). This would mean that the breeding-season in the Lesser Sunda Islands coincides completely with the presumed migration to Java, a matterthat will require a lot of explanation before it can be accepted. The status of A. minor inJava certainly deserves further investigation. The specimens examined by me range in
dates from 24 April to 24 July, rather earlier than as given by Van Bemmel. All nine birds(with one possible exception, of which I am not sure) are in heavy moult. All records arefrom high elevations (850-2000 m). On the Lesser Sunda Islands, A. minor is not in thefirst place a mountain bird, ranging from sea-level to perhaps 1500 m. As far as I amaware, Rensch (1928b) was the first to record the species from Java (“eine große Serie...
vom Berge Tjerimai”); the first to collect it in Java probably was H. W. van der Weele(four specimens from Tirtasari, West Java, April/May 1910). Dr Somadikarta (in litt.,
11.iii.1988) has provided me with a list of the MZB-material, on which van Bemmel’sdata were based: 1, June 1928, G. Tjerimai, West Java; 1, July 1939, Tengger, East Java; 1,
May, 3, June and 1, August 1941, Bandjarwangi, Tjikadjang, West Java; 1, August 1941,
G. Lawoe, Middle Java. This shows that the species may be found in all parts of Java.
White & Bruce (1986: 390) do not recognise subspecies, but I would hesitate to accept their conclusion without further investigation. Absence of geographical variation
would seem to contradict Deignan’s opinion (1955) that this is a relict species, in theMees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

211

process of being replaced by its close relative A. panayensis wherever the two come into
contact. On the other hand, the occurrence of the peculiar endemic subspecies A. panayensis gusti on Bali, and of other well-marked subspecies on the West Sumatra islands,
the Nicobars and the Andamans, indicates that A. panayensis is certainly not a recent
colonist in these regions. This makes it less likely that Deignan was right in believing A.
panayensis to be an expanding species. To verify Deignan’s statement that A. p. strigata
is “a larger and stronger race” than A. minor, which was at the basis of his theory thatthe former is replacing the latter, I measured some specimens of A. p. strigata from Javaand of A. p. gusti from Bali. The measurements show that the differences are negligible.
Actually, A. minor is a little longer-winged than A. p. strigata, but the latter exceeds A.
minor a trifle in length of the tail and the bill. Deignan’s speculative, and therefore stimulating, paper has made clear how much more research is needed, before the relationships between the closely related species (or subspecies?) A. panayensis and A. minor
may be understood. Mayr (1944a: 143) regarded A. metallica as the closest relative of A.
minor, but his opinion has not been shared by later authors.
The table of measurements suggests that specimens of A. minor from Flores are
smaller than specimens from Timor, and agree with birds from Sumba, the small size of
which had been noted previously (cf. Hellmayr, 1914: 43; Mayr, 1944a: 143). But the series are short and, as Mayr observed, the difference is not sufficient for subspecific separation. It also deserves notice that birds from Java average a little larger than birds from
Flores, providing further evidence that they do not belong to the same population.
Of the six RMNH specimens from Timor, one  was collected by H.A. Lorentz in
1909; the other five are the syntypes of A. minor: four of these have been collected by S.
Müller, in 1828/1829, and the fifth one (the unsexed bird) by F. Péron in 1801/1803.
Measurements:
Flores (A. minor)
4
5
Timor (A. minor)4(?)
Binongko (A. minor)Java (A. minor)
4

wing

tail

tarsusentire culmen

exposed culmen

96-100(98.0)
92-98(95.3)

51-58(54.3)
51-55½(53.5)

19-21(20.1)
19.2-21(20.0)

18.2-20.4(19.4)
18-20.4(19.2)

13.2-15(14.3)
13.7–15(14.4)

101-107(103.5)105
106

56-60½(58.6)
55
59

20-21(20.7)
21.4
20

19.2-21(20.1)
20.1
20.2

13.6-15.4(14.5)
14.6
15.2

100

59

20.4

19

14.5

55-60
(57.8)
49-57(52.5)

19.8-21(20.5)
20.6-21.6(21.2)

18.6-20(19.2)
19-21(20.0)

14.2-15(14.7)
14.4-16(15.2)

58-62½(60.0)

20.9-22.7(21.5)

21-24(22.3)

15-18.2(16.6)

56, 5751

20.3, 21.2
21.2

20.0, 20.0
19

14.9, 15.0
1498-102(100.3)5
97-102(98.6)
Java (A. panayensis strigata)10 
95-101(97.7)
Bali (A. panayensis gusti)
2
99, 102
-

212

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Gracula religiosa venerata BonaparteG[racula] venerata Bonaparte, 1850, Consp. Gen. Av. I: 422 – ex Sumbava.
Gracula venerata mertensi Rensch, 1928, Orn. Mber. 36: 48 – Sita (6-800 m) Westflores.
Collectors.— Semmelink, Allen, Martens (1), Weber (1, ZMA), ten Kate, Everett, Endih (2), Rensch (5),de Jong (2), Verheijen (1, MCZ), Verheijen/Schmutz.
Material.— (?), 1862, Larantoeka (Semmelink, RMNH cat. no. 4); (?), 26/28.xi.1888, Bari (Weber, ZMA
no.29985); 2 (?) ad., 24/27.vi.1891, Koting (Ten Kate, RMNH cat. nos. 5 and 6); , 15.i.1969, Nunang, 650
m (Schmutz, RMNH no. 85124); , vii/viii.1969, Nunang, 650 m (Schmutz, RMNH no. 85126); , 9.
vii.1971, Narang, Todo, 150 m (Verheijen, RMNH no. 85127); , , 12.x.1971, Nunang, 650 m (Schmutz,
RMNH nos. 85125, 85123).
Eggs (fig. 17c): c/2, 3.xi.1960, Soa (RMNH no. 71774); c/1, ca. 11.xii.1960, Soa (RMNH no. 71775); c/1,
x.1961, Soa (RMNH no. 71776); c/2, 20.ii.1962, Mataloko (RMNH no. 71777); c/3, 14.iii.1962, Mataloko(RMNH no. 71778). The eggs are sky-blue, moderately glossy, somewhat sparsely mottled with brown;some eggs almost plain.
Measurements and weights:

RMNH no. 71774
RMNH no. 71775
RMNH no. 71776
RMNH no. 71778

37.4 × 26.5
39.1 × 26.8
41.8 × 27.0
40.3 × 27.3
37.3 × 26.5
38.3 × 26.3
38.5 × 24.8

- very large blow-holes
1.01840.98380.86780.9440
damagedNotes.— Modest as the available material is, it supports the conclusion previously
reached by White & Bruce (1986: 393) that mertensi is not tenable, either on the basis ofmeasurements, or on supposed differences in gloss of the plumage.
Measurements:
Flores3
2
4 (?)
Sumbawa

Alor



(?)


wing
161, 176,
180
166, 173
170, 171172, 173
168
171
169
170

tail
73, 77tarsus41, 41entire culmen
42, 38

exposed culmen
31.5, 33

77, 79½
72, 72,75, 8369
73
74
73

38, 3943, 40, 42,
42.5
41
38
39
41

40, 37.4
41, 40, 39,
38
40
42
41.3
40

33, 31.2
34, 32, 35,
32
32
33.2
32.5
32.5

Oriolus chinensis broderipi BonaparteO[riolus] broderipi Bonaparte, 1850, Consp. Gen. Av. I: 348 – Ins. Sumbava.
Oriolus sumbawensis Schlegel, 1857, Handl. Dierk. 1: 479, Vogelen pl. II fig. 20 – Sumbawa.
Collectors.— Semmelink, Allen, Weber, ten Kate (2), Rensch (3), de Jong (7), Verheijen (1, MCZ), Verheijen/Schmutz.
Material.— (?) ad., 1862, Larantoeka (Semmelink, RMNH cat. no. 6); (?) im., 1862, Larantoeka (Semmelink, RMNH cat. no. 7); (?), 26.xi.1888, Bari (Weber, collector’s no. 494a, RMNH); , 23.xi.1968, Nunang (Schmutz, RMNH no. 97119); , 17.xii.1968, Nunang (Schmutz, RMNH no. 66140); (?) nestling,
23.xii.1969, Todo (Verheijen, RMNH no. 65349); , 1969, Nunang, 650 m (Schmutz, RMNH no. 85164);

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

213

, 19.ix.1970, Kisol (Verheijen, RMNH no. 65350); 2 , 5.iii.1971, Rana Loba, Borong, 50 m (Verheijen,
RMNH nos. 66141, 85161); , 12.x.1971, Nunang, 650 m (Schmutz, RMNH no. 85163); , 25.x.1971,
Nunang, 650 m (Schmutz, RMNH no. 85162); (?) juv., 26.xi.1975, Nunang, 650 m (Schmutz, RMNH no.
81083); 3 , , 28.xi.1975, Nunang, 650 m (Schmutz, RMNH nos. 81079-81082); , 18.xii.1975, Nunang,
650 m (Schmutz, RMNH no. 81078). Adults of both sexes have the iris blood-red, bill dark pink, legs
black. Immatures: iris brown, bill dusky (gradually changing to dark pink), legs black.
Eggs (fig. 17d): c/2, 12.x.1955, Ruteng (RMNH no. 71779); c/1, 17.v.1957, Ruteng (RMNH no. 71780);
c/1, 4.vi.1959, Potjong (RMNH no. 71781); c/2, 20.vi.1959, Poéng (RMNH no. 71782); c/3, 5.xi.1959, locality unknown (RMNH no. 71783); c/1, 17.vi.1962, Mataloko (RMNH no. 71784). The eggs are glossy
white or very pale pinkish chamois, with widely spaced, sharply defined black dots.
Measurements and weights:

RMNH no. 71779

31.5 × 23.4
very large holes,
33.0 × 23.4
not weighed
RMNH no. 71780
33.6 × 24.2
0.5233
RMNH no. 71781
32.2 × 22.70.5007
RMNH no. 71782
33.7 × 23.5
0.548233.8 × 23.5
0.5425RMNH no. 71783
36.9 × 25.4
0.655137.7 × 25.1
0.677438.5 × 25.2
0.6744
These eggs average only slightly larger, but are conspicuously heavier, than eggs of O. c. maculatus from
Java, as recorded by Hellebrekers & Hoogerwerf (1967: 160). Clutch no. 71783 is particularly large andheavy.

Notes.— Greenway (1962: 130) cites the name as broderipii (note the difference inspelling), from Bonaparte (1852), but the description in the Conspectus has clear priority. Both descriptions are, of course, based on the same specimen.
Büttikofer (1892: 194) recorded 2  ad. from Koting, collected by Ten Kate. These are
no longer present in the collection, but there is an unsexed specimen, received at thesame time from the same collector, from Trong, Adonara (where Ten Kate stayed from
13/16.v.1891). I do not know why Büttikofer omitted this specimen (which has an original label) from his paper. The species had not yet been recorded from Adonara, although its occurrence was to be expected.
As it seems to have been generally overlooked, I list here the objective synonym
Oriolus sumbawensis Schlegel. The name broderipi is based on two mounted specimensfrom Bima, Sumbawa (Forsten coll., RMNH cat. nos. 1, 2; cf. Schlegel, 1867: 106), bothhaving written underneath on the socle, in Temminck’s handwriting: Oriolus Sumbawanus Temm sp. Nov. The specimen figured by Schlegel appears to be cat. no. 2, which
is marked as a  (the other one has no indication of its sex), and is the deeper coloured
one with an orange-yellow mantle.
Unexpectedly, there is no difference in plumage between fully adult males and females: they have a deep cadmium-yellow, almost orange-yellow plumage, and theremiges are glossy black. Duller birds, the yellow of the dorsal surface tinged withgreen, and the inner remiges with greenish edges, are immature, further, in the skins,characterized by wholly or partly dusky bills. These immature birds may already havewell-developed gonads. This subspecies is larger than the adjacent O. c. maculatus (asnoted above, the eggs are also larger). The material suggests a slight sexual differencein size, the males averaging a little larger than the females.

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Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Measurements:
7
5

wing
152-161(157.4)150-155(152.6)tail
101-108(105.8)98-105(102.3)tarsus
27.5-29.2(28.5)
27-28.2(27.7)

entire culmen
36-39.3(38.0)
37.3-39.2(38.4)

exposed culmen
32.5-36(34.6)
33.5-35.4(34.6)

Dicrurus hottentottus bimaensis Wallace
Dicrourus bimaënsis Wallace, 1864, Proc. Zool. Lond. (1863): 492 – Lombock, Sumbawa, and Flores.
[Dicrourus] bimaensis Bonaparte, 1850, Consp. Gen. Av. 1: 352 – nomen nudum (Sumbava).
Dicrurus hottentottus renschi Vaurie, 1949, Bull. Amer. Mus. Nat. Hist. 93: 298 – Tambora Mountain, 3000
feet, Sumbawa.
Collectors.— Semmelink, Allen, Weber (3), Everett (6), Rensch (3), de Jong (5), Pfeffer (1), Verheijen/
Schmutz.
Material.— (?), 1862, Larantoeka (Semmelink, RMNH cat. no. 4); (?), 23/25.xi.1888, Reo (Weber, RMNH
cat. no. 5); , 24.x.1968, Nunang (Schmutz, RMNH no. 66142); (?), 3.iii.1971, Rana Loba, Borong (Verheijen, RMNH no. 66143); (?), 25.v.1971, Manggarai (Verheijen, RMNH no. 66144); , 17.vi.1973, WaéRéca, Borong (Verheijen, RMNH no. 85284); , 24.vi.1976, Poco Mulu (Schmutz, RMNH no. 97180).
Eggs (figs 17e, 17ee): 18 clutches of c/1 (2 ×), c/2 (8 ×), c/3 (5 ×) and c/4 (3 ×), collected in the months
October (12), November (5) and December (1). There are two quite different types of egg: one with lightbrown markings on a cream ground, the other with a pure white ground colour, sparsely marked withsmall black primary spots, and somewhat larger light violet-grey secondary spots. There is no need toquestion the identification of the eggs, as this kind of variation is known to occur in the Dicruridae.
Some measurements and weights: RMNH no. 71785

RMNH no. 71786
RMNH no. 71787
RMNH no. 71788
RMNH no. 71789

RMNH no. 71790

RMNH no. 71791

RMNH no. 71792
RMNH no. 71793
RMNH no. 71794

30.2 × 21.7
30.3 × 21.2
31.3 × 21.3
29.0 × 21.0
28.3 × 21.0
28.7 × 21.0
29.4 × 21.3
29.6 × 21.4
29.0 × 20.5
29.3 × 20.6
29.6 × 20.6
29.8 × 20.8
29.0 × 20.5
30.4 × 20.5
30.5 × 20.3
27.3 × 20.5
27.6 × 20.6
28.6 × 20.8
31.7 × 20.8
31.8 × 20.7
27.8 × 20.4
27.9 × 20.5
29.2 × 21.3

0.34880.36720.35610.41540.38900.36330.38210.38970.36570.38340.37460.38780.31780.33980.32980.33530.33450.3696

Notes.— Two authors have studied the geographical variation of D. hottentottus in
the Lesser Sunda Islands: first Rensch (1928a, 1931a, 1931b), and later Vaurie (1949).
Rensch (1928a) observed that specimens from Lombok are larger than “typische D. h.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

215

bimaensis (Bp.) von Sumbawa”, and he named the former D. h. vicinus. Rensch believed
birds from Sumbawa and Flores to be identical. Although he first accepted Sumbawa as
the type-locality of bimaensis (as just quoted), he later restricted the type-locality to
Flores (cf. Rensch, 1931a: 589).
Vaurie had an entirely different opinion on the geographical variation in the region,in that he regarded birds from Lombok, Flores, Pantar, Alor and Goenoengapi as consubspecific, and large, and only birds from Sumbawa as small. On the basis of Rensch’s
restriction, he accepted Flores as the type-locality of bimaensis, which name he therefore
used for the birds from all these islands. This left the supposedley smaller subspecies
from Sumbawa without a name, which he provided. Judging by the sequence in which
he listed the islands inhabited by bimaensis (repeated by Vaurie, 1962: 151), Vaurie confused Goenoeng Api, Sangeang, off Sumbawa, with the island Goenoeng Api in the
Banda Sea (where D. hottentottus does not occur, cf. van Bemmel & Hoogerwerf, 1940).
In Vaurie’s classification, specimens from Sangeang should, on the basis of geographical nearness, belong to the Sumbawa subspecies rather than to bimaensis sensu Vaurie.
Since then, I pointed out that there had been no need to shift the type locality of bimaensis, and confirmed the adult specimen from Bima, Sumbawa, listed as type by Finsch(1901b: 196), as lectotype (cf. Mees, 1965: 194-195).
The classification proposed by Vaurie, with the range of one subspecies interrupted
by that of another, is not very satisfactory. Although the material available is not exactly
rich, I have tried to verify whether it conforms to Vaurie’s conclusions as regards variation. In this connection it may be noted that Vaurie’s material was not particularly richeither. Excluding several specimens which, with dull plumage and some white edging
on the middle of the underparts, are clearly juveniles, the material (none of which was
examined by Vaurie), provides the following
Measurements:

wing

LombokCat.10 (?)
140
Cat.11 (?)
146
Cat. 9 (?)
149
Cat. 8 (?)
151
Cat. 7 (?)
152
SumbawaCat. 1 (?)
141(lectotype of bimaensis)
Flores66142 143
97180 143
85284  im.
132
Cat. 5 (?)
13866143 (?)
139
66144 (?)
139
Cat. 4 (?)141*) tip damaged.

tail
(OTF)

tail
(CTF)

tarsusculmenculmenfrom skull from nostril

125
127
118
124
125

116
118
112
107
112

26
26
27
26.3
26

35.5
37
38.2
37
36

25
23.6
25.3
23.8
22.5

106

100

–

33

20.7

110
110
109
106
112
116
111

102
100
104
99
103
107
100

27
27
24
27
25
26
25.8

33.6
33.3
32.9
35
34.8
33.2+
34

22.8
21.5
20.3
22.3
21.0
20.5 *)
21

As far as the evidence provided by this material goes, it clearly supports Renschand not Vaurie: no difference between birds from Sumbawa and Flores, and specimens

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Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

from Lombok distinctly larger, with longer tails. Thus, D. h. vicinus is a valid subspecies,
confined to Lombok, and the chain of islands from Sumbawa to Alor is inhabited by D.
h. bimaensis. In this connection it is worth mentioning that Hoogerwerf (1956) found D.
hottentottus fairly common on Komodo, between Sumbawa and Flores.
Vaurie (1962: 150) cites the original description of D. h. bimaensis as “Dicrorus (sic)bimaënsis Wallace...”. But in the original description the spelling of the generic name is,in both places where it appears (Wallace, 1864: 485, 492), Dicrourus (in those days theusual spelling). The citation is correct in Vaurie’s (1949: 296) earlier publication.
The substantial and complex agglomerate of forms united by Vaurie (1949) under
the specific name of D. hottentottus, is an obvious candidate for subdivision into several
species. An attempt at this has been made by White & Bruce (1986: 314-315), who separated the forms inhabiting the Lesser Sunda Islands under the specific name D. densus.
Bruce concluded the discussion with the statement that a detailed study was in preparation, to be presented later. I prefer to await the publication of this study before accepting the proposed changes.
Artamus leucorhynchus leucorhynchus (Linnaeus)
Lanius leucoryn[chus] Linnaeus, 1771, Mantissa Plant.: 524 – in Manillis.
Artamus leucorrhynchus, var. celebensis Brüggemann, 1876, Abh. Naturwiss. Ver. Bremen 5: 69 – Celebes.
Collectors.— Semmelink, Allen, ten Kate, Everett (2), Rensch (4), Verheijen.
Material.— (?), 1862, Larantoeka (Semmelink, RMNH cat. no. 38); (?) juv., 28/30.i.1891, Endeh (ten Kate,RMNH cat. no. 39); , large gonads, 25.x.1970, Ruteng, captured on its nest (Verheijen, RMNH no.
85224).
Eggs: c/3, 29.x.1958, Lamba, Todo (RMNH no. 71801); c/1, 27.x.1959, Ruteng (RMNH no. 71802); c/1,
14.ix.1961, Ruteng (RMNH no. 71803); c/3, 7.ix.1962, Ruteng (RMNH no. 71804); c/3, 30.viii.1964,
Ruteng (RMNH no. 71805); c/3, c/2, 20.ix.1964, Ruteng (RMNH nos. 71806, 71807); c/3, 18.x.1964,
Ruteng (RMNH no. 71808); c/4, 7.xi.1964, Ruteng (RMNH no. 71809).
Some measurements and weights: RMNH no. 71801

RMNH no.71804

RMNH no. 71806

24.0 × 18.3
24.4 × 18.3
24.5 × 18.3
23.3 × 18.0
23.6 × 17.9
23.9 × 18.5
24.8 × 18.3
25.0 × 18.6
25.1 × 18.2

