io s m b

THE NATURAL

HISTORY MUSEUM

2 1 AUG 2012

PURCHASED

TRING LIBRARY

Forktail 28 (2012)

f

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Cover picture: Camiguin Flawk Owl, new species, Camiguin Sur, Philippines, 8 June 201 1

by Rob Hutchinson

ISSN 0950-1746

© Oriental Bird Club 2012

FORKTAIL

Number 28, 2012

Acting Editor

N. J. Collar

Associate Editors

Jeremy Bird, David Buckingham, Stuart Butchart, Will Duckworth, Eben Goodale, John Pilgrim and Jack Tordoff

THE NATURAL

HISTORY MUSEUM

2 I AUG 2012

PURCHASED

TRING LIBRARY

CONTENTS

P. C. RASMUSSEN, D. N. S. ALLEN, N. J. COLLAR, B. DeMEULEMEESTER, R. 0. HUTCHINSON, P. G. C. JAKOSALEM,

R. S. KENNEDY, F. R. LAMBERT & L. M. PAGUNTALAN

Vocal divergence and new species in the Philippine Hawk Owl Ninox philippensis complex . 1

M. MONIRUL H. KHAN & NASIM AZIZ

Bird species diversity in five protected areas of Bangladesh . 21

MARY ROSE C. POSA & DAVID ALEXANDER MARQUES

Peat swamp forest birds of the Tuanan research station, Central Kalimantan, Indonesia, with notes on habitat specialists . 29

MUHAMMAD NAEEM AWAN, HASSAN ALI & DAVID C. LEE

An annotated checklist of birds and conservation issues in Salkhala Game Reserve, an isolated Important Bird Area

in Azad Kashmir, Pakistan . 38

SIMON P. MAHOOD & JONATHAN C. EAMES

A review of the status of Collared Laughingthrush Garrulax yersini and Grey-crowned Crocias Crocias langbianis . 44

TILLTOPFER

Seasonal changes in plumage coloration of Orange Bullfinches Pyrrhula aurantiaca . 49

D. KHAMCHA & G. A. GALE

The use of tree-fall gaps by a forest interior avian frugivore in a tropical evergreen forest . 53

SAYAM U. CHOWDHURY, ALEXANDER C. LEES & PAUL M. THOMPSON

Status and distribution of the endangered Baer's Pochard Aythya baeri in Bangladesh . 57

ZHIXIN ZHOU, YUE SUN, LU DONG, CANWEI XIA, HUW LLOYD & YANYUN ZHANG

Breeding biology of Asian House Martin Delichon dasypus in a high-elevation area . 62

DZUNG-YUN YANG, CHUNGYU CHIANG & YU-CHENG HSU

Sex bias in a wintering population of Dunlin Calidris alpina in central Taiwan . 67

JEREMY P. BIRD, BERRY MULLIGAN, ROURS VANN, PHILIP D, ROUND & JAMES J. GILROY

Habitat associations of the Manchurian Reed Warbler Acrocephalus tangorum wintering on the Tonle Sap floodplain

and an evaluation of its conservation status . 71

COLIN R. TRAINOR, PHILIPPE VERBELEN & RON E. JOHNSTONE

The avifauna of Alor and Pantar, Lesser Sundas, Indonesia . 77

H. L. WRIGHT, N. J. COLLAR, I. R. LAKE, BOU VORSAK & P. M. DOLMAN

Foraging ecology of sympatric White-shouldered Ibis Pseudibis davisoni and Giant Ibis Thaumatibis gigantea

in northern Cambodia . 93

LE MANH HUNG, MARK B. ROBBINS, NATHAN H. RICE & ERICK A. GARCIA-TREJO

Survey of the avifauna at Muong Nhe Nature Reserve, Dien Bien province, Vietnam . 101

T. E. MARTIN, D. J. KELLY, N. T. KEOGH, D. HERIYADI, H. A. SINGER & G. A. BLACKBURN

The avifauna of Lambusango Forest Reserve, Buton Island, south-east Sulawesi, with additional sightings from

southern Buton . 107

PHILIP D. ROUND, JOHN M. HOBDAY, RUNGSRIT KANJANAVANIT & JAMES S. STEWARD

A nesting pair of Gecinulus woodpeckers in a likely zone of intergradation between Pale-headed Woodpecker G. grantia

and Bamboo Woodpecker G. viridis . 1 1 3

PAUL J. LEADER & GEOFF J. CAREY

Zappey's Flycatcher Cyanoptila cumatilis, a forgotten Chinese breeding endemic . 121

ALAIN HENNACHE, SIMON P. MAHOOD, JONATHAN C. EAMES & ETTORE RANDI

Lophura hatinhensis is an invalid taxon .

129

Forktail 28 (2012)

Short Notes

SIMON P. MAHOOD, DAVID P. EDWARDS & FELICITY A. EDWARDS

Bar-winged Wren Babbler Spelaeornis troglodytoides: a first record for Vietnam, with speculation for

1 7 further avifaunal additions . 1 36

ZHU LEI, ZHANG JUN, QIU JING, WEI QIAN, DONG LEI & SUN YUE-HUA

Long-tailed Duck Clangula hyemalis and Red-breasted Goose Branta ruficollis : two new birds for Sichuan,

with a review of their distribution in China . 138

DELIP K. DAS

Western Hoolock Hoolock hoolock preying on chicks of Greater Racket-tailed Drongo Dicrurus paradiseus

in Lawachara National Park, Bangladesh . 142

FRANK E. RHEINDT & JAMES A. EATON

Notes on the life-history and taxonomy of Muscicapa dauurica umbrosa, an overlooked Bornean canopy bird . 144

YANG LIU & SERGEY PYZHJANOV

Apparent inter- and intraspecific brood-parasitism in a nest of Tufted Duck Aythya fuligula . 147

DAVID J. KELLY & NICOLA M. MARPLES

Annual survival rate and mean life-span of Lemon-bellied White-eyes Zosterops chloris flavissimus on Kaledupa island,

Wakatobi, south-east Sulawesi, Indonesia . 148

CHANG-YONG CHOI & HYUN-YOUNG NAM

Migrating dragonflies: famine relief for resident Peregrine Falcons Falco peregrinus on islands . ) . 1 49

DONG-WON KIM, CHANG-WAN KANG, HWA-JUNG KIM, YOUNG-SOO KWON & JIN-YOUNG PARK

Breeding of the Japanese Murrelet Synthliboramphus wumizusume in South Korea . 151

NORIMASA SUGITA, TOSHITAKA N. SUZUKI, CRAIG A. BARNETT & KEISUKE UEDA

An intraspecific adult killing in female Japanese Great Tits Parus major minor . 1 53

VLADIMIR DINETS

Nesting Fork-tailed Swifts Apus pacificus in north-eastern Vietnam . 155

ADAM C. STEIN & GALINA NOSACHENKO

Notes on the 2009 autumn crane migration in Muraviovka zakaznik, Amur oblast, Russian Federation . 1 56

JAMES W. BURNHAM & ERIC M. WOOD

Woolly-necked Stork Ciconia episcopus at Napahai wetland, Yunnan, China . 1 58

TIM ROBINSON

First records of Javan Munia Lonchura leucogastroides in Peninsular Malaysia . 1 59

GERARD E. RYAN

Brahminy Kites Haliastur indus fishing with Irrawaddy dolphins Orcaella brevirostris in the Mekong River . 161

MOHAMMAD IRHAM, E. MEIJAARD & S. (BAS) van BALEN

New information on the distribution of White-fronted Microhierax latifrons and Black-thighed Falconets

M. fringillarius in Kalimantan, Indonesia . 162

WEN-LOUNG LIN, SI-MIN LIN & HUI-YUN TSENG

Breeding ecology of the Northern Boobook Ninox japonica totogo in central Taiwan . 164

SIMON P. MAHOOD & JAMES A. EATON

The vocalisations of Red-collared Woodpecker Picus rabieri . 1 67

JAMES A. FITZSIMONS, ERIK MEIJAARD, IWAN HUNOWU, DEWI PRAWIRADILAGA, JANELLE L. THOMAS & JOHNY S.TASIRIN

Diet of the Speckled Boobook Ninox punctulata in north Sulawesi, Indonesia . 169

TOSHITAKA N. SUZUKI

Mobbing to death of a Japanese Long-eared Bat Plecotus sacrimontis by two species of tit . 171

Guidelines for contributors

inside back cover

Forktail 28 (2012)

Editorial notes

The re-styling of Forktail last year was designed in large part to

allow for a larger number of contributions to the journal, and the

upshot was that the issue for 201 1 appeared rather slim by

comparison with those of the previous five years. In fact with

1 1 full papers and 21 short notes it was an average size issue,

the mean number of full papers and short notes for the years

2000-2010 being 1 1.7 and 16.8 respectively. This year,

somewhat by contrast, the journal carries 18 papers and 19 short

notes, and the space gains of last year's makeover have proved

invaluable. The trend seems to be universal: all ornithological

journals are growing in size in response to the increasing

quantity of material that is competing for a place in them. In a

club as small and as prudently managed as OBC, however, the

budgetary constraints on publications are significant and I

express my gratitude to OBC Council for its approval of the

considerable costs that this relatively large and colour-rich issue

has incurred.

The other costs that the journal incurs are on the time of its

acting and associate editors, all of whose positions are strictly

honorary. Assuming that contributions to Forktail will continue

to increase, the burden of editorship will need further

redistribution, not only in terms of more or new human

shoulders but also in relation to the annual editorial time-frame.

Having only recently established 31 March as the deadline for

the receipt of material for publication in the same year, we now

propose to bring this deadline forward to

14 February

so as to allow an extra six weeks for the various necessary

processes of moving from manuscript to publication — a six-

month period, given the publication date of mid-August which

Forktail now rigidly observes.

I once more thank Jez Bird, Dave Buckingham, Stuart

Butchart, Will Duckworth, Eben Goodale, John Pilgrim and Jack

Tordoff for their help through the year, Brian and Margaret Sykes

for their unswerving support, and Peter Creed for his super-fast,

efficient and stylish layout of the end product.

N. J. Collar

Forktail 28 (2012)

Plate. Species in the Philippine Hawk Owl Ninox philippensis complex: (1) Mindoro Hawk Owl N. mindorensis (a, dark individual: b, pale

individual); (2) Mindanao Hawk Owl N. spilocephala; (3) Luzon Hawk Owl N. philippensis', (4) Cebu Hawk Owl, new species; (5) Romblon Hawk Owl

Ninox spilonota, new subspecies from Tablas; (6) Romblon Hawk Owl, nominotypical from Sibuyan; (7) Camiguin Hawk Owl, new species; (8) Sulu

Hawk Owl Ninox reyi. Original painting by John Gale.

FORKTAIL 28 (2012): 1-20

Vocal divergence and new species

in the Philippine Hawk Owl Ninox philippensis complex

P. C. RASMUSSEN, D. N. S. ALLEN, N. J. COLLAR, B. DeMEULEMEESTER, R. 0. HUTCHINSON,

P. G. C. JAKOSALEM, R. S. KENNEDY, F. R. LAMBERT & L. M. PAGUNTALAN

We show, based on morphology and especially vocalisations, that the Philippine Hawk Owl Ninox philippensis requires treatment as seven

allopatric species and at least one additional subspecies. Morphological distinctions between three groups of taxa are striking, and although

taxa within one major group are relatively similar in plumage they vary rather consistently in size and proportions. It has not been possible

until now to resolve the species limits in this complex due mainly to the lack of sound recordings of key taxa, a problem now rectified.

Vocalisations differ significantly between all seven species, the limits of which are incongruent with all previous taxonomies. Taxa from

Mindoro (mindorensis), Mindanao [spilocephala), Camiguin Sur (named herein), and the Sulu Islands ( reyi ) exhibit especially great vocal

differences from all other taxa along with smaller but consistent differences in plumage and morphometries. Although specimens have

been in museum collections for many years, two of these species and one subspecies have heretofore remained undescribed, and we

formally name these taxa for science. The recommended species-level treatment and English names of the N. philippensis complex are:

Luzon Hawk Owl N. philippensis; Mindanao Hawk Owl N. spilocephala; Mindoro Hawk Owl N. mindorensis; Sulu Hawk Owl Ninox reyi; Romblon

Hawk Owl N. spilonota; Camiguin Hawk Owl new species; and Cebu Hawk Owl new species.

INTRODUCTION

During the heyday of ornithological discovery in the Philippines

(approximately 1850-1910), as many as seven species were

recognised in the islands’ endemic hawk owl complex (e.g.

McGregor 1909-1910). Since 1945, however, it has been treated

as a single polytypic species, Philippine Hawk Owl Ninox

philippensis (Delacour & Mayr 1945), recently considered to

contain eight subspecies (Dickinson 2003). These subspecies group

into three ’distinctive plumage types: one with all-streaked

underparts and plain crown ( philippensis , proximo., ticaoensis and

centralis of Luzon and many other islands); one with mottled or

barred breast, streaked lower underparts, and spotted crown

(. spilocephala of Mindanao); and one with barred to nearly plain

underparts (the ‘unstreaked’ group: mindorensis of Mindoro,

spilonota of several small islands, and reyi of the Sulu Islands) (Collar

& Rasmussen 1998; for main islands and distribution of taxa see

Figure 1). This last group of three described races is highly disjunct,

and the distribution of the race spilonota as currently defined in

particular is biogeographically peculiar because the Cebu

population is surrounded by members of th e philippensis group, and

the Camiguin Sur population is from a small island off northern

Mindanao and well away from other taxa in spilonota.

Within the unstreaked group there are marked differences in

size and proportions, as well as more subtle distinctions in plumage.

However, it has not been possible to resolve relationships between

these racial groups based on morphology owing to intra-island

plumage variation and, for some taxa, small sample size. Sound

recordings until recently were available only for taxa from a few

islands, and most were incomplete and of poor quality. Given the

complexity of the vocal repertoire in this group, the small sample

of recordings previously available precluded further analysis of

species limits. The sample was, however, adequate to establish that

Mindoro mindorensis differs profoundly in vocalisations from

Luzon nominotypical philippensis, prompting the separation of

Mindoro Hawk Owl N. mindorensis (Konig etal. 1999). However,

this cursory treatment left other unstreaked forms united with N.

philippensis, although in the absence of acoustic data it seemed

plausible that they could separate out as a single species for which

the name with priority was Ninox reyi (Collar & Rasmussen 1 998).

Within this species, however, there were clearly multiple

undescribed taxa based on morphology, although the populations

on Cebu and Tablas were feared possibly extinct (Collar &

Rasmussen 1998, Collar etal. 1999).

There the matter unsatisfactorily rested, in the absence of

adequate or sometimes any vocal evidence from all the taxa,

including the four insular populations comprising the form

spilonota. However, recent fieldwork has resulted in nearly complete

sampling with extensive, high-quality recordings of the vocal

repertoire of the key island populations of Ninox philippensis sensu

lato. Taxa that differ in plumage also differ in vocalisations, so much

so that their treatment as conspecific in a group with innate

vocalisations such as owls is untenable. Surprisingly, however, some

unstreaked taxa that resemble each other closely are also divergent

in vocalisations, and cannot be maintained as taxa below the species

level. We propose here that seven vocally well-defined allopatric

species are involved in the Philippine Hawk Owl complex, of which

two are new species (described herein) with distinctly divergent

vocalisations. Because individuals from four islands now known to

pertain to four different taxa were included in the description of

N. spilonota, we designate a lectotype and hence type locality for

this ambiguous name. We also describe an additional island taxon

here that shows only moderately distinctive vocalisations and

morphology, and which we consider better treated at the subspecies

level.

METHODS

Acoustic analyses

We analysed sound recordings of all taxa known or suspected to be

critical to an analysis of species-level taxonomy of the Philippine

Hawk Owl {sensu lato). The majority of recordings we used were

made by ROH during trips specifically targeting islands that hold

morphologically 'distinctive Ninox taxa for which we previously

lacked or had poor representation of vocalisations. Several other

recordings were made by co-authors and others, and most of these

are available in full on AVoCet (avocet.zoology.msu.edu, AY). (To

access individual numbered recordings on AVoCet, use e.g. http:/

/avocet.zoology.msu.edu/recordings/14561.) A few recordings

were assembled from other sound archives (Macaulay Library,

http://macaulaylibrary.org/, ML; National Sound Archive,

London, http://www.bl.uk/nsa, NSA; xeno-canto, http://

www.xeno-canto.org/, XC).

2

P. C. RASMUSSEN etal.

Forktail 28 (2012)

Figure 1. Map of the known

distribution of taxa ofthe Philippine

Hawk Owl Ninox philippensis ( sensu

lato). Taxa and groups are Philippine

Hawk Owl Ninox philippensis ( sensu

stricto ) nominotypical philippensis

group, of Luzon, Samar, Leyte, and

small surrounding smaller islands;

centralis, of Panay, Negros, Bohol,

Siquijor (unlabelled small island

between southern Negros and

Bohol) and surrounding smaller

islands; proximo, of Masbate; and

ticaoensis, of Ticao; Mindanao Hawk

Owl N. spilocephala: Mindanao and

smaller surrounding islands except

Camiguin Sur; Sulu Hawk Owl N. reyi:

larger islands of Sulu and Tawi Tawi

provinces; Camiguin Hawk Owl new

species; Mindoro Hawk Owl N.

mindorensis: Mindoro; Romblon

Hawk Owl N. spilonota: Tablas and

Sibuyan islands, Romblon Province;

Cebu Hawk Owl new species.

By island (listed alphabetically), the number of recordings used

is listed below, with recordist (initials used for co-authors) and

(where applicable) abbreviation for sound archive where recording

is held (for recordings by FRL first uploaded to AVoCet but also on

xeno-canto, only the AV number is provided here): Biliran, 1 (RSK:

ML#38695); Bohol, 2, (F. Verbelen [FV]: AV#8971 -8972);

Camiguin Sur, 19 (ROH: 13554-13557, AV#13559, AV#13567,

AV# 13575, AV# 13577, AV# 13593, AV# 13598, AV# 13602,

AV#13605, AV#13609, AV#136l4-136l 5, AV#13618,

AV#13622; LMP: AV#13552-13553); Cebu, 13 (DNSA:

XC#79316, AV#1 1320-1 1322; LMP: AV#10469- 10470,

AV#10805; PGCJ: AV#10804, AV#10806; BD: AV#12609-

12612); Leyte, 2 (RSK: ML#38671, ML#38674); Luzon, 34 (PCR:

AV#2 168-2 174; FV: AV#8970; FRL: XC#30725-30728; ROH:

AV# 1 2420, AV# 12450-1 2454, AV# 13551, AV# 1 3648- 1 35 54; D.

Edwards: XC#35238; P. Noakes: XC#40819, 40821, XC#40823,

XC#40825-40826, XC#40828; G. Wagner: XC#23 1 16); Masbate,

1 (LMP: AV#14563); Mindanao, 23 (FRL: AV#8056-8057,

AV#8088, AV#81 1 1-8113; ROH: AV# 12455- 12467; S. Harrap:

NSA Wildlife ref. #132605-132606; B. F. King: NSA Wildlife ref.

#54931); Mindoro, 6 (ROH: AV#11507, AV#13655; P. Morris:

NSA#65216 W1CDR0000309 BD24, NSA#W1CDR0000307

BD1, NSA#W1CDR0000309 BD21, NSA#W1CDR0000309

BD24); Negros, 5 (FRL: AV#10664, AV#10699-10701,

AV# 10800- 10801); Sibuyan, 8 (ROH: AV#13637, AV#1364l-

13647); Siquijor, 2 (DNSA: AV#14564- 14565); Tablas, 15

(DNSA: AV#10803,AV#1 1323-1 1324; BD: AV#11325; ROH:

AV#1 1508-1 1515, AV# 12606- 12608); Tawi Tawi, 17 (DNSA:

AV#10802; ROH AV#14566-14581). Recordings are highly

variable in length, quality and documentation, but we have extensive,

Forktail 28 (2012) Vocal divergence and new species in the Philippine Hawk Owl Ninox philippensis complex

3

good to excellent material from all the above islands except Biliran,

Leyte and Masbate; material for Siquijor is rather inadequate.

Sounds were studied, measured and graphed in Raven Pro 1.3

(Raven 2012). Measurements taken (where possible) for each

recording were maximum number of notes/strophe, maximum and

minimum note length per recording; minimum inter-note spacing;

maximum and minimum fundamental frequencies (one each per

recording), maximum frequency difference within a single note and

between strophes; and maximum note bandwidth at a single point.

Principal components analyses (PCAs) were done in SYSTAT 13

(SYSTAT 2012) using these measurements except for maximum

note bandwidth (excluded because hisses were difficult to measure

precisely). After an initial run that showed that all recordings from

islands populated by nominotypical philippensis or centralis grouped

together, these groups were combined in the analysis of all groups.

Also, because inclusion of atypical song sequences resulted in much

greater variance in early analyses, only songs considered to be typical

were included in further analyses. Thus, typical songs of all taxon

groups (with philippensis and centralis combined) were used in the

main PCA. However, because of the great vocal differences of

Camiguin Sur, reyi , mindorensis and spilocephala from each other

and all other (‘core’) groups (those with non-extreme song

characteristics), the core groups ( philippensis plus centralis , Tablas

spilonota, Sibuyan spilonota, and Cebu) were not well resolved.

Therefore, a subsequent PCA was undertaken using just the core

groups, and for this recordings from the different islands were

graphed separately.

For taxa (all but Camiguin Sur, reyi, mindorensis and

spilocephala ) that typically give lengthy series starting with single

notes and building after a few minutes to a multi-note climax, only

one of each of the above measurements was taken. It should be noted

that many recordings and analyses of this type of strophe are likely

to be incomplete, as the recordist may have begun recording only

after hearing the bird, or the initial notes may be very soft and

cannot be picked up by a recording. For taxa that typically give a

series of short strophes each of which climaxes individually, each

of the above measurements was taken for each good-quality strophe.

It must also be borne in mind that many recordings made by a single

recordist on a single night, or even possibly over multiple nights at

the same locality, are likely to be of the same individual owls, leading

to some potential pseudoreplication in our analyses (a problem we

could not avoid but which is unlikely to bear on the outcome).

Moreover, many recordings will have been made after playback, and

this has not typically been documented by the recordist. Many

recordings are duetted between pair members, while others appear

to be counter-singing between birds in neighbouring territories; it

is not easy on present knowledge to distinguish these. However,

the ability to do so is not critical to our analyses, as the differences

between taxa here considered species are so marked. We chose the

above measurements because they are little affected by such

problems. However, given the great variation among taxa in

vocalisations, few if any song characteristics are shared among all

taxa, so choosing appropriate measurements was challenging.

In the vocal transcriptions presented in Results, notation

follows Rasmussen & Anderton (2005). Lower case signifies

relatively low volume compared to SMALL CAPITALS then to FULL

CAPITALS, which is much louder than lower case. The forward

slash / signifies a rise in frequency and the backslash \ a frequency

drop between elements. Run-together syllables signify no pause,

an apostrophe (’) extremely short breaks (e.g. in a trill), a hyphen (-

) indicates a very short pause, a comma a mid-length break, and an

open underscore _ a still longer pause. Ellipsis (...) is used to indicate

the continuation of a vocalisation as previously transcribed, not to

indicate pauses or fading out.

Sonagrams (spectrograms) prepared in Raven Pro 3.1 are

presented for each taxon along with the corresponding waveform

(oscillogram, in Raven units, which are unique to Raven software

and hence not indicated on the Y axis), which shows power versus

time, and hence allows visualisation of rhythm better than the

sonagram alone. The corresponding spectrum plots power versus

frequency, hence allowing visualisation of power peaks. The area

of the sonagram highlighted in grey is that on which the spectrum

is constructed.

Mensural analyses

We studied specimens of Ninox philippensis (. sensu lato ) held (in

alphabetical order of acronym) in the American Museum of Natural

History, New York (AMNH); Academy of Natural Sciences,

Philadelphia (ANSP); Natural History Museum, Tring, UK

(BMNH); Carnegie Museum of Natural History, Pittsburgh

(CM); Cincinnati Museum of Natural History, Cincinnati

(CMNH); Delaware Museum of Natural History, Greenville

(DMNH); Field Museum of Natural History, Chicago (FMNH);

Institut Royal des Sciences Naturelles, Brussels (IRSNB); Museum

of Comparative Zoology, Boston (MCZ); Bell Museum of Natural

History, Minneapolis (MMNH); Museum National d’Histoire

Naturelle, Paris (MNHN); Naturalis, Leiden (NCB); Philippine

National Museum, Manila (PNM); Royal Ontario Museum,

Toronto (ROM); Senckenberg Forschungsinstitut und

Naturmuseum, Frankfurt (SFN); Staatliches Museum fur

Tierkunde, Dresden (SMTD); Staatliches Naturhistorisches

Museum, Braunschweig (SNHM); University of Michigan

Museum of Zoology, Ann Arbor (UMMZ); National Museum of

Natural History, Washington, DC (USNM); Peabody Museum,

Yale University, New Haven (YPM); and Museum fiir Naturkunde,

Berlin (ZMB). All specimens of this complex available at the above

museums during our visits were studied, and most of them

photographed and measured. In total, 177 specimens of the

following established taxa were included in mensural analyses: 40

philippensis (5 Catanduanes, 5 Leyte, 2 Lubang, 26 Luzon, 1

Marinduque, 1 Polillo);4 proxima (Masbate); 1 ticaoensis (Ticao);

36 centralis (2 Bohol, 27 Negros, 7 Siquijor); 16 spilonota (1 Tablas,

12 Sibuyan, 1 Cebu, 2 Camiguin Sur); 43 spilocephala (10 Basilan,

31 Mindanao, 2 Siargao); 25 mindorensis (Mindoro); and 12 reyi

(1 Siasi, 9 Tawi Tawi [8 main island, 1 Bongao], 1 Sibutu, 1 Sulu).

Not all measurements were available for all specimens (see below),

hence numbers in certain analyses are smaller.

For specimens from most of the above collections, PCR

measured a wide array of characters, and then after preliminary

analyses chose the following as most useful: culmen from cere;

upper mandible height at cere; auriculars maximum length

(including filamentous extensions); tail length (measured by

inserting ruler between two central rectrices); tarsus length;

unfeathered (bristled) portion of tarsus; length of middle claw;

wing length (flattened), and maximum width of dark and light

bands on central portion of one central rectrix. Broken, heavily worn

or incompletely grown feathers were not measured. NJC also

measured culmen from skull for specimens from some museums.

Univariate statistics and PCAs were run in SYSTAT 12. Sexes were

combined for analyses, because most specimens of key taxa were

not labelled as to sex. Although it would be possible to examine

sexual size dimorphism in the better-represented taxa, and it may

be significant, we leave that to future studies and do not consider

the interpretation of our results to hinge on the matter.

Colour and pattern analyses

Plumage characters were documented at major collections with

holdings of multiple key taxa. Photographs were taken of nearly all

specimens examined and were used for later comparisons, but only

as a general guide and mnemonic.

In addition to traditional assessment of species limits under the

Biological Species Concept, we apply the system proposed by Tobias

4

P. C. RASMUSSEN etal.

Forktail 28 (2012)

et al. (2010) to measure the degree of phenotypic differentiation

between taxa. In this system an exceptional difference (a radically

different coloration, pattern or vocalisation) scores 4, a major

character (pronounced difference in body part colour or pattern,

measurement or vocalisation) 3, a medium character (clear

difference reflected, e.g., by a distinct hue rather than different

colour) 2, and a minor character (weak difference, e.g. a change in

shade) 1 ; a threshold score of 7 is required for taxa to be considered

separate species, but only three plumage characters, two vocal

characters, two biometric characters (assessed for effect size using

Cohen’s d where 0.2-2 is treated as minor, 2-5 medium, 5-10

major and >10 exceptional) and one behavioural or ecological

character may be counted (Tobias etal. 2010). However, we observe

that, in the case of nightbirds where vocalisations are crucial for

species recognition, this system may not give enough weight to

single key vocalisations and may give too much weight to plumage

characteristics, which in owls often show relatively high degrees of

individual variability.

Because the original diagnoses of heretofore named taxa placed

in spilonota and reyi were invariably based on inadequate material,

often involving single specimens and without reference to the most

similar taxa, and because spilonota proves to contain four different

taxa, we provide new diagnoses based (where possible) on larger

samples. We are constrained by the paucity of specimens from Cebu,

Tablas and Camiguin Sur, but good photographs now exist of these

and other taxa. Photographs often show features no longer existing

or never apparent in specimens, so we rely strongly on them to

supplement specimen material in these diagnoses. We have few data

on immatures, so these are not included in the diagnoses. In general

immatures are less well-marked and fluffier than adults.

Owing to the generally small sample sizes and poor labelling of

key taxa we were unable to analyse whether sexual dichromatism exists

in these owls. However, we suspect that it may, as in some other Ninox

owls. In photographs that show what is almost certainly a singing

pair for each ‘unstreaked’ taxon, one individual has noticeable whitish

streaking on the lower underparts, while the other is plain or barred

below. Whether this is a simple sexual difference is, however, unclear,

given that the white-streaked birds are in a minority among

specimens. Further fieldwork is needed to clarify this issue.

RESULTS

Acoustic analyses

The following are vocal comparisons by taxon or island (see

Methods for conventions used). For simplicity, we focus on a few

individual recordings and then summarise variations. Overall song

patterns and quality are summarised in Table 1, and univariate

statistics for measurements of songs in Table 2.

N. p. philippensis (Luzon)

(AV#2168, AV#2171, Luzon). — The non-duetted song of N. p.

philippensis (Table 1, Figure 2a) begins quietly, with single, short,

rather sharp cuk notes (each note comprising a short upslurred then

rapidly downslurred element, upper frequency limits c.0.9 kHz,

frequency range c.0.6 kHz, note duration c.0. 11s) spaced c.2-2.5 s

apart (at first slightly more than 2 s, then gradually accelerating

slightly to just under 2 s apart). The series becomes louder, and after

several single notes the bird then begins to add in soft, lower-pitched

(0.66 kHz) preliminary notes, single or doubled, as in boo, /CUK

and bu-bu,/CUK. There are several iterations of this, in which the

first element becomes progressively louder, but the main subsequent

element is still greatly accented, more strident and yapping, and

much broader-band, and the couplets or triplets are separated by

pauses of just over 1 s. This then changes into a four-element

repeating motif in which the first note is mellow and low as before,

and the subsequent three notes are broader-band, louder and

sharper, the last slightly softer and tailing off in pitch. In the four-

element motif, the second and third notes are farther apart than

the third and fourth, hence the rhythm seems halting; in addition

the third note is primarily upturned, while the second and third

are primarily downturned, and the third note has a slightly lower

maximum frequency. The four-note strophes are c.2 s in duration.

The series becomes slightly higher-pitched overall towards the end,

just exceeding 1 kHz, and the pauses between motifs are c.2 s long.

The series lasts 75-95 s and ends suddenly, or becomes intermittent,

with longer pauses (up to 8 s) between motifs, e.g.:

cuk_cuk_cuK_cuK_cuK_boo,/CUK_boo,/CUK_bu-bu,/CUK_bu-

bu,/CUK_boo,/CUK,CRIK-COok_boo,/CUK,CRIK-COok ... boo,/

C UK, CRIK- CO ok...

Shorter versions and song fragments (SOM la. 12453) may be

given, as may long series of single note types, and excited duets with

multiple birds joining in that may appear to have more than four

notes per motif (SOM lb. 13551).

N. p. centralis (Bohol, Negsros)

(AV#8971, Bohol; AV#10699-10700, Negros). — Song on both

islands is similar to that of N. p. philippensis. A Bohol recording

(SOM lc.8972) of a duet is similar to duets of N. p. philippensis.

AV# 10699, from Negros, has much longer (c.5 s) pauses between

initial notes than similar song strophes studied for nominotypical

philippensis, while those of AV# 10700 (SOM ld-e. 10700) are c.2 s,

as in nominotypical philippensis , and the strophes last 120-168 s.

N. p. centralis (SSquijor)

Song (SOM 1 f . 1 12a) is similar to that of nominotypical

philippensis, except that it is hoarser and the later strophes contain

more (sometimes several more) notes, especially in duets, which

have many short hoarse notes (SOM lg.99).

N. p. spilocephala (Mindanao)

The long song of spilocephala (Table 1, Figure 2d, continued as

SOM 1 j) differs dramatically from all others in consisting of very

slow, long, mellow, dove-like notes, singly or in couplets. The

strophe starts simple and becomes somewhat more complex, but

slows down considerably. The song (e.g. AV# 12462) starts with

single mellow, slightly slurred hUUUu notes that eventually

become longer, with the addition of an initial very soft low segment

to the note (now oo/hUU\Ju ), then adding a short final element

after a short pause (now oo/hUUUu, Huh). The fundamental

frequency is 0.37-0.53 kHz, with only the main ( hUUUu ) note

reaching the highest frequencies. All notes are slightly slurred and

convex (especially the main note) and have the same quality. Early

notes in a strophe are 0.4 s and the last couplets are 1.3 kHz. The

notes have one strong harmonic. Pauses between early notes are

c.2. 5 s and between later couplets c.4 s. The entire strophe may last

over 190 s. A transcription is: hUU\Ju_hUUUu_ ...

h UUUuJj UUUu_oo/hUUUu_oo/hUUUu_ ... oo/hUUUu_oo/

hUUUu_oo/ hUUUu, huu_oo/ hUUUu, huu oo/hUUUu oo/

hUUUu, huu ... oo/hUUUu, huu oo/hUUUu, huu oo /

hUUUu, huu ...

Duets (e.g. AV# 14263, Figure 2e) may be much shorter, e.g. 11s,

with the primary singer starting with just a few single notes that

quickly lead into oo/hUUUu, huu couplets like those described above.

The duetting bird sporadically adds in slightly but noticeably higher

couplets in which the first, stressed note is mostly upslurred and the

second, slightly softer note is mostly downslurred, transliterated as

IVAUUUU, JVUUUu. The lowest frequency of this couplet is 0.36

and the highest c.0.73. The couplet length is 1.2 s. The single

noticeable harmonic produced by this second individual reaches c. 1 .3

kHz. These same WA UUUU, IVUUUu couplets may be given with

another distant bird giving this same type instead of the oo/hUUUu,

Forktail 28 (2012) Vocal divergence and new species in the Philippine Hawk Owl Ninox philippensis complex

5

Table 1 . Qualitative summary characteristics of songs of each taxon and/or island population of Ninox philippensis ( sensu loto) studied.

Song characteristic

Table 2. Univariate summary statistics [mean±SD (range, n)] of measurements of song characteristics of each taxon and/or island population of

Ninox philippensis (sensu lato).

Song characteristics

Max. interstrophe

6

P.C. RASMUSSEN etol.

Forktail 28 (2012)

Figure 2. Typical songs of each major taxon in the Philippine Hawk Owl Ninox philippensis ( sensu lato ) complex. For each song, the first row is

the waveform, the second is the sonagram, and the third is the spectrum: (a): Philippine Hawk Owl N. p. philippensis, probably non-duetted long

song of single bird singing in different directions, hence the variations in amplitude late in the song series; AV#21 71, PCR, Luzon; (b): Camiguin

Hawk Owl Ninox new species 1 , duetted series of short strophes; AV#1 3554, ROH, Camiguin Sur; (c): Camiguin Hawk Owl Ninox new species 1 ,

long song by single bird; AV#1 3622 ROH, Camiguin Sur; (d): Mindanao Hawk Owl N. spilocephala, long song by single bird (series continued in

SOM 1 j); NSA Wildlife ref. #132605, S. Harrap, Mindanao; (e): Mindanao Hawk Owl N. spilocephala, short duetted songs; AV#12461, ROH, Mindanao;

(f): Sulu Hawk Owl N. reyi, series of short probably non-duetted strophes; AV#1 4577, ROH, Tawi Tawi; (g): Mindoro Hawk Owl, first half of long

duet (continued in SOM 1 1); AV#1 1 507, ROH, Mindoro; (h): long song of single Romblon Hawk Owl Ninox spilonota new subspecies; AV#10803,

DNS A, Tablas; (i-j): long song of single Romblon Hawk Owl N. spilonota on Sibuyan, AV#1 3637, ROH, Sibuyan; (k): long song of single Cebu Hawk

Owl Ninox new species 2, AV#1 261 2, BD, Cebu; (I): duetted song of Cebu Hawk Owl Ninox new species 2; AV#1 0805, LMP, Cebu.

a)

1 . . . „z

95 100 105 110 115 120

time (s)

.HI . in . Ui . m

OS It) , , 1.S 2.0 25

kHz

h . . — 4U-

120 125 >30 IBS KO

time (s)

i 1 i .ui .in .in .in

b)

c)

Camiguin Sur

10 IS 20 25

•IS SO SS 60

BS 40 4S SO

d)

e)

f)

05 1.0 15 20 25

Forktail 28 (2012) Vocal divergence and new species in the Philippine Hawk Owl Ninox philippensis complex

7

Huu couplets, or the latter type may grade into the former type (e.g.

AV#8088). Otherversions include (AV#805 6) aseries oflong dove-

like notes ending in a stressed barking upslur, a version in which the

first note of each couplet is more accented and staccato (AV#81 1 1),

or single short gruff notes. None of the vocalisations of spilocephala

closely resembles those of any other taxon in the group, although

there is some resemblance to the much shorter, sharper, two-note

portion of- the long song of philippensis.