0.23110.22820.23280.23670.22720.2385- (damaged)
0.26240.2471

Notes.— In their large size, birds from the Lesser Sunda Islands, east to Flores(Alor?) correspond with birds from Sulawesi (Celebes) and the smaller islands in the
Flores Sea; they are larger than birds from Java (cf. measurements published by Mees,
1986: 150), and (abruptly?) larger than birds from Timor and Roti. Therefore, the geographic variation of the species in this region is by no means without interest, eventhough it is difficult to express it in nomenclature.
In a previous publication, I observed that the irregular pattern of variation makes itvery difficult to distinguish subspecies in the whole region from the Andamans to New
Guinea and Australia. An exception must be made for A. l. musschenbroeki from theMees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

217

Tanimbar Islands, which is conspicuously darker in plumage than all the surrounding
populations, and on this basis deserves nomenclatural recognition. Its bill is also rather
larger and thicker than that of surrounding populations.
Measurements:


wing
145

tail
65

tarsus17.6

entire culmen
24

exposed culmen
19.8

Corvus florensis Büttikofer
Corvus florensis Büttikofer, 1894, in Weber: Zool. Ergebnisse 3: 304 – Maumeri, Flores.
Collectors.— Weber, Everett (1), Verheijen (2, MCZ), Verheijen/Schmutz.
Material.— (?) ad., xii.1888, Maumeri (Weber, RMNH cat. no. 1, holotype); (?), 19.vii.1969, Sok-Rutung,West-Manggarai (Schmutz, RMNH no. 66247); , small gonads, 20.vii.1969, Sokrutung, 50 m (Schmutz,
RMNH no. 97150);  with large gonads, 25.ix.1971, Sésok, 1000 m (Schmutz, RMNH no. 85140).
Eggs (fig. 18a): c/3, 28.xii.1946, Rekas (RMNH no. 71816); c/3, of which two badly damaged, 1.i.1953,
Waé Rambung (RMNH no. 71817); c/3, 28.ix.1955, (RMNH no. 73524); c/2, xii.1955, Bénténg Djawa,
Djinggor (RMNH no. 71818); c/3, 27.xi.1956, W. Nandá (RMNH no. 71819); c/2, xii.1958, Heret (RMNH
no. 71820); c/2, 5.xi.1959, Djinggor (RMNH no. 71821); c/1, 1.xii.1959, Djinggor (RMNH no. 71822); c/2,
i.1962, Soa (RMNH no. 71823).
Measurements and weights:

RMNH no. 71816

RMNH no. 71817

RMNH no. 73524

RMNH no. 71818
RMNH no. 71819

RMNH no. 71820
RMNH no. 71821
RMNH no. 71822
RMNH no. 71823

38.0 × 27.4
39.8 × 28.3
41.7 × 28.6
39.3 × 27.0
42.6 × 28.1
42.9 × 27.8
38.8 × 26.7
39.1 × 28.4
42.7 × 28.6
34.7 × 27.4
36.2 × 27.6
40.5 × 29.1
41.2 × 27.6
42.2 × 27.1
38.0 × 26.8
40.5 × 26.3
39.2 × 26.5
39.8 × 26.0
43.2 × 28.0
42.7 × 28.1
43.2 × 27.7

0.83670.89180.91921.003
0.82580.93381.000
0.81960.826
0.86970.87490.83390.828
0.83370.8436
0.7963
1.09410.8931
0.9623

Notes.— There has been speculation about the affinities of this little-known species,
with the suggestion that it might be a geographical representative of C. enca (cf. Meinertzhagen, 1926: 73). Rensch (1931a: 588) enumerated morphological differences, which
point away from a close relationship between these two species. Vaurie (1958) agreed
with Meinertzhagen in considering C. florensis a derivative of C. enca, but he drew attention to the high tail/wing ratio (72.5%) of his one specimen. The four specimens measured by me confirm this (70.1-73.4%). Jollie (1978: 98-99) regarded C. florensis as a remnant of an early radiation and called it: “a fine ancestral type”, but he did not further

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Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

elaborate this, and as it stands, this remark is not very useful. That is a pity, for his opinion that C. florensis is an ancestral type, is the opposite of the opinions of Meinertzhagen
and Vaurie, that it is a derivative of the C. enca group. These conflicting opinions showthat the relationships of C. florensis remain obscure. The statement by Haffer (in Glutz
von Blotzheim, 1993: 1657): “C. florensis auf Flores ist ein vertreter von C. enca” is far too
definite and is presented without any explanation, although probably based on Vaurie.
The eggs (previously undescribed) support the separate position of C. florensis: theground-colour is white, unlike the eggs of C. enca, C. macrorhynchus and the Australian
species, which have eggs with a blue-green ground-colour. One has to go as far as Africa to find an other member of the genus with eggs which lack green: C. capensis. The
eggs of C. capensis look, however, quite different from those of C. florensis, as the former
has the ground-colour almost concealed by large brown freckles. Schönwetter’s (1983:
696) description gives the impression that Corvus enca celebensis (a synonym of C. e. enca,cf. Oortwijn, 1987) would normally have eggs with a whitish ground-colour. That is not
so, all eggs of C. enca examined by me (subspecies enca and compilator) have a bluishgreen ground-colour.
C. enca violaceus from Ceram, which is more likely a separate species, may have eggs
with a white ground-colour, but there was some doubt about the identification (Stresemann, 1914a: 153; Schönwetter, 1983: 721). The green ground-colour of crows’ eggs
fades with age, and some of the very pale eggs described by Schönwetter were old, andhad probably been exposed to sunlight.
Measurements:


(?)
(?)

wing
227
224
226
228

tail
166
157
166
160

tarsus48
46
46
49

bill (from forehead feathers)
51½
50
51
50

Corvus macrorhynchos macrorhynchos Wagler
C[orvus] macrorhynchos Wagler, 1827, Syst. Av., Corvus, sp. 3 – Nova-Hollandia, Nova Guinea, et in insulis Sumatra et Java = Java.
Cornix timorensis Bonaparte, 1853, Compt. Rend. Acad. Sci. Paris 37: 829 – Timor (reference not verified).
Corvus coronoides inoptatus Rensch, 1928, Orn. Mber. 36: 7 – Rana Mesé, Flores.
Collectors.— Allen, Weber (1), Everett (2), Rensch (5), de Jong (1), Verheijen (1, MCZ), Verheijen/
Schmutz.
Material. –(?) ad., 23/27.xii.1888, Endeh (Weber, RMNH cat. no. 15);  im., 23.i.1968, Pongkor (Verheijen, RMNH no. 65375); (?), 8.v.1969, Nisar (Schmutz, RMNH no. 66246); , 6.viii.1969, Dalo, Rahong(Verheijen, RMNH no. 65376); (?), 16.viii.1969, Dalo (Verheijen, RMNH no. 85141).
Eggs (fig. 18b): c/2 28.xi.1955, Ruteng (RMNH no. 71810); c/5, 14.xi.1957, Mano (RMNH no. 71811); c/3,
1958, Lamba, Todo (RMNH no. 71812); c/5, 30.xi.1958, Potjong (RMNH no. 71813); c/4, 20.xii.1958,
Wesang (RMNH no. 71814); c/3, 16.xi.1960, Soa (RMNH no. 71815).
Measurements and weights:

RMNH no. 71814

41.2 × 29.0
42.3 × 29.2
43.5 × 28.8
43.9 × 28.8

1.1265
1.23461.24201.1422

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

219

9. Unconfirmed and erroneous records
Threskiornis melanocephalus (Latham)
The inclusion of Flores and Timor in the range of this species on the distributional
map in Hancock et al. (1992: 218) is an error, not supported by their text. On the other
hand, East Java has mistakenly been excluded on the map, for the Brantas Delta has
been known as a breeding place for half a century and in their text, the authors report
recent breeding in that area.
Circus assimilis Jardine & Selby
Marchant & Higgins (1993: 95-96) list this harrier as occurring in the Lesser Sunda
Islands, which, as a generalisation, is acceptable, but it has been recorded there from
Sumba and Timor only, so that their distributional map showing it as ranging over thewhole chain of islands, from Lombok to Wetar, is definitely in error.
Falco longipennis longipennis Swainson
As has been mentioned on a previous page, the record of nominate F. l. longipennis
from Flores (Stresemann & Amadon, 1979: 417) was a mistake.
Arenaria interpres interpres (Linnaeus)
White & Bruce (1986: 167) report this migrant wader from Flores on the basis of: “anoverlooked specimen from Flores (RMNH)”, but that is an error. It concerns RMNH cat.
no. 69 from Adonara recorded by Mees (1976); the error could originate as in the cardindex of the collection the locality is given as “Flores”; underneath the socle of themounted bird is, however, written: “tué 8.1880, Adonare, Pte. E. de Flores”. It is therefore the same individual on which the Adonara record is based.
Trichoglossus euteles (Temminck)
This parrot was supposedly collected by Allen; for a discussion and the rejection of
the record, see p. 96.
Collocalia salangana subsp.
Verheijen (1964: 199) listed, besides Collocalia esculenta, two swiftlets from Flores,
under the names C. vanikorensis dammermani and C. inexpectata subsp. The first of theseis now called C. fuciphaga, the second might refer to the species now known as C. salangana and its occurrence on Flores is quite likely, but requires verification. See also thediscussion under C. fuciphaga in the main text.
Alcedo meninting meninting Horsfield
In Rensch’s (1931a: 634) list, through an evident slip, this species is included instead

220

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)of Ceyx rufidorsa. The mistake was copied by Kuroda (1936: 735). It is probably due toRensch’s error, that Pfeffer (1958: 72) considered it justified to claim that he had “souvent rencontré” A. meninting!
Eurystomus orientalis pacificus Latham
The inclusion of this subspecies into the avifauna of Flores by White & Bruce (1986:
285-287), followed by Forshaw (1993: 92), is due to the assumption that resident birds of
the Lesser Sunda Islands would belong to this mainly or wholly Australian subspecies(see the discussion in the main text). E. o. pacificus is strongly migratory, and one would
almost expect it to occur on the Lesser Sunda Islands as a winter visitor, but as yet there
is no evidence. Father Schmutz paid special attention to the species, concentrating on
birds looking dull in the field, but all specimens procured by him were nominate E. o.
orientalis with somewhat dull plumage because of immaturity.
Zoothera peronii (Vieillot)
There is in the RMNH-collection an unsexed specimen of Z. peronii, with on its label:“v. Lansberge, 1882, Flores”. The label is not an original one, but was written by Finsch,ca. 1900, when the mounted bird was turned back into a study skin, and it received theRMNH cat. no. 6. With its history of mounting and dismounting, there must have been
ample opportunity for a mix-up. Hoping to find evidence for this, I checked Büttikofer’s list (see p. 11), where I found the bird, s. n. Geocichla rubiginosa, already with theprovenance Flores. The record was published by Finsch (1901b: 264), who did not doubt
it, as indeed he had no reason to, given the imperfect knowledge of the distribution of
the species at that time and the fact that there is no other evidence that Colfs’s labelling,sketchy as it is, is unreliable.
There are three specimens from Timor in the collection, also formerly mounted andlabelled with “v. Lansberge, 1881”. These, therefore, belong to a consignment received
a year earlier, of which there is no list.
On the basis of the material at that time available in the collection, Finsch (l. c.) concluded that the species shows no significant geographical variation and that, therefore,
Geocichla audacis, described from Damar two years before, was a synonym.
This was followed by Hartert (1904: 208-209): “After comparing our magnificent seriesof 34 audacis with 10 Timor birds (5 in Tring and 5 in London) I am at a loss to understand Dr Finsch’s statement, that Timor examples are of the same colour as audacis. G.
peronii peronii is above yellowish cinnamon, audacis chestnut- or rufous-cinnamon, andalso the chest and sides differ in the same way”. Later authors have accepted Hartert’s
conclusion, based on such an impressive material, rather than Finsch’s, and retained
audacis as a subspecies, claimed to be much darker and more rufous than nominate
peronii (Mayr, 1944a: 155; White & Bruce, 1986: 333). Mayr stated that all birds from
Dutch Timor including four specimens from Atapupu, belonged to typical peronii, andadded that a single male from Dilly was indistinguishable from a series of audacis, so
that: “The range of this form includes probably all eastern Timor”. Now it should be
remembered that Atapupu and Dilli are both situated on the north coast of Timor, notseparated by any obvious geographical barriers, and barely 100 km apart. As Mayrconsidered the subspecies well-differentiated (“much darker and more rufous”), thisMees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

221

implies an abrupt boundary between Atapupu and Dilli, which in a lowland bird witha presumably more or less continuous distribution is not very likely. Hartert considered
the whole of Timor to be inhabited by the nominate race. Mayr restricted its range towestern Timor, and added eastern Timor to the range of audacis. Hartert mentions having examined BM material and (knowing that Wallace has collected in eastern Timor,
and that some of his specimens are in the BM) it occurred to me that it would be worth
checking whether some of the five BM specimens which Hartert acknowledges having
examined, were from eastern Timor.
Mr Walters (in litt., 3.ix.1992) has informed me very fully on the BM material. I
quote here the most important passages of his letter: “We have a total of 8 specimens of
nominate Geocichla (Zoothera) peronii from Timor, all of which should have been here at
the time Hartert was writing this description, as they were all registered prior to 1899...
I can offer no ... suggestion as to why he apparently only examined 5. Two of the specimens are from “E. Timor”. Both were collected by A.R. Wallace. We also have an Everett
specimen from Atapupu. All the others are merely from “Timor”. Unfortunately, there
does not seem to be any way of determining whether Hartert saw these 3 skins. I havecompared the series with those of audacis and can see very little difference in colour. A
couple of the skins from Timor (including the Everett specimen) appear to be very marginally paler than those of audacis, but most (including the two Wallace specimens)
seem to me to be indistinguishable.... I would be inclined to agree that audacis is probably not valid, if there is any difference it would seem to be a very slight clinal variation”.
Thus, Mr Walters’s examination of the BM specimens supports my own conclusion,
based on the material in Leiden, that, in spite of the contrary conclusions arrived at by
Hartert and Mayr, audacis is not a valid subspecies.
Phylloscopus borealis borealis (Blasius)
Ticehurst (1938: 129) claimed to have seen: “quite typical borealis from Sumbawa,
Flors and Timor”. As mentioned on a previous page, all specimens examined by me arereferable to xanthodryas, and I consider that the occurrence of nominate borealis on Flores(and other Lesser Sunda Islands) requires confirmation.
The specimen from Scott Reef, Western Australia (14°03’S, 121°46’E) identified by
McKean (1980) as belonging to the nominate race, should be re-examined. The winglength of this specimen, an unsexed immature bird (WAM no. A 16285), is 66 mm (Johnstone, in litt., 8.v.1998): this is inconclusive, but is within the range of variation of xanthodryas. There is now also a record from Broome, mainland of Western Australia (Hassell, 1998). The bird was captured, examined, photographed, and released; the need fortaking detailed measurements is stressed twice in the text, yet, unbelievably, not a single measurement is presented. I have tried to obtain those measurements, but no reply
has been received.
Lonchura malacca ferruginosa (Sparrman)
L. m. ferruginosa was listed for Flores by Wallace (1864: 486, s. n. Munia ferruginea),
but certainly in error (see p. 10). See also Rensch (1931a: 596 footnote): “Munia ferruginea
wurde von Wallace ... fälschlicherweise für Flores angegeben”.

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10. Species and subspecies with type locality Flores
Accipiter novaehollandiae sylvestris Wallace, 1864
Accipiter virgatus quinquefasciatus Mees, 1984
Spizaetus (cirrhatus) floris (Hartert, 1898)
Turnix maculosa floresiana Sutter, 1955
Rallus pectoralis exsul (Hartert, 1898)
Rallus philippensis wilkinsoni (Mathews, 1911)
Treron floris Wallace, 1864
Ptilinopus cinctus albocinctus Wallace, 1864
Ptilinopus melanospilus melanauchen (Salvadori,
1875)
Trichoglossus (haematodus) weberi (Büttikofer,
1894)
Loriculus flosculus Wallace, 1864
Geoffroyus geoffroyi floresianus Salvadori, 1891
Tanygnathus megalorynchos floris Hartert, 1924
Otus magicus albiventris (Sharpe, 1875)
Otus alfredi (Hartert, 1897)
Otus silvicola (Wallace, 1864)
Ninox scutulata florensis (Wallace, 1864)
Alcedo atthis floresiana Sharpe, 1892
Pelargopsis capensis floresiana Sharpe, 1870
Mirafra javanica parva Swinhoe, 1871
Anthus novaeseelandiae albidus Stresemann, 1912
Coracina (novaehollandiae) floris (Sharpe, 1879)
Brachypteryx montana floris Hartert, 1897
Pnoepyga pusilla everetti Rothschild, 1897
Tesia everetii everetti (Hartert, 1897)
Orthotomus cucullatus everetti (Hartert, 1897)

Phylloscopus presbytes floris (Hartert, 1898)
Seicercus montis floris (Hartert, 1897)
Culicicapa ceylonensis sejuncta Hartert, 1897
Rhinomyias oscillans (Hartert, 1897)
Rhipidura diluta diluta Wallace, 1864
Monarcha sacerdotum Mees, 1973
Terpsiphone paradisi floris Büttikofer, 1894
Pachycephala fulvotincta fulvotincta Wallace, 1864
Pachycephala nudigula nudigula Hartert, 1897
Dicaeum annae (Büttikofer, 1894)
Dicaeum igniferum Wallace, 1864
Dicaeum sanguinolentum rhodopygiale Rensch,
1928
Anthreptes malacensis convergens Rensch, 1929
Zosterops palpebrosa unica Hartert, 1897
Lophozosterops superciliaris superciliaris (Hartert,
1897)
Lophozosterops dohertyi subcristata Hartert, 1897
Heleia crassirostris crassirostris (Hartert, 1897)
Lichmera lombokia fumidigula (Rensch, 1928)
Philemon buceroides neglectus (Büttikofer, 1891)
Amandava amandava flavidiventris (Wallace, 1864)(“Timor and Flores”)
Erythrura hyperythra obscura (Rensch, 1928)
Lonchura molucca propinqua (Sharpe, 1890)
Lonchura pallida pallida (Wallace, 1864) (“Lombock and Flores”)
Corvus florensis Büttikofer, 1894

This makes a total of 50 forms, described over a period of 120 years (1864-1984), by adozen authors. I have excluded forms described without indication of a holotype from a
range including Flores, of which the type locality has subsequently been restricted toanother island. Synonyms have also been excluded from this enumeration; they can befound easily in the main text. Of one or two subspecies included in this list (Tanygnathus
megalorynchos floris, Erythrura hyperythra obscura), the validity requires confirmation. Interestingly, only one of the birds described from Flores is a winter visitor (Ninox scutulata florensis), all others are residents. In sequence of the number of forms described, thelist of authors is: Hartert (16), Wallace (11), Sharpe (5), Büttikofer (5), Rensch (4), Salvadori (2), Mees (2), Mathews, Rothschild, Stresemann, Sutter and Swinhoe (1 each).
As a historical curiosity I want to recall that Hartert & Goodson (1918), arbitrarily
and without consideration of historical probability, designated Flores as the type localityof Ducula aenea, on the basis of the following argument: “The specific name aenea was
first given to Brisson’s ‘Columba moluccensis’, said to have been brought from the Moluccan Islands. As this species does not occur on the Moluccas, we propose as the restricted
locality for the name aenea Flores, which seems to be the nearest place to the Moluccas
where it occurs. Even if this should be considered incorrect, no great harm can be doneby this action, as the birds from the Greater and Lesser Sunda Islands, Sumatra, Borneo,

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223

Java, Lombok, Flores and Sumba are inseparable”. The argument, such as it is, holds
water only when it was certain that the subspecific identity of the type of aenea has beencorrectly established, for D. aenea actually does reach the Moluccas (D. a. sulana: Sula Islands), and Sulawesi (Celebes) (D. a. paulina) is also nearer the Moluccas than is Flores.
The type locality has been corrected to Manila by Stresemann (1952: 514, 520, 523).
It is easy to compile from the above list, a shorter list of 22 species and subspecies
endemic to Flores. Of course, a few species and subspecies at present thought to be confined to Flores may in future still be found on Sumbawa and perhaps other islands.
Accipiter virgatus quinquefasciatus
Rallus pectoralis exsul
Trichoglossus (haematodus) weberi
Loriculus flosculus
Tanygnathus megalorynchos floris
Otus magicus albiventris
Otus alfredi
Brachypteryx montana floris
Pnoepyga pusilla everetti
Tesia everetii everetti
Orthotomus cucullatus everetti

Phylloscopus presbytes floris
Seicercus montis floris
Culicicapa ceylonensis sejuncta
Rhinomyias oscillans
Monarcha sacerdotum
Pachycephala nudigula nudigula
Dicaeum sanguinolentum rhodopygiale
Lophozosterops superciliaris superciliaris
Lophozosterops dohertyi subcristata
Heleia crassirostris crassirostris
Corvus florensis

Even though a few forms (e. g., Heleia c. crassirostris) may require confirmation,these 22 forms, several of which are very well-differentiated, prove a considerable degree of endemism, in an island which is, after all, only a link in the middle of a closelyknit chain.
11. Zoogeography
The zoogeography of the Lesser Sunda Islands has been studied in particular byRensch (1928d, 1930, 1936) and Mayr (1944a, 1944b). Stresemann, in his classical workon the avifauna of Celebes (1939/1941), also paid attention to the Lesser Sunda Islands,
especially Flores, in their relation to Celebes. Here I shall try to give a zoogeographicalanalysis of the avifauna of Flores.
Species endemic to Flores (6)Trichoglossus (haematodus) weberi
Loriculus flosculus
Otus alfredi
Rhinomyias oscillans
Monarcha sacerdotum
Corvus florensis

Species endemic to Flores and Sumbawa (10) (with an asterisk, species are marked,
which have different subspecies on each island)
Spizaetus (limnaeetus) floris
Otus silvicola
Pericrocotus lansbergei
Tesia everetti*
Dicaeum annae

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Rhipidura diluta*
Pachycephala nudigula*
Lophozosterops superciliaris*
Lophozosterops dohertyi*
Heleia crassirostris*

Species endemic to Flores and Sumba (1)Coracina dohertyi

Species endemic to Flores and Timor (1)Phylloscopus presbytes*

Species endemic to Flores and Lombok (1)Ducula (lacernulata) sasakensis

Species endemic to Lombok, Sumbawa and Flores (2)Halcyon fulgida
Lichmera lombokia*

Species endemic to Flores, Sumbawa, Sumba and Lomblen (Komodo, Rintja) (1)Zosterops wallacei

Species endemic to Sumbawa, Flores, Pantar and Alor (1)Dicaeum igniferum

Of peculiar interest are species which one would expect to occur on Flores, butwhich apparently do not.
Lanius schach: Lombok, Sumbawa, Alor, Timor, Wetar, etc.
Brachypteryx leucophrys: Lombok, Sumbawa, Timor.

Other species skip Flores in a different way:
Halcyon australasia: Lombok, Sumba, Timor, Wetar and eastwards to the Tanimbar Islands.
Dicaeum maugei: Lombok, Roti, Semau, Timor.
Acrocephalus stentoreus: Lombok, Sumbawa, Sumba, Timor.
Milvus migrans: Lombok, Sumba, Timor.

As was to be expected on purely geographical grounds, it is the avifauna of Sumbawa which is closest to that of Flores. They share 10 endemic species, and although sixof these have different subspecies on Flores and Sumbawa, these subspecies are only
slightly differentiated, suggesting that the isolation is not a long one. On the other hand,in a few instances there is a strong differentiation: Trichoglossus haematodus forsteni on
Sumbawa vs. T. (haematodus) weberi on Flores, Phylloscopus trivirgatus trivirgatus on
Sumbawa vs. P. presbytes floris on Flores, Lonchura punctulata nisoria on Sumbawa vs. L.
p. blasii on Flores. In addition, the following resident species of Sumbawa are not known
from Flores: Lanius schach, Brachypteryx leucophrys, Acrocephalus stentoreus.
In the above enumeration, the emphasis has been on endemic and near-endemic
species of the Lesser Sunda Islands. For a balanced view, however, also subspecies ofmore widely distributed species have to be considered. When that is done, it is at once

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225

clear how strong the relation is between Flores and the south-eastern Sundaland (Java,
Bali). Many Sundaland birds have penetrated along the chain of Lesser Sunda Islands
to, and beyond, Flores. The affinity between Flores and Sumba becomes greater thanjust the one shared endemic species, through such species as Gallinula chloropus, Ceyx
rufidorsa, Culicicapa ceylonensis, Terpsiphone paradisi, Parus major. These species have notreached Timor, but others have.
Species of presumed Australian/Papuan origin and affinities are: Cacatua sulphurea,
Tanygnathus megalorynchos, Geoffroyus geoffroyi, Monarcha trivirgatus, Monarcha sacerdotum, Pachycephala fulvotincta, Lichmera indistincta, Lichmera lombokia, Philemon buceroides.
A problem is where to draw the line: from the distribution of their congeners, one would
deduce an Australian origin for Gerygone sulphurea and Artamus leucorhynchus, but these
species are widely distributed in south-east Asia, making a reconstruction of their history speculative.
Stresemann’s (1939: 321) suggestion that grassland birds have colonized Sulawesi(Celebes) and the Lesser Sunda Islands from south-eastern Asia, through Taiwan and
the Philippines, around the heavily forested Malay Peninsula, Sumatra and Borneo,
finds little support (cf. Elanus caeruleus, Circaetus gallicus, Tyto longimembris, Caprimulgus
affinis); it is more likely that, during Pleistocene periods of low sea-level, including thelast glacial, a direct connection through the exposed Sunda Shelf was possible.1213)
In the avifauna of Flores, there is little evidence for a close relation with Sulawesi(Celebes). Mayr (1944a) has forcibly argued against land bridges between Flores andSulawesi, projected by some of his predecessors. And, as noted, for an explanation of theornithogeography of the region, no land bridge is needed. With the passing of time, andthe discovery of fossil elephants as far east as Sulawesi and Timor, the land bridge papers have temporarily gone up again (Audley-Charles & Hooijer, 1973; Hooijer, 1974)
but once more have failed to find general acceptance, as there is too much contrary evidence. The occurrence of elephants and other large herbivores on Sulawesi and the Lesser Sunda Islands has now to be explained in a different way, not with complete landbridges. There is general agreement that during Pleistocene periods of low sea-level
there was much more land between Java, Sulawesi, and the Lesser Sunda Islands, witha corresponding narrowing of sea straits separating them. The controversy over how
well elephants are able to swim is an old one (with some extreme views expressed by the
opposing camps), but swimming remains the only explanation for the wide distributionof elephants, deer and buffaloes east of Wallace’s Line (cf. de Vos et al., 1994).1314)
No species of bird is represented on Flores by more than one resident subspecies.
The only island in the chain of the Lesser Sunda Islands, in which intra-island geographical variation has been recorded is Timor, the largest island.

13)

Although, over half a century after its publication, it is clear that not all Stresemann’s (1939) conclusions are acceptable, his work, written in the lucid and persuasive style for which he was justly known,
remains a major contribution to the zoogeography of the region. It is incomprehensible that Dickinson
et al. (1991), in their study of the zoogeography of the Philippines, do not even mention it, and haveomitted it from their extensive bibliography.
14)
But the present-day example given by these authors, of large herds of deer and buffaloes crossing the
100 km wide Gulf of Bone in southern Sulawesi, cannot have been meant seriously, especially the partabout these hoofed animals loading a food supply for the crossing on their heads. The story may haveoriginated in some floating carcasses covered in seaweed having been seen.

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According to Mayr (1944a: 130), five species show geographic variation on Timor, as
follows:

Coturnix ypsilophora
Trichoglossus iris
Zoothera peronii
Turdus poliocephalus
Philemon inornatus

Western Timor
raaltenii
iris
peronii
schlegelii
inornatus

Eastern Timorcf. castanea
rubripileum
audacissterlingi
robustus

However, I do not accept the validity of two of these cases (Coturnix ypsilophora andZoothera peronii). White & Bruce (1986: 397) have also rejected a third one (Philemon inornatus), but the measurements published by Mayr (1944a: 165) show that the size difference between inornatus and robustus is real enough. Admittedly, the occurrence of anapparently steep cline in measurements in a lowland bird, on a comparatively small
island, is unexpected and will be worth further study.
Recently added to the group is Ptilinopus regina: P. r. flavicollis (or P. r. ewingii) inwestern Timor, and P. r. roseicollis (intermediate between flavicollis and xanthogaster)from eastern Timor (cf. Johnstone, 1981). So long as only a single specimen of roseicollis
is known from Timor, the possibility that it is a vagrant and not a resident, cannot be
ruled out. Therefore, this case requires confirmation.
Mayr (1944a: 170) drew attention to another phenomenon occurring on Timor, andnot unknown elsewhere: an increase of size corresponding with an increase in altitude.
This has its theoretical basis in Bergmann’s Rule. On the other hand, I find it difficult tobelieve that this can have much influence in birds: 1000 m difference in altitude, neednot mean more than a real distance of three to four kilometers and in a continuous
population it is unlikely that much real genetical difference could build up over such asmall distance (although the case of Zosterops borbonica comes to mind as an example
that differences do occur over a short distance; cf. Gill, 1973). The matter is of course
different, when there is a break in habitat, for example lowland and alpine grasslands,
separated by a belt of forest.
Particularly fascinating I find the occurrence of differences in vertical range between
adjacent subspecies; in this paper Phylloscopus presbytes, Rhipidura rufifrons and Lichmera
lombokia. As stated in the systematic account, I do not include Dicaeum sanguinolentum
here, for I regard Dicaeum wilhelminae as a separate species.
A total of 214 forms has now been recorded from Flores. The number of species isslightly lower, as of some species more than one subspecies occurs, usually a resident
one and a migrant visitor.
Dividing the birds into the categories shown in the register, we get:Residents and presumed residents
Migrant visitors from the North
Migrant visitors and vagrants from the South
Tropical marine birds which may breed in the neighbourhood of
Flores, but are unlikely to do so on the mainland
Uncertain

155
28
14
4
13

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227

The last group, marked as uncertain, consists almost entirely of freshwater birds
which may breed on Flores, but there is as yet no evidence that they do, neither are theyknown to breed on any of the other Lesser Sunda Islands (this is the case with 2 speciesof shags and 8 species of herons). Of the remaining species, two, Gallinula tenebrosa andPetrochelidon nigricans, may be no more than vagrants from Timor, where they breed.
The case of Lichmera indistincta is different, in that it is known to occur, and undoubtedly to reproduce, on small islands near Flores, and indeed, its occurrence on Flores asa resident may still be confirmed.
Naturally, the question arises of how well the avifauna of Flores is now known
quantitatively. My obvious guess is that few resident land and freshwater birds remain
to be discovered, although the facts that some presumably resident land birds (Loriculus
flosculus, Monarcha trivirgatus, Dicaeum agile) have not been recorded for a century (andthat at least Monarcha trivirgatus and Dicaeum agile are not likely to be extinct), and that
Dicaeum sanguinolentum remains known from only three specimens, show that evenhere surprises remain possible. There are recent field-observations of all four species
mentioned in this paragraph, with Dicaeum sanguinolentum being recorded as common
above 1000 m (Verhoeye & Holmes, 1998).
With migrant wading birds of the families Charadriidae and Scolopacidae it is anentirely different matter. Evidence from other islands shows that at least some 15 more
species of these families ought to occur as regular migrant visitors.
The family Laridae (Sterninae) should also contribute half a dozen species.
From the rather sparse records of waders in Wallacea, White (1975a) drew the conclusion that the region is not an important wintering ground for waders. Although heexpressed an awareness of collector’s bias in favour of resident, indigenous birds, I
consider that he has grossly underestimated this bias; indeed, it is rumoured that somecollectors were expressly instructed not to waste time paying attention to migrant waders. As I mentioned in an earlier paper, such birds, when recorded at all, were dismissed
in a footnote (cf. Mees, 1976). In contradistinction to what White assumed, there areplenty of reefs, beaches, and mudflats in Wallacea, which can and do support waders. I
realise that White distinguished between birds actually wintering in Wallacea, andtransients (what reaches Australia has to pass through Wallacea), but he underestimated both categories.
Finally, it must not be overlooked that these migrants are seasonal and that for thatreason expeditions active in the northern summer were not likely to meet with larger
numbers.
Addendum.— The Sumbawa records published by Butchart et al. (1996) and Johnstone et al. (1996) make some modifications to this chapter necessary. The species ofparticular interest in relation to Flores are Ducula (lacernulata) sasakensis, Coracina dohertyi, Seicercus montis and Rhinomyias oscillans.
Ducula sasakensis was the only member of the category Lombok-Flores, not Sumbawa; with its (predictable) occurrence on Sumbawa the hiatus has been filled and D.
sasakensis joins the next category: Lombok-Sumbawa-Flores. Coracina dohertyi was listedwith the unique distribution Flores-Sumba; with its discovery on Sumbawa, its patternof distribution has become more natural. Butchart et al. suggest that birds from Sumbawa are subspecifically separable, but that remains to be verified. The Sumbawa recordof Seicercus montis, although not unlikely, is unsatisfacory, being based on what seems

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to have been little more than a glimpse by one (or more than one?) unnamed observer.
Rhinomyias oscillans: an interesting and properly-documented record of a species hitherto thought to be endemic to Flores.
These records emphasise the already known close affinity between the avifaina of
Flores and Sumbawa.
12. List of localities
Many of the numerous localities mentioned in the main text are not found on ordinary maps, and very likely quite a few are not found on any map. With the help of Father Verheijen, and of the map of Manggarai by Fr. Janssen (ca. 1953) which he sent me,
I have localized most of these, if not exactly, at least approximately.
The Daerah (county) of Manggarai is (or was until recently) divided in Kedaluans,perhaps best translated as communities or districts, at the head of which stood a Dalu
(feodal headman). A Kedaluan, however, is more than just an administrative division,
it is also a social and linguistic unit.
The map (fig. 4) shows the Kedaluans of Manggarai (after Verheijen, 1967). When ofa minor locality the Kedaluan in which it is situated, is known, that already gives a fairidea of its position. Often a Kedaluan takes its name from its head village (cf. Pongkor,
Ruteng, Todo).
A particular difficulty lies in the transcription of geographical names. This is aworld-wide problem: for example, the introduction of a new system of transcription ofChinese names, has made the geography of that country permanently inaccessible tome. On Flores, the situation is not so bad as, at least, the Latin alphabet has been used.
Even so, for the island here called Palué, the names and transcriptions Roesa Radja,
Rusa Radja, Paloweh, Paloë, Paloe and Paluë have been used. In the title of Verheijen’s(1961) article, the name is spelled Paluë, but in the text consistently Palué. Father Verheijen informed me that the linguistically most correct transcription would probably be
Palu’e. Besides differences in transcription, there have, in the past forty years, also been
several changes in spelling. Following Malay (in Indonesia called Indonesian), theDutch oe was replaced by u, so that, for example, the places Roeteng and Larantoekachanged their names to Ruteng and Larantuka. Later there were other changes, theworst one that of tj to c. It is a moot question, whether it is advisable to accept spelling
rules introduced for modern Malay uncritically for other languages. The change of thename Mborong to Borong probably also comes in this class.
I had been unable to find the place Mataloko, where so much material was collected, on the map, until Father Verheijen informed me that it is the same as Toda-Belu.
As this is a paper about birds, not about linguistics or geography, I have felt free tobe opportunistic. Therefore, I have not tried to “modernize” names as they appear onlabels.
A few common words: Golo = hill, sometimes also mountain-top; Lareng = placeagainst a mountain, part of a mountain-slope; Lingko = the name of a special Manggarese communal garden, which is used at intervals of from 7-10 years, and in the otheryears lies fallow and becomes covered with dense secondary growth; Poco or Potjo =mountain, mountain peak; Puar = forest; Rana = lake; Waé = water, including creek,
river, sometimes lake; Ulu = source, upper course, origin of a stream.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)
Aimere, 0-200 m
BadjarBadjawa, 1200 m
Badjo, near Nisar
Bara-Latji, 200 m
Bari
Bénténg Djawa
Boa Waé
Borong, coast (on maps Mborong!)
Celebesbaai = Hading-baai
Ceréng, 300 m, see Tjeréng
Dalo, Rahong
DampékDéngé, Todo
Deno
Desoe, 3300 ft. (Everett)Djinggor, ca. 300 m, north-east of Waé-Nénda
Djoneng
Endeh
Ero
Geli Moetoe
Golo Léhot
Golo -Karot, Borong
Golo Mbélar
Golo Rucuk
Groot Bastaard = Geliting Besar
Gurung, Langkas, Rahong, 900 m
Heret
Hochwald zwischen Paku und Sok-Rutung
Hotju, Ruteng, 1500 m
Joneng, beach and dunes, see Djoneng
Kandang, 900 m
Karot, Borong, 50 m
Kedindi, coast
Kenari (west coast), 50 m
Kisol, Borong, 175 m
Koting = Kotting
Kuwu, Rahong
Labuanbadjo = Laboean Badjo
Lai, Palué
Lalang, Todo
Lamba, Todo
Lamé
Larantoeka
Laréng Pongkor, 1100 m
Lawir Lamba-Leda
Léma, 900m
Lemboi, 250 m
Lenteng
Léong, Méngé
Léwé
Liang Bitu
Lingko Cehet

229
Lingko Laring Pongkor (also Lareng)
Lingko-Moak, Langkas
Lingko Ncilor
Lingko Ngkiong
Lingko Ros
Ling Ndela, 600 m
Lita, 50 m
Lo Kong, 900 m
Longko, Wangkung, Rahong, 900 m
Look = Loé = Looh, 5 m
Look-Mabacone
Lumu
Madjung
Mano
Maro-Kama = Marokama, Maro Kama, ca. 10 km
East of Borong
Mataloko = Toda-Belu
Mata-Waé
MaumeriMbawa
Mbépé, 1700 m
Mboera = Mburak
Mborong = Borong, coast
Méléng
Méngé
Mengé-Ruda
Mongu Luwa
Montjok, Lamba-Leda, 600-800 m
Mt. Repok, above 3500 ft, an Everett locality near
Nanga Ramau recently visited by Butchart etal. (1996) and shown on their fig. 1.
Mukun
Naga, 150m, 250m
Nanga-Lili = Nangalili, 20 m
Nanga Ramau = Nanga Ramut
Nanga Rema
Nantal, Rahong
Narang, Todo, 150 m
Nempong
Ngalor-Roga, Langkas, Rahong
Nggoang, 850 m
Nggoér, 5 m
Nggolong-Tédé, Ruteng, 1900 m
Ngkiong, L. Ngkiong
Nilo
Nisar, west Manggarai, beach, 50/100 m, 180 m
Ntéwéng, Mano
Nunang, 650 m, 700 m
Nunuk
Orong, 550 m [nb. Orong, Wélak, Orong, Tjongkar]
Paku, 350-400 m
Paku, Mbelawang-Bach, 300 m (brook?)