N. p. mindorensis (Mindoro)

The song (Table 1, Figure 2g, continued in SOM ll) is distinctly

high-pitched, and composed ol long series of thin, slightly

descending squealed whistles fiiiiew (each c.0.8 s long) separated

by pauses of c. 1 s, then gradually changing into descending squealed

whistles that become hoarse raspy downslurred screeches, e.g.

jiiikshreiv , the whistle-screeches often preceded by several very short

high-sounding tittering toots, 0.1 -0.3 s. The screeches and toots

45 . 50 55 60

T tt V't H 1

♦ ■» ♦ ♦ 4 ¥-■+*- — -

fihvvrvmHir

fcrr

r ' ' TTtTT' 1 11T

i)

05 W 15 Z0 2

kHz

-4 - 4 - 4 - 4 - 4-

Sibuyan

1

1JS 140

h 1 1 'i

k)

10 is

ill ! " HI 'nil ' ih m U

i -i > ■> k 4 4 4 ^ 1 4 4 4 f f H

i ID)) i) nn i) m in m ) o n

4 >4 »4 4 4 >4 ‘4 >4 Mf lM If f 4

65 70 75 80 as

time (s)

j it M i \) i> h \ \) |) m (n i\

i» >4 M it t

8

P.C. RASMUSSEN etal.

Forktail 28 (2012)

are duetted, both birds often chiming in simultaneously giving both

note types, which may overlap closely. Whistled notes may rise

slightly at the start before falling again, and others may be entirely

screeched. Duetted song may also consist of much longer series

almost entirely of long whistled notes, some of them strongly

frequency-modulated (SOM lm-n).

The song oi mindorensis bears no resemblance to those of

philippensis , spilocephala , reyi or Camiguin Sur birds. It somewhat

resembles songs of Tablas, Cebu and Sibuyan birds (as noted by

Allen [2006], except that no recordings were then available for

Sibuyan) in being composed of thin whistles and hoarser rasps,

but it is very different in frequency and structure, especially in its

much longer-drawn whistles and screeches, the near-constant

frequencies of the long whistled notes, presence of high tittering

toots, much harsher screeches, and lack of rhythmic repetition of

lower notes.

N. p. spilonota (Tafoias and Sibuyan)

The song of Tablas birds (Table 1, Figure 2h. 10803) consists of

long slow series of single short, steeply falling wistful whiny whistles

\FYew separated by pauses of several seconds, and later in a strophe

often changing into a hoarse version \FYuRsh, then into two-note

and finally three- to (occasionally) four-note versions, e.g. (tut-)TUT-

TIY\lEw-hut near the end. Hoarser note-types have a few widely

spaced harmonics (SOM lo). The maximum fundamental

frequency is 2 kHz, and that only at the start of strongly

downslurred notes, while most of the notes are 1-1.7 kHz.

Song in Sibuyan (Table 1, Figs. 2i— j. 13637) is similar to that in

Tablas, but sounds slightly more croaking and is even slower, with

many introductory notes nearly level, barely downslurred, and with

strong harmonics. The later notes in a strophe are typically clearly

four-noted, and each note is more similar to the others except the

longer, stressed, downslurred third note, e.g. TOOT, TOOT, TIYUEW,

TOOT.

N. p. spilonota (Cebu)

In Cebu, song (Table 1 , Figure 2k) is highly variable in rhythm and

note type, although in quality notes are of two main types, gruff

staccato chucks and plaintive short downslurs, and (less often)

metallic treefrog-like upslurred abrupt bwick! notes (several of

which may be given in quick succession), low clear abrupt duit!

bell-tones (sometimes quickly doubled), and hoarse white-noise

screeches lasting c.0. 6 s (Figure 21, SOM lq-r). Overall, compared

to Tablas and Sibuyan birds, to which it is most similar, the song in

Cebu is much faster (but relatively low-pitched, with notes mostly

below 1.5 kHz and even down to 0.8 kHz) and lacks the longpauses

between notes within a strophe so characteristic of these other taxa.

Non-duetted strophes are long series of short IKYeut notes

(separated by pauses of only c.0.9 s), running into djuk, /

KYEUr_djuk , IKYeut... series and even four-noted, more complex

versions. Duets are medium-length strophes starting with several

low soft djuk and gwick notes, running into djuk, \KYEUr-givuck,

djuk, YKYEUr-gwuck , etc., petering out after c.30 s and then starting

again after a pause of a few seconds. Despite similarities to Tablas

and Sibuyan birds the numerous notable and consistent differences

exhibited by Cebu birds are very striking.

N. p. spilonota (Camiguin Sur)

(AV# 13554, 13556-13557). — Duetted strophes (the common type

available to us; Table 1, Figure 2b) differ greatly from those of all

other known taxa. They begin with a few sporadic, very soft, very

low (0.28 kHz) mid-length gruff notes, then turn into a low, very

hoarse growl (e.g. 0.9 s) that quickly leads into rapid laughing mellow

barks (0.36-0.69 kHz), recalling the chorused yapping of distant

mid-sized dogs. The duetted notes typically overlap only partially,

so the strophe sounds faster than is each individual’s contribution.

and the rhythm is non-uniform and jerky. Each note (excepting the

growl) is strongly convex (highest part in the middle), and note shape

is quite uniform; individual note length is 0.10-0.15 s, and pauses

between note-pairs in the first two-thirds ol the main strophe can

be as long as 0.1 s, then to 0.2 s between this part (signified by CUR

notes below) and the accented ending ( BOO notes). Strophes

(including introductory grace notes) last 5-1 1.5 s and are separated

irom each other by 7-18 s. Strophes slow down slightly near the

end, with the final few notes slightly longer, louder and higher-

pitched. There are often a few stray loud notes just following the

well-defined strophe (SOM lh. 13557). Harmonics are prominent

throughout the entire strophe following the growl, which has very

broad bandwidth. The highest detectable harmonics on available

recordings are at c.2.6 kHz, and there are up to five bands of

harmonics above the fundamental frequency, although only the first

harmonic (peaking at J.25 kHz) is very prominent. Growls do not

occur elsewhere within the strophe but the introductory growls are

duetted, with the second individual joining in just after the first, or

they may be shorter growls separated by short pauses, e.g.:

(huk__huk__)burrrrrrrr’CUR,CUR ’r,CUR ’ur, CUR, CUR ’ur,

CUR ’ur, CUR dr, CUR, CUR ’r,CUR r, CUR r, CUR’ ur, CUR’ CUR-/

BOO-BOOR-BU’U _ (boo)

Lone birds (Figure 2c, SOM 1 i) also give shorter, low-intensity

series of c.6 boo notes, the series lasting c.2 s, and the fourth note

highest and stressed (e.g. AV# 13602), or long series starting with

single notes, then two, the first stressed, then three, first still stressed,

then grading into strophes of up to seven low hoots followed by up

to six higher stressed hoots, the first of the higher notes being most

stressed, but all evenly spaced and of approximately the same length

(e.g. AV# 13622).

N. p. reyi (Tawi laws)

(AV# 10802). — The song of this taxon is extraordinarily different

in tone from those of all other known taxa (Table 1, Figure 2f),

being a hollow wooden knocking, usually in duetted short strophes

of a few to several seconds, separated from other such strophes by

short pauses (1-2 s). Each strophe starts with several quiet low

clucks at the rate of 1/0.1 s, then gets louder and higher-pitched,

although each note has broad bandwidth; the strophe then slows

down, often dropping in pitch and becoming more emphatic and

higher again. The duet has a stuttering rhythm overall and there

are no harmonics. Each note has broad bandwidth, the earlier ones

c.45 kHz and the louder, higher ones c.80 kHz, and note length is

c.0.0 1 s for earlier notes and 0.02 s for most others, some perhaps

as long as 0.03 s, e.g. kt-kt-kt-kt-kt-kt-kt-/KRT-KRT-KRT-KRT-

KR T\ -KLAK-KLAK-KLAK-KR T-KR T-KR T-KR T-KR T-KR 7V

TOK'.-TOK!

Strophes may be much shorter, e.g. kt-kt-kt-KR T-/T OK!- T OK!,

and successive strophes in a series can vary considerably in length

(SOM lk). They can also be more uniform, but still with slightly

higher pitch and emphasis near the end, or they can be mostly

uniform kt- series until a sudden speeding up and more clattery

pattern at the end. This clattery effect occurs when clacking notes

are introduced by a very short low grace note that increases the

hollow knocking quality. The latter notes are evidently responsible

for the taxon’s local name of ‘lukluk’ (Allen 1998).

This form also gives a simple song of hollow triplets continued

for longperiods, e.g. tluk’tluk’tluk_tluk’tluk’tluk ... tluk’tluk’tluk. The

motifs are c.0.7 s in duration and separated by usually 3-5 s

(commonly 4.5 s), and consist of three principal elements, each

preceded by a very short, lower, introductory element. The

frequency range of the principal elements is about 0. 5-0.9 kHz.

This song type somewhat resembles in quality the typical song of

Cinnabar Hawk Owl Ninox ios of Sulawesi, although that species’s

song is distinctly two-noted (King 2005, Hutchinson et al. 2006),

and no songs comparable to the other types of N. reyi are known.

Forktail 28 (2012) Vocal divergence and new species in the Philippine Hawk Owl Ninox philippensis complex

9

Very occasionally a longer, very hoarse growl or hoarse hoot is

interspersed with the clacking notes.

Principal components analysis

The PCA using all groups (with philippensis and centralis combined;

Table 3, Figure 3a) shows that Tawi Tawi reyi, Camiguin Sur

spilonota , Mindanao spilocephala and Mindoro mindorensis are

highly differentiated from each other and from ‘core’ groups (those

with intermediate characteristics) on the vocal characteristics

measured. Factor 1 contrasts mainly frequency and note length with

number of notes, so factor scores for individuals of the high-pitched,

long-noted mindorensis are uniformly higher on this axis than other

groups, while those of the short-noted, low-pitched Camiguin Sur

spilonota and (to a lesser extent) reyi are lowest on this axis. Factor 2

is more difficult to interpret, but it contrasts mainly frequency

differences within and between notes with length of shortest notes

and length of shortest internote pauses; on this axis reyi and

Camiguin Sur spilonota separate out clearly from each other, and

spilocephala separates out cleanly from the philippensis group.

The PCA using only core groups (with all island forms of

philippensis and centralis graphed separately, and including Tablas

spilonota , Sibuyan spilonota and Cebu spilonota ; Table 3, Figure 3b)

shows that all philippensis/ centralis forms group together, with the

two Siquijor recordings somewhat marginal, while Cebu, Tablas and

Sibuyan are all fairly similar to each other on vocal measurements,

Tablas birds being somewhat intermediate between those of Cebu

and Sibuyan. Factor 1 is mainly a contrast between frequency and

note length versus number of notes/strophe and degree of inter¬

strophe frequency change, while Factor 2 mainly contrasts number

Table 3. S ummary results of principal components analyses of

measurements of vocal characteristics for all groups and core groups

only ( philippensis and centralis, Tablas spilonota, Sibuyan spilonota, and

Cebu).

of notes, maximum note length, and frequency change within notes

versus minimum note length and minimum pause length.

Summary of vocalisations

To summarise the main points of the vocal evidence reviewed above

and in Tables 1 and 2, the Camiguin Sur population and reyi from

Tawi Tawi give many more notes per strophe than do other forms.

Tawi Tawi reyi is strikingly divergent from all others in giving almost

exclusively very short percussive toneless notes in extremely rapid,

rhythmic strophes. Camiguin Sur birds and, to a lesser extent,

Mindanao spilocephala are much lower-pitched than other taxa,

while Mindoro mindorensis is much higher-pitched than others. The

song of Mindanao spilocephala is unique in several ways, including

the stressed first note in the couplets, low number of notes per

strophe, and consistently mellow tone. Songs of nominotypical

philippensis and centralis are similar to each other; they are in the

middle of the pitch range for the entire group, and differ from other

taxa in the halting, regular rhythm of later notes, strongly convex

note shape, without whistles and normally without growls, and

mellow barking quality. Recordings from Siquijor appear to show

rather more distinctive vocalisations, but further and higher-quality

material is required before taxonomic conclusions can be drawn.

While Cebu birds are similar in frequency range and number of notes

to Tablas and Sibuyan spilonota, their songs have several peculiarities

including the rapid, continuous series, varied unmatched note types,

and often erratic delivery. Tablas and Sibuyan spilonota , while by no

means identical vocally, do share most characteristics, and on present

data appear to represent distinctive races. The other taxa summarised

here all show an extraordinary degree of vocal differentiation in a

group of birds for which vocal communication is innate and of

paramount importance in species recognition.

Mensural analyses

Our sample of Ninox philippensis sensu lato, with all island

populations kept separate (Table 4), shows that, while we do not

have adequate sample sizes of most taxa for significance testing,

mensural differences between philippensis, centralis, proxima and

ticaoensis as curirently recognised are not striking. Flowever, in a PCA

of nominotypical philippensis, centralis , proxima and ticaoensis

(Figure 4, Table 6), centralis from Siquijor separates out from almost

all centralis from Negros and Bohol on Factor 1, a size axis, while

the sole ticaoensis specimen included falls at the extreme small end

of this axis. Masbate proxima is medium-sized on this axis (Figure

4, Table 6).

Measurements (Table 4) also show that all island populations of

spilocephala are quite similar in size and proportions. We therefore

combine these poorly if at all differentiated forms for subsequent

analyses. Table 5, in which only clearly differentiated taxa (named

or unnamed) are included, shows that there is considerable size

variation, with philippensis, spilocephala and mindorensis all being

small, while reyi and the four spilonota populations from Camiguin

Key

Q) spilocephala

O Camiguin

9 'eyi

□ philippensis

□ Cebu

| Sibuyan

Tablas

^ mindorensis

Factor 1

Figure 3. Results of PCAs of

measurements of vocal characteristics

for (a) all groups (philippensis and

centralis combined) and (b) core

groups (philippensis and centralis from

different islands graphed separately;

Tablas spilonota, Sibuyan spilonota,

and Cebu).

10

P. C. RASMUSSEN etal.

Forktail 28 (2012)

Sur, Tablas, Sibuyan and Cebu are all large in most measures. PCAs

(Figure 5, Table 6) consistently show this same pattern. On Factor

1 , which is a strong size axis, taxa are either small or large, although a

few individuals in both groups are intermediate in size. On Factor 2,

the highest-loading variables are the two tail-banding measurements

(Figure 5a, Table 6). On this axis ^philippensis, with its broadly banded

tail and to a lesser extent the Cebu bird, score high, in contrast to

narrow tail-banded taxa including spilocephala, mindorensis and the

Camiguin Sur birds. Although spilocephala is small, it has the longest

(and densest) ear-covert extensions of all taxa (readily visible in

photographs of live birds; Figure 6b), with philippensis not too

different; all the larger taxa have relatively shorter and less prominent

ear-covert extensions, which are rarely apparent in photographs. In

fact, auricular length is much the highest-loading variable on Factor

3 of the PCA (Figure 5b, Table 6), on which spilocephala loads

highest, with considerable overlap with philippensis and a few Sibuyan

birds.

Figure 4. Results of PCAs of skin specimen measurements of Ninox

philippensis philippensis, N. p. centralis, N. p. proximo and N. p. ticaoensis,

with all islands shown with different symbols but previous subspecies

groupings within dashed polygons. Summary statistics in Table 6.

Table 4. Summary statistics for univariate measurements [mean±SD (range, n); (in mm)] of island populations of Ninox philippensis sensu lato,

using previously recognised racial divisions. Not included here are mindorensis and taxa formerly united in spilonota. Ht = height; L = length.

Variable

Forktail 28 (2012) Vocal divergence and new species in the Philippine Hawk Owl Ninox philippensis complex

11

Table 5. Summary statistics for univariate measurements [mean±SD (range, n); (in mm)] of major taxon groups of Ninox philippensis (sensu lato).

reyi 24.2±1.0 13.7±1.1 8.1±0.3 21.0±5.2 97.5±4.7 35.6±2.5 17.7±2.4 12.5±0.6 186.7±8.2 6.6±1.2 3.2±0.8

(23.0-26.0,12) (10.7-15.2,11) (7.7-8.5,11) (15.7-31.3,10) (88.7-107.6(12) (31.9-39.4(11) (14.4-21.1,8) (11.5-13.5,11) (172-195,15) (5.2-9.2,11) (2.4-4.3,11)

Key

O spilocephala

O Camiguin

• reyi

□ philippensis

□ Cebu

■ Sibuyan

■^Tablas

— If mindorensis

Figure 5. Results of PCAs of skin

specimen measurements of Ninox

philippensis ( sensu lato), with all

major taxa shown with different

symbols and those with more than

two specimens grouped within

dashed polygons. Summary

statistics in Table 6.

Unfortunately, sample sizes of all three new taxa formally

described in this analysis are very small, and therefore we can only

guess at the ranges of their measurements. However, the sample size

for Sibuyan is moderate, and while the single Tablas specimen

included falls just within the range of measurements for Sibuyan

birds (on the small side for all but one measure: Table 4), it

consistently falls outside all Sibuyan birds, on the small side of Factor

1 (a size axis), in the PCA (Figure 5). This could either indicate

minimal or moderate difference in size between these two

morphologically and acoustically fairly similar taxa. There is great

plumage variation in photographs of Cebu birds, and we assume

they also vary in size as with other taxa. The only Cebu specimen

known is as large as the smallest Sibuyan birds but has a

proportionately long tarsus, the measurement of which has been

independently verified (Table 4, Figure 5). Camiguin Surbirds may

vary less in size and plumage than other taxa, based on the two

specimens studied and photographs of at least three further

individuals.

Colour and pattern analyses

Most taxa in the. Ninox philippensis ( sensu lato) complex are at least

moderately variable in plumage, particularly in the case of most of

the small island forms formerly united within spilonota or reyi.

Given the small sample sizes available for most of these taxa,

delineating diagnostic plumage characteristics is problematic.

Plumage or structural characters that are unique to a single taxon

are scarce. It may not even be possible to attribute every individual

Table 6. Summary results of principal components analyses of

measurements of morphological characteristics for all groups and

philippensis ( sensu stricio) island taxa only (philippensis, centralis,

proximo, ticaoensis). Ht = height; I = length; w = width.

All groups philippensis only

Component

variance explained

12

P. C. RASMUSSEN etal.

Forktail 28 (2012)

Figure 6. Photographs of (a) Luzon Hawk Owl N. philippensis, Dolores, Quezon, Luzon, 1 6 January 201 2 (R. J. Quisumbing); (b) Mindanao Hawk

Owl N. spilocephala, PICOP, Mindanao, February 201 1 (R. O. Hutchinson); (c) Mindoro Hawk Owl N. mindorensis, Sablayan, Mindoro, December

2008 (James Eaton); (d) Romblon Hawk Owl N. spilonota, Tablas, 4 March 201 2 (Marc Thibault); (e) Camiguin Hawk Owl (new species), Camiguin

Sur, 8 June 201 1 (R. O. Hutchinson); (f) Cebu Hawk Owl (new species), Cebu, 3 January 2012 (Christian Artuso); (g) Sulu Hawk Owl N. reyi, Tawi-

Tawi, January 2012 (R. O. Hutchinson).

Forktail 28 (2012) Vocal divergence and new species in the Philippine Hawk Owl Ninox philippensis complex

13

to taxon without recourse to measurements or knowledge of island

of origin. However, on a combination of characters, each of the

taxa we recognise below can be readily identified.