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Palué = Paloë = Paluë = Paloweh = Rusa Radja
Pangabatang
Pangkor = Pongkor?
Papagaran
PocoLareng, 1400-1600 m.
Poco Nernancang, Ruteng, ca. 1500 m
Poéng, Méngé
Raé
Rahong, ca. 1000 m, 900 m
Rana-Ka, mountain lake, 2200 m
Rana Kulan, Biting, 400 m
Rana Loba, Borong
Rana Mesé, 1200-1400 m
Rekas, 450 m, Kempo
Rembong
Rempo
Rentung Lelak
Réo (Weber)
Repok, see Mt. Repok
Riung
Robo
Roka, Wangkung
Rokka, near Endeh
Rowang, Ruteng, 1200 m
Rua, Ruteng
Rusa Radja = Paluë
Ruteng/Kumba
sawah Lanar
Sésok, 610, 900 m
Sikka = Sika
Siru, 50 m, near Lita
Sita, 700 m
Soa Soknar
Sok-Rutung = Sokrutung, 50 m
Suma, Langkas
Tado
Tado Walok, Ruteng, ca. 1700 m
Tendo, Ruteng

Tjantjar, Rahong
Tjara, Rahong
Tjeréng, 600 m
TjololTjumbi, Rahong
Todabelu = Todabeloe = Toda-Belu = Mataloko
Todo, 700 m
Tulang, Potjong
Ulu Ros
Ulu Tukenikit = Ulu Tuké Nikit
Ulu Wae Wua
Waé Djamal
Waé Golo Lolo
Waé Guang
Waé Hiam
Waé Laci
Waé Lega
Waé-Mésé, Kandang, Nisar
Waé-Mulu
Waé-Ntjuang, Langkas, 900 m = Wai-Ntjuang
Waé Radja, 20 m
Waé Rambung
Waé Rembong
Waé Rempo
Waé Rétja = Waé Réca, Borong
Waé-Rukus = Ulu Waé Rukus
Waé Rungget
Waé Sano, 650 m
Waé Tua, 400 m
Waé-Wako, 150-200 m
Wai Moké, south coast of middle Flores
Wangkung = Wang keng, Rahong
Watuneso
Weleng, Ruteng
Wesang, 700 m, Lamba-Léda
Wewak, 400 m
Wodja, Palué

13. Postscript
As mentioned in the introduction, the tortured history of this paper began many
years ago and progress was slow. Originally intended to include only discussions ofsome of the more interesting species collected by the Fathers, it soon became clear that
only a full report would do justice to their work. At the time of my retirement (1991), allmeasurements of bird specimens and a proportion of those of the eggs had been taken,
and a draft of the text had been written. In the following years, after my departure forAustralia, access to literature was limited and there was very little progress.
A stay in Leiden in 1995 gave me an opportunity to study more literature, somematerial that had been received after my departure, and to complete the measuring of
the eggs. Sections of the text were supplemented or revised. Short descriptions of theMees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

231

eggs were added, but this task was not completed and could not be completed later forthe obvious reason that I had no longer access to the collections. When at the end of thisvisit I left Leiden for good, the work was in a computer, and my colleague Dr J. van derLand most generously offered to take care of it, and to make any further additions andcorrections required. Perhaps predictably this went less well than I had hoped and delays led to an increasing doubt that my paper would ever be published.
In this period several papers appeared which would have made extensive re-writing necessary, but for the reason just stated I lacked the motivation. The first of these
papers to reach me, through the courtesy of Dr Dekker, was the one by Butchart et al.
(1996). Dr Dekker informed me that before its publication there had been no contact
with Leiden, and that he had been unaware that it was in preparation. In its introduction it states somewhat scathingly that: “Two missionaries made anecdotal observations on Flores”, a strange remark when their text and their list of references show thatthey had full access to Father Schmutz’s notes. The paper by Ottow & Verheijen (1969)is also mentioned, but Father Verheijen’s other publications are conveniently ignored.
These comments are not intended as unkindness, but merely to emphasise the need for
the Flores collections in Leiden to be made more widely known, to end such underestimation of their extent and value.
All these years, my manuscript was getting more and more out of date. It was also
as I despaired of ever seeing it published, that complete and partial copies were madeavailable to several persons showing an interest: in that way I hoped that my efforts
would not be entirely wasted.
In the introduction I have discussed the question of how to treat additions to the
Flores avifauna based on undocumented field-observations, but there is every reason to
return to this subject. In the West Indies, Bond (1963) was plagued by it forty years ago.
He took a very strict view, which is, and was even then, probably right for the West
Indies, but not necessarily for the less known Lesser Sunda Islands. Obviously, many
common migrants and other regular visitors had not yet been recorded for the simple
reason that nobody had bothered about them, so that records in these groups were confined to the odd specimen taken accidentally. There is no need, indeed it would beslightly silly, to apply to observations of such birds the standards required in the West
Indies (leave alone Europe and North America). My problem with field-observations
was a different one, that arose only in the nineties. Indeed, the beginning was inauspicious: Verhoeye & King (1990) and King (1990) recorded four species as “new to Flores”;
of these four, three had already been recorded many years before by the Fathers, and I
had no difficulty accepting the remaining one (Hieraaetus kieneri) as it was not unexpected, being of a species previously known from Sumbawa. However, my paper isabout collections, from which measurements were taken, and which provided the basis
for systematic and zoogeographical discussions. Clearly field-observations would notcontribute much to these particular subjects.
The supplementary list of birds from Flores, as presented by Verhoeye & Holmes(1998), with some corrections made by me, follows. Bulweria bulwerii, Puffinus leucomelas, Puffinus pacificus, Oceanites oeceanicus, Oceanodroma matusudairae, Phaethon lepturus, Fregata minor, Sula dactylatra, Sula sula, Plegadis falcinellus, Falco peregrinus, Nettapus
pulchellus, Anas querquedula, Aythya australis, Fulica atra, Pluvialis squatarola, Charadrius
dubius, Charadrius mongolus, Charadrius peronii, Numenius minutus, Numenius madagas-

232

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

cariensis, Tringa totanus, Tringa stagnatilis, Tringa brevipes, Xenus cinereus, Arenaria interpres, Calidris acuminata, Calidris ferruginea, Glareola maldivarum, Stercorarius pomarinus,
Stercorarius parasiticus, Chlidonias leucopterus, Gelochilidon nilotica, Sterna hirundo, Sterna
sumatrana, Sterna anaethetus, Sterna fuscata, Ducula bicolor, Caloenas nicobarica, Apus affinis, Alcedo beryllina.
Although this list of 41 species is impressive, perhaps its most striking feature isthat there is not a single Passerine bird in it, confirming that this group of birds is now
well known.
As was to be expected, most additions are of migrant waders from the Northern
Hemisphere, and widely-ranging tropical seabirds: neither of these two categories is ofmuch zoogeographical interest and they do not require a discussion. This leaves thefollowing records of note. It should be kept in mind that all these records were presented without any details of identification, so that an evaluation is impossible.
Nettapus pulchellus: unexpected, as records were hitherto confined to Australia andsouthern New Guinea.
Plegadis falcinellus: previously known from Java, Sulawesi (Celebes) and Australia,
so that the record from Flores fills a gap of its range as hitherto known.
Falco peregrinus: this species would be expected as a migrant visitor from the North,already known from Java and Timor, but the observers state expressly that the bird seen
was dark-breasted, suggesting the subspecies ernesti, which might be a resident.
Charadrius dubius was to be expected, but in view of the late date (29.v.1993) and thefact that no less than eight individuals were observed, some basic information, such aswhether the birds were in summer or winter plumage, and an assurance that there wasno possibility of confusion with C. peronii, would have been welcome. Descriptions of
the plumages of this species in the literature are often misleading, as I pointed out before(Mees, 1982a, 1984b). In my previous review (1982a: 54) I claimed that the tropical subspecies (jerdoni and nominate dubius) have even in the immature plumage a black pectoral band. This was based on information supplied by Deignan in Smythies (1960 and
subsequent editions); I had not personally examined any juvenile birds. Since then, however, I received a juvenile specimen of jerdoni on loan from the Bombay Natural History
Society, and this bird has a brown breast band, proving the existence of a juvenile plumage. I cannot say whether nominate dubius has a similar juvenile plumage, but consider
it very likely. Another small error in my 1982a paper is the statement that there are no
specimens of the subspecies jerdoni in the Leiden collection: several years after the publication of that paper I discovered that, although there are no skins, there are several
mounted specimens in the old collection which, somehow, I had managed to overlook.
Ducula bicolor: Additional observations supporting Hoogerwerf’s questionable
record are welcome. It is perhaps not superfluous to mention here that when I do not
accept a record this does not necessarily mean that it is erroneous, but only that the evidence presented is not of sufficient quality to be completely convincing (Hoogerwerf
himself listed the record with a query). In the present case I cannot help adding that thenotes provided by Hoogerwerf are better than those presented by Verhoeye & Holmes.
Apus affinis is a genuine addition to the resident avifauna of Flores; whether it has
been overlooked previously or is a recent colonist is a point not discussed by the observers.
Alcedo beryllina: a single individual as observed in the mangrove at Riung on 5 andMees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

233

7.viii.1997, by Pilgrim. This species is well distributed on Sumbawa. It was either a migrant from Sumbawa or belonged to a local breeding population.
14. Bibliography and references
Anon., 1998. European news.— Brit. Birds 91: 38-57.
Alström, P., 1990. Calls of Pacific Golden Plovers.— Brit. Birds 83: 70-72.
Amadon, D., 1941. Notes on some Australian birds of prey.— Emu 40: 365-384.
Amadon, D., 1943. Birds collected during the Whitney South Sea Expedition. 52. Notes on some nonpasserine genera, 3.— Amer. Mus. Novit. 1237: 1-22.
Amadon, D., 1959. Remarks on the subspecies of the Grass Owl, Tyto capensis.— J. Bombay Nat. Hist.
Soc. 56: 344-346.
Amadon, D. & J. Bull, 1988. Hawks and owls of the World, an annotated list of species.— Proc. West.
Found. Vert. Zool. 3: 295-357.
Amadon, D. & G. Jewett, Jr., 1946. Notes on Philippine birds.— Auk 63: 541-559.
Amadon, D. & G. Woolfenden, 1952. Notes on the Mathews collection of Australian birds. The orderCiconiiformes.— Amer. Mus. Novit. 1564: 1-16.
Amaral, A. do, 1964. Gender of generic names ending in -ops.— Bull. Zool. Nomencl. 21: 212-215, also218-221.
Andrew, P., 1986. Notes on some birds of Timor.— Kukila 2: 92-95.
Ash, J.S., 1984. Bird observations on Bali.— Bull. Brit. Orn. Cl. 104: 24-35.
Assem, J. van den., 1960. Vogelwaarnemingen in Nieuw-Guinea.— Ardea 48: 178-182.
Audley-Charles, M.G. & D.A. Hooijer, 1973. Relations of Pleistocene migrations of pygmy stegodonts toisland arc tectonics in eastern Indonesia.— Nature 241: 197-198.
Baker, R.H., 1951. The avifauna of Micronesia, its origin, evolution, and distribution.— Univ. Kansas
Publ., Mus. Nat. Hist. 3: 1-359.
Balen, S. van & A.R. Compost, 1989. Overlooked evidence of the Short-toed Eagle Circaetus gallicus on
Java.— Kukila 4: 44-46.
Barbagli, F. & C. Violani, 1997. Canaries in Tuscany.— Boll. Mus. Reg. Sci. Nat. Torino 15: 25-33.
Bartels, M., 1914. Über eine anscheinend neue Art Dicaeum von Java.— Orn. Mber. 22: 125-126.
Bartels, M., 1923. Neues aus Java.— Orn. Mber. 31: 54-57.
Bartels Jr., M. & E. Stresemann, 1929. Systematische Übersicht der bisher von Java nachgewiesenen
Vögel.— Treubia 11: 89-146.
Bastide, J.G. la, 1941. De Kaneelkleurige Reiger, Ixobrychus cinnamomeus (Gm.).— Irena 1: 22-24.
Beehler, B.M., T.K. Pratt & D.A. Zimmerman, 1986. Birds of New Guinea. Princeton University Press: xv+ 293 p., 55 pl.
Bemmel, A.C.V. van, 1938. Twee snippen.— Trop. Nat. 27: 138-140.
Bemmel, A.C.V. van, 1939. Eine neue Monarcha von der Insel Boano.— Orn. Mber. 47: 152-153.
Bemmel, A.C.V. van, 1940a. Ornithologische Notizen I-III.— Treubia 17: 333-335.
Bemmel, A.C.V. van, 1940b. Een zeldzame trekvogel.— Trop. Nat. 29: 35-36.
Bemmel, A.C.V. van, 1948. A faunal list of the birds of the Moluccan Islands.— Treubia 19: 323-402.
Bemmel, A.C.V. van & A. Hoogerwerf, 1940. The birds of Goenoeng Api. -– Treubia 17: 421-472, pl.
17-34.
Bemmel, A.CV. van & K. H. Voous, 1951. On the birds of the islands of Muna and Buton, S. E. Celebes.—
Treubia 21: 27-104.
Berlioz, J., 1946. Oiseaux de la Réunion. Faune de l’Empire Français IV: 83 p.
Bingham, C.F., 1880. Additional notes on the birds of Tenasserim and specially on those of the Thoungyeen valley.— Stray Feathers 9: 138-198.
Blakers, M., S.J.J.F. Davies & P.N. Reilly, 1984. The atlas of Australian birds. Melbourne University Press:
xlvi + 738 p.
Blasius, W., 1883. Ueber neue und zweifelhafte Vögel von Celebes.— J. f. Orn. 31: 113-162.
Blasius, W., 1886. Beiträge zur Kenntnis der Vogelfauna von Celebes. II.— Zeitschr. ges. Orn. 3: 81-179.
Blasius, W., 1888. Die Vögel von Gross-Sanghir.— Ornis 4: 527-646, Tafl. III, IV.

234

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Boie, F., 1844. Auszüge aus dem “System der Ornithologie”.— Isis (1844): Col. 164-198.
Boles, W.E., 1988. Comments on the subspecies of Australian native and introduced finches.— Emu 88:
20-24.
Bonaparte, C.L., 1852. Descriptions of two new species of oriole.— Proc. Zool. Soc. Lond. 18 (1850):
279.
Bond, J., 1963. Eighth supplement to the Check-list of birds of the West Indies (1956). Acad. Nat. Sci.
Philadelphia: 11 p.
Brisson, D., 1760. Ornithologie 3. Paris: 734 + xci p.
Broek, M. van den, 1991. Zeldzame vogel vermoord door twee onderzoekers.— Volkskrant, 4 November 1991.
Brouwer, G.A., 1954. Historische gegevens over onze vroegere ornithologen en over de avifauna van
Nederland.— Ardea 41:1-225.
Brown, L. & D. Amadon, 1968. Eagles, hawks and falcons of the world. Country Life Books / Feltham,Middlesex; 2 vols: 1-945.
Brüggemann, F., 1876. Beiträge zur Ornithologie von Celebes und Sangir.— Abh. naturwiss. Ver. Bremen
5: 35-102, Taf. III, IV.
Bühler, A. & E. Sutter, 1951. Sumba-Expedition des Museums für Volkenkunde und des Naturhistorischen Museums in Basel, 1949. Reisebericht und allgemeine Einführung.— Verh. Naturf. Ges. Basel 62: 181-217, Taf. I-VI.
Butchart, S.H.M., T.M. Brooks, C.W.N. Davies, G. Dharmaputra, G.C.L. Dutson, J.C. Lowen & A. Sahu,
1996. The conservation status of forest birds on Flores and Sumbawa, Indonesia.— Bird Conservation International 6: 335-370.
Büttikofer, J., 1891. On a collection of birds from Flores, Samao and Timor.— Notes Leyden Mus. 13:
210-216.
Büttikofer, J., 1892. On a collection of birds from the islands of Flores, Sumba and Roti.— Notes LeydenMus. 14: 193-206.
Büttikofer, J., 1894. Ornithologische Sammlungen aus Celebes, Saleyer und Flores.— Zool. Ergebnisse 3:
269-306, pl. XVII, XVIII.
Burton, J.A., 1973. Owls of the world. Peter Lowe, Eurobook: 1-216.
Cabanis, J. & F. Heine, 1860. Museum Heineanum. II: Schreivögel: 1-176.
Cade, T.J., 1982. The falcons of the world. Collins, London, etc.: 1-183 + index.
Cain, A.J., 1955. A revision of Trichoglossus haematodus and of the Australian Platycercine parrots.— Ibis
97: 432-479.
Chasen, F.N., 1935. A handlist of Malaysian birds.— Bull. Raffles Mus. 11: xx + 389 p.
Chasen, F.N. & A. Hoogerwerf, 1941. The birds of the Netherlands Indian Mt. Leuser expedition 1937 tonorth Sumatra.— Treubia 18, Suppl.: 1-125, pl. 1-8, map.
Cheng, T.H. 1976. Distributional list of Chinese birds. Science Publishing House, Beijing: xvii + 1218 p.
(in Chinese).
Cheng, T.H., 1987. A synopsis of the avifauna of China. Beijing: xvi + 1223 p.
Cheng, T.H., et al.,1978. Fauna Sinica. Series Vertebrata. Aves 4: Galliformes. Academia Sinica, Peking:vi + 203 p., 10 pl.
Christides, L. & W.E. Boles, 1994. The taxonomy and species of birds of Australia and its territories.
Monograph 2. RAOU, Melbourne: iv + 112 p.
Condon, H.T., 1975. Checklist of the birds of Australia. Part 1, Non-Passerines. RAOU, Melbourne: xx +311 p.
Connors, P.G., 1983. Taxonomy, distribution, and evolution of golden plovers (Pluvialis dominica andPluvialis fulva).— Auk 100: 607-620.
Connors, P.G., B.J. McCaffary & J.L. Maron, 1993. Speciation in golden-plovers, Pluvialis dominica and P.
fulva: evidence from the breeding grounds.— Auk 110: 9-20.
Coomans de Ruiter, L., 1936. Oölogische en biologische aanteekeningen van eenige roofvogels in dewesterafdeeling van Borneo.— Org. Cl. Ned. Vogelk. 9: 34-52.
Coomans de Ruiter, L., 1947. Vogelwaarnemingen gedurende een tocht door het Toradjagebergte.— Lev.
Natuur 50: 61-66.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

235

Coomans de Ruiter, L. 1948a. Nog enkele opmerkingen naar aanleiding van de “Avifauna van Batavia
en omstreken” van A. Hoogerwerf en wijlen Jhr Mr G.F.H.W. Rengers Hora Siccama.— Ardea 36:
61-70, pl. III.
Coomans de Ruiter, L., 1948b. Waarnemingen van steltlopers (Gressores) in Zuid-Celebes, in het bijzonder van het blauw-witte reigertje, Notophoyx picata (Gould).— Limosa 21: 69-83.
Coomans de Ruiter, L., 1951. Vogels van het dal van de Bodjo-rivier (Zuid-Celebes).— Ardea 39: 261-318,
pl. IV-VIII.
Coomans de Ruiter, L., 1954. Trekvogels van Sulawesi (Celebes).— Penggemar Alam (Trop. Nat.) 34:
67-96.
Coomans de Ruiter, L., W.C. van Heurn & W.K. Kraak, 1948. Beteekenis en etymologie van de wetenschappelijke namen der Nederlandsche vogels. Club Ned. Vogelk.: xii + 166 p.
Coomans de Ruiter, L. & L.L.A. Maurenbrecher, 1948. Stadsvogels van Makassar (Zuid-Celebes).— Ardea 36: 163-198.
Cooper, R.P., 1975. Wilson’s Promontary National Park and its avifauna. Bird Obervers Club, Melbourne:
vi + 168 p.
Cramp, S. & K.E.L. Simmons (eds.), 1977. Handbook of the birds of Europe, the Middle East and North
Africa I. Oxford University Press: ix + 722 p., 108 pl.
Cramp, S., et al., 1983. Handbook of the birds of Europe, the Middle East and North Africa III - Waders
to gullss. Oxford University Press: viii + 1063 p., 84 pl.
Cramp, S., et al., 1988. Handbook of the birds of Europe, the Middle East and North Africa V - Tyrantflycatchers to thrushes. Oxford University Press: viii + 913 p., 105 pl.
d’Albertis, L.M., 1880. New Guinea: what I did and what I saw. Sampson Low, Marston, Searle & Rivington, London: vol. 1: i-x, 1-424; vol. 2: i-x, 1-406.
Dammerman, K.W., 1929. On the zoogeography of Java.— Treubia 11: 1-88.
David, N. & M. Gosselin, 2000. The supposed significance of originally capitalized species-groupnames.— Bull. Brit. Orn. Cl. 120: 261-266.
David, N. & M. Gosselin, 2002. Gender agreement of avian species names.— Bull. Brit. Orn. Cl. 122: 1449.
Deignan, H.G., 1951. A new race of the Hawk-Owl, Ninox scutulata, from the Philippines.— Proc. Biol.
Soc. Wash. 64: 41-42.
Deignan, H.G., 1955. The glossy starlings (Aplonis) of Borneo.— Sarawak Mus. J. 6 (1954): 129-132.
Deignan, H.G., 1960. Harry Cusheir Raven’s travels in eastern Borneo.— Sarawak Mus. J. 9 (1959): 267269.
Deignan, H.G., 1961. Type specimens of birds in the United States National Museum.— U. S. Nat. Mus.
Bull. 221: ix + 718 p.
Deignan, H.G., 1963. Checklist of the birds of Thailand.— U. S. Nat. Mus. Bull. 226: i-x, 1-263.
Deignan, H.G., 1964. Subfamily Timaliinae. In: E. Mayr & R.A. Paynter (eds.): Check-list of birds of theworld 10: 240-427.
Delacour, J., 1942-1943. The bush-warblers of the genera Cettia and Bradypterus with notes on allied
genera and species.— Ibis (14) 6: 509-519; 85: 27-40.
Delacour, J., 1946. Notes on the taxonomy of the birds of Malaysia.— Zoologica 31: 1-8.
Delacour, J., 1947a. The subspecific name of the Malayan Green Imperial Pigeon (Docula aenea).— Auk
64: 129.
Delacour, J., 1947b. Birds of Malaysia. Macmillan, New York: xvi + 382 p.
Delacour, J., 1954. The waterfowl of world, 1. Country Life, London: 284 p., XV pl. + frontesp.
Delacour, J., 1956. The waterfowl of the world 2. Country Life, London: 232 p., XXIII pl. + frontesp.
Delacour, J., 1966. Guide des oiseaux de la Nouvelle-Calédonie et de ses dépendances. Neuchâtel: 172 p.
Delacour, J. & E. Mayr, 1946. Birds of the Philippines. Macmillan, New York: xv + 309 p.
Diamond, A.W., 1987. Studies of Mascarene island birds. Cambridge: 458 p.
Diamond, J.M., 1972. Avifauna of the eastern highlands of New Guinea. Publ. Nuttall Orn. Cl. 12: vii +438 p.
Dickinson, E.C., 1989. A review of smaller Philippine swiftlets of the genus Collocalia.— Forktail 5:
23-34.

236

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Dickinson, E.C., R.S. Kennedy & K.C. Parkes, 1991. The birds of the Philippines.— B.O.U. Check-list No.
12: 507 p.
Duncan, F.M., 1937. On the dates of publication of the Society’s ‘Proceedings’, etc.— Proc. Zool. Soc.
Lond. 107A: 71-84.
duPont, J.E., 1971a. Notes on Philippine birds (No. 1).— Nemouria 3: 1-6.
duPont, J.E., 1971b. Philippine birds. Delaware Mus. Monogr. 2: × + 480 p.
Eck, S., 1977. Ergänzendes über die Vögel der Insel Taliabu, Sula-Inseln.— Zool. Abh. Dresden 34: 127133.
Eck, S. & H. Busse, 1977. Eulen. 2., erg. Aufl. Neue Brehm-Bücherei 469: 196 p.
Erritzoe, J., 1995. A small collection of birds from the Philippines with notes on body mass, distribution
and habitat.— Nemouria 40: 1-15.
Escott, C.J. & D.A. Holmes, 1980. The avifauna of Sulawesi, Indonesia: faunistic notes and additions.—
Bull. Brit. Orn. Cl. 100: 189-194.
Eskell, R. & S. Garnett, 1979. Notes on the colours of the legs, wings and flanks of the Dusky Moorhen
Gallinula tenebrosa.— Emu 79: 143-146.
Etchécopar, R.D. & F. Hüe, 1978. Les oiseaux de Chine. Non passereaux. Paris: 586 p.
Finsch, O., 1868. Die Papageien 2. Leiden: viii + 999 p.
Finsch, O., 1898a. On the identity of Muscicapula westermanni, Sharpe and M. melanoleuca, Hodgs.—
Notes Leyden Mus. 20: 93-96.
Finsch, O., 1898b. Ueber Scops magicus (S. Müll.) und die verwandten Arten.— Notes Leyden Mus. 20:
163-184, pl. 9, 10.
Finsch, O, 1900. Zur Catalogisirung der ornithologischen Abtheilung.— Notes Leyden Mus. 22: 75-161.
Finsch, O., 1901a. Ueber eine dritte Sendung Vogelbälge aus Central-Borneo (Mahakkam), gesammelt
von Herrn Dr A.W. Nieuwenhuis.— Notes Leyden Mus. 22: 163-178.
Finsch, O., 1901b. Zur Catalogisirung der Ornithologischen Abtheilung V.— Notes Leyden Mus. 22: 193224.
Finsch, O., 1901c. Systematische Uebersicht der Vögel der Südwest-Inseln.— Notes Leyden Mus. 22:
225-309.
Finsch, O., 1905. Dr A.W. Nieuwenhuis’: Forschungsreisen in Niederländisch Borneo. Ornithologische
Ergebnisse, hauptsächlich vom oberen Mahakam und Kajan.— Notes Leyden Mus. 26: 1-154.
Follett, W.I. & L.J. Dempster, 1963. Request for amendment of the International Code of Zoological Nomenclature to provide a single gender for all generic names ending in -ops.— Bull. Zool. Nomencl.
20: 73.
Ford, J.R., 1981. Hybridization and migration in Australian populations of the Little and Rufous-breasted Bronze-Cuckoos.— Emu 81: 209-222.
Ford, J.R., 1983. Taxonomic notes on some mangrove-inhabiting birds in Australia.— Rec. W. Austr.
Mus. 10: 381-415.
Forshaw, J.M., 1973. Parrots of the world. Lansdowne, Melbourne: 584 p.
Forshaw, J.M., 1983. Kingfishers and related birds. Alcedinidae: Ceryle to Cittura. Lansdowne Ed., Melbourne, etc.: 1-284, inc. pl. I-XXVI.
Forshaw, J.M., 1985. Kingfishers and related birds. Alcedinidae: Halcyon to Tanysiptera. Lansdowne Ed.,
Sydney, etc.: 285-564, incl. pl. XXVII-LII.
Forshaw, J.M., 1993. Kingfishers and related birds. Leptosomatidae, Coraciidae, Upupidae, Phoeniculidae. Lansdowne Ed., Sydney, etc.: 1-252, incl. pl. LXXIX-C.
Friedmann, H., 1968. The evolutionary history of the avian genus Chrysococcyx. U. S. Nat. Mus. Bull. 265:i-viii, 1-137.
Frith, H.J. (editor), 1976. Complete book of Australian birds. Reader’s Digest, Sydney: 615 p.
Frith, H.J. (editor), 1982. Pigeons and doves of Australia. Rigby, Adelaide, etc.: 304 p.
Frith, H.J. & W.B. Hitchcock, 1974. Birds. In H.J. Frith & J.H. Calaby (eds.): Fauna survey of the Port
Essington district, Cobourg Peninsula, Northern Territory of Australia.— CSIRO Div. Wildl. Res.
Techn. Pap. 28: 109-178.
Galbraith, I.C.J., 1956. Variation, relationships and evolution in the Pachycephala pectoralis superspecies(Aves, Muscicapidae).— Bull. Brit. Mus. (Nat. Hist.), Zool. 4: 131-222.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

237

Galbraith, I.C.J., 1967. The Black-tailed and Robust Whistlers Pachycephala melanura as a species distinct
from the Golden Whistler P. pectoralis.— Emu 66: 289-294.
Galbraith, I.C.J., 1974. Pachycephalidae. In B.P. Hall (ed.): Birds of the Harold Hall Australian expeditions, 1962-70: 244-264.
Gill, B.J., 1983. Morphology and migration of Chrysococcyx lucidus, an Australasian cuckoo.— N.Z.J.
Zool. 10: 371-381.
Gill, F.B., 1973. Intra-island variation in the Mascarene white-eye Zosterops borbonica. Orn. Monogr.
(A.O.U.) 12: 66 p.
Glutz von Blotzheim, U.N., 1975. Handbuch der Vögel Mitteleuropas 6. Akademische Verlagsgesellschaft, Wiesbaden: 840 p.
Glutz von Blotzheim, U.N., 1982. Handbuch der Vögel Mitteleuropas 8/II. Akademische Verlagsgesellschaft, Wiesbaden: 709-1270.
Glutz von Blotzheim, U.N., 1985. Handbuch der Vögel Mitteleuropas 10/II. Akademische Verlagsgesellschaft, Wiesbaden: 509-1184.
Glutz von Blotzheim, U.N., 1993. Handbuch der Vögel Mitteleuropas 13/III. AULA-Verlag, Wiesbaden:
1367-2178.
Goodwin, D., 1970. Pigeons and doves of the world. (2nd. ed.). London: vi + 446 p.
Goodwin, D., 1982. Estrildid finches of the world. London & Oxford: 328 p.
Gould, J., 1838a. Synops. Birds Austr. III: pl. with text.
Gould, J., 1838b. Characters of a large number of new species of Australian birds.— Proc. Zool. Soc.
London 5 (1837): 138-157.
Gould, J., 1848. The birds of Australia I. Author/London: no pagination.
Gould, J., 1860. The birds of Asia, vol. I. Author/London: no pagination.
Gould, J., 1865. Handbook of the birds of Australia I. Author/London: viii + 636 p.
Green, R.H., 1977. Birds of Tasmania. Launceston: xiv + 68 p.
Green, R.H., 1993. Birds of Tasmania (4th ed.). Potoroo Publishing, Launceston: xvi + 84 p.
Greenway, J.C., 1962. Family Oriolidae. In E. Mayr & J.C. Greenway (eds.): Check-list of birds of theworld 15: 122-137.
Greenway, J.C., 1973. Type specimens of birds in the American Museum of Natural History. Part 1.
Tinamidae...Rallidae.— Bull. Amer. Mus. Nat. Hist. 150: 207-345.
Greenway, J.C., 1978. Type specimens of birds in the American Museum of Natural History. Part 2.
Otididae... Picidae.— Bull. Amer. Mus. Nat. Hist. 161: 1-305.
Griffin, J., 1964. Comments on the gender of generic names ending in -ops.— Bull. Zool. Nomencl. 21:
217-218.
Gyldenstolpe, N., 1955. Birds collected by Dr Sten Bergman during his expedition to Dutch New Guinea
1948-1949.— Ark. f. Zool. (2) 8: 183-397, pl. I-VI.
Hancock, J. & H. Elliott, 1978. The herons of the world. London Editions: 304 p.
Hancock, J. & J. Kushlan, 1984. The herons handbook. Croom Helm, London & Sydney: 288 p.
Hancock, J.A., J.A. Kushlan & M.P. Kahl, 1992. Storks, ibises and spoonbills of the world. Academic
Press, London etc.: vi + 385 p.
Harrison, C.J.O. & S.A. Parker, 1967. The eggs of Woodford’s Rail, Rouget’s Rail, and the Malayan Banded Crake.— Bull. Brit. Orn. Cl. 87: 14-16.
Hartert, E., 1896. An account of the collections of birds made by Mr William Doherty in the Eastern Archipelago.— Novit. Zool. 3: 537-590, pl. xi. xii.
Hartert, E., 1897a. Descriptions of seven new species of birds and one new subspecies from Flores, andof one new subspecies from Djampea, all collected by Mr Alfred Everett.— Novit. Zool. 4: 170-172.
Hartert, E., 1897b. On the birds collected by Mr Everett in south Flores. Part I.— Novit. Zool. 4: 513528.
Hartert, E., 1898a. On the birds collected by Mr Everett in south Flores. Part II.— Novit. Zool. 5: 42-50.
Hartert, E., 1898b. List of birds collected in Timor by Mr Alfred Everett.— Novit. Zool. 5: 111-124.
Hartert, E., 1898c. On the birds of Lomblen, Pantar, and Alor.— Novit. Zool. 5: 455-465.
Hartert, E., 1898d. Account of the birds collected in Sumba by Alfred Everett and his native hunters.—
Novit. Zool. 5: 466-476.

238

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Hartert, E., 1904. The birds of the South-West Islands Wetter, Roma, Kisser, Letti and Moa.— Novit.
Zool. 11: 174-221.
Hartert, E., 1906. On the birds of the island of Babber.— Novit. Zool. 13: 288-302.
Hartert, E., 1922. Die Vögel der paläarktischen Fauna III (Heft 18, 19): 2149-2328, I-XII.
Hartert, E., 1924. Notes on some birds from Buru.— Novit. Zool. 31: 104-111.
Hartert, E., 1929. Birds collected during the Whitney South Sea expedition. VIII. Notes on birds from theSolomon Islands.— Amer. Mus. Novit. 364:
Hartert, E., 1932. Liste der Vögel aus Neuguinea im Buitenzorger Museum in Java.— Nova Guinea 15:
435-484.
Hartert, E., & A.T. Goodson, 1918. Notes on pigeons.— Novit. Zool. 25: 346-348.
Hartert, E. & F. Steinbacher, 1938. Die Vögel der Paläarktischen Fauna, Ergänzungsband (Heft 6-7): 481602.
Harvey, W.G. & D.A. Holmes, 1976. Additions to the avifaunas of Sumatra and Kalimantan, Indonesia.— Bull. Brit. Orn. Cl. 96: 90-92.
Hassell, C., 1998. Arctic Warbler at Broome: first mainland record.— W. Austr. Bird Notes 85: 1-2.
Hellebrekers, W.Ph.J. & A. Hoogerwerf, 1967. A further contribution to our oological knowledge of theisland of Java (Indonesia).— Zool. Verh. 88: 1-164.
Hellmayr, C.E., 1914. Die Avifauna von Timor. In Haniel, Zoologie von Timor 1: i-vi, 1-112, Taf. 1.
Hemming, F. (ed.), 1953. Copenhagen decisions on zoological nomenclature: xxix + 135 p.
Heurn, W.C. van, 1931. Reis-indrukken van een bioloog op Timor en Flores.— Tijdschr. K.N.A.G. (2)48:
924-930.
Heurn, W.C. van, 1932. Vluchtige zoölogische waarnemingen, gedaan tijdens een kort verblijf op Timoren Flores, April-Mei, 1930.— Nat. Tijdschr. Ned. Ind. 92: 64-84.
Higgins, P.J. & S.J.J.F. Davies (eds.), 1996. Handbook of Australian, New Zealand & Antarctic birds, 3 Snipe to pigeons. Oxford University Press, Melbourne: 1028 p., 60 pls.
Holmes, D.A., 1977. Faunistic notes and further additions to the Sumatran avifauna.— Bull. Brit. Orn.
Cl. 97: 68-71.
Holsworth, W.N., 1965. Nesting success of the Osprey on Rottnest Island, Western Australia.— W. Austr. Nat. 10: 13-15.
Hoogerwerf, A., 1935. Ornithologische merkwaardigheden in de Brantas-delta.— Trop. Nat. 24: 89-96.
Hoogerwerf, A., 1937. Enkele biologische aanteekeningen over de kleine zilverreiger, Egretta garzetta
nigripes.— Limosa 10: 1-11.
Hoogerwerf, A., 1939. Naar den Vuurberg. Een bezoek aan het natuurmonument Goenoeng Api in deBanda Zee.— 3 Jaren Indisch Natuurleven, Elfde Verslag (1936-1938), Ned.-Ind. Ver. tot Natuurbesch., Batavia: 316-333.
Hoogerwerf, A., 1948a. Contribution to the knowledge of the distribution of birds on the island of
Java.— Treubia 19: 83-137.
Hoogerwerf, A., 1948b. Enkele aanvullingen op de “Avifauna van Batavia en omstreken”.— Ardea 36:
52-61, pl. II.
Hoogerwerf, A., 1949a. Bijdrage tot de oölogie van Java. Buitenzorg: 279 p., xiX pl.
Hoogerwerf, A., 1949b. De avifauna van Tjibodas en omgeving (Java). Buitenzorg: 158 p., Xxi pl.
Hoogerwerf, A., 1954a. Phalacrocorax sulcirostris territori (Math.) als broedvogel in West-Java gedurende
1952.— Limosa 26 (1953): 71-79, pl. 3.
Hoogerwerf, A., 1954b. Rapport over een naar Komodo, Padar en Rintja ( Kleine Sunda Eilanden) gemaakte dienstreis van 21 Mei - 6 Juli 1953. Bogor: 263 p.
Hoogerwerf, A., 1956. Iets over de vogels van de eilanden Komodo, Padar en Rintja, het land van Varanus komodoensis.— Limosa 28 (1955): 96-112.
Hoogerwerf, A., 1962. Some ornithological notes on the smaller islands around Java (with the description of seven new subspecies).— Ardea 50: 180-206.
Hoogerwerf, A., 1963a. A new race of the Spotted Munia, Lonchura punctulata (Linn.).— Bull. Brit. Orn.
Cl. 83: 36-40.
Hoogerwerf, A., 1963b. Notes on Ducula aenea (Linn.), particularly the population of the Kangean Archipelago, and Ducula problematica Rensch from Sumba.— Bull. Brit. Orn. Cl. 83: 120-125.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

239

Hoogerwerf, A., 1963c. Notes on Ducula rosacea (Temminck) living on the Karimundjawa and Kangean
Archipelagos.— Bull. Brit. Orn. Cl. 83: 150-152.
Hoogerwerf, A., 1963d. Notes on Indonesian birds with special reference to the avifauna of Java and thesurrounding islands (II).— Treubia 26: 133-157.
Hoogerwerf, A., 1964. Notes sur le gobe-mouches á nuque noire Hypothymis azurea notamment dessous-espèces javana et prophata de l’Archipel Indonesien.— L’Ois. & Rev. Fr. d’Orn. 34: 210-219.
Hoogerwerf, A., 1965. Notes on Indonesian birds with special reference to the avifauna of Java and thesurrounding small islands (III).— Treubia 26: 211-291.
Hoogerwerf, A., 1966. On some strange birds from the islands of Komodo and Rintja between Sumbawa
and Flores (Lesser Sunda Islands).— Misc. Rep. Yamashina Inst. 4: 488-497.
Hoogerwerf, A., 1969/1971. On the ornithology of the rhino sanctuary Udjung Kulon in West Java (Indonesia).— Nat. Hist Bull. Siam Soc. 23: 9-65, 447-500; 24: 79-135.
Hooijer, D.A., 1975. Quaternary mammals west and east of Wallace’s Line.— Neth. J. Zool. 25: 46-56.
Horsfield, T. & F. Moore, 1858. A catalogue of the birds in the museum of the Hon. East-India Company
II. London: xix-xxx, 453-752, i-v, i-ix.
Hoyo, J. del, A. Elliott & J. Sargatal (eds), 1997. Handbook of the birds of the world, 4. Lynx Edicions,
Barecelona: 1-679.
Hoyo, J. del, A. Elliott & J. Sargatal (eds), 1999. Handbook of birds of the world, 5. Lynx Edicions, Barecelona: 1-759.
ICZN, 1961. Code international de nomenclature zoologique / International code of zoological nomenclature: xvii + 176 p.
ICZN, 1985. Code international de nomenclature zoologique / International code of zoological nomenclature (3. ed.): xx + 338 p.
Janssen, W., ca. 1953. Peta Daerah Manggarai, Flores-Barat, dengan keterangan-keterangan.
Jany, E., 1955. Neuer Fund von Porzana fusca fusca (L.) auf Flores.— J. f. Orn. 96: 220.
Johnsgard, P., 1979. Order Anseriformes. In E. Mayr & G.W. Cottrell (eds.): Check-list of birds of theworld (2nd ed.) 1: 425-506.
Johnstone, R.E., 1981. Notes on the distribution, ecology and taxonomy of the Red-crowned Pigeon(Ptilinopus regina) and Torres Strait Pigeon (Ducula bicolor) in Western Australia.— Rec. W. Austr.
Mus. 9: 7-22.
Johnstone, R.E., 1984. Notes on the distribution and ecology of the Green-winged Pigeon Chalcophaps
indica (Linnaeus) in Western Australia, with comments on subspeciation in the Australian region.—
W. Austr. Nat. 16: 4-8.
Johnstone, R.E., 1994. Observations of seabirds and shorebirds in Nusa Tenggara (Lesser Sundas) andMoluccas (Indonesia).— W. Austr. Nat. 19: 339-349.
Johnstone, R.E., P. Jepson, S.H.M. Butchart, J.C. Lowen & D. Prawiradilaga, 1996. The birds of Sumbawa,
Moyo and Sangeang islands, Nusa Tenggara, Indonesia.— Rec. W. Austr. Mus. 18: 157-178.
Johnstone, R.E. & G. M. Storr, 1998. Handbook of Western Australian birds I. Non-Passeriformes. Western Australian Museum, Perth: x + 436 p.
Jollie, M., 1978. Phylogeny of the species of Corvus.— Biologist 60: 73-108.
Jonkers, B. & H. Roersma, 1990. New subspecies of Lonchura spectabilis from East Sepik Province, Papua
New Guinea.— Dutch Birding 12: 22-25.
Junge, G.C.A., 1936. Fauna Simalurensis - Aves.— Temminckia 1: 1-74, pl. I-III.
Junge, G.C.A., 1937a. Further notes on the birds of Simalur.— Temminckia 2: 197-202.
Junge, G.C.A., 1937b. The birds of south New Guinea. Part I. Non Passeres.— Nova Guinea (n.s.) 1: 125188, pl. I-II.
Junge, G.C.A., 1938. On a collection of birds from Enggano.— Treubia 16: 339-356.
Junge, G.C.A., 1948. Notes on some Sumatran birds.— Zool. Meded. 29: 311-326.
Junge, G.C.A., 1953. Zoological results of the Dutch New Guinea expedition, 1939, No. 5. The birds.—
Zool. Verh. 20: 1-77.
Junge, G.C.A., 1954. Ornithologisch onderzoek in de Indische Archipel.— Ardea 41: 301-336.
Junge, G.C.A., 1956. New bird records from Biak Island.— Zool. Meded. 34: 231-237.
Kate, H.F.C. ten, 1894. Verslag eener reis in de Timorgroep en Polynesië. E.J. Brill, Leiden: 311 p., 12 pl.,1 kaart.