The main plumage and other external features that differ

between taxa in the Ninox philippensis ( sensu lato ) complex are

summarised in Table 7. While plumage differences between N.

philippensis ( sensu stricto ) and all other taxa are straightforward and

consistent, those between all other taxa are less obvious and are

only diagnostic in combination, although the conformation of the

streaks of the lower underparts of spilocephalus is highly distinct

except from a very few individuals of reyi. The profound differences

in vocalisations demonstrated above are entirely inconsistent with

treatment as a single species, but these differences are largely

congruent with additional evidence from mensural analyses and

plumage and soft-part colours. Under the Biological Species

Concept, we consider that Ninox philippensis ( sensu lato ) represents

seven species, ofwhich two formerly treated under the race spilonota

are undescribed species and a third is an undescribed subspecies.

Under the Tobias criteria (Tobias et al. 2010), all unstreaked forms

differ from the taxa in N. philippensis sensu stricto by their

exceptional vocalisations (score 4) and unstreaked versus streaked

plumage (score 3), plus various other characters not necessary to

enumerate here; total at least 7.

Diagnoses

N. p. philippensis

Our evidence indicates clearly that the streak-breasted taxa form a

highly consistent group very weakly differentiated in plumage,

morphometries and voice, and without attempting to score these

characters, all of which appear to be minor, we propose that the taxa

philippensis, proximo., centralis and ticaoensis be considered a single

species, Luzon Hawk Owl Ninox philippensis. Indeed, given the

variability in plumage, size and voice we feel that the case for

subspecific separation in this widespread northern and central

Philippine species is not solid. The Masbate rac cproxima (diagnosed

on a sample size of two as ‘similar to philippensis but larger and with

a relatively shorter tail’, but also darker above, with less pronounced

pale tail-bars, reduced pale spots on wing-coverts and wings, and

coarser, darker stripes below: Mayr 1945) is particularly doubtful,

and we subsume it here in the nominotypical. The small dark Ticao

race ticaoensis (diagnosed in a comparison with proxima on a sample

size of three as having ‘the upperparts darker, the ventral streaking

sharper and darker, and the light barring on the tail narrower’:

duPont 1972), is at best marginally distinct. Birds from Siquijor,

bearing the name centralis (diagnosed as ‘much larger than proxima

and with a long tail’, with no rufous or tawny tones in the ‘dark

earth brown’ of the upperparts, much reduced pale spotting on the

scapulars and upperwing-coverts, ill-defined streaking below with

an ochraceous wash to the white feathers: Mayr 1945), are certainly

larger than and, on our sample, vocally distinguishable from other

island populations currently placed with centralis , so we restrict

centralis to the Siquijor population (for which the name was

established) and place populations from other islands except Ticao

in the nominotypical, but further study is needed.

Diagnosis (from large series of specimens and many photos of

live birds; see Plate and Figure 6a). — Size small, with compact, sleek

plumage. Head has prominent if small and/or buff-tinged whitish

supercilium. Auriculars are uniform plain dark brown, concolorous

with or slightly paler than crown, with fairly long, prominent

filamentous extensions; facial disk not well defined. Above, uniform

brown above from crown to rump, lacking barring or speckling.

Scapular patch with some largely white to pale buff feathers with

dark brown edgings, or large white/buff patch on mid-distal portion

of dark brown feather; barring or mottling lacking. Larger outer

wing-coverts have large white spots, usually one but sometimes two

per feather; barring is lacking. Tail banding is variable in intensity

but most have fairly weakly marked pale bands that are much

narrower than the dark bands. Below, throat narrowly white, with

fine dark streaks; underparts prominently streaked dark brown or

rufescent-brown, either crisply or indistinctly, on a white or rufous-

tinged background. The streaks are not edged with darker colour

or intruded by spots or bars. The streaks become more distinct,

narrower, and longer on the central lower underparts, where they

typically cover the central half to third of the feather, the remainder

being white or whitish, and the undertail-coverts are white without

markings or with a few faint and/or narrow dark markings. Bare

parts: irides bright yellow, often appearing slightly paler, more

lemon-yellow on outer ring; narrow orbital ring pale tan, not

conspicuous; cere and bill pale olive-yellow, the culmen ridge and

tip slightly yellower; tarsi and toes bright yellow, claws almost

entirely black, the bases paler. Vocalisations are long, mid-pitched

series of strongly convex barking notes starting with single notes

and ending with about four halting notes, the first note softest,

without whistles and usually without growls.

Table 7. Main qualitative plumage and soft-part character differences between taxa in the Ninox philippensis ( sensu lato ) complex.

14

P. C. RASMUSSEN etal.

Forktail 28 (2012)

A/, p. spiloeephala

Tweeddale (1879) established this taxon simply on the basis of the

'bright rufous’ to ‘pale tawny rufous’ spotting on the forehead and

crown, a feature absent in the otherwise similar form from Luzon

with which he compared his extensive (6 males, 1 3 females) material

from Mindanao, noting that this material was variable in coloration

above and pattern below. He also observed, without proposing these

points as possessing diagnostic force, that male spiloeephala were

longer-winged than male philippensis , but that spiloeephala by his

measurements was shorter-legged than R. B. Sharpes of philippensis.

However, Mayr (1945) pointed out that spiloeephala is ‘the principal

connecting link’ between the extremes represented by philippensis

on the one side and mindorensis on the other, and in this he was

clearly referring to the fact that spiloeephala possesses underparts

which conflate the streaking of philippensis and the barring of

mindorensis.

Specimen material confirms the apparent intermediacy of

plumage for spiloeephala , which shows brown and white streaking

in varying degree on the belly, as in philippensis and its Visayan

representatives. However, this streaking is only vague on the breast,

which is interrupted by a disorganised and inconsistent pattern of

bars and mottling in many specimens. In addition, the streaking of

the lower underparts differs from philippensis in being dark-edged

and irregular (see Diagnosis below). The spotting and stippling on

the crown (which extends onto the mantle) is similar between

spiloeephala and the bar-breasted forms.

Thus the form spiloeephala differs significantly from all the

others in plumage, morphometries and voice, and we propose

that this also be considered a species under the name Mindanao

Hawk Owl. Its mam song and other vocalisations are highly

distinct from that of any other form (Tobias score 3). Moreover,

in its combination of barred breast and streaked belly, it displays

what we judge to be a major character difference from any other

taxon (score 3). It also has exceptionally long, full auricular

extensions (from data in Table 5 effect size vs philippensis — 0.74;

score 1).

Diagnosis (from large series of specimens and photos of at

least three live birds; see Plate and Figure 6b). — Size small,

with especially short tail. Plumage compact and sleek. Head has

a very small, indistinct or absent white patch between eye

and supercilia. Auriculars appear nearly uniform dark brown but

they are in fact vaguely speckled and/or barred; filamentous

auricular extensions normally very long, full and profuse, more so

than for any other taxon. Above, crown, nape and upper mantle of

adults is strongly and usually evenly spotted with buff, the spots

becoming weak on the upper mantle and disappearing. In an

immature, the crown is unspotted brown but there is a collar of

weak pale buff spots on the upper mantle. Mid- to lower mantle to

rump is uniform dark brown. Uppertail-coverts may have a little

vague fine pale barring. Scapular patch similar to those in

philippensis group. Larger wing-coverts have both large white spots

and narrow buff barring. Tail is typically more distinctly banded,

with narrower dark bands, than for philippensis. Below, throat is

narrowly white with medium-broad, distinct blackish-brown shaft-

streaks, becoming broader on upper breast. Sides of neck and breast

are weakly barred and spotted dark brown and buff, and central

breast is mostly streaked. Lower underparts have mostly

longitudinal dark markings but these are typically somewhat

rounded, with dark chevron tips and irregular dark markings edging

the paler brown internal markings. Undertail-coverts white, often

with tiny dark spots. Bare parts: irides pale yellow; narrow bare

orbital ring tan-brown, not conspicuous; cere and bill pale olive,

more yellow on culmen ridge near tip; feet bright yellow, claws

nearly all black, paler at bases. Vocalisations are low-pitched,

mellow, slow, and two-noted, with first note of each couplet

stressed.

N. p. mindorensis

Ogilvie Grant (1896a) characterised this form as ‘nearly allied to

N. spiloeephala... in having the top of the head and nape barred [sic]

with buff’ but ‘the whole of the underparts... tawny buff,

transversely barred with brown, while in... N. spiloeephala , though

the breast is generally like that of the present species, the belly and

flanks are always white, with longitudinal reddish-brown shaft-

stripes’. He distinguished mindorensis from spilonota (which the

describers had listed as including the Mindoro population) simply

by its ‘much smaller size’.

Our examination of specimens confirms this basic diagnosis.

In addition, as noted above, the barring on the tail is denser in

mindorensis than in spiloeephala. Konig et al. (1999) mistakenly

suggested that mindorensis is larger and that the number of its tail-

bars is fewer than the Philippine Hawk Owl sensu lato.

Dutson et al. (1992) and Brooks et al. (1995) were the first in

the modern era to realise that the form mindorensis is so different

from Luzon birds that it appears to represent a separate species. After

hearing tapes by P. Morris made in 1999, Konig et al. (1999)

concurred and established it as Mindoro Hawk Owl N. mindorensis.

However, they made this move without knowing the songs of other

unstreaked forms, which they retained with N. philippensis ( sensu lato).

Only now is it confirmed that mindorensis possesses a highly distinctive

voice, with few similarities to those of the other unstreaked forms.

Compared to any other unstreaked form, mindorensis is

distinctly smaller (see Table 5: wing and tail shorter, with effect

size vs reyi -2.79 and -3.39 respectively; Tobias score 2), with a

weaker pale throat than Camiguin Sur and Cebu birds and reyi but

more prominent than in spilonota (score 1), especially narrow

underparts barring (score 1), high-pitched song (score 2), and very

short toots in climax song (score 2). There are other unique vocal

characteristics as well (not scored), such as the unusually long notes.

Diagnosis (from large series of specimens and photos of

numerous live individuals; see Plate and Figure 6c). — Size small,

with small bill, tarsus mostly feathered. Head moderately patterned,

with small short white supercilia; facial disk rather pale brown, small,

with moderately distinct rim, and usually rather short filamentous

auricular extensions. Above, crown and nape finely barred dark

brown and buff to whitish, but bars broken resulting in overall

appearance of speckling. Speckling grades out on upper mantle, and

rest of central mantle uniform, with vague fine barring at edge of

mantle and on rump. Scapular patches are extensively white with

irregular dark margins and dark spots. Wing-coverts are extensively

and finely barred, with dark bands predominating, and with white

spots/bands on outer webs of larger coverts that have tiny dark spots

and scrawls within the white portions. Primaries have prominent

broad dark brown bands and narrower buff bands on outer webs,

and very vaguely banded inner webs. Tertials virtually unbanded

and with very fine, weak pale mottling, appearing uniform brown.

Tail banding is obscure and both dark and light bands appear narrow.

Below, throat-patch pale buff, not very clear-cut but more so than

for Sibuyan and Tablas birds. Underparts highly variable but

typically the most finely barred of any taxon in the N philippensis

[sensu lato ) complex. Underparts vary in tone from pale buff to

saturated, and may have fairly prominent white streaking below. Bare

parts: irides have mustard-yellow inner ring, brighter lemon-yellow

outer ring. Bill and cere pale dull yellow, tinged greenish on sides.

Tarsi and toes dull olive -yellow, claws black, paler near bases.

Vocalisations are high-pitched long whistles often ending with a

screech, and often starting with tittering high toots.

N. p. spilonota from Tablas and Sibuyan

Bourns & Worcester (1894) gave the name spilonota to unstreaked

birds from ‘Cebu, Sibuyan, Tablas, Mindoro’, but their description

made no comparison with any other Ninox taxon, so the diagnostic

features of spilonota are not immediately obvious. The description

Forktail 28 (2012) Vocal divergence and new species in the Philippine Hawk Owl Ninox philippensis complex

15

refers to an overall colour of ‘fulvous brown’ above, with light rufous

speckling from head to mantle and wing-coverts, a tail ‘nearly black

with nine narrow transverse bands of light rufous brown’,

underparts ‘rufous brown... many feathers of breast and abdomen

with fulvous brown spots’. McGregor (1905) noted that the

scapulars, normally white in many Ninox , are reduced to ‘bars of

pale fulvous’ in the specimen to which he had access. We now verify

that this character holds in the far more extensive material we have

examined from Sibuyan.

On Sibuyan and Tablas, spilonota is diagnosed by its plain head

without white supercilia (score 2), lack of notably paler throat (score

3), lack of white spots in scapulars (score 2), and distinctive song

pattern and quality (score 3).

Diagnosis (from several Sibuyan specimens and two photos of

a live bird, and one Tablas specimen and two photos of a live bird;

see Plate and Figure 6d). — Size large, with large bill. Plumage rather

lax, and general appearance very plain. Head plain brown, lacking

pale supercilia or any strongly contrasting pattern, and with

indistinct facial disk. Above, finely speckled buff on dark brown

crown and nape, mantle unmarked, scapulars and tertials entirely

and distinctly barred dark brown and buff, without large white

scapular spots; uppertail rather narrowly but distinctly barred dark,

with narrower buff bars. Below, no prominent pale throat -patch.

Underparts typically have ochraceous background, either lightly

and indistinctly marked with large darker brown broken bars and

spots, sometimes looking almost plain ochraceous-brown, or

completely barred below, with fairly even, dark brown bars on

ochraceous background; rarely has fairly prominent white streaking

on lower underparts. Bare parts: irides bright yellow, not differing

between inner and outer rings; narrow bare orbital skin dull tan-

brown, not standing out from surrounding feathers; bill and cere

pale olive -yellow, yellowest along culmen and at tip. Vocalisations

are short falling whistles, often hoarse, starting singly then changing

into a three-note (Tablas) or four-note (Sibuyan) version.

Populations of both islands are covered by this diagnosis;

however, Tablas birds differ from those on Sibuyan and require a

name.

Ninox spilonota fisheri subsp. n.

Romblon Hawk Owl

Diagnosis. — Smaller than N. s. spilonota. Vocalisations differ from

those of N. s. spilonota as indicated above.

Holotype. — USNM 314875, female, Badajos [now San

Agustin] , Tablas, 1 8 September 1 892, collected by D. C. Worcester

and F. S. Bourns; Ex. Menage Collection no. 373. Culmen from

cere 15.5 mm, wing 188 mm, tarsus 30.2 mm, tail 96.1 mm.

Description of holotype. — Medium-large, weakly marked round-

headed owl, largely barred above and nearly plain ochraceous below.

Head. — S mall, with short, inconspicuous partially concealed

whitish supercilium extending to about half-way over eye;

auriculars appear rather dark brown with inconspicuous buff

spotting, their filamentous extensions fairly weak and short;

forehead and crown dark brown moderately narrowly and distinctly

barred and spotted. Forehead to hindcrown and sides of neck

profusely covered with small, closely spaced buff spots and short

buff bars. Throat pale buff overall (appearing Munseli 2.5Y 8/4),

each larger throat feather with a narrow darker brown shaft-streak

and often with narrow whitish outer webs and/or bases. Rictal

bristles largely pale buff, with black rachis that extend well beyond

the barbs, some exceeding the bill tip in length. Upperparts. —

Barring/spotting on crown becomes less regular and less

conspicuous on sides of neck and nape and grades into a rather

vague pattern on upper mantle of broad dark brown and narrower

buff (Munseli 10YR 7/6) banding. Scapulars lack any white but

have bold, broad but irregular rich buff (Munseli 10YR 8/6) and

dark brown banding, the dark brown banding with narrow blackish

edgings. Lower mantle and rump are essentially unstreaked warm

dark brown (Munseli 10YR 3/4). There is a single all-white

abnormal-appearing, badly worn feather in the upper mantle.

Wings . — Small external coverts of carpal area nearly uniform warm

dark brown, with very vague lighter buff speckling. Median wing-

coverts fairly distinctly barred with narrow buff bars and broad

dark brown; greater coverts yet more distinctly and broadly banded.

Tertials very broadly and evenly banded rather dark brown with

narrow, widely spaced, moderately distinct dull buff bars. Primaries

have broad dark and narrower dull bull banding with narrower,

irregular dull buff uniform dark brown inner webs. Underparts. —

Pale buff throat becomes darker, richer buff on lower throat and

grades into rich ochre underparts (Munseli 7.5YR 6/8). Medium

brown shaft-streaks on lower throat grade into rather indistinct,

broken but fairly broad medium brown bars on upper breast, which

break up and fade out on lower breast. Except for flank feathers

overlying thighs, which are indistinctly banded medium brown,

lower underparts are nearly solid rich ochre 6/8, with a few tiny

darker spots and vague mottling. Lower centre of belly has some

feathers with a few broad whitish tips, and undertail-coverts are

buffy-white with narrow buff tips. Tibial and tarsal feathering is

ochre-buff mottled whitish, and extends more than half-way to

distal end of tarsus. Tail. — Uppertail-coverts are dark brown with

a few narrow ill-marked dull buff bars. Uppertail has moderately

prominent broad dark brown and rather narrow pale dull buff

bands. Counting from base of feather (with uppertail-coverts

moved aside), there are 9 clearly discernible pale bands and 10 dark

ones. Bare parts. — In the dried specimen cere and bill are fairly

dark horn, with a paler, more orange -yellow culmen ridge. Distal

end of tarsus and toes have sparse, stiff, pale bristles. Toes and claws

on dried specimen are medium horn, tips of claws slightly darker.

Remarks. — Appears to be quite closely related to the

nominotypical form on Sibuyan, but differences at least in size and

vocalisations dictate recognition as a new subspecies.

Etymology. — We name this form in honour of the late Tim

Fisher, dedicated Philippine ornithologist and co-author of

Kennedy ei-u/.(2000).

N. p. spilonota from Camiguin Sur

Birds on Camiguin Sur are unique in their boldly, broadly barred

underparts (Tobias score 2), the combination of no white

supercilium with an extensive white throat (score 2), grey to whitish

eyes (score 3), and mostly pale claws (no score available; would be

1 ). Vocally they are highly distinctive as well, with their low-pitched,

rapid, multi-noted hooting duets (score 3 or 4). This taxon has

always been treated within the race spilonota , but clearly under both

the traditional Biological Species Concept and the scoring system

employed here it requires a name and recognition at the level of

species.

Ninox leventisi sp. n.