240

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Keast, J.A., 1958. Variation and speciation in the Australian Campephagidae (Passeres).— Austr. J. Zool. 6:
248-267.
Keast, J.A., 1961. Bird speciation on the Australian Continent.— Bull. Mus. Comp. Zool. 123: 303-495.
Keve, A., 1978. Revision der Unterarten des Feldsperlings [Passer montanus (Linné, 1758)].— Zool. Abh.
Dresden 34: 245-273.
King, B., 1989. The avian genera Tesia and Urosphena.— Bull. Brit. Orn. Cl. 109: 162-166.
King, B., 1990. Metallic Pigeon Columba vitiensis sighting on Flores.— Kukila 5: 74.
Kooiman, J.G., 1940a. Zeldzame trekvogels.— Trop. Nat. 29: 122-123.
Kooiman, J.G., 1940b. Mededeelingen over het voorkomen in Oost-Java van enkele voor dit gewest nog
niet in de literatuur genoemde vogels.— Ardea 29: 98-108.
Kooij, H. van der & B. Voslamber, 1997. Aantalsontwikkeling van de Grote Zilverreiger Egretta alba in
Nederland sinds 1970 in een Europees perspectief.— Limosa 70: 119-125.
Kuroda, N., 1933. Birds of the Island of Java. I: Passeres. Tokyo: ixv, 1-370, pl. I-XIV.
Kuroda, N., 1936. Birds of the Island of Java II: Non-Passeres. Tokyo: i-vi, 371-794, pl. XV-XXXIV.
Lack, D., 1971. Ecological isolation in birds. Blackwell, Oxford & Edinburgh: xi + 404 p.
Lane, B.A., 1987. Shorebirds in Australia: ix + 187 p.
Lansdown, R.V., 1987. Recent extensions of breeding range of the Yellow Bittern Ixobrychus sinensis.—
Forktail 3: 61-63.
Lansdown, R.V., 1992. The status and conservation of the Sumatran Heron Ardea sumatrana.— Heron Conservation Newsletter 5.
LeFevre, R.H., 1962. The birds of northern Shantung Province, China. York, Pennsylvania: 151 p.
Lieftinck, M.A., 1950. Bij het nest van Monarcha trivirgata nigrimentum G.R. Gray.— Ardea 38: 238-239, pl.
IV-VII.
Linnaeus, C., 1766. Systema naturae (ed. 12), 1. Holmiae: 532 p.
Lynes, H., 1930. Review of the genus Cisticola.— Ibis (12) 6, Suppl.: i-ii, 1-673 and Pictures.
McCracken, K.G. & F.H. Sheldon, 1998. Molecular and osteological heron phylogenesis: sources of incongruence.— Auk 115: 127-141.
McKean, J.L., 1980. The first record of the Arctic Warbler Phylloscopus borealis from Australia.— W. Austr.
Nat. 14: 200.
McKean, J.L., 1982. Some notes on the birds of Sulawesi (Celebes).— Papua New Guinea Bird Soc. Newsl.
191-192: 12-17.
McKean, J.L., 1989. Taxonomic relationships of the White-rumped Kingfisher Caridonax fulgidus.— Kukila
4: 43-44.
McKean, J. L., I. J. Mason & L. W, O’Connor, 1975. Birds not previously recorded from Timor.— Emu 75:
62-64.
Marchant, J., 1916. Alfred Russel Wallace: letters and reminiscences. London. I: xii + 320 p., II: viii + 291
p.
Marchant, S. & P.J. Higgins (eds.), 1993. Handbook of Australian, New Zealand and Antarctic birds, 2 Raptors to lapwings. Oxford University Press, Melbourne: 984 p., 68 pl.
Marin, M. & F.H. Sheldon, 1987. Some new nesting records of padi-dwelling birds in Sabah, East Malaysia(North Borneo).— Bull. Brit. Orn. Cl. 107: 23-25.
Marle, J.G. van, 1940a. Aanteekeningen omtrent de vogels van de Minahasa (N.O. Celebes).— Limosa 13:
65-70.
Marle, J.G. van, 1940b. Aanteekeningen omtrent de vogels van de Minahasa (N.O. Celebes) II.–– Limosa
13: 119-124.
Marle, J.G. van & K.H. Voous, 1988. The birds of Sumatra, an annotated check-list.— B.O.U. Check-list No.
10: 265 p.
Marshall, J.T., 1978. Systematics of smaller Asian night birds based on voice. A.O.U. Orn. Monogr. 25: i-v,
1-58.
Martens, E. von, 1876. Die Preussische Expedition nach Ost-Asien. Zool. I: i-xii, 1-412.
Mason, I.J. & R.I. Forrester, 1996. Geographical differentiation in the Channel-billed Cuckoo Scythrops novaehollandiae Latham, with description of two new subspecies from Sulawesi and the Bismarck Archipelago.— Emu 96: 217-228.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

241

Mason, I.J & J.L. McKean, 1982. Taxonomic status of the Timor cuckoo-shrike Coracina personata Müller.—
Bull. Brit. Orn. Cl. 102: 127-129.
Mathews, G.M., 1910-1912. The birds of Australia 1. Witherby & Co.,London: xiv + 301 p.
Mathews, G.M., 1912. A reference-list to the birds of Australia.— Novit. Zool. 18: 171-656.
Mathews, G.M., 1913. A list of the birds of Australia. Witherby & Co., London: xxvii + 453 p.
Mathews, G.M., 1914a. A list of the birds of Melville Island, Northern Territory, Australia.— Ibis (10) 2:
91-132.
Mathews, G.M., 1914b. On the species and subspecies of the genus Fregata.— Austral Av. Rec. 2: 117-121.
Mathews, G.M., 1915-1916. The birds of Australia 5.— Witherby & Co., London: xi + 440 p.
Mathews, G.M., 1918. On Pachycephala melanura Gould. –– Austral Av. Rec. 3: 134-138.
Mathews, G.M., 1927. Systema avium Australasianarum. Part I. B.O.U., London: i-xvii, 1-426.
Mathews, G.M., 1930. Systema avium Australasianarum. Part II. Taylor and Francis, London: i-iv, 4271047.
Mauersberger, G., 1983. Ungelöste taxonomische Probleme der mongolischen Avifauna.— Mitt. Zool.
Mus. Berlin 59, Suppl.: Ann. Orn. 7: 47-83.
Mayr, E., 1937. Birds collected during the Whitney South Sea expedition XXXIII. Notes on New Guinea
birds. I.— Amer. Mus. Novit. 1915: 1-19.
Mayr, E., 1941a. Ueber einige Raubvügel der Kleinen Sunda-Inseln.— Orn. Mber. 49: 42-47.
Mayr, E., 1941b. List of New Guinea birds. American Museum of Natural History, New York: xi + 260 p.
Mayr, E., 1943. Notes on Australian birds (II).— Emu 43: 3-17.
Mayr, E., 1944a. The birds of Timor and Sumba.— Bull. Amer. Mus. Nat. Hist. 83: 123-194.
Mayr, E., 1944b. Timor and the colonization of Australia by birds.— Emu 44: 113-130.
Mayr, E., 1945. Birds collected during the Whitney South Sea expedition. 55. Notes on the birds of northern
Melanesia. 1.— Amer. Mus. Novit. 1294: 1-12.
Mayr, E., 1967. Family Zosteropidae. In R.A. Paynter (ed.): Check-list of birds of the world 12: 289-326.
Mayr, E., 1979. Family Pittidae. In M.A. Traylor (ed.): Check-list of birds of the world 8: 310-329.
Mayr, E. & D. Amadon, 1947. A review of the Dicaeidae.— Amer. Mus. Novit. 1360: 1-32.
Mayr, E., R.A. Paynter & M.A. Traylor, 1968. Family Estrildidae. In R.A. Paynter (ed.): Check-list of birdsof the world 14: 306-389.
Mayr, E. & F. Vuilleumier, 1983. New species of birds described from 1966 to 1975.— J. f. Orn. 124: 217232.
Medway, Lord & D.R. Wells, 1976. The birds of the Malay Peninsula 5. Witherby, London: xxxi + 448 p., 24
pl.
Mees, G.F., 1954. De geluiden van Cacomantis variolosus sepulcralis (S. Müller) en Cacomantis merulinus lanceolatus (S. Müller). –– Ardea 42: 346–349.
Mees, G.F., 1957. A systematic review of the Indo-Australian Zosteropidae (Part I).— Zool. Verh. 35: 1204.
Mees, G.F., 1961a. A systematic review of the Indo-Australian Zosteropidae (Part II).— Zool. Verh. 50: 1168, pl. I-IV.
Mees, G.F., 1961b. An annotated catalogue of a collection of bird-skins from West Pilbara, Western Australia.— J. Roy. Soc. W. Austr. 44: 97-143.
Mees, G.F., 1964a. A revision of the Australian owls (Strigidae and Tytonidae).— Zool. Verh. 65: 1-62.
Mees, G.F., 1964b. Notes on two small collections of birds from New Guinea.— Zool. Verh. 66: 1-37.
Mees, G.F., 1964c. Geographical variation and distribution of some birds from Western Australia.— J. Roy.
Soc. W. Austr. 47: 91-96.
Mees, G.F., 1965. The avifauna of Misool.— Nova Guinea, Zool. 31: 139-203, pl. III-VI.
Mees, G.F., 1969. A systematic review of the Indo-Australian Zosteropidae (Part III).— Zool. Verh. 102: 1390.
Mees, G.F., 1970. Notes on some birds from the island of Formosa (Taiwan).— Zool. Meded. 44: 285-304.
Mees, G.F., 1971a. The Philippine subspecies of Centropus bengalensis (Gmelin) (Aves, Cuculidae). ––Zool. Meded. 45: 189-191.
Mees, G.F., 1971b. Systematic and faunistic remarks on birds from Borneo and Java.— Zool. Meded. 45:
225-244.

242

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Mees, G.F., 1972. Die Vögel der Insel Gebe.— Zool. Meded. 46: 69-89.
Mees, G.F., 1973. Description of a new member of the Monarcha trivirgata-group from Flores, Lesser
Sunda Islands.— Zool. Meded. 46: 179-181.
Mees, G.F., 1975a. Status of Ardea novaehollandiae in the Lesser Sunda Islands.— Bull. Brit. Orn. Cl. 95:
128.
Mees, G.F., 1975b. A list of the birds known from Roti and adjacent islets (Lesser Sunda Islands).— Zool.
Meded. 49: 115-140.
Mees, G.F., 1976. Some birds recorded from Timor by Salomon Müller in 1828-29.— Emu 76: 150-151.
Mees, G.F., 1977a. Geographical variation of Caprimulgus macrurus Horsfield (Aves, Caprimulgidae).—Zool. Verh. 155: 1-47.
Mees, G.F., 1977b. The subspecies of Chlidonias hybridus (Pallas), their breeding distribution and migrations (Aves, Laridae, Sterninae).— Zool. Verh. 157: 1-64.
Mees, G.F., 1981. The sparrow-hawks (Accipiter) of the Andaman Islands.— J. Bombay Nat. Hist. Soc. 77(1980): 371-412.
Mees, G.F., 1982a. Birds from the lowlands of southern New Guinea (Merauke and Koembe).— Zool.
Verh. 191: 1-188.
Mees, G.F., 1982b. Bird records from the Moluccas.— Zool. Meded. 56: 91-111.
Mees, G.F., 1982c. Review. [R. Schodde & I.J. Mason: Nocturnal birds of Australia, 1981].— Emu 82: 182184; Corrigendum, Emu 84: 63.
Mees, G.F., 1984a. A new subspecies of Accipiter virgatus (Temminck) from Flores, Lesser Sunda Islands,
Indonesia (Aves: Accipitridae).— Zool. Meded. 58: 313-321.
Mees, G.F., 1984b. Boekbespreking. [Cramp, Handbook of the birds of Europe, etc. III, 1983].— Ardea 72:
243-245.
Mees, G.F., 1985. Caprimulgus macrurus Horsfield and related forms, a re-evaluation.— Proc. Kon. Ned.
Akad. Wetensch. C 88: 419-428.
Mees, G.F., 1986. A list of the birds recorded from Bangka Island, Indonesia.— Zool. Verh. 232: 1-176.
Mees, G.F., 1989. Remarks on the ornithological parts of Horsfield’s ‘Zoological Researches in Java’.—
Proc. Kon. Ned. Akad. Wetensch. C 92: 367-378.
Mees, G.F., 1990. Historisches über Anthus gustavi Swinhoe.— Mitt. Zool. Mus. Berlin 66, Suppl. Ann.
Orn. 14: 137-141.
Mees, G.F., 1992. Review. [E.C. Dickinson, R.S. Kennedy & K.C. Parkes: The birds of the Philippines,
1991].— Ibis 134: 385-386.
Mees, G.F., 1994. Vogelkundig onderzoek op Nieuw–Guinea in 1828. –– Zool. Bijdr. 40: 1-64.
Mees, G.F., 1996. Geographical variation of birds in Java. Publ. Nuttall Orn. Cl. 126: viii + 119 p.
Meinertzhagen, R., 1912. On the birds of Mauritius.— Ibis (9) 6: 82-108.
Meinertzhagen, R., 1926. Introduction to a review of the genus Corvus.— Novit. Zool. 33: 57-121.
Meise, W., 1929. Die Vögel von Djampea und benachbarten Inseln nach einer Sammlung Baron Plessens(I).— J.f. Orn. 77: 431-480.
Meise, W., 1941. Ueber die Vogelwelt von Noesa Penida bei Bali nach einer Sammlung von Baron Viktor
von Plessen.— J.f. Orn. 89: 345-376.
Meyer, A.B., 1894. Beschreibung einiger neuen Vögel aus dem Ostindischen Archipel.— J.f. Orn. 42: 8983.
Meyer, A.B. & L.W. Wiglesworth, 1898. The birds of Celebes. Friedländer/Berlin: xxxii + 130 + 962 p.
Mivart, St.G., 1896. A monograph of the lories or brush-tongued parrots, composing the family Loriidae.
R.H. Porter, London: liii + 193 p., Lxi pls.
Moeliker, C.W. & C.J. Heij, 1995. The resdiscovery of Monarcha boanensis (Aves: Monarchidae) from
Boano Island, Indonesia.— Deinsea, Rotterdam 2: 123-143.
Moeliker, C.W. & C.J. Heij, 1995. Black-chinned monarchs Monarcha boanensis on Boano Island.— OBC
Bull. 24: 49-52.
Moreau, R.E., 1967. Family Zosteropidae (African and Indian Ocean taxa). In R.A. Paynter (ed.): Checklist of birds of the world 12: 326-337.
Müller, S., 1842/1847. Bijdragen tot de kennis van Timor en eenige naburige eilanden.— Verh. Nat. Gesch.
Ned. Overz. Bez., Land- en Volkenkunde: 131-320.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

243

Musschenbroek, S.C.J.W. van, 1883. Dagboek van Dr H.A. Bernstein’s laatste reis, van Ternate naar
Nieuw-Guinea, Salawati en Batanta, 17 October 1864-19 April 1865. ‘s-Gravenhage: 258 p.
Nadler, T., 1976. Die Zwergseeschwalbe. Neue Brehm-Bücherei 495: 136 p.
Nicoll, M.J., 1906. On the birds collected and observed during the voyage of the ‘Valhalla’, R.Y.S., from
November 1905 to May 1906.— Ibis (8) 6: 666-712, pl. XXI.
Oberholser, H.C., 1909. A revision of the kingfisher genus Ramphalcyon (Pelargopsis).— Proc. U.S. Nat.
Mus. 35: 657-680.
Oberholser, H.C., 1911. A monograph of the flycatcher genera Hypothymis and Cyanonympha.— Proc.
U.S. Nat. Mus. 39: 585-615.
Olsen, P. & T.G. Marples, 1993. Geographic variation in egg size, clutch size and date of laying of Australian raptors (Falconiformes and Strigiformes).— Emu 93: 167-179.
Oort, E.D. van, 1907. Catalogue ostéologique des oiseaux.— Mus. Hist. Nat. Pays-Bas 10(1): i-viii, 1-384.
Oort, E.D. van, 1910. List of a collection of birds from western Java and from Krakatau.— Notes LeydenMus. 32: 105-166.
Oortwijn, R.G.M., 1987. Geographic variation and subspecies of Corvus enca (Horsfield, 1821).— Zool.
Meded. 61: 31-51.
Ottow, J. & J.A.J. Verheijen, 1969. Zur Lebensweise der Kuckucke von Flores.— J.f. Orn. 110: 27-29.
Parker, S.A., 1981. Prolegomenon to further studies in the Chrysococcyx ‘malayanus’ group (Aves, Cuculidae).— Zool. Verh. 187: 1-56, pl. 1-2.
Parker, S.A, H.J. Eckert & G.B. Ragless, 1985. An annotated checklist of the birds of South Australia. Part
2A: Waterfowl. Adelaide: viii + 25 p.
Parkes, K.C., 1958. Taxonomy and nomenclature of three species of Lonchura (Aves: Estrildinae).— Proc.
U.S. Nat. Mus. 108: 279-293.
Parkes, K.C., 1971. Taxonomic and distributional notes on Philippine birds.— Nemouria 4: 1-67.
Payne, R.B., 1979. Family Ardeidae. In E. Mayr & G.W. Cottrell (eds.), Check-list of birds of the world 1(2nd ed.): 193-244.
Payne, R.B. & C.J. Risley, 1976. Systematic and evolutionary relationships among the herons (Ardeidae).— Misc. Publ. Mus. Zool. Univ. Michigan 150: iv + 115 p.
Paynter, R.A., 1963. Birds from Flores, Lesser Sunda Islands.— Breviora 182: 1-5.
Peters, J.L., 1937. Check-list of birds of the world 3: xiii + 311 p.
Peters, J.L., 1940. Check-list of birds of the world 4: xii + 291 p.
Peters, J.L., 1960. Family Hirundinidae. In E. Mayr & J.C. Greenway (eds.), Check-list of birds of theworld 9: 80-129.
Peters, J.L. & E. Mayr, 1960. Family of Campephagidae. In E. Mayr & J.C. Greenway (eds.), Check-list ofBirds of the world 9: 167-221.
Pfeffer, P., 1958. Répartition écologique de l’avifaune des îles Flores, Rintja et Komodo (Petites îles de laSonde).— Ois. & Rev. Fr. d’Orn. 28: 59-72.
Pfeffer, P., 1960. Etude d’une collection d’oiseaux de Borneo.— Ois. & Rev. Fr. d’Orn. 30: 191-218.
Pfeffer, P., 1965. Auf den Inseln des Drachen. Schwabenverlag/Stuttgart: 158 p.
Pilgrim, J., 1998. Preliminary report on the Flores ‘97 expedition.— Ibis 140: 559.
Pulle, A.W., 1910. In memorian Gijsbertus Adriaan Johan van der Sande, Officier van Gezondheid 1e
klasse.— Marineblad, Febr. 1910: 586a-d. (personalia 65).
Rand, A.L., 1942. Results of the Archbold expeditions. No. 43. Birds of the 1938-1939 New Guinea expedition.— Bull. Amer. Mus. Nat. Hist. 79: 425-515.
Rand, A.L., 1970. Species formation in the blue monarch flycatchers genus Hypothymis.— Nat. Hist. Bull.
Siam Soc. 23: 353-365.
Rand, A.L. & E.T. Gilliard, 1967. Handbook of New Guinea birds. Weidenfeld & Nicholson, London: ×+ 612 p.
Rasmussen, P.C., 1998. A new scops-owl from Great Nicobar Island.— Bull. Brit. Orn. Cl. 118: 141-153.
Reichenow, A., 1882. Conspectus Psittacorum. Systematische Übersicht aller bekannten Papageienarten.
Berlin: 234 p.
Rensch, B., 1928a. Neue Vogelrassen von den Kleinen Sunda-Inseln I.— Orn. Mber. 36: 6-10.
Rensch, B., 1928b. Neue Vogelrassen von den Kleinen Sunda-Inseln II.— Orn. Mber. 36: 47-49.

244

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Rensch, B., 1928c. Neue Vogelrassen von den Kleinen Sunda-Inseln III.— Orn. Mber. 36: 80-81.
Rensch, B., 1928d. Die Kleinen Sunda-Inseln als indo-australisches Mischgebiet.— Sitzb. Ges. naturf. Fr.
Berlin (1928): 1-7.
Rensch, B., 1930. Eine biologische Reise nach den Kleinen Sunda-Inseln. Borntraeger, Berlin: xii + 236
p.
Rensch, B., 1931a. Die Vogelwelt von Lombok, Sumbawa und Flores.— Mitt. Zool. Mus. Berlin 17: 451637.
Rensch, B., 1931b. Ueber einige Vogelsammlungen des Buitenzorger Museums von den Kleinen SundaInseln.— Treubia 13: 371-400.
Rensch, B., 1936. Die Geschichte des Sundabogens. Borntraeger, Berlin: viii + 318 p.
Restall, R.L., 1992. A new sub-species of the Spice Bird Lonchura punctulata from Borneo.— Avicult. Mag.
98: 115-118.
Restall, R.L., 1995. Proposed additions to the genus Lonchura (Estrildinae).— Bull. Brit. Orn. Cl. 115: 140157.
Ridgway, R., 1914. The birds of North and Middle America. Part VI.— U.S. Nat. Mus. Bull. 50(6): xx +882 p., XXXVI pl.
Riley, J.H., 1918. Two new genera and eight new birds from Celebes. –– Proc. Biol. Soc. Wash. 31: 155–
160.
Ripley, S.D., 1941a. Notes on a collection of birds from northern Celebes.— Occ. Pap. Boston Soc. Nat.
Hist. 8: 343-358.
Ripley, S.D., 1941b. Notes on the genus Coracina.— Auk 58: 381-395.
Ripley, S.D., 1942a. A review of the species Anas castanea.— Auk 59: 90-99.
Ripley, S.D., 1942b. The species Eurystomus orientalis.— Proc. Biol. Soc. Wash. 55: 169-176.
Ripley, S.D., 1944. The bird fauna of the West Sumatra islands.— Bull. Mus. Comp. Zool. 94: 305-430, pl.
1-2.
Ripley, S.D., 1952. A note on the species Muscicapa westermanni.— Proc. Biol. Soc. Wash. 65: 71-74.
Ripley, S.D., 1959. Birds from Djailolo, Halmahera.— Postilla 41: 1-8.
Ripley, S.D., 1977. Rails of the world. Boston: 406 p., pls.
Ripley, S.D. & B.M. Beehler, 1987. Species status of the Malaysian three-toed kingfishers (Ceyx) – a re-assessment.— Bull. Brit. Orn. Cl. 107: 145-151.
Roberts, T.J. & B. King, 1986. Vocalizations of the owls of the genus Otus in Pakistan.— Ornis Scand. 17:
299-305.
Robinson, H.C. & C.B. Kloss, 1921-1924. The birds of south-west and peninsular Siam.— J. Nat. Hist.
Soc. Siam 5: 1-397.
Robinson, H.C. & C.B. Kloss, 1924a. On a large collection of birds chiefly from West Sumatra made by
Mr E. Jacobson.— J. Fed. Malay St. Mus. 11: 189-347.
Robinson, H.C. & C.B. Kloss, 1924b. A nominal list of the birds collected in Java.— Treubia 5: 267-298.
Robinson, H.C. & C.B. Kloss, 1927. Some notes on bird names. –– J. Fed. Malay St. Mus. 13: 213-216.
Rosenberg, H. von, 1881. Ein Jäger-Eldorado.— Zool. Garten 22: 164-168.
Rothschild, W., 1897. Description of a new hill-wren from Flores.— Novit. Zool. 4: 168. [Pnoepyga everetti].
Rothschild, W., 1927. Corrections and criticisms to the article on the avifauna of Yunnan.— Novit. Zool.
33: 398-400.
Rüppell, E., 1845. Systematische Uebersicht der Vögel Nord-Ost-Afrika’s. Frankfurt a.M.: viii + 140 p.,
50 pl.
Sabrosky, C.W., 1964. Comments on the gender of generic names ending in -ops.— Bull. Zool. Nomencl.
21: 215-217.
Salomonsen, F., 1960. Notes on flowerpeckers (Aves, Dicaeidae) 2. The primitive species of the genus
Dicaeum.— Amer. Mus. Novit. 1991: 1-38.
Salomonsen, F., 1961. Notes on flowerpeckers (Aves, Dicaeidae) 4. Dicaeum igniferum and its derivatives.— Amer. Mus. Novit. 2057: 1-35.
Salomonsen, F., 1983. Revision of the Melanesian swiftlets (Apodidae, Aves) and their conspecificforms in the Indo-Australian and Polynesian region.— Kong. Danske Vidensk. Selsk., Biol. Skr. 23(5): 1-112.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

245

Salvadori, T., 1874. Catalogo sistematico degli uccelli di Borneo.— Ann. Mus. Genova 5 i-lii, 1-431.
Salvadori, T., 1879. Prodromus ornithologiae Papuasiae et Moluccarum. VII. Passeres. HirundinidaeMuscicapidae.— Ann. Mus. Genova 14: 490-508.
Salvadori, T., 1881. Ornitologia della Papuasia e delle Molucche 2: xvi + 707 p.
Salvadori, T., 1891. Catalogue of the birds in the British Museum 20: xvii + 659 p., XVIII pl.
Salvadori, T., 1893. Catalogue of the birds in the British Museum 21: xix + 676p., XV pl.
Salvadori, T., 1895. Catalogue of the birds in the British Museum 27: xv + 438 p., 19 pl.
Salvin, O., 1882. A catalogue of the collection of birds formed by the late Hugh Edwin Strickland, M.A.
University Press, Cambridge: xvi + 652 p.
Schauensee, R.M. de, 1957. One some avian types, principally Gould’s, in the collection of the Academy.— Proc. Acad. Nat. Sci. Philad. 109: 123-246.
Schauensee, R.M. de & J.E. du Pont, 1962. Birds from the Philippine Islands.— Proc. Acad. Nat. Sci.
Philad. 114: 149-173.
Schifter, H., 1992. Die Vögel aus dem Nachlaß Ferdinand Lucas Bauer im Naturhistorischen Museum
Wien und die darunder befindlichen Typen. –– Ann. Naturhist. Mus. Wien 93 B: 41–57, Taf. 1-3.
Schlegel, H., 1862. Aquilae.— Mus. Hist. Nat. Pays-Bas 2 (mon. 3): 1-24.
Schlegel, H., 1863a. Ardeae.— Mus. Hist. Nat. Pays-Bas 5 (mon. 16): 1-64.
Schlegel, H., 1863b. Alcedines.— Mus. Hist. Nat. Pays-Bas 3 (mon. 17): 1-52.
Schlegel, H., 1864a. Sternae.— Mus. Hist. Nat. Pays-Bas 6 (mon. 24): 1-44.
Schlegel, H., 1864b. Cuculi.— Mus. Hist. Nat. Pays-Bas 1 (mon. 25): 1-85.
Schlegel, H., 1864c. Scolopaces.— Mus. Hist. Nat. Pays-Bas 5 (mon. 27): 1-112.
Schlegel, H., 1864d. De vogels van Nederlandsch Indië / Les oiseaux des Indes Neérlandaises. De ijsvogels: 1-70, pl. 1-16.
Schlegel, H., 1865a. Ciconiae.— Mus. Hist. Nat. Pays-Bas 5 (mon. 28): 1-26.
Schlegel, H., 1865b. Cursores.— Mus. Hist. Nat. Pays-Bas 4 (mon. 29): 1-80.
Schlegel, H., 1866a. Anseres.— Mus. Hist. Nat. Pays-Bas 6 (mon. 31): 1-122.
Schlegel, H., 1866b. Observations zoologiques.— Ned. Tijdschr. Dierk. 3: 181-213.
Schlegel, H., 1866c. De vogels van Nederlandsch Indië / Les oiseaux des Indes Neérlandaises. De
valkvogels. Accipitres: 1-84, pl. 1-28.
Schlegel, H., 1873. Columbae.— Mus. Hist. Nat. Pays Bas 4 (mon. 35): 1-180.
Schlegel, H., 1874a. Revue de la collection des brèves (Pitta) faisant partie du Musée des Pays-Bas.—Mus. Hist. Nat. Pays-Bas 3 (mon. 37): 1-19.
Schlegel, H., 1874b. Revue de la collection des Alcedines faisant partie du Musée des Pays-Bas.— Mus.
Hist. Nat. Pays-Bas 3 (mon. 39): 1-47.
Schlegel, H., 1880. Megapodii.— Mus. Hist. Nat. Pays-Bas 8 (mon. 41): 52-86.
Schmutz, E., MS (1977). Die Vögel der Manggarai (Flores).
Schodde, R. & I.J. Mason, 1980. Nocturnal birds of Australia. Lansdowne Editions, Melbourne, etc.: 135p., ill.
Schodde, R. & I. J. Mason, 1998. Directory of Australian birds. Passerines. CSIRO Publishing, Collingwood, Vic.: x + 851 p.
Schodde, R. & G. F. van Tets, 1981. First record of the Brown Hawk-Owl Ninox scutulata from Australasia.–– Emu 81: 171.
Schönwetter, M., 1934. Vogeleier von Timor und Waigeu.— Orn. Mber. 42: 40-43.
Schönwetter, M., 1962. Handbuch der Oologie I (Lief. 6): 321-384.
Schönwetter, M., 1988. Handbuch der Oologie IV (Lief. 44): 193-384.
Severin, T., 1999. The Spice Islands voyage in search of Wallace. Abacus, London: 302 p. (reprint; firstpublished in 1997).
Sharland, M., 1958. Tasmanian birds. Angus & Robertson, Sydney, etc.: xv + 175 p.
Sharpe, R.B., 1868-1871. A monograph of the Alcedinidae. London: 120 pl. with text.
Sharpe, R.B., 1877. On birds collected by Professor J. B. Steere in the Philippine Archipelago.— Trans.
Linn. Soc. Lond., Zool. 1: 307-355, pl. 46-54.
Sharpe, R.B., 1885. Catalogue of the birds in the British Museum 10: xiii + 682 p., XII pls.
Sharpe, R.B., 1890. Catalogue of the birds in the British Museum 13: xvi + 702 p., 15 pl.

246

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Sharpe, R.B., 1892. Family Alcedinidae.— Catalogue of the birds in the British Museum 17: 93-312.
Sharpe, R.B., 1896. Catalogue of the birds in the British Museum 24: xii + 794 p., VII pl.
Sheldon, F.H., 1985. The taxonomy and biogeography of the thick-billed flowerpecker complex in Borneo.— Auk 102: 606-612.
Shelley, G.E., 1891. Family Cuculidae.— Cat. Birds Brit. Mus. 19: 209-434.
Sibley, C.G. & B.L. Monroe, Jr., 1990. Distribution and taxonomy of birds of the world: xxiv + 1111 p.
Siebers, H.C., 1930. Fauna Buruana. Aves.— Treubia 7, Suppl.: 164-303.
Simpson, K., 1972. Birds in Bass Strait. A. H. & A. W. Reed for BHP, Sydney, etc.: 112 p.
Smythies, B.E., 1957. An annotated checklist of the birds of Borneo.— Sarawak Mus. J. 7: v-xv, 523-818,
map.
Smythies, B.E., 1960. The birds of Borneo. Oliver & Boyd, Edinburgh/London: xvi + 562 p.
Smythies, B.E., 1968. The birds of Borneo (2nd ed.): xx + 593 p.
Smythies, B.E., 1981. The birds of Borneo (3rd ed., revised by the Earl of Cranbrook): xiv + 473 p.
Smythies, B.E., 1986. The birds of Burma. 3rd (revised) ed.: xxxii + 432 p., XXXII pl.
Smythies, B.E., 1999. The birds of Borneo (4th ed., revised by G.W.H. Davison). Natural History Publications (Borneo): xii + 853 p.
Snow, D.W. & R.E. Moreau, 1955. The gender of Zosterops.— Bull. Brit. Orn. Cl. 75: 44.
Sody, H.J.V., 1954. Waarnemingen over de trek van de Aziatische Watersnip, Capella stenura (Bp.).— Ardea 42: 341-343.
Somadikarta, S., 1986. Collocalia linchi Horsfield & Moore - a revision.— Bull. Brit. Orn. Cl. 106: 32-40.
Somadikarta, S. & D.A. Holmes, 1979. An influx of Australian pelicans Pelecanus conspicillatus in Indonesia.— Bull. Brit. Orn. Cl. 99: 154.
Somadikarta, S. & M. Noerdjito, 1982. The Lesser Whistling Duck Dendrocygna javanica (Horsfield) in
Flores.— Bull. Brit. Orn. Cl. 102: 94-95.
Sousa, J.A. de, 1883. Uma collecçao de aves de Timor.— Bol. Soc. Geogr., Lisboa 9: 453-459. (non vidi).
Steinbacher, J., 1979. Family Threskiornithidae. In E. Mayr & G.W. Cottrell (eds.): Check-list of birds ofthe world (2nd. ed.) 1: 253-268.
Stokes, T., 1988. A review of the birds of Christmas Island, Indian Ocean.— Austr. Nat. Parks Wildl.
Service, Occ. Pap. 16: viii + 39 p.
Stone, W. & G.M. Mathews, 1913. A list of the species of Australian birds described by John Gould, withthe location of the type-specimens.— Austral Av. Rec. 1: 129-180.
Storr, G., 1973. List of Queensland birds.— W.Austr. Mus. Spec. Publ. 5: iii + 177 p.
Storr, G.M., 1984. Revised list of Queensland birds.— Rec. W. Austr. Mus. Suppl. 19: 1-189.
Stresemann, E., 1913a. Ornithologische Miszellen aus dem indo- australischen Gebiet. II. Teil.— Novit.
Zool. 20: 289-324.
Stresemann, E., 1913b. Die Vögel von Bali.— Novit. Zool. 20: 325-387, pl. II.
Stresemann, E., 1914a. Die Vögel von Seran (Ceram).— Novit. Zool. 21: 24-153, pl. III-V.
Stresemann, E., 1914b. Was ist Collocalia fuciphaga (Thunb.)? – Verh. Orn. Ges. Bayern 12: 1-12.
Stresemann, E., 1914c. Beiträge zur Kenntnis der Avifauna von Buru.— Novit. Zool. 21: 358-400.
Stresemann, E., 1920. Avifauna Macedonica. Dultz & Co., München: xxiv + 270 p., VI Taf.
Stresemann, E., 1922. Fregata minor minor (Gm.) an der Küste von Timor.— Orn. Mber. 30: 113-114.
Stresemann, E., 1925. Beiträge zur Ornithologie der indo-australischen Region.— Mitt. Zool. Mus. Berlin
12: 177-195.
Stresemann, E., 1930. Eine Vogelsammlung vom Vulkan Papandajan (West-Java).— Treubia 12: 425-430.
Stresemann, E., 1931. Vorläufiges über die ornithologischen Ergebnisse der Expedition Heinrich 193031. V. Zur Ornithologie von Halmahera und Batjan.— Orn. Mber. 39: 167-171.
Stresemann, E., 1936. A nominal list of the birds of Celebes.— Ibis (13) 6: 356-369.
Stresemann, E., 1939. Die Vögel von Celebes. Teil I und II.— J.f. Orn. 87: 299-425.
Stresemann, E., 1940a. Die Vögel von Celebes. Teil III. Systematik und Biologie.— J.f. Orn. 88: 1-135,
389-487.
Stresemann, E., 1940b. Zur kenntnis der Wespenbussarde (Pernis).— Arch. f. Naturg. (N.F.) 9: 137-193.
Stresemann, E., 1941. Die Vögel von Celebes. Teil III. Systematik und Biologie (Schluss).— J.f. Orn. 89:
1-102.

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

247

Stresemann, E., 1951. Type localities of Australian birds collected by the ‘Expedition Baudin’ (18011803).— Emu 51: 65-70.
Stresemann, E., 1952. On the birds collected by Pierre Poivre in Canton, Manila, India and Madagascar(1751-1756).— Ibis 94: 499-523.
Stresemann, E. & D. Amadon, 1979. Order Falconiformes. In E. Mayr & G.W. Cottrell (eds.): Check-listof birds of the world 1 (2nd ed.): 271-425.
Stresemann, E. & V. Stresemann, 1966. Die Mauser der Vögel.— J. f. Orn. 107, Sonderheft: i-viii, 1-445.
Sutter, E., 1955. Über die Mauser einiger Laufhühnchen und die Rassen von Turnix maculosa und sylvatica im indo-australischen Gebiet.— Verh. Naaturf. Ges. Basel 66: 85-139.
Swann, H.K., 1922. A synopsis of the Accipitres (Diurnal birds of prey), 2nd. ed., London: viii + 233 + 1 p.
Temminck, C.J., 1839. Tableau Méthodique.— Recueil d’Ois. 1 (livr. 102): 1-109.
Thompson, M.C., 1966. Birds from North Borneo.— Univ. Kansas Publ., Mus. Nat. Hist. 17: 377-433.
Ticehurst, C.B., 1938. A systematic review of the genus Phylloscopus.— British Museum, London: viii +193 p., II pl.
Trainor, C.R., 2002a. The birds of Adonara, Lesser Sundas, Indonesia.— Forktail 18: 93-100.
Trainor, C.R., 2002b. Status and habitat associations of birds on Lembata Island, Wallacea, Indonesia,
with reference to a simple technique for avifaunal survey on small islands.— Bird Conservation
Internattional 12: 365-381.
Trainor, C.R. & Dwi Lesmana, 2000. Exploding volcanoes, unique birds, gigantic rats and elegant ikat:
identifying sites of international biodiversity significance on Flores, Nusa Tanggara PKA/
BirdLifeWWF. Bogor. Rep. 11: xvi + 269 p.
Turbott, E.G. (convener), 1990. Checklist of the birds of New Zealand (3rd ed.): xv + 247 p.
Vaurie, C., 1949. A revision of the bird family Dicruridae.— Bull. Amer. Mus. Nat. Hist. 93: 199-342.
Vaurie, C., 1951. Notes on some Asiatic swallows.— Amer. Mus. Novit. 1529: 1-47.
Vaurie, C., 1952. A review of the bird genus Rhinomyias (Muscicapini).— Amer. Mus. Novit. 1570: 1-36.
Vaurie, C., 1957. Systematic notes on Palearctic birds. No. 25. Motacillidae: the genus Motacilla.— Amer.
Mus. Novit. 1832: 1-16.
Vaurie, C., 1958. Remarks on some Corvidae of Indo-Malaya and the Australian region.— Amer. Mus.
Novit. 1915: 1-13.
Vaurie, C., 1959. The birds of the Palearctic fauna. Passeriformes. Witherby, London: xii + 762 p.
Vaurie, C., 1962. Family Dicruridae. In E. Mayr & J.C. Greenway (eds.), Check-list of birds of the world
15: 137-157.
Vaurie, C., 1965. The birds of the Palearctic fauna. Non-Passeriformes. Witherby, London: i-xx, 1-763.
Vaurie, C. & D. Amadon, 1962. Notes on the honey buzzards of eastern Asia.— Amer. Mus. Novit. 2111:
1-11.
Verheijen, J.A.J., 1956 (1957?). Een ral repareert haar ei.— Limosa 29: 149-150.
Verheijen, J.A.J., 1961. Some notes on the birds of the island of Paluë, Flores, observed from 13th April to
5th May 1960.— Ardea 49: 183-187.
Verheijen, J.A.J., 1963. Bird-names in Manggarai, Flores, Indonesia.— Anthropos 58: 677-718.
Verheijen, J.A.J., 1964. Breeding season on the island of Flores, Indonesia.— Ardea 52: 194-201.
Verheijen, J.A.J., 1967. Kamus Manggarai I. Manggarai-Indonesia. Martinus Nijhoff, ‘s-Gravenhage:
xxviii + 772 p.
Verheijen, J.A.J., 1971. Red-necked Phalarope Phalaropus lobatus in the island of Flores, Indonesia.— Ardea 59: 64.
Verheijen, J.A.J., 1976. Some data on the avifauna of the island of Roti, Lesser Sunda Islands, Indonesia.— Zool. Meded. 50: 1-21.
Verhoeye, J. & D.A. Holmes, 1999. The birds of the islands of Flores – A review.— Kukila 10: 3-59.
Verhoeye, J. & B. King, 1990. Notes on three raptor species new to Flores.— Kukila 5: 65-68.
Vigors, N.A. & T. Horsfield, 1827. A description of the Australian birds in the collection of the Linnean
Society; with an attempt at arranging them according to their natural affinities.— Trans Linn. Soc.
London 15: 170-331.
Violani, C., 1979. Uccelli di Honolulu raccolti da Luigi Maria d’Albertis.— Ann. Mus. Civ. Genova 82;
172-183.