Camiguin Hawk Owl

Diagnosis (from FMNH 284397 and 399384 and photographs oi

at least three living birds; see Plate, Figure 6e, and journal front

cover). — Size large, with long, deep bill. Plumage appears full and

lax compared to other members of the N. philippensis complex,

except that ear-coverts have only short filamentous extensions

visible only at very close range. Face drab warm brown, the

auriculars barred but appearing plain at a distance; lacks pale

supercilium but may have small white patch above bill extending

just over eye; face shows no contrast with rest of head, and facial

disk border is not well marked. Above, crown very distinctly and

regularly barred buff and dark brown, more narrowly and obscurely

barred on upper mantle, the barring becoming vague and irregular

on lower mantle to rump. Wing-coverts are entirely boldly barred

dark brown, buff and white. Scapular patch includes feathers with

16

P. C. RASMUSSEN etal.

Forktail 28 (2012)

large unbarred white areas on outer web, just tip and base being

barred. Primaries are dark brown, the outer webs mainly dark buff

edged darker brown, with narrow pale buff bars. Tertials are

mottled and incompletely but prominently banded. Uppertail has

rather narrow dark and pale bars. Below, large bright white throat-

patch with a few narrow dark streaks extends to top of breast, with

some largely white feathers in uppermost central breast; white

throat-patch normally mostly hidden, but shown to be extensively

white in song display. Sides of neck and entire underparts heavily

and distinctly banded dark brown and buff. Underparts from breast

to abdomen have some irregular white or light buff banding;

banding below is heavier, broader and more distinct, with generally

darker, warmer brown tone on underparts than other taxa.

Undertail-coverts white with extensive dark barring and spotting.

Undertail irregularly banded with broad dark and narrow buffy

bands, banding sometimes obsolete. Bare parts: irides grey to

whitish or very pale yellow-green ( versus yellow in all other taxa);

narrow but distinct bare eye-ring mustard-yellow; bill and cere

mustard-yellow, more olive-green at base of bill; legs and feet

mustard-yellow, tarsi less extensively feathered than in most other

taxa in complex, mostly thinly bristle-covered; claws mostly pale,

only the tips fairly dark (claws mostly dark in other taxa, judging

from photos of live birds). Vocalisations are very low-pitched,

typically short strophes repeated after brief pauses, with many rapid,

irregular, barking notes per strophe.

Holotype. — FMNH 284397, female, Catarman Mountain,

Catarman, Camiguin province, Camiguin Sur Island, 17June 1968,

D. S. Rabor/W. S. Anguila; edge of second-growth, approx. 1,5002

(2,000-4,0002 on label; Balete et al. [2006] state 1, 5002, c. 450 m).

‘Irish [sic] dark white’. Culmen from cere 16.2 mm, wing 187 mm,

tarsus 37.8 mm, tail 90 mm.

Paratype. — FMNH 399384, female, Mt Timpong, Matugnao,

Mahinog, 13 June 1969, D. S. Rabor. Specimens of other species

taken on 13 June 1969 on Mt. Timpong were from 3,1502 ore. 950

m (Balete et al. 2006). Culmen from cere 16.3 mm, wing 181 mm,

tarsus 32.6 mm, tail 105.4 mm.

Description of holotype. — Crown dark brown, distinctly and

regularly barred with buff from forehead to nape and sides of neck,

the barring becoming less distinct on upper mantle and obsolete

on lower mantle. Scapulars moderately distinctly barred, outer

scapulars largely white with dark brown tips and bases. Wing-

coverts heavily barred with buff, the greater coverts barred mostly

white. Tertials moderately barred buff on dark brown. Uppertail

has broad dark and narrow dull buff bands. No pale supercilium,

and facial disk dark brown, weakly barred, and weakly defined.

Throat extensively bright white, the white extending well onto the

upper central breast. Rest of underparts heavily and distinctly but

irregularly banded dark brown on buff, with some short broad white

bars on central and lower underparts. Tarsal feathering dull brown

with narrow darker bars.

Remarks. — This is evidently the first known owl with grey or

whitish eyes. It is vocally highly distinct, normally giving a rapid,

low-pitched duet, the two birds facing and leaning towards each

other while very close together, with white throat puffed out and

pulsating, wings drooped.

Etymology. — We name this new species for Anastasios P.

Leventis, whose generous long-term commitment to BirdLife

International has been crucial in the stable development of the

organisation, and whose particular support for NJC has allowed

him to work extensively on Philippine birds and conservation issues

over the past decade.

N. p. spilonota from Cebu

The Cebu population of N. p. spilonota is diagnosed from Tablas

and Sibuyan birds, to which it is in vocal, morphological and

geographical terms most closely related, by its combination of fairly

prominent white supercilium and barred wing-coverts (Tobias score

1), barred back (score 1), prominent (when singing) white throat

strongly edged dark (score 3), relatively small bill (score at least 1),

long tarsus (score at least 1), and fast-paced song (score 2) with

multiple unmatched note types(score 2). Because n=l in the

morphometric sample (Table 4) we cannot generate an effect size

but the differences in bill and leg length are striking. We consider

that, under both the traditional Biological Species Concept and

the scoring system utilised here, this heretofore unnamed taxon is

a new species.

Ninox rumseyi sp. n.

Cebu Hawk Ow!

Diagnosis (based on holotype, BMNH 1955. 6. N30. 4747, and

photographs of at least nine living birds; see Plate and Figure 6f). —

Size medium-large, with long tarsi (based on type specimen), but

relatively small-billed for its size class. Prominently marked round-

headed owl, mostly barred above, weakly marked or uniform below,

with white or near-white vent. Plumage compact and sleek, as

with most other taxa in complex; ear-coverts have only short

filamentous extensions. Head shows prominent short white or

whitish supercilia; plain dark ear-coverts contrast with prominent

well-marked white throat-patch bordered below and on sides by

conspicuous dark markings (apparent in life, especially when

singing); crown strongly marked with rows of pale, almost

contiguous speckles between dark bars. Above, including wing-

coverts and tertials, strongly and narrowly barred dark brown

and bull, the barring varying from strong to obscure on lower

mantle. Scapular spots are mostly white, with narrow dark

brown edgings and warm buff between the white and brown areas.

Below, weakly and irregularly speckled or marked with short

broken bars on pinkish-buff to fulvous background, usually heaviest

on breast; sometimes with broad white streaking on lower

underparts and/or flanks, or virtually unmarked pinkish-buff to

rich fulvous below, with or without indistinct white streaking on

lower underparts. Undertail-coverts white (rarely pale buff), usually

with very narrow dark fringes to longest feathers. Central rectrices

have relatively broad, distinct dark bands. Bare parts: irides pale

to bright lemon-yellow, the inner and outer rings not differing;

narrow eye-ring indistinct and dull brown, not standing out from

adjacent feathers; bill and cere dull pale olive, tip and culmenridge

yellower; legs and feet bright yellow, tarsi about half-feathered,

otherwise thinly bristle-covered; claws mostly black, only the

proximal portions paler. Vocalisations are mid-pitched, irregular

strophes delivered rapidly and irregularly with multiple note types.

Holotype. — BMNH 1955.6.N.20.4747, female, Cebu, 17

March 1888, collected F. S. Bourns and D. C. Worcester. Ex.

Norwich Castle Museum. Culmen from cere 13.1 mm, wing 195

mm, tarsus 38.4 mm, tail 98.5 mm.

Description of holotype. — Crown very dark brown, heavily

speckled with buff in rows that resemble bars; weak short pale

supercilium; dark brown auriculars. Rear crown and nape heavily

speckled with rows of pale buff spots, becoming more barred on

upper mantle, then weaker and more mottled on central mantle,

which is nearly uniformly dark brown; inner scapulars are more

distinctly barred dark brown and buff, and outer scapulars are largely

white with very dark brown edgings, golden-buff between the

edgings and white inner portions; wing-coverts lightly barred, more

prominently so on greater coverts and tertials; uppertail broadly

barred very dark brown and narrowly barred dull buff. Throat

white, but mostly hidden due to preparation style. Breast pale

ochraceous, moderately barred and mottled dark brown, the bars

breaking up into small dull brown speckles on lower breast, flanks,

and even smaller, fewer speckles on central lower underparts. Tarsal

feathering and undertail-coverts buffy white, with a few irregular

dull brown speckles.

Forktail 28 (2012) Vocal divergence and new species in the Philippine Hawk Owl Ninox philippensis complex

17

Remarks. — The holotype is the sole known specimen in

existence. Photographs show this species to be highly variable in

underparts colour and pattern, less so in other characters. In

plumage closest to the much smaller N. mindorensis, from which it

most obviously differs in the more widely spaced bars below (where

these exist). Vocally fairly distinctive, although recognisably closer

to N. spilonota and N. mindorensis than to other taxa in song

characteristics and quality. Thought possibly extinct (Brooks et al.

1995b) until its rediscovery in 1998 (Pagantalan & Jakosalem

2008).

Etymology. — We name this new species for Stephen J. Rumsey,

dedicated conservationist and ornithologist who has given great

support of BirdLife International but who has also helped promote

research and conservation on the island of Cebu.

N. p. reyi

This was the third form of resident Ninox to be described from the

Philippines; but Tweeddale’s (‘1878’ = 1879 fide Dickinson et al.

1991) description of spilocephala apparently was too recent for

Oustalet (1880) to be aware of it. Thus Oustalet (1880) only had

philippensis for comparison, finding that reyi is immediately

distinguished by its larger size, longer wings (reaching beyond the

tail-tip), and russet (‘ roux ’) plumage with transverse brown bars

on head and shoulders. Mayr (1945), having no specimens to hand

and evidently misreading or ignoring Oustalet as well as Sharpe’s

(1897) Latin account of Ninox everetti (a synonym of reyi),

misattributed Sulu birds to the spilocephala group, and duPont &

Rabor (1973) mysteriously failed to remarkon this, despite having

eight reyi to hand by 1972 (material in DMNH).

The form reyi is distinguished from all other taxa by its very

pronounced, even barring on head and breast (Tobias score 2),

combination of tiny white supercilia with large bright white throat

(score 2), and its extraordinarily distinct song (4) as well as other

unscored song pattern characters.

Diagnosis (from numerous specimens and photos of three live

individuals; see Plate and Figure 6g). — Size large, with large, broad

bill. Head shows tiny white patch in supercilium, concentrically

dark-barred ear-coverts, and relatively well-defined facial disk.

Above, usually distinctly, regularly and rather narrowly barred dark

brown on whitish or buffy background; barring becomes muted

on upper mantle, and mantle vaguely barred, mottled and

vermiculated. Outer scapulars largely or at least partly white, with

intricate dark patterning intruding to vane on some; wing-coverts

prominently banded, the banding on tertials more muted.

Uppertail with prominent dark bands of medium width. Below,

throat -patch bright white and very well defined, visible even in

perched birds looking down. Breast distinctly and evenly dark-

barred, but lower underparts highly variable, often being broadly

white-streaked or even irregularly dark-mottled on white

background; equally the lower underparts can be fulvous, either

fairly plain or regularly dark-barred. Some individuals appear almost

black-and-white barred overall, lacking warm tones; others

(especially young birds) resemble spilocephala in underparts pattern,

but the dark-streaking pattern on the lower underparts is not as

distinct or regular as in spilocephala. Bare parts: irides bright yellow,

often paler on outer ring; bill dull greenish-yellow; feet and toes

mustard-yellow, claws black except for paler bases. Vocalisations

are very short rhythmic strophes of rapid toneless clucks, accented

at the end.

Lectotype designation

The description of Ninox spilonotus (spelling thus) Bourns &

Worcester 1894 was explicitly based on specimens from Cebu,

Sibuyan, Tablas and Mindoro. The description does not indicate

how many specimens were available, but the only specimens of the

last three of these taxa at the NMNH (formerly USNM) are now

considered the syntypes. We consider that Bourns & Worcester’s

(1894) spilonotus was based on four diagnosable taxa, and hence

designation of a lectotype is necessary. The first specimen listed,

from Cebu, is nowBMNH 1 955. 6. N. 20.4747, and is not currently

considered a syntype (it does not bear a type label, and is not listed

in Warren & Harrison 1973). However, Recommendation 74D in

the current code (ICZN 2000; http://www.nhm.ac.uk/hosted-

sites/iczn/code/index.jsp?nfv=true&article=74) states that the

lectotype should be ‘...preferably of the institution containing the

largest number of syntypes of the nominal species-group taxon...’.

On this basis we exclude the BMNH Cebu specimen from further

consideration as the lectotype, because NMNH has three syntypes.

Subsequent to the description of spilonota, Ninox mindorensis

Ogilvie Grant 1896 was described as a new species, based on a

BMNH specimen (97.6.14.51) collected by J. Whitehead. This

excludes the NMNH Mindoro specimen (USNM 314876) from

further consideration as the lectotype of spilonotus. Therefore, we

are left to choose between the two NMNH specimens, USNM

314877 from Sibuyan and USNM 314875 from Tablas. Neither

specimen was figured at the time, and both agree approximately

equally well with the description. We therefore arbitrarily fix

USNM 314877 from Sibuyan, which precedes Tablas in the

‘Habitat’ list on the original description, as the lectotype of Ninox

spilonotus. This furthermore implies that BMNH

1955.6.N. 20.4747, USNM 314876, and USNM 314875 are

paralectotypes of the name Ninox spilonotus. Note, however, that

the first and last of these specimens are herein designated as types

of a new species and a new subspecies, respectively.

DISCUSSION

Earlier treatments of taxa in the complex

Treatments of the Philippine Ninox philippensis complex have

inevitably varied over time, and separate into two periods of activity,

1940-1945 and 1990-2000. First, Peters (1940) grouped the taxa

into three species, ( 1 ) monotypic N. philippensis from Luzon,

Marinduque, Masbate, Ticao, Guimaras, Negros, Leyte and

Siquijor; (2) monotypic N. spilonota from Mindoro, Tablas, Sibuyan

and Cebu; and (3) polytypic N. spilocephala, consisting of N. s.

mindorensis from Mindoro, nominotypical spilocephala from

Mindanao and Basilan, N. s. reyi from Sulu and Bongao (Sulu

Islands), and N. s. everetti from Siasi (Sulu Islands). Shortly

afterwards, Delacour & Mayr (1945) united these six forms (plus

two ,proxima and centralis, added by Mayr) as races of one species,

Ninox philippensis, falling into three groups based on simple shared

plumage characters (some of which were mistaken): (1) philippensis

group (‘upperparts plain, underparts boldly striped’) — iV. p.

philippensis (Luzon, possibly also Marinduque, Samar and Leyte),

N. p. proxima (Ticao and Masbate) and N. p. centralis (Siquijor;

apparently also Panay, Guimaras, Negros); (2) spilocephala group

(‘head and neck spotted or barred, underparts striped or

variegated’) — N. p. spilocephala (Mindanao, Basilan), N. p. reyi

(Sulu, Tawi Tawi, Bongao) and N. p. everetti (Siasi); and (3)

mindorensis group (‘head and neck spotted or barred; underparts

entirely vermiculated or barred’) — N. p. mindorensis (Mindoro) and

N. p. spilonota (Tablas, Sibuyan, Cebu).

Sibley & Monroe (1990) accepted these groupings but

rationalised them geographically north-south, making the

mindorensis group (which, owing to chronological precedent, they

called the spilonota group) second and spilocephala third. However,

Dickinson et al. (1991) recast the arrangement (adding ticaoensis,

described in 1972) rather more radically to give: N. p. philippensis

(Luzon, Polillo, Catanduanes, Marinduque, Samar, Buad, Biliran

and Leyte); N. p. spilocephala (Dinagat, Siargao, Mindanao and

Basilan); N. p. reyi (Sulu, Siasi, Tawi Tawi and the adjacent Sanga

18

P.C. RASMUSSEN etal.

Forktail 28 (2012)

Sanga and Bongao, and Sibutu); N. p. centralis (Semirara, Carabao,

Boracay, Panay, Guimaras, Negros, Siquijor and Bohol); N. p.

spilonota (Tablas, Sibuyan, Cebu and Camiguin Sur); N. p.proxima

(Masbate); N. p. ticaoensis (Ticao); and N. p. mindorensis

(Mindoro). Kennedy et al. (2000) retained this treatment in spite

of the queries by Collar & Rasmussen (1998) and the split of

mindorensis by Konig et al. (1999).

On the basis of our data (and reiterating our doubt about the

validity of ticaoensis ) we propose the following arrangement (ranges

derived from Dickinson et al. 1991, Dickinson 2003):

Luzon Hawk Owl Ninox philippensis

N. p. philippensis Biliran, Bohol, Boracay, Buad, Carabao,

Catanduanes, Guimaras, Leyte, Lubang, Luzon,

Marinduque, Masbate, Negros, Panay, Polillo, Samar,

Semirara

N p. ticaoensis Ticao

N. p. centralis Siquijor

Mindanao Hawk Owl Ninox spilocephala

Mindanao, Dinagat, Siargao, Basilan

Mindoro Hawk Owl Ninox mindorensis

Mindoro

Romblon Hawk Owl Ninox spilonota

N. s. spilonota Sibuyan

N s. fisher i Tablas

Cebu Hawk Owl Ninox rumseyi

Cebu

Camiguin Hawk Owl Ninox leventisi

Camiguin Sur

Sulu Hawk Owl Ninox reyi

Sulu, Siasi, Tawi Tawi and the adjacent Sanga Sanga and

Bongao, Sibutu

Figure 7 maps the newly defined species limits of the N. philippensis

( sensu lato ) complex. We use ‘Luzon’ rather than ‘Philippine’ as the

common name for N. philippensis ( sensu stricto ) to avoid confusion

with N. philippensis ( sensu lato). For N. spilonota ( sensu stricto) we

use ‘Romblon’ because, although no member of this taxon is

definitely known to occur on Romblon Island proper (there are

unconfirmed reports), both Sibuyan and Tablas are part of

Romblon province, so this seems the most appropriate geographical

name for this species.

Relationships

How can we explain the curious circumstance in which rather

similar-plumaged unstreaked Ninox populations are scattered on

six well-separated islands or island groups within the Philippine

archipelago while streak-breasted (or part-streak-breasted)

populations occupy the largest islands ? One possibility, consistent

with the speculation in Collar & Rasmussen (1998) that all

unstreaked forms are monophyletic, is that they represent the

remnants of an earlier stock of Ninox which occupied the

archipelago before being displaced from almost all islands which

had a Pleistocene land-bridge by a later invasion of streak-breasted

birds. Militating against this notion is the fact that the streak¬

breasted birds are generally smaller than the unstreaked forms.

Even so, Ninox spilonota of Tablas and Sibuyan, N. rumseyi of

Cebu and N. mindorensis of Mindoro seem to form a monophyletic

group. Their distribution mirrors that of some Philippine Hypsipetes

bulbuls, of which the forms on Mindoro, Tablas, Romblon and

Cebu are more closely related to each other than to the bulbuls of

Negros, Panay, Greater Luzon and Greater Mindanao (Oliveros &

Moyle 2010). Vocally spilonota, rumseyi and mindorensis all share

certain characteristics including descending thin, squealed whistles

and croaky hisses, and all are at least relatively high-pitched. Ninox

mindorensis often particularly resembles the much larger rumseyi

Figure 7. Map of species of the Ninox philippensis sensu lato species

complex as recognised herein.

of Cebu in plumage pattern, although it is typically more narrowly

and clearly barred below, and has the throat-patch duller and less

conspicuous. Ninox rumseyi, which is geographically surrounded

by Ninox philippensis, can only be diagnosed morphologically from

mindorensis and spilonota on a combination of characters, although

its song is moderately autapomorphic. Both forms of Ninox

spilonota lack both strong head pattern and markedly paler throat-

patch, but one specimen ofiV. r. spilonota from Sibuyan (FMNH

358295) shows a white-streaked underparts pattern similar to many

N. reyi specimens, and thus recalls a weakly marked N. spilocephala.

The relationships of Mindanao’s spilocephala remain unclear.

Its different voice and certain other features (e.g. well-developed

auriculars) are inconsistent with its being the link (Mayr 1945)

between the plain-headed, streak-breasted philippensis and the

unstreaked taxa. On the other hand its Sulu replacement reyi,

although completely different from any other Philippine taxon in

voice, shows plumage characters in some individuals (dark teardrop¬

shaped streaking on lower underparts) reminiscent of spilocephala,

hinting at a shared ancestry. Given reyi' s geographical isolation from

all taxa except spilocephala, this inference is more parsimonious than

the notion that reyi is more closely related to a more widespread

unstreaked form. (The possibility that reyi is more closely related

to Ninox ios of Sulawesi, based on the similarity noted above in the

quality if not the pattern of their songs, is not supported by their

very dissimilar morphologies; see Rasmussen 1999.)

The derivation of Ninox leventisi of Camiguin Sur, despite the

fact that this island lies just off Mindanao, is no less problematic.

Although at least superficially similar in plumage to other

unstreaked taxa, leventisi is the most highly autapomorphic of all

taxa (although vocally N. reyi is even more so). In fact, leventisi

Forktail 28 (201 2) Vocal divergence and new species in the Philippine Hawk Owl Ninox philippensis complex

19

and spilocephala are strikingly dissimilar to each other, in size,

plumage texture and fullness, development of filamentous auricular

extensions, pattern of lower underparts, and soft-part colours. The

only character that links leventisi with spilocephala to the exclusion

of other taxa is the low pitch of their songs, albeit these are very

different in other ways. Indeed, the long solo song of leventisi

(SOM) has closer resemblances to that of philippensis than to any

other taxon, and it may well be more closely related to philippensis

than to the geographically closer spilocephala or the

morphologically closer spilonota.

Resolution will have to await molecular analysis. For the

moment, we note a high level of concordance between the species

limits proposed here and zones of turnover proposed by Peterson

(2006). A separate issue is that of generic limits of Ninox, which at

least on morphology is probably a non-monophyletic group

(Rasmussen 1999). A recent molecular phylogeny (Wink et al.

2009) shows a monop hyletic Ninox but taxon sampling therein was

not dense. Although we think that all the species formerly united

under Ninox philippensis are indeed fairly closely related, this

remains to be established, most likely by molecular analyses.

The conservation status of these various forms needs full

evaluation elsewhere. However, it should be noted at once that five

species — mindorensis , spilonota, rumseyi, leventisi and reyi — are

likely to be at risk. On Mindoro, the Sablayan Penal Colony holds

the largest remaining area of lowland forest on Mindoro, and is

critically important for populations of lowland bird species endemic

to Mindoro (Brooks etal. 1995a, Mallari cZTz/. 2001). The Mindoro

Hawk Owl is also known from forest in Mts Iglit-Baco National

Park (Gonzalez & Dans 1998), Naujan Lake National Park (Ticsay

& Ledesma 1998), and Mt Halcon between 1,150-1,250 m

(ROH). Satellite images taken in the late 1980s suggested Tablas

then held two forest parcels together covering under 0.5 km2

(Goodman & Ingle 1993) and recent visits confirm that the amount

of habitat there is tiny, although the species occurs in tall secondary

as well as primary forest on the island (Allen 2006, DNSA, ROH).

Meanwhile, although forest covered over half of Sibuyan’s 233 km2

land surface at the start of the 1 990s, logging was proceeding apace

and, in the absence of intervention, all lowland areas were predicted

to be cleared within a few years (Goodman & Ingle 1993); illegal

logging inside Mt Guiting-guiting Natural Park continues (M.

Wallbank verbally 2009 to DNSA). On Cebu, where it was

rediscovered in 1998 after a gap of 110 years, the total population

of Cebu Hawk Owls may be just 192 pairs (and possibly many

fewer) scattered among 1 1 forest patches ( Jakosalem et al. in press).

On Camiguin Sur forest covers just the centre of the small island,

and numbers of the species cannot be high (Heaney & Tabaranza

2006). Finally, of the six islands in the Sulu archipelago from which

the owl has been recorded, the current status of forest on Siasi

appears on satellite photos to show that almost the entire island’s

forest has been replaced by orchards; Sulu (Jolo) has apparently

also been largely converted except around the volcanic cones; Sanga-

Sanga’s last forest patch was cleared in 1992-1993; and Bongao

retains only a small patch of forest on its sacred mountain (DNSA,

ROH). Sibutu may have significant areas of secondary forest,

although that seen by DNSA in 1995 was of low stature. The main

island of Tawi Tawi still has much secondary and some primary

forest (Dutson etal. 1996, DNSA, ROH). However, on Tawi Tawi

N. reyi is also found in forest edge, mature mangroves and large

trees in the vicinity of villages.

ACKNOWLEDGEMENTS

Work on early drafts of this paper in the mid-1990s benefited greatly from

information supplied by T. M. Brooks, G. C. L. Dutson, S. J. Harrap, B. F.

King, P. A.J. Morris, R. Ranft, R. J. Timmins and M. P. Walters. More recently

we have had the help of f. A. Eaton, C. Espanola, the late T. Fisher, P. C.

Gonzales, N. Icarangal, Jr., H. C. Miranda, Jr., M. Lagerqvist, F. E. Rheindt, I.

Sarenas, P. Simpson, M. Thibault, C. Tipp and P. Verbelen. DNSA's work on

Tawi Tawi was greatly facilitated by Governor Hadji Sadikul Sahali and his

family, and by Eddie Ali Dean. Some of DNSA’s studies on Tablas and Tawi

Tawi were sponsored by ZGAP (Zoologische Gesellschaft fur Arten- und

Populationsschutz). For permission to examine specimens in their care and other

assistance (institutions given in full in main text) we thank staff including P. R.

Sweet, T. Trombone and M. Shanley (AMNH), L. Joseph (ANSP), R. P. Prys-

Jones and M. P. Adams (BMNH), the late B. C. Livezey (CM), G. K. Hess and

J. Woods (DMNH), D. E. Willard (FMNH), G. Lenglet (IRSNB), the late R.

A. Paynter, A. Pirie and J. Trimble (MCZ), R. Zink (MMNH), C. Voisin

(MNHN), R. W. R. J. Dekker (NCB), L. Alvarez (PNM), B. Millen (ROM),

G. Mayr (SFN), the late S. Eck (SMTD), J. Hinshaw (UMMZ), C. Milensky

(USNM), N. Rice (YPM) and S. Frahnert (ZMB). For permission to use their

photographs in Figure 6 we thank C. Artuso, J. Eaton, R. J. Quisumbing and

M. Thibault. We also thank the National Geographic Society Committee on

Research and Exploration for financial support to PCR (NGS CRE 8919-11).

Two referees, one of them Lawrence Heaney, the other anonymous, made very

helpful comments to improve the manuscript. The colour plate is an original

work by John Gale commissioned for this paper.

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20

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581-591.

P. C. RASMUSSEN, Dept, of Zoology and Michigan State

University Museum, Michigan State University, East Lansing,

Ml 48824, USA; and Bird Croup, Department of Zoology,

Natural History Museum, Akeman St, Tring, Herts HP23 6AP,

UK. Email rasmus39@gmail.com

D. N. S. ALLEN, 97 Sussex Way, London N7 6RU, UK

N. J. COLLAR, BirdLife International, Wellbrook Court, Girton

Road, Cambridge CB3 0NA, UK; and Bird Group, Department

of Zoology, Natural History Museum, Akeman St, Tring, Herts

HP23 6AP, UK

B. DeMEULEMEESTER, Koningin Astridlaan 154, 9000 Gent,

Belgium. Email bram_dm@yahoo.co.uk

R. O. HUTCHINSON, 26 Sutton Avedue, Chellaston, Derby DE73

6RJ, UK. Email robhutchinson@birdtourasia.com

P. G. C. JAKOSALEM, Philippines Biodiversity Conservation

Foundation, Inc. c/o Negros Forest Ecological Foundation, Inc.

South Capitol Road, Bacolod City, 6100 Philippines

R. S. KENNEDY, Frederick and Amey Geier Collections and

Research Center, Museum of Natural History & Science,

Cincinnati Museum Center, 1301 Western Ave, Cincinnati,

OH 45203; and Ornithology Department, Museum of

Comparative Zoology, Harvard University, 26 Oxford St,

Cambridge, MA 02138; current address 18 Riverview Rd,

Durham NH 03824, USA

F. R. LAMBERT, El 802 A Perdana Condo, Jalan PJU 8/1,

Damansara Perdana, Petaling Jaya, 47820 Selangor, Malaysia

L. M. PAGUNTALAN, Philippines Biodiversity Conservation

Foundation, Inc. c/o Negros Forest Ecological Foundation, Inc.

South Capitol Road, Bacolod City, Philippines

Supplementary online material

To be found under Forktail 28 page on www.orientalbirdclub.org.

SOM 1. Additional songs of taxa in the Philippine Hawk Owl Ninox

pbilippensis ( sensu Into ) complex. SOM la.12453: short isolated duetted

segment of song of Philippine Hawk Owl N. p. pbilippensis, AV# 12453, ROH,

Luzon; SOM lb. 1355 1 : long song by a single bird, joined near the end with

another in a presumed duet; Philippine Hawk Owl N. p. pbilippensis ,

AV# 1 355 1 . ROH, Luzon. SOM 1 c.8972: series of short duetted strophes by

Philippine Hawk Owl N. p. centralis , F. Verbelen, Bohol; SOM ld-e. 10700;

long series by Philippine Hawk Owl N. p. centralis , FRL, Negros; SOM

If. 1 12a: long series by Philippine Hawk Owl/V. p. centralis , DNSA, Siquijor;

SOM lg.99: three short duetted strophes by Philippine Hawk Owl N. p.

centralis , DNSA, Siquijor, with many hoarse notes. SOM lh. 13557: duetted

series of short strophes with stray notes between main strophes; Camiguin

Hawk Owl Ninox sp. 1. SOM li: single Camiguin Hawk Owl Ninox sp. 1

singing, giving several more notes/strophe than in Figure 2c. SOM lj:

continuation of long song of Mindanao Hawk Owl N. spilocepbala , starting

where Figure 2d ends. SOM lk: duet by Sulu Hawk Owl N. reyi , series of

short variable strophes. SOM 11: continuation of duet of Mindoro Hawk Owl

N. mindorensis, starting where Figure 2gends; SOM lm-n: whistled duet of

Mindoro Hawk Owl N. mindorensis , with many highly frequency-modulated

notes. SOM lo: Variant song and note-types of Romblon Hawk Owl Ninox

spilonota newssp. on Tablas; SOM lp.l 1510: duetted song type of Romblon

Hawk Owl Ninox spilonota on Sibuyan. SOM lq-r: various alternate song

types of Cebu Hawk Owl Ninox new sp. 2.

FORKTAIL 28 (2012): 21-28

Bird species diversity in five protected areas

of Bangladesh

M. MONIRUL H. KHAN & NASIM AZIZ

Birds of five protected areas (Lawachara, Satchari, Rema-Kalenga, Chunati and Teknaf ) in north-east and south-east Bangladesh were studied

during 2005-2008 by strip transect sampling and opportunistic surveys. Of 239 species of birds recorded, 189 were residents, 39 winter

visitors, 6 summer visitors and 5 vagrants; 40 (17%) were 'Very Common', 66 (28%) 'Common', 48 (20%) 'Fairly Common' and 85 (35%)

'Uncommon'. Population densities of eight species, selected as indicators of forest condition, were estimated; two understorey species, Red

Junglefowl Gallus gallus and Puff-throated Babbler Pellorneum ruficeps, increased in the study period, perhaps owing to understorey

regeneration increasing the carrying capacity and nesting sites, plus (for the junglefowl) reduced hunting pressure. However, illegal logging

of timber trees continues in some areas, probably causing the decline of Oriental Pied Hornbill Anthracoceros albirostris. Densities of the

other five indicators (Red-headed Trogon Harpactes erythrocephalus, Greater Racket-tailed Drongo Dicrurus paradiseus, White-rumped Shama

Copsychus malabaricus, Hill Myna Gracula religiosa and White-crested Laughingthrush Garrulax leucolophus) remained more or less unchanged.

INTRODUCTION

Bangladesh has a network of protected areas under the jurisdiction

of the Forest Department, but very little information is available

on the birds of these areas. Moreover, there had been no attempt to

use birds as indicators of ecological change in the protected areas,

although birds are one of the best indicators of such change

(Morrison 1986, Temple & Wiens 1989, Browder et al. 2002).

Figure 1 . Forested areas of Bangladesh showing the locations of five

NSP (Nishorgo Support Project) sites where the survey was conducted.

Panchagarh

Thakurgaon

® Nilphamari

Dinajpur

0

Ranggi

® Kurigran

r ©

Gaibandha

© \

Joypurhat

©

Chapai Nawabganj

©

Nator

Naogaon Bogra

0

Sherpur

© . , Netrakona

Jamalpur ©

Sunamgani

© T

Sylket

W

4“

Forest Types

Mixed Evergreen Forest

■ Deciduous Forest

1 Mangrove Forest

Others

© District Headquarter

.. . River or Sea

District Boundary

100

100 Kilometer

Scale 1:3,80,000

91

Different species of bird occur in different vertical strata of an area

and are adapted to specific types of plant and animal food.

Therefore, avian population density and species diversity reflect

the changes in their habitat conditions, and indicate the health of

different vertical strata of the forest. Determining the extent of

change in ecological systems is critical to the long-term conservation

of biotic diversity (Canterbury et al. 2000).

In its 147,570 km2 Bangladesh harbours nearly 700 species of

bird, which is 50% of the total of the Indian subcontinent and 7%

of the world’s total (Harvey 1990, Khan 2008). However, the

natural forests and other wilderness areas of the country are under

great pressure from legal and illegal overexploitation, together with

improper management. Forest cover has been reduced by more than

50% since the 1970s (IUCN-Bangladesh 2000). Bangladesh has

less than 0.02 ha of forest land per person — one of the lowest forest-

to-population ratios in the world (IUCN-Bangladesh 2000). The

urgent need to conserve the remaining tracts and their inhabitants,

O O

Table 1. Strip transect locations in five NSP sites where the bird survey

was conducted.

22

M. MONIRUL H. KHAN & NASIM AZIZ

Forktail 28 (2012)

while simultaneously developing non-consumptive uses, gave rise

to the Forest Department’s five-year Nishorgo Support Project

(NSP), which was implemented by the International Resources

Group (IRG) of Washington DC, together with its local partners.

The work reported here was supported by NSP, which used

the results as a tool to understand the impact of newly introduced

co-management on live protected areas. In our survey, the temporal

changes of population densities of eight indicator species of bird

were studied so that the ecological health of five protected areas

could better be monitored. All species of bird were recorded,

together with their relative abundance. The study was conducted

on a participatory basis, involving birdwatchers, local communities

and Forest Department officials in order that everybody would

become aware of what is happening to the birds in these sites.

There are 28 protected areas in Bangladesh with a total area of

8,718.87 km:, covering only 5.9% of the total area of Bangladesh.

All the five protected areas (Lawachara, Satchari, Rema-Kalenga,

Chunati and Teknaf; Fig. 1, Table 1) where the bird survey was

conducted are mainly covered by mixed evergreen forests, but the

areas also have some bushy and bamboo-covered areas.

METHODS

Survey methods were selected on the basis ol simplicity and

effectiveness so that local stakeholders, including people with little

or no education, could participate. Strip transect sampling

(Buckland et al. 2001) and opportunistic survey methods were

selected. The survey was conducted in the breeding season

(February- August), so that there would be more bird activity. The

baseline survey was conducted in 2005, and was repeated in 2006,

2007 and 2008, in the same season and following the same transects,

so that the results could be compared across different years.

Strip transect samplingwas found most suitable to estimate the

population densities of eight indicator species of bird (see below

for choice rationale). In this method the observers slowly walked

(c.1.5 km/hr) in relatively straight lines through the study areas

and counted the indicator birds from both sides. The observation-

range on each side (half-width of the strip) was 20 m, which was

found suitable in terms of visibility. Thus the strip width was

40 m. The initial location of the bird was recorded. The

observation-range was measured by an infrared rangefinder

(Bushnell Yardage Pro). Any bird sighted beyond the pre-decided

observation-range or coming from behind the observer was not

counted. The survey was conducted in early mornings and late

afternoons when the birds were most active. This method assumes

that all birds in the strip are recorded. Transects were located mainly

in the rich parts of the protected areas where the bird habitats were

least disturbed. Depending on reserve size, the total lengths of

transects in each site varied from 3.00 to 6.96 km (Table 1), but

transects were periodically repeated 12-15 times per year.

The eight indicator species of bird, suggested at a meeting of

experts organised in IRG, Dhaka, were: Red Junglefowl Gallus

gallus , Oriental Pied Hornbill Anthracoceros albirostris, Red-headed

Trogon Harpactes erythrocephalus , Greater Racket-tailed Drongo

Dicrurus paradiseus , White-rumped Shama Copsy clous malabaricus ,

Hill Myna Gracula religiosa , White-crested Laughingthrush

Garrulax leucolophus and Puff-throated Babbler Pellorneum

ruficeps. They were selected because: (1) they are primarily forest

birds; (2) they live in different vertical strata of the forest

(junglefowl, laughingthrush and babbler in lower stratum; trogon,

drongo and shama in middle stratum; and hornbill and myna in

upper stratum); (3) they are noisy (and thus less likely to be missed

during counts); and (4) they are breeding residents. It was assumed

that the improvement or degradation of the forest condition would

have a direct impact on the feeding and breeding of these species,

which in turn would show changes in population densities

(Morrison 1986, Temple & Wiens 1989, Canterbury et al. 2000,

Browder et al. 2002, Lammertink et al. 2009).

Any important or interesting observation or information

relating to diet, foraging guild and threats was recorded

opportunistically at any time in the field. The birds were identified

from Ali & Ripley (1987), Grimmett et al. (1998) and Rasmussen

& Anderton (2005). Relative abundance was assessed by classifying

sighting frequencies into four: ‘Very Common’ — recorded on 76-

100% observation-days, ‘Common’ — recorded on 51-75%

observation-days, ‘Fairly Common’ — recorded on 26-50%

observation-days, and ‘Uncommon’ — recorded on 25% or fewer

observation-days.

RESULTS

During the avian breeding season (February- August) of four

consecutive years (2005-2008), 239 species of bird were recorded

in the five protected areas, of which 189 were residents, 39 winter

visitors, 6 summer visitors and 5 vagrants (see Appendix). The

relative abundance shows that 40 (17%) species were ‘Very

Common’, 66 (28%) ‘Common’, 48 (20%) ‘Fairly Common’ and

85 (35%) ‘Uncommon’. Based on principal diet 131 (55%) species

were insectivorous, 32 (13%) carnivorous, 29 (12%) omnivorous,

23 (10%) granivorous and herbivorous, 17 (7%) frugivorous, and

7 (3%) nectarivorous. Based on principal foraging guild 100 (42%)

species were middle-canopy foragers, 78 (33%) ground foragers,

36 (15%) upper-canopy foragers, and 25 (10%) undergrowth and

bush foragers.

The site lists show that the total number of species and the total

number of primarily forest species for Lawachara were, respectively,

167 and 90 (54%), Satchari 1 53 and 84 (55%), Rema-Kalenga 206

and 91 (44%), Chunati 162 and 53 (33%), and Teknaf 188 and 73

(39%). The percentages indicate how the ratio of total species to

total forest species varied across the five sites.

Annual mean density estimates from 2005 to 2008 revealed that

two understorey indicator species (Red Junglefowl and Puff-

throated Babbler) increased at every site and at all sites combined

(Fig. 2). Oriental Pied Hornbill declined and the other five

indicator species remained more or less unchanged (Fig. 2).

Three species of global conservation concern were recorded:

White-rumped Vulture Gyps bengalensis (Critically Endangered;

nesting colony found in Rema-Kalenga), Lesser Adjutant Leptoptilos

javanicus (Vulnerable; one adult in Rema-Kalenga, 10 March 2008)

and White-cheeked Partridge Arborophila atrogularis (Near

Threatened; parent and hatchling in Lawachara, 15 March 2008).

Eleven nationally threatened species (according to IUCN-

Bangladesh 2000; since the list is out of date the White-rumped

Vulture is not included) were recorded: Kalij Pheasant Lophura

leucomelanos, Grey Peacock Pheasant Polyplectron bicalcaratum ,

Oriental Pied Hornbill, Red-headed Trogon, Dollarbird

Eurystomus orientalis , Spot-bellied Eagle Owl Bubo nipalensis.

Figure 2. Comparison of the population density of eight indicator bird

species across five NSP sites during 2005-2008.

■ In 2005

□ In 2006

□ In 2007

□ In 2008

Forktail 28 (2012)

Bird species diversity in five protected areas of Bangladesh

23

Brown Fish Owl Ketupa zeylonensis. Tawny Fish Owl Ketupa

flavipes, Malayan Night Heron Gorsachius melanolophus , Lesser

Adjutant and Streaked Spiderhunter Aracbnothera magna.

DISCUSSION

The total bird species (239) recorded in five NSP sites represents

over 30% of the birds recorded in Bangladesh (Khan 2008, Siddiqui

et al. 2008), and almost 18% recorded in the Indian Subcontinent

(Grewal et al. 2002). The species were recorded mainly during

transects while counting indicator birds, so some may have been

missed either because they occurred away from the transects or

because of observer inattention. The cryptic nature of some birds

might also have contributed to the crudeness of the relative

abundance.

An increase in density of two indicator species (Red Junglefowl

and Puff-throated Babbler) would suggest that the forest

understorey is regenerating and increasing the carrying capacity and

nesting sites for these two species. Community patrolling and

awareness, and other programmes conducted by NSP has played a

key role in reducing understorey firewood exploitation and hunting

pressure. However, White-crested Laughingthrush did not respond

like the junglefowl and babbler, possibly having different limiting

factors; it is a rare species in Bangladesh and was found only in

Chunati, which is more open compared to four other NSP sites.

Illegal removal of timber trees and forest fruits still persists, and

probably caused the decline of the Oriental Pied Hornbill over the

four years. This bird lives in the upper canopy and suffers severely

if large trees are removed from the forest.

Habitat loss remains the main threat to birds in all five NSP

sites. Illegal felling of trees and bamboo, and conversion of natural

forests to monoculture plantations and agricultural, fields, were

witnessed during the survey. Hunting and trapping of birds,

together with the taking of nestlings as cagebirds (particularly

mynas, parakeets and hornbills), is another threat. Large-scale illegal

harvest of forest fruits, particularly Artocarpifs chaplasha and Bixa

sp., is a growing threat for frugivorous birds. Moreover, the number

of visitors in Lawachara and Satchari is becoming excessive. These

threats should be reduced in order to maintain a healthy status of

birds in the study’s five protected areas.

ACKNOWLEDGEMENTS

The authors are grateful to Nishorgo Support Project, Bangladesh Forest

Department, together with United States Agency for International

Development, International Resources Group (IRG) and its partners in

Bangladesh (CODEC, RDRS, NACOM and IUCN) for providing necessary

support to this work. Sincere thanks to Philip DeCosse and Paul Thompson,

who provided strong support and advice during the survey. Also thanks to all

volunteers (particularly Dhrubo Kundu, Shafiqur Rahman, Ahsanul Haque,

Mustafezur Rahman, Samiul Mohsanin, Shehab Rayhan and Jewel Ahmed)

and local participants.

REFERENCES

Ali, S. & Ripley, S. D. (1987) Compact handbook of the birds of India and

Pakistan together with those of Bangladesh, Nepal, Bhutan and Sri Lanka.

1 0 vols. Delhi, India: Oxford University Press.

Browder, S. F. Johnson, D. H. & Ball, I. J. (2002) Assemblages of

breeding birds as indicators of grassland condition. Ecol. Indicators 2:

257-270.

Buckland, S.T., Anderson, D. R., Burnham, K. P., Laake, J. L., Borchers, D. L. &

Thomas, L. (2001) Introduction to distance sampling: estimating

abundance of biological populations. Oxford, UK: Oxford University

Press.

Canterbury, G. E„ Martin, T. E„ Petit, D. R„ Petit, L. J. & Bradford, D. F. (2000)

Bird communities and habitat as ecological indicators of forest

condition in regional monitoring. Conserv. Biol. 14(2): 544-558.

Grewal, B., Harvey, B. & Pfister, 0. (2002) A photographic guide to the birds of

India including Nepal, Sri Lanka, The Maldives, Pakistan, Bangladesh and

Bhutan. London, UK: Christopher Flelm.

Grimmett, R„ Inskipp, C.& Inskipp, T. (1 998) Birds of the Indian subcontinent.

Delhi, India: Oxford University Press.

Harvey, W. G. (1990) Birds in Bangladesh. Dhaka, Bangladesh: University Press

Limited.

lUCN-Bangladesh (2000) Red book of threatened birds of Bangladesh. Dhaka,

Bangladesh: IUCN -The World Conservation Union.

Khan, M. M. H. (2008) Protected areas of Bangladesh: a guide to wildlife. Dhaka,

Bangladesh: Nishorgo Program, Bangladesh Forest Department.

Lammertink, M., Prawiradilaga, D. M., Setiorini, U., Naing, T. Z., Duckworth,

J. W„ & Menken, S. B. J. (2009) Global population decline of the Great

Slaty Woodpecker (Mulleripicus pulverulentus). Biol. Conserv. 142: 1 66—

179.

Morrison, M. L. (1986) Bird populations as indicators of environmental

changes. Current Ornith. 3:429-451.

Rasmussen, P. C. & Anderton, J. C. (2005) Birds of South Asia: the Ripley guide.

Vols. 1 and 2. Washington DC, USA: Smithsonian Institute, and

Barcelona, Spain: Lynx Edicions.

Siddiqui, K. U., Islam, M. A., Kabir, S. M. H., Ahmad, M., Ahmed, A. T. A.,

Rahman, A. K. A., Haque, E. U., Ahmed, Z. U., Begum, Z. N.T., Hassan, M.

A., Khondker, M. & Rahman, M. M., eds. (2008) Encyclopedia of flora and

fauna of Bangladesh, 26, Birds. Dhaka, Bangladesh: Asiatic Society of

Bangladesh.

Temple, S. A. & Wiens, J. A. (1989) Bird populations and environmental

changes: can birds be bio-indicators? Amer. Birds 43: 260-270.

M. Monirul H. KHAN, Deportment of Zoology, Jahanglrnagor

University, Savor, Dhoko 1342, Bangladesh. Email:

mmhkhan@hotmail.com

Nasim AZIZ, International Resources Group - Bangladesh,

House # 68, Road # 1, Block # 1, Banani, Dhaka 1213,

Bangladesh. Present address: lUCN-Bangladesh, House # 1 1,

Road # 138, Gulshan 1, Dhaka 1212, Bangladesh. Email:

nasim.aziz@iucn.org

24

M. MONIRUL H. KHAN & NASIM AZIZ

Forktail 28 (2012)

Appendix

List of bird species recorded in five NSP sites in the breeding season (February-August) during 2005-2008

Distribution: Wl - wide (all NSP sites), L - Lawachara, S - Satchari, RK - Rema-Kalenga, C - Chunati, and T - Teknaf. * Primarily forest species.

Forktail 28 (2012)

Bird species diversity in five protected areas of Bangladesh

25

26

M. MONIRUL H. KHAN & NASIM AZIZ

Forktail 28 (2012)

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Bird species diversity in five protected areas of Bangladesh

27

28

M. MONIRUL H. KHAN & NASIM AZIZ

Forktail 28 (2012)

Principal

FORKTAIL 28 (2012): 29-37

Peat swamp forest birds of the Tuanan research station.

Central Kalimantan, Indonesia, with notes on

habitat specialists

MARY ROSE C. POSA & DAVID ALEXANDER MARQUES

The avifauna of tropical peat swamp forest has not been well documented, even though it is an extensive habitat in parts of South-East

Asia. We conducted surveys using various methods at the Tuanan research station and surrounding areas in Central Kalimantan, Indonesian

Borneo. These observations resulted in a list of 138 bird species and numerous noteworthy records. Although more depauperate than

lowland rainforest on mineral soils, peat swamp forest is an important habitat for many threatened and Near Threatened bird species,

especially habitat specialists such as Hook-billed Bulbul Setornis criniger and Grey-breasted Babbler Malacopteron albogulare. We also

recorded in selectively logged peat swamp several high-profile, globally threatened species such as Crestless Fireback Lophura

erythrophthalma, Storm's Stork, Ciconia stormi, Great Slaty Woodpecker Mulleripicus pulverulentus, Black Hornbill Anthracoceros malayanus

and Wrinkled Hornbill Aceroscorrugatus. In view of its importance to certain species, peat swamp forest should be afforded more protection,

especially in light of the recent rapid loss of this habitat to land conversion and forest fires.

INTRODUCTION

Borneo is the third largest tropical island in the world and particularly

rich in biodiversity, with 630 recorded bird species (Mann 2008).

Geopolitically, the island is divided into Brunei Darussalam, the

Malaysian states of Sabah and Sarawak, and the four Kalimantan

provinces of Indonesia. Most of the ornithological work on Borneo

has been conducted in the northern part of the island, and

Kalimantan remains poorly studied, with only a few avifaunal lists

published specifically for this region (e.g. Holmes & Burton 1987,

Holmes 1997). In Central Kalimantan, the second largest province

of Indonesian Borneo with a land area of 1 54,564 km2, bird surveys

have largely concentrated in a few well-known areas such as Tanjung

Puting National Park and Barito Ulu (Bohap & Galidikas 1987, Nash

& Nash 1988, Dutson et al. 1991, Wilkinson et al. 1991). The

extensive tropical swamp forests dominating the southern lowland

plains of Central Kalimantan have largely been ignored and

unexplored, even though these habitats constitute one of the seven

biogeographic zones of Borneo and have an important influence on

species distribution (MacKinnon et al. 1996).

Tropical peat swamp forest (PSF) is a unique wetland ecosystem

that develops in areas where waterlogging prevents the complete

decomposition of plant debris, which over time accumulates as peat

soils (Anderson 1983). PSF occurs throughout the tropics but

reaches its greatest extent and depth in South-East Asia, especially

in the lowlands of Sumatra and Kalimantan (Rieley et al. 1996).

PSFs are characterised by periodic flooding, nutrient limitation and

high acidity due to the leaching of organic compounds. PSF trees

are adapted to tolerate nutrient deficiency, unstable substrate and

fluctuating water levels, and in this respect exhibit structural

features such as stilt roots and pneumatophores. The overall

primary productivity and biodiversity levels in this nutrient-

deficient forest type are lower than in lowland forest on mineral

soils (Bruenig& Droste 1995).

The neglect of PSF by biologists might result either from its

relatively depauperate flora and fauna or from the difficult access

and working conditions brought about by the boggy soils and dense

understorey vegetation, which severely hamper movement and

visibility. However, recent research indicates that PSF may harbour

a considerable proportion of the South-East Asian fauna (Posa et

al. 2011). Thus, there is an urgent need for more information on

the flora and fauna of this unique ecosystem, as its destruction has

accelerated in recent years. Nearly half of the PSF in Peninsular

Malaysia, Borneo and Sumatra has been lost since 1990 (Miettinen

& Liew 2010). Many areas have already been converted into oil palm

and paper pulp plantations and much of what remains under forest

cover has been selectively logged (Miettinen & Liew 2010). Such

disturbance renders PSF extremely prone to forest fires, since peat

itself is combustible when dry (Page et al. 2009). Fire is now one of

the major drivers of PSF loss and conversion to degraded land.

In this paper, we present the first avifaunal list for the Tuanan

research station in the Mawas Conservation Area and surrounding

areas based on field observations, mistnetting and camera trapping

conducted in Central Kalimantan from 2009 and 2010.

STUDY AREA AND METHODS

The Mawas Conservation Area comprises a 3,000 km2 area managed

by the Borneo Orangutan Survival Foundation located east of the

Kapuas river, about 55 km from Palangkaraya, the capital of Central

Kalimantan (Figure 1). Here, the Tuanan research station (2°09,06"S

114°26'26"E) was established in 2003 for long-term Orangutan

behavioural studies. It comprises a 9.45 km2 grid-based trail system

situated on peat of varying thickness up to 2 m. The forest was

subjected to selective commercial logging in the early 1990s,

followed by illegal logging (van Schaik et al. 2005). However, there

has been no systematic logging since 2002 (Vogel et al. 2009).

Despite this disturbance, the forest supports a relatively high density

of Bornean Orangutans Pongo pygmaeus wurmbii , Bornean Agile

Gibbons Hylobates agilis albibaris and other globally threatened

mammals (van Schaik & Brockman 2005, pers. obs.). Central

Kalimantan has a humid tropical climate, with very little variability

in temperature. The wet season normally occurs during the north¬

west monsoon in November to April: the climate is drier during

June to August. There is some climatic variation associated with the

El Nino Southern Oscillation cycle, which affects the duration and

severity of the seasons. During strong El Nino events, southern

Kalimantan can experience prolonged drought conditions (Page et

al. 2009). The mean average annual rainfall measured from 2004 to

2007 at the Tuanan research station was 2,678 mm, with an average

monthly rainfall of 223 mm (Wartmann 2008). Outside of the

Mawas Conservation Area, the PSF has been heavily disturbed by

humans. Drainage canals dug in the early 1990s for the Indonesian

government’s Mega-Rice Project have disrupted the natural

hydrology of the area, making it extremely prone to fire (Page et al.

2009). As a result, large areas have been subjected to one or more

fires and are now dominated by ferns and other low-growing plants.

30

MARY ROSE C. POSA & DAVID ALEXANDER MARQUES

Forktail 28 (2012)

Figure 1 . Map of the ex-Mega-Rice Project area with remaining peat

swamp forest cover (in black). Approximate location of the Tuanan

research station indicated. Straight solid grey lines are canals that were

dug for drainage.

Bird observations were made in the intact PSF around the

Tuanan research station as well as in surrounding degraded areas

using a variety of methods. DM mad e ad libitum observations from

November to December 2009, while MRCP conducted standard

10 minute, 25 m radius point-count surveys from August 2009 to

July 2010 (Posa 2011). In addition, MRCP also used mistnets and

camera traps to survey Tuanan for a total of 2,535 net hours and

3,924 trap nights respectively (details of methodology in Posa

2011). We included in our list only species that were confirmed by

visual sightings or distinctive calls, mistnetting and camera trapping.

With the exception of Bornean Ground Cuckoo Carpococcyx

radiatus (following Collar & Long 1996), nomenclature follows

the 2009 checklist of the Oriental Bird Club (sequence of

Dickinson 2003) available online at http://orientalbirdimages.org/

new-obc-checklist.html.

RESULTS

We recorded a total of 138 bird species from all survey methods in

the intact and degraded PSF habitats in and around the Mawas

Conservation Area (Appendix). Four of these were migratory

species, and two were most likely introduced recently to Borneo.

Thus, 1 32 resident birds were found in the PSF. Twenty-six species

were observed only along canals or rivers and in the non-forested

regrowth vegetation in areas that had previously been burned. The

rest of the species were observed in logged PSF and remnant forest

fragments. Several species of particular interest were observed,

including seven globally threatened, 31 Near Threatened species

as well as two PSF specialists (Hook-billed Bulbul Setornis criniger

and Grey-breasted Babbler Malacopteron albogulare) and three

Bornean endemics (Bornean Ground Cuckoo, Bornean Bristlehead

Pityriasis gymnocephala and Dusky Munia Lonchura fuscans).

We captured a total of 293 birds from 28 species in mistnets

(see Appendix), with recaptures (21 individuals) comprising 7.2%.

Two species that were recorded only by mistnetting were Blue-eared

Kingfisher Alcedo meninting and Oriental Cuckoo Cuculus

saturatus. Camera traps took 45 photographs of birds from ten

species, of which we were unable to identify three. Three species

were detected only through photography, namely Black Partridge

Melanoperdix niger , Crestless Fireback Lophura erythrophthalma

and Bornean Ground Cuckoo.

Significant records

Species accounts are given for threatened species, endemics and

habitat specialists and a few other notable records. We give the

species conservation status (Endangered, Vulnerable, Near

Threatened, Least Concern) based on the 2010 Red List by the

International Union for Conservation of Nature (IUCN 2010).

Black Partridge Melanoperdix niger

Vulnerable. A lone female was photographed in July 2010 in the

early morning. An uncommon and local resident on Borneo, this

species is poorly known, as it is shy and secretive. It has been

recorded from swamp forest in Gunung Palung National Park

(Laman et al. 1996).

CrestSess Fireback Lophura erythrophthalma

Vulnerable. Only detected by camera traps in intact forest, but the

commonest bird species recorded using this method (29 of 45

photographs of birds). Pictures of individuals or pairs were taken

between dawn and dusk, but the majority of photos was taken

before 08h00. Other researchers reported occasional encounters

on man-made transects, but in general this species is very elusive.

Storm's Stork Ciconia stormi

Endangered. Recorded three times by camera traps and two seen

flying over the research station in the early morning in June 2010.

This species has also been reported from other swamp forests,

includingpeat swamp (Laman et al. 1996, Danielsen etal. 1997, Page

et al. 1997). It is considered very rare throughout its range, but our

record and the recent one in Thailand, where it was thought to be

extinct (Cutter et al. 2007), indicates that camera trapping is a very

useful method in detecting this species’s presence in forested areas.

Lesser Adjutant Leptoptilos javanicus

Vulnerable. A few to a dozen individuals were spotted at various

times in drained and deforested areas while travelling on canals in

May-July 2010.

Wallace's Hawk Eagle Spizaetus nanus

Vulnerable. An adult was observed perched in the PSF interior in

December 2009.

Cinnamon-headed Green Pigeon Treron fulvieollis

Near Threatened. Individuals and a group of three were seen feeding

in fruiting trees lining canals in the deforested area in September

2009. This species was not recorded in intact forest, but was possibly

overlooked. It has also been recorded from PSF in Sebangau (Page

etal. 1997) and was reportedly the most abundant green pigeon in

the wooded areas of the Barito' region and southern Kalimantan by

Holmes & Burton (1987).

Long-tailed Parakeet Psittacula longieauda

Near Threatened. One bird was observed in PSF in December and

groups of more than a dozen birds were seen on dead remnant trees

in degraded areas in August to November 2009.

Forktail 28 (2012)

Peat swamp forest birds of theTuanan research station, Central Kalimantan, Indonesia

31

Chestnut-bellied Malkoha Phaenicophaeus sumatranus

Near Threatened. Surprisingly, a commonly encountered bird in

the PSF interior habitat, travelling in singles, pairs or groups of

three birds. Reported as uncommon throughout Bornean lowland

and hill dipterocarp forests (Mann 2008).

Bornean Ground Cuckoo Carpococcyx radiatus

Near Threatened. Endemic. One individual was photographed

following a Sun Bear Helarctos malayanus in December 2010.

Described as a rare resident of lowland forests (Mann 2008). It has

been recorded in a few other places in Central Kalimantan in

alluvial and swamp habitats (Long & Collar 2002, Fredriksson &

Nijman 2004).

Black Hornbil! Anthracoceros malayanus

Near Threatened. Small groups of up to eight individuals were

uncommonly encountered in intact forest.

Wrinkled Hornbil! Aceros conugatus

Near Threatened. One adult male was observed in PSF in

November 2009.

Red-crowned Barbet Megalaima rafflesii

Near Threatened. The commonest barbet in PSF and disturbed

forest around Tuanan with up to eight individuals recorded in a

day.

Great Slaty Woodpecker Mulleripicus pulverulentus

Vulnerable. Noisy conspicuous groups were uncommonly

encountered in intact forest and also observed in forest fragments

in the degraded area. This species has also been reported from PSF

in West Kalimantan (Laman etal. 1996).

Bornean Bristlehead Pityriasis gymnocephala

Near Threatened. Bornean endemic. Considered to be rare and

uncommon on the island (Mann 2008). Individuals and small

groups of up to five birds were uncommonly encountered in intact

PSF and forest fragments in the degraded area. This species has

been reported from PSF in Sarawak (Laman et al. 2006) and the

‘swamp’ forests ofTanjungPuting (Nash & Nash 1988). Smythies

(1981) suggested that it may prefer PSF, but Witt & Sheldon

(1994) refuted this.

Hook-billed Bulbul Setornis criniger

Vulnerable. PSF habitat specialist. Fairly common in intact forest,

often travelling in small groups. Their call has been described as a

rattling series of notes (3.4 kHz, 1 1 notes) or a soft crrrk (Myers

2009) or harsh alarm cuurrk (MacKinnon & Phillipps 2008). These

bulbuls are quite easy to detect in the PSF understorey because of

these calls, which we were able to record (Figure 2; www.xeno-

canto.org catalogue number XC74801). We also caught six

individuals in mistnets. This species has been observed in other

nutrient-poor forests in Borneo and was described as a local lowland

resident (Mann 2008). In Central Kalimantan, it has been recorded

in swamp forest in Sebangau (Pag eetal. 1997) and TanjungPuting,

as well as frequently encountered in kerangas at Barito UIu (Dutson

etal. 1991). It has been suggested that Setornis criniger is intolerant

of habitat degradation (Dutson et al. 1991), but our observations

indicate that it can persist in large blocks of selectively logged PSF.

However, the rapid loss of this habitat will continue to threaten

this species.

Grey-breasted Babbler Malaeopteron albogularis

Near Threatened. PSF habitat specialist. This species is rare except

in poor soil habitats such as PSF, kerangas and ultrabasic forests

(Sheldon etal. 2001). It is often overlooked because of its skulking

habits, absence from mixed-species flocks and infrequent

vocalisations. We observed it only on three occasions in the

understorey. However, it was the fourth most commonly

mistnetted species (19 out of 272 individuals), supporting the

suggestion by some authors that this method can reveal its presence

(Sheldon 1987, Dutson et al. 1991). Thus it may be overlooked

even in habitats where it is fairly common. Birds from Barito Ulu

and T anjung Puting are described as having white lores, while those

from north Borneo have yellow lores (Dutson etal. 1991, Sheldon

et al. 2001), but see Collar (201 1). The lores of the adult birds

captured at Tuanan were white but consistently have a few rufous

feathers on the edge near the forehead. We captured one juvenile

with an inflated gape on 14 June 2010. It had similar coloration to

the adult, except for a yellow lower mandible (grey in adult), brown

iris (red in adult) and pink legs (grey in adult).

Crimson-breasted Flowerpecker Prionochilus percussus

Least Concern. Observed on four days in November-December

2009 and on three days in March-June 2010 with a maximum of

four individuals. This species is supposed to be a rare lowland

resident, especially in northern Borneo (Mann 2008), but

identification issues may mask its true occurrence. No Yellow-

rumped Flowerpeckers P. xanthopygius were observed in the

Tuanan PSF, supporting the statement by Holmes & Burton

(1987): ‘ xanthopygius occurs with percussus in upper Barito Sep

1986 but not with it in southern lowlands of Kalimantan’.

Scarlet-breasted Flowerpecker Prionochilus thoracicus

Near Threatened. Single adult males were observed on two separate

occasions in the canopy of the PSF interior. An uncommon and

local bird on Borneo (Mann 2008). It seems to prefer poor soil

habitats (Sheldon et al. 2001).

Figure 2. Sonagram of the crrrk or

cuurrk call typical for Hook-billed

Bulbul Setornis criniger.

32

MARY ROSE C. POSA & DAVID ALEXANDER MARQUES

Forktail 28 (2012)

Dusky Munia Lonchura fuscans

Least Concern. Bornean endemic. Not uncommon in disturbed

forest, forest edge and forest regrowth areas around the Tuanan

site.

DISCUSSION

While only one study (Gaither 1994) has made a direct comparison

and shown that bird diversity in PSF is lower than in lowland

rainforest on mineral soils, we reach the same conclusion from our

survey in this nutrient-poor environment. We observed only 132

of Borneo’s 398 resident bird species in PSF, including only

three of at least 41 Bornean endemics, whereas other lowland sites

around Borneo show higher species richness — even up to twice

those numbers in eastern Sabah where forests are unusually rich

(e.g. Lambert 1992, Johns 1996, Cleary etal. 2007, Edwards etal.

2011). Aside from fewer numbers of species and endemics, total

abundance of observed birds is also very low. However, if we

compare the bird species (102) found in the Tuanan PSF (excluding

degraded and riverine areas) with other sites on Borneo containing

PSF (Appendix), the numbers of species are roughly similar. Laman

et al. (1996) reported 104 resident species from Gunung Palung

National Park, which contains about 400 ha of swamp forest in a

mosaic with lowland dipterocarp and upland forests. Tanjung

Puting National Park, which is composed of 50% PSF in mosaic

with freshwater swamp and heath forests, has 111 resident

species reported in its ‘swamp forest’ by Nash & Nash (1988),

although these authors surveyed peat basin margins and not

true PSF. Our list shares 97 species (41%) with the lowland

habitats of the Cabang Panti research site in Gunung Palung (61

reported from PSF and an additional 36 reported from lowland

dipterocarp forest) and 123 species (56%) with Tanjung Puting,

including 104 species reported from swamps by Nash & Nash

(1988) with an additional 19 reported by Bohap & Galdikas

(1987). Page et al. (1997) reported 150 species of birds observed

over three years from various habitats, including both forest and

riverine sedge swamp, at the Sungei Sebangau catchment in Central

Kalimantan. However, because they did not provide a complete

list of species, we cannot directly compare their results with ours.

Mistnetting has been conducted in PSF at only a few other sites

on Borneo. Gaither (1994) captured 34 species in Gunung Palung

but did not provide a complete species list. In Sarawak, sporadic

mistnetting from 1996 to 1999 in a previously logged 20-ha patch

of PSF at the UNIMAS Campus near Kuching revealed 68 resident

species (Tuen & Darub 1999, Rahman & Tuen 2006). However,

most of these (31) were represented by one or two captures only,

including M. albogulare. They also failed to detect S. criniger, which

suggests the site is heavily degraded and fragmented. In PSF at Loagan

Bunut National Park, mistnetting during a short 10-day survey

yielded 18 resident species, including S. criniger but not M.

albogulare; observations produced an additional 12 species (Laman

etal. 2006). In Sabah, six-days of mistnetting in primary PSF of the

Klias Forest Reserve (Sheldon etal. 2004) yielded 28 species. While

it is more difficult to set up mistnets in PSF than in dryland forests,

this method is very effective and, thus, useful for studying understorey

species, particularly in revealing the presence of M. albogulare.

However, with the mistnetting bias towards understorey species and

the low number of recorded species compared with point count

sampling, researchers should be judicious in the use of mistnetting

for rapid assessments of avian biodiversity in PSF (Remsen & Good

1996).

Camera trapping of birds has usually been incidental to surveys

of terrestrial mammals, but it is starting to be explored as a viable

method for sampling large ground-dwelling bird species (O’Brien

& Kinnaird 2008). Although low numbers of birds were recorded

with this method, it is a valuable method for detecting rare and

elusive species such as L. erythrophthalma and C. stormi in addition

to the other standardised procedures.

Kalimantan’s PSF may represent a stronghold for S. criniger and

M. albogulare , as large, albeit disturbed, tracts of this habitat still

remain. These species are most likely to be declining in

Peninsular Malaysia and Sumatra, where a greater percentage of

PSF has been converted to plantations and other non-forest land uses

(Miettinen & Liew 2010). Their current status needs to be assessed

and monitored in light of the recent rapid loss of PSF habitats. More

research is needed on the local distribution of bird species in PSF,

and specific microhabitat requirements of these species in PSF needs

to be elucidated. Despite low bird diversity in PSF, the occurrence of

the PSF specialists M. albogulare and S. criniger together with other

threatened and Near Threatened species underlines the urgent need

for continued efforts in the Mawas Conservation Area to prevent

further habitat loss and hunting.

ACKNOWLEDGEMENTS

We thank the Borneo Orangutan Survival Foundation (BOS), especially BOS

Jakarta and BOS-MAWAS Palangka Raya, Universitas Palangka Raya and

Universitas Nasional, Jakarta, for their facilitation and logistical support. We

are also grateful for field assistance from Tono, Ulang, Yundiata, Agus and

Andrika. Research permission was granted by the Indonesian State Ministry

of Research and Technology (RISTEK), Indonesian Institute of Science

(LIPI), Direktorat Jenderal PHKA, Balai Konservasi Sumber Daya Alam and

BAPPEDA Palangkaraya and the BOS Scientific Advisory Board. We thank

Drs. Fred Sheldon and Nigel Collar for their extensive comments on earlier

drafts of the manuscript. DM would like to thank Maria A. van Noordwijk,

Carel P. van Schaik and Michael Kriitzen from the Anthropological Institute

& Museum, University of Zurich, Switzerland, for their support. MRCP was

supported by the Singapore-Delft Water Alliance peatland research

programme (R 264-00 T004-272).

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Mary Rose C. POSA, Department of Biological Sciences,

National University of Singapore, 1 4 Science Drive 4, Singapore

1 17543; dbspmrc@nus.edu. sg

David Alexander MARQUES, Zentralstrasse 6, 8003 Zurich,

Switzerland; dmargues@bluemail.ch

34

MARY ROSE C. POSA & DAVID ALEXANDER MARQUES

Forktail 28 (2012)

Appendix

List of bird species recorded at Tuanan and surrounding areas and their occurrence in other sites containing peat swamp

forest habitat

Abbreviations: VU - Vulnerable; EN - Endangered; NT - Near-threatened; LC - Least Concern; I - introduced to Borneo; M - migrant to Borneo;

PSF - intact peat swamp forest; DIS - disturbed and non-forest habitat; RIV - riverine forest. Species marked with x* were recorded in 'swamp'

(including peat swamp) habitat in other sites, x - habitat type not specified or recorded from non-swamp habitat, * - caught in mistnet.

TanjungPuting, Unimas, Sarawak

Central (mistnetting in

Kalimantan Berbak, Sumatra Loagan Bunut regrowth, Tuen &

Tuanan record (Bohap & Galdikas Gunung Palung, (Silvius & National Park, Darub1999,

Status habitat 1987; Nash& West Kalimantan Verheugt 1986, Sarawak Rahman

Species (IUCN 2010) (this study) Nash 1988) (Laman etal. 1996) Hornskov 1987) (Gumalefa/. 2008) & Tuen 2006)

Forktail 28 (2012)

Peat swamp forest birds of theTuanan research station, Central Kalimantan, Indonesia

35

36

MARY ROSE C. POSA & DAVID ALEXANDER MARQUES

Forktail 28 (2012)

TanjungPuting, Unimas, Sarawak

Central (mistnetting in

Kalimantan Berbak, Sumatra Loagan Bunut regrowth, Tuen &

Tuanan record (Bohap & Galdikas Gunung Palung, (Silvius & National Park, Darubl999,

Status habitat 1987; Nash & West Kalimantan Verheugt 1986, Sarawak Rahman

Forktail 28 (2012)

Peat swamp forest birds of theTuanan research station, Central Kalimantan, Indonesia

37

FORKTAIL 28 (2012): 38-43

An annotated checklist of birds and conservation issues

in Salkhala Game Reserve, an isolated

Important Bird Area in Azad Kashmir, Pakistan

MUHAMMAD NAEEM AWAN, HASSAN ALI & DAVID C. LEE

Salkhala Game Reserve (SGR) in Azad Kashmir, Pakistan, lies within an Important Bird Area (IBA) of the Western Himalayas Endemic Bird

Area. The conservation status of the reserve and its birds is poorly known due to political instability in the disputed territory of Kashmir and

the relative remoteness of the site. The findings of a bird survey undertaken from May 2007 to April 2008 are documented here. In total, 101

species were recorded including 45 resident species, 48 breeding migrants and six winter migrants. There were significant records of the

globally threatened Western Tragopan Tragopan melanocephalus, the Near Threatened Pallid Harrier Circus macrourus and European Roller

Coraciasgarrulus, and the restricted-range Kashmir Nuthatch Sitta cashmirensis and Spectacled Finch Callacanthis burtoni. Kashmir Flycatcher

Ficedula subrubra and Cheer Pheasant Catreus wallichi were not recorded in the IBA, with the latter species now possibly locally extirpated.

An annotated checklist of the species recorded is presented along with measures of relative abundance. Habitat fragmentation, degradation

and clearance through the collection of fuel and timber, forest fire, livestock grazing, collection of non-timber forest products and

unsustainable use of pastures are the major threats to the wildlife of SGR. These conservation issues are discussed briefly along with

recommendations for the future management of the reserve.

INTRODUCTION

Located in the Neelum valley within the Western Himalayas

Endemic Bird Area (EBA; Stattersfield etal. 1998), Salkhala Game

Reserve (SGR) forms part of the Salkhala Wildlife Sanctuary

Important Bird Area (IBA; Chan etal. 2004). It is classified as such

owing to the presence of three globally threatened IBA trigger

species: Western Tragopan Tragopan melanocephalus , Cheer

Pheasant Catreus wallichi and Kashmir Flycatcher Ficedula subrubra

(BirdLife International 2011a). All three species are listed as

Vulnerable (IUCN 2011).

Western Tragopan is distributed in five separate populations in

the Western Himalayas of Pakistan and India (BirdLife

International 2001). Previous work in the Neelum Valley

established its presence in SGR (Mirza et al. 1978, Islam 1982)

and recorded it as ‘common’ and at densities of 0.8- 1.6 birds/km2

(Mirza et al. 1978). More recently, it has been recorded as locally

rare’ in the region (Hassan 2004). It is found in mixed coniferous

forest, often with a dense understorey, from as low as 1,350 m and

up to 2,800 m in winter, and from 2,400 m to 3,600 m in summer

(Gaston et al. 1983, Islam & Crawford 1987, Ramesh 2003).

Cheer Pheasant is patchily distributed, owing to its association

with early successional habitats, between 1,200 and 3,000 m

throughout the southern foothills of the Himalayas (Gaston et al.

1981, Garson 1983, Kaul 1993). In Pakistan, it is found in the

mountains of eastern North-West Frontier Province and Azad

Kashmir (Roberts 1991). A previous survey in SGR flushed 20

individuals (Mirza 1978) but, despite a recent record of 126 birds

in Jhelum Valley, Azad Kashmir (Awan etal. 2004), there have been

no reports of the species in SGR since.

Kashmir Flycatcher has a very restricted distribution in

northern India and parts of Pakistan, and occurs as a scarce and

apparently irregular summer breeding migrant in the side valleys

of Kashmir and the Pir Panjal range of northern Pakistan, with one

record from Sind, southern Pakistan (BirdLife International 2001).

It breeds between 1,800 and 2,300 m where there is predominantly

deciduous vegetation (Roberts 1992). In 1983, one breeding pair

with newly fledged young was recorded at 2,100 m in SGR (Roberts

1992).

In addition to the three IBA trigger species, the site is important

for a number of mammal species, including Kashmir Musk-deer

Moschus chrysogaster and Kashmir Gray Langur Semnopithecus ajax

(both Endangered), Himalayan Black Bear Ursus thibetanus

(Vulnerable), and Leopard Panthera pardus and Himalayan Goral

Naemorhedus goral (both Near Threatened) (Dar 2006, IUCN

2011).

There are six villages with a total population of about 6,000

people adjacent to SGR (Awan 2008). These communities depend

on the natural resources of the area, entering the reserve to graze

their cattle, cut trees for timber and collect firewood. Trunks of

older trees are sometimes partially burnt to make them easier to

cut. In addition to the loss of tree cover, these activities cause much

damage to the forest understorey of the reserve (Awan 2008).

Salkhala Game Reserve is situated at the ceasefire line between

Pakistan and India and, consequently, cross-border conflict between

1989 and 2003 prevented the completion of any field studies in

the area during that time. This, coupled with its relative remoteness,

means there have been few recent ornithological surveys in the

reserve (Islam 1982). This survey is the first to consider all bird

species in SGR and was conducted to provide a checklist for the

site, measures of relative abundance for key species, and a current

understanding of the conservation issues in the reserve after a

comparatively long period of isolation.

METHODS

Salkhala Game Reserve (34°33'N 73°50'E), Neelum Valley, is

located 80 km north-west of Muzaffarabad in the Himalayan

foothills of Azad Kashmir, Pakistan (Figure 1). Covering 810

hectares at 1,320-3,150 m elevation, it was notified as a Game

Reserve in 1982 and is classified as an IUCN Category IV protected

area (Dudley 2008). The reserve lies within the Himalayan