248

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Voous, K.H., 1948. Notes on a collection of Javanese birds.— Limosa 21: 85-100.
Voous, K.H., 1950. The races of the Yellow Wagtail (Motacilla flava) in the Indo-Australian Archipelago.— Treubia 20: 647-656.
Voous, K.H., 1967. Australian Pelican recorded in Indonesia.— Emu 67: 124.
Voous, K.H., 1995. In de ban van vogels. Scheffers, Utrecht: 605 p.
Voous, K.H. & J.G. van Marle, 1949. The distributional history of Coracina in the Indo-Australian archipelago.— Bijdr. Dierk. 28: 513-529.
Vorderman, A.G., 1888. Het journaal van Albert Colfs. Batavia.
Vorderman, A.G., 1895a. Bijdrage tot de kennis der avifauna van het eiland Noordwachter (Java Zee).—Nat. Tijdschr. Ned.-Ind. 54: 317-326.
Vorderman, A. G., 1895b. Lombok-vogels.— Nat. Tijdschr. Ned.-Ind. 54: 327-353.
Vos, J. de, P.Y. Sondaar, G.D. van den Bergh & F. Aziz, 1994. The Homo bearing deposits of Java and its
ecological context.— Courier Forschungs-Institut Senckenberg 171: 129-140.
Wakefield, N.A., 1958. Erroneous or doubtful records of birds from Gippsland, Victoria.— Emu 58:
271-276.
Walden, A., 1872. A list of the birds known to inhabit the island of Celebes.— Trans. Zool. Soc. Lond. 8:23-108, pl. III-X.
Wallace, A.R., 1864. A list of the birds inhabiting the islands of Timor, Flores and Lombock, with descriptions of the new species.— Proc. Zool. Soc. Lond. (1863): 480-497, pl. XXXIX.
Wallace, A.R., 1869. The Malay Archipelago. London. I: xxiv + 478 p., II: 524 p.
Warner, D.W., 1951. A new race of the cuckoo, Chalcites lucidus, from the New Hebrides Islands.— Auk
68: 106-107.
Warren, R.L.M., 1966. Type-specimens of birds in the British Museum (Natural History). Vol. 1. Nonpasserines. London: ix + 320 p.
Warren, R.L.M. & C.J.O. Harrison, 1971. Type-specimens of birds in the British Museum (Natural History) 2: Passerines. London: vi + 628 p.
Watling, D., 1982. Birds of Fiji, Tonga and Samoa. Millwood Press, Wellington, New Zealand: 176 p.
Watson, G.E. & E. Mayr, 1986. Family Monarchidae. Subfamily Rhipidurinae. In: E. Mayr & G.W. Cottrell (eds.): Check-list of birds of the world 11: 530-556.
Watson, G.E., M.A. Traylor & E. Mayr, 1986a. Family Sylviidae. In E. Mayr & G.W. Cottrell (eds.): Checklist of birds of the world 11: 3-294.
Watson, G.E., M.A. Traylor & E. Mayr, 1986b. Family Monarchidae, Subfamily Monarchinae. In E. Mayr
& G.W. Cottrell (eds.): Check-list of birds of the world 11: 464-526.
Wattel, J., 1973. Geographical variation in the genus Accipiter. Publ. Nuttall Orn. Club 13: vi + 231 p.
Weber, M., 1890. Zoologische Ergebnisse einer Reise in Niederländisch Ost-Indien I. Einleitung: i-xi.
Wells, D.R., 1999. The birds of the Thai-Malay Peninsula, 1: Non-Passeres. Academic Press, San Diego,
etc.: liii + 648 p.
Wells, D.R. & Lord Medway, 1976. Taxonomic and faunistic notes on birds of the Malay Peninsula.—
Bull. Brit. Orn. Cl. 96: 20-34.
White, C.M.N., 1936. Note on the Australian tree-martin (Petrochelidon nigricans).— Bull. Brit. Orn. Cl.
56: 90-92.
White, C.M.N., 1963. A revised check list of African flycatchers, tits, tree creepers, sunbirds, white-eyes,
honey eaters, buntings, finches, weavers and waxbills. Lusaka: vi + 218 p.
White, C.M.N., 1973a. Migrant Pernis apivorus in the Indonesian Archipelago.— Bull. Brit. Orn. Cl. 93:
116.
White, C.M.N., 1973b. Night herons in Wallacea.— Bull. Brit. Orn. Cl. 93: 175-176.
White, C.M.N., 1974. Three water birds of Wallacea.— Bull. Brit. Orn. Cl. 94: 9-11.
White, C.M.N., 1975a. Migration of Palaearctic waders in Wallacea.— Emu 75: 37-39.
White, C.M.N., 1975b. Further notes on birds of Wallacea.— Bull. Brit. Orn. Cl. 95: 106-109.
White, C.M.N., 1976a. Migration of Palaearctic non-passerine birds in Wallacea.— Emu 76: 79-82.
White, C.M.N., 1976b. Comments on the Dusky Moorhen Gallinula tenebrosa.— Bull. Brit. Orn. Cl. 96:
125-128.
White, C.M.N., 1977. Notes on some non-passerine birds of Wallacea.— Bull. Brit. Orn. Cl. 97: 99-103.

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White, C.M.N. & M. D. Bruce, 1986. The birds of Wallacea. B.O.U. Check-list No. 7: 524 p.
Whittell, H.M., 1954. The literature of Australian birds. Paterson Brokensha, Perth: xi + 116 + 788 p.
Widodo, W., J. H. Cox & P. C. Rasmussen, 1999. Rediscovery of the Flores Scops-owl Otus alfredi on
Flores, Indonesia and reaffirmation of its specific status.— Forktail 15: 15-23.
Wiegant, W. & B. van Helvoort, 1987. First sighting of Tachybaptus novaehollandiae on Bali.— Kukila 3:
50-51.
Wurmb, F. von, 1781. De nacht-uil van ‘t eiland Java.— Verh. Bat. Gen. 3: 371-375.
Wynne, O.E., 1969. Bibliographical key-names of birds of the world, to authors and those commemorated: iii + 246 p.
Ziswiler, V., H.R. Güttinger & H. Bregulla, 1972. Monographie der Gattung Erythrura Swainson, 1837
(Aves, Passeres, Estrildidae).— Bonner Zool. Monogr. 2: 1-158.

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13. Systematic index
The meaning of the letters given under the heading ‘status’ is as follows: M = marine: tropical sea-bird, which may breed in the region; N = migrant (winter-visitor) fromthe North; R = presumed resident; R* = resident, breeding recorded; S = migrant (winter-visitor) from the South.
status page
Podicipedidae
Podiceps ruficollis vulcanorum
RenschR
23
Podiceps novaehollandiae
novaehollandiae StephensS?
24
Pelecanidae
Pelecanus conspicillatus Temminck
S
25
Sulidae
Sula leucogaster plotus (Forster)M
26
Phalacrocoracidae
Phalacrocorax melanoleucos
melanoleucos (Vieillot)?
26
Phalacrocorax sulcirostris (Brandt)?
27
Fregatidae
Fregata ariel ariel (G.R. Gray)
M
27
ArdeidaeArdea sumatrana Raffles
R*
28
Ardea purpurea manilensis Meyen?
29
Ardea alba modesta (J.E. Gray)?
30
Ardea novaehollandiae LathamR*
30
Egretta intermedia (Wagler)?
31
Egretta garzetta nigripes (Temminck) ?
31
Egretta sacra sacra (Gmelin)R*
31
Bubulcus ibis coromandus (Boddaert) ?
32
Ardeola speciosa speciosa (Horsfield) ?
32
Butorides striatus javanicus (Horsfield) R
33
Nycticorax nycticorax nycticorax(Linnaeus)?
33
Nycticorax caledonicus hilli Mathews ?
34
Ixobrychus sinensis (Gmelin)N
35
Ixobrychus cinnamomeus (Gmelin)R*
36
Ciconiidae
Ciconia episcopus neglecta (Finsch)
R*
37
Plataleidae
Platalea regia Gould
S
38
Accipitridae
Pandion haliaetus cristatus (Vieillot) R
38
Aviceda subcristata timorlaoensis(A.B. Meyer)R
44Pernis ptilorhynchus orientalis
Taczanowski
N
45
Elanus caeruleus hypoleucus Gould
R*
46
Haliastur indus intermedius Gurney R*
47
Haliaeetus leucogaster (Gmelin)R
48

Circaetus gallicus gallicus (Gmelin)R
Accipiter soloensis (Horsfield)
N
Accipiter novaehollandiae sylvestris
WallaceR*
Accipiter fasciatus wallacii (Sharpe)
R*
Accipiter virgatus quinquefasciatus
Mees
R
Accipiter gularis (Temminck &Schlegel)N
Hieraaetus fasciatus renschi
Stresemann
R
Hieraaetus kienerii formosus
StresemannR
Spizaetus (cirrhatus) floris (Hartert)
R*
Falconidae
Falco moluccensis subsp.
R*
Falco cenchroides cenchroides Vigors& HorsfieldS
Falco longipennis hanieli Hellmayr
R*
AnatidaeDendrocygna arcuata arcuata(Horsfield)R
Dendrocygna javanica (Horsfield)
R
Anas superciliosa rogersi Mathews
R*
Anas gibberifrons gibberifrons
S. Müller
R*
Megapodiidae
Megapodius reinwardt reinwardt
DumontR*
Phasianidae
Coturnix ypsilophora raaltenii(S. Müller)
R*
Coturnix chinensis chinensis (Linnaeus) R*
Gallus varius (Shaw)R*
Turnicidae
Turnix suscitator powelli Guillemard R*
Turnix maculosa floresiana Sutter
R*
RallidaeRallus pectoralis exsul (Hartert)
R*
Rallus philippensis wilkinsoni(Mathews)
R*
Rallina fasciata (Raffles)
R*
Porzana pusilla pusilla (Pallas)
R?
Porzana fusca fusca (Linnaeus)
R

48
49
49
50
51
51
52
52
52
53
54
55

56
56
57
57

58

60
60
62
62
63
64
65
67
68
69

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)
Porzana cinerea cinerea (Vieillot)
R*
Amaurornis phoenicurus leucomelanus(S. Müller)
R*
Gallicrex cinerea (Gmelin)N
Gallinula tenebrosa frontata Wallace ?
Gallinula chloropus orientalis Horsfield R*
Porphyrio porphyrio indicus Horsfield R*
Jacanidae
Irediparra gallinacea gallinacea(Temminck)
R
Rostratulidae
Rostratula benghalensis benghalensis(Linnaeus)
R*
Charadriidae
Pluvialis fulva (Gmelin)N
Charadrius leschenaultii leschenaultii
LessonN
Charadrius veredus Gould
N
Scolopacidae
Numenius phaeopus variegatus(Scopoli)
N
Limosa lapponica bauerí Naumann
N
Tringa hypoleucos Linnaeus
N
Tringa glareola Linnaeus
N
Tringa nebularia (Gunnerus)
N
Gallinago stenura (Bonaparte)N
Gallinago megala SwinhoeN
Calidris ruficollis (Pallas)
N
Calidris alba (Pallas)
N
Himantopus himantopus leucocephalus
Gould
R
Phalaropidae
Phalaropus lobatus (Linnaeus)
N
Burhinidae
Esacus magnirostris (Vieillot)
R*
Glareolidae
Stiltia isabella (Vieillot)
S
LaridaeSterna albifrons sinensis Gmelin
M
Sterna bergii cristata Stephens
M
Chlidonias hybridus subsp.
S
Columbidae
Chalcophaps indica indica (Linnaeus) R*
Treron floris WallaceR
Ptilinopus cinctus albocinctus Wallace R
Ptilinopus regina flavicollis Bonaparte R
Ptilinopus melanospilus melanauchen(Salvadori)R*
Ducula aenea polia (Oberholser)
R*
Ducula rosacea (Temminck)
R
Ducula (lacernulata) sasakensis(Hartert)
R
Columba vitiensis metallica Temminck R

70
71
73
74
76
76

77

77
77
78
78

79
79
80
80
80
80
81
81
82
82
82
83
83
83
84
84
85
86
86
86
87
87
89
90
90

251
Macropygia unchall unchall (Wagler) R
Macropygia ruficeps orientalis Hartert R
Macropygia emiliana emiliana
Bonaparte
R*
Streptopelia bitorquata bitorquata(Temminck)
R
Streptopelia chinensis tigrina(Temminck)
R*
Geopelia striata maugei (Temminck) R*
Psittacidae
Trichoglossus (haematodus) weberi(Büttikofer)R*
Cacatua sulphurea occidentalis Hartert R*
Tanygnathus megalorynchos floris
HartertR
Loriculus flosculus Wallace(R)
Geoffroyus geoffroyi floresianus
Salvadori
R*
Cuculidae
Cuculus saturatus horsfieldi Moore
N
Cuculus saturatus lepidus S. Müller
R
Cuculus pallidus (Latham)S
Cacomantis variolosus sepulcralis(S. Müller)
R
Chrysococcyx basalis (Horsfield)
S
Chrysococcyx lucidus plagosus(Latham)S
Chrysococcyx minutillus subsp.
R*
Eudynamys scolopacea malayana
Cabanis & Heine
R
Scythrops novaehollandiae Latham
R
Centropus bengalensis sarasinorum
Stresemann
R*
Tytonidae
Tyto alba javanica (Gmelin)R*
Tyto longimembris longimembris(Jerdon)R
Strigidae
Otus magicus albiventris (Sharpe)
R*
Otus alfredi (Hartert)
Otus silvicola (Wallace)R*
Ninox scutulata florensis (Wallace)N
Caprimulgidae
Caprimulgus macrurus schlegelii Meyer R*
Caprimulgus affinis affinis Horsfield R*
Apodidae
Collocalia fuciphaga fuciphaga(Thunberg)
R*
Collocalia esculenta sumbawaeStresemann
R*
Apus pacificus pacificus (Latham)
N
Alcedinidae
Alcedo atthis floresiana Sharpe
R

91
91
92
94
94
95

95
96
97
98
99
100
100
101
101
104
105
105
107
107
107
108
110
111
111
113
115
116
117

120
121
123
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252
Ceyx rufidorsa Strickland
R
Pelargopsis capensis floresiana Sharpe R
Halcyon chloris chloris (Boddaert)
R*
Halcyon sancta sancta
Vigors & HorsfieldS
Halcyon fulgida Gould
R*
Meropidae
Merops ornatus Latham
S
Merops philippinus Linnaeus
R*
Coraciidae
Eurystomus orientalis orientalis(Linnaeus)
R*
PicidaeDendrocopos moluccensis grandis(Hargitt)
R*
Pittidae
Pitta elegans concinna Gould
R*
Pitta elegans maria Hartert
S
Alaudidae
Mirafra javanica parva Swinhoe
R
Hirundinidae
Hirundo rustica gutturalis Scopoli
N
Hirundo tahitica javanica Sparrman
R*
Cecropis striolata striolata (Schlegel) R*
Petrochelidon nigricans timoriensis
Sharpe?
Motacillidae
Motacilla flava simillima Hartert
N
Motacilla caspica subsp.
N
Anthus novaeseelandiae albidus
Stresemann
R*
Anthus gustavi Swinhoe
N
Campephagidae
Coracina novaehollandiae floris(Sharpe)
R*
Coracina novaehollandiae
novaehollandiae (Latham)
S
Coracina novaehollandiae subpallida
Mathews
S
Coracina dohertyi (Hartert)
R
Lalage sueurii sueurii (Vieillot)
R*
Pericrocotus lansbergei Büttikofer
R
LaniidaeLanius cristatus superciliosus Latham N
TurdidaeBrachypteryx montana floris Hartert R
Saxicola caprata fruticola HorsfieldR*
Zoothera interpres (Temminck)
R
Zoothera dohertyi (Hartert)
R*
Zoothera andromedae (Temminck)
R*
Turdus obscurus Gmelin
N
Timaliidae
Pnoepyga pusilla everetti Rothschild R

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)
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125
126
127
128
129
130

130

132
133
134
137
138
138
139
140
141
141
142
143

143
144
146
147
148
148
149
149
149
151
151
152
153
153

Sylviidae
Tesia everetti everetti (Hartert)
R*
Cisticola juncidis fuscicapilla Wallace R*
Cisticola exilis lineocapilla Gould
R*
Orthotomus cucullatus everetti(Hartert)
R*
Phylloscopus borealis xanthodryas(Swinhoe)N
Phylloscopus presbytes floris (Hartert) R
Seicercus montis floris (Hartert)
R
Maluridae
Gerygone sulphurea sulphurea
WallaceR*
Muscicapidae - Muscicapinae
Ficedula westermanni hasselti (Finsch) R
Ficedula hyperythra vulcani Robinson R
Ficedula dumetoria dumetoria(Wallace)
R
Culicicapa ceylonensis sejuncta
HartertR*
Rhinomyias oscillans (Hartert)
R
Muscicapidae - Rhipidurinae
Rhipidura diluta diluta Wallace
R*
Rhipidura rufifrons semicollaris
S. Müller
R*
Muscicapidae - Monarchinae
Monarcha trivirgatus trivirgatus(Temminck)
R
Monarcha trivirgatus wellsi(Ogilvie-Grant)
R
Monarcha trivirgatus nigrimentum
G.R. GrayR
Monarcha (trivirgatus) boanensis
van Bemmel
R
Monarcha sacerdotum MeesR
Monarcha cinerascens cinerascens(Temminck)
R*
Hypothymis azurea symmixta
Stresemann
R*
Terpsiphone paradisi floris Büttikofer R*
Pachycephalidae
Pachycephala fulvotincta fulvotincta
WallaceR*
Pachycephala nudigula nudigula
HartertR
Paridae
Parus major cinereus Vieillot
R*
Dicaeidae
Dicaeum annae (Büttikofer)R*
Dicaeum agile tinctum (Mayr)R
Dicaeum igniferum Wallace
R
Dicaeum sanguinolentum rhodopygiale
Rensch
R

154
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161

162
162
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164
164
165
166

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168
169
170
171
171
173

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Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)
Nectariniidae
Anthreptes malacensis convergens
RenschR*
Nectarinia jugularis ornata (Lesson)
R*
Nectarinia solaris solaris Temminck
R*
Zosteropidae
Zosterops palpebrosa unica Hartert
R*
Zosterops montana montana
Bonaparte
R*
Zosterops chloris intermedia Wallace R*
Zosterops wallacei Finsch
R*
Lophozosterops superciliaris superciliaris(Hartert)
R
Lophozosterops dohertyi subcristata
HartertR*
Heleia crassirostris crassirostris(Hartert)
R*
Meliphagidae
Lichmera indistincta limbata (S. Müller) ?
Lichmera lombokia fumidigula(Rensch)R
Philemon buceroides neglectus(Büttikofer)
R*
Estrildidae
Amandava amandava flavidiventris(Wallace)
R*

183
184
184
185
188
189
191
191
192
193
194
195
196

197

Received: 15.ix.2005
Accepted: 15.v.2006
Edited: A.C. van Bruggen & J. van der Land

253
Poephila guttata guttata (Vieillot)
R
Erythrura hyperythra obscura(Rensch)R
Lonchura molucca propinqua(Sharpe)R*
Lonchura punctulata blasii(Stresemann)R*
Lonchura quinticolor (Vieillot)
R*
Lonchura pallida pallida (Wallace)R*
Ploceidae
Passer montanus malaccensis Dubois R*
Sturnidae
Aplonis minor minor (Bonaparte)
R*
Gracula religiosa venerata Bonaparte R*
Oriolidae
Oriolus chinensis broderipi Bonaparte R*
Dicruridae
Dicrurus hottentottus bimaensis
WallaceR*
Artamidae
Artamus leucorhynchus leucorhynchus(Linnaeus)
R*
CorvidaeCorvus florensis Büttikofer
R*
Corvus macrorhynchos macrorhynchos
Wagler
R*

198
198
199
200
207
208
209
209
212
212

214

216
217
218

254

Mees. Avifauna of Flores. Zool. Med. Leiden 80 (2006)

Figs 11-18 on pp. 254-260 show eggs at approximately natural size of Flores birds in the
Leiden Museum.
Fig. 12. – a, Accipiter novaehollandiae sylvestris, RMNH 70070, c/2; b, A. fasciatus wallacii, RMNH 70066,c/2; c, Spizaetus (cirrhatus) floris, RMNH 70075, c/1; d, Falco moluccensis, RMNH 70096, c/4.
Fig. 13. – a, Falco moluccensis, RMNH 70099, c/3; b, F. longipennis hanieli, RMNH 70097, c/1; c, Rallus
pectoralis exsul, RMNH 70315, c/3; cc, do., RMNH 70316, c/3; d, R. philippinensis wilkinsoni, RMNH 70350,
c/6; e, Porzana c. cinerea, RMNH 70325, c/4; f, Trichoglossus (haematodes) weberi, RMNH 70578, c/2; g,
Cacatua sulphurea occidentalis, RMNH 70573, c/1; gg, do., RMNH 70575, c/1; ggg, do., RMNH 70576,
c/1.
Fig. 14. – a, Otus magicus albiventris, RMNH 70606, c/2; b, O. silvicola, RMNH 70610, c/2; c, Halcyon c.
chloris, RMNH 70637, c/4; d, H. fulgida, RMNH 70644, c/2; e, Pitta elegans concinna, RMNH 70683, c/3.
Fig. 15. – a, Coracina novaehollandiae floris, RMNH 70756, c/2; aa, do., RMNH 70755, c/2; b, Lalage s.
sueurii, RMNH 70759, c/3; c, Zoothera dohertyi, RMNH 70762, c/1; d, Z. andromedae, RMNH 70769, c/2;e, Tesia e. everetti, RMNH 70886, c/2; f, Cisticola exilis lineocapilla, RMNH 70970, c/4; g, Orthotomus
cucullatus everetti, RMNH 375a (field number), c/3; h, Culicicapa ceylonensis sejuncta, RMNH 71098, c/1;
i, Monarcha c. cinerascens, RMNH 71115, c/1; j, Terpsiphone paradisi floris, RMNH 71145, c/2.
Fig. 16. – a, Pachycephala f. fulvotincta, RMNH 71155, c/2; aa, do., RMNH 71165, c/2; b, Dicaeum annae,
RMNH 71210, c/2; c, Anthreptes malaccensis convergens, RMNH 71339, c/2; d, Nectarinia jugularis ornata,
RMNH 71291, c/2; dd, do., RMNH 71289, c/2; e, N. s. solaris, RMNH 71319, c/2; ee, do., RMNH 71326,
c/2; f, Zosterops palpebrosa unica, RMNH 71455, c/2; g, Z. wallacei, RMNH 71526, c/2; h, Lophozosterops
dohertyi subcristata, RMNH 71608, c/2; i, Heleia crassirostris, RMNH 71642, c/3.
Fig. 17. – a, Philemon buceroides neglectus, RMNH 71691, c/3; b, Aplonis m. minor, RMNH 71763, c/3; c,Gracula religiosa venerata, RMNH 71774, c/2; d, Oriolus chinensis broderipi, RMNH 71782, c/2; e, Dicrurus
hottentotus bimaensis, RMNH 71789, c/4; ee, do., RMNH 71800, c/4.
Fig. 18. – a, Corvus florensis, RMNH 71816, c/3; b, C. macrorhynchus, RMNH 71814, c/4.

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Fig. 12

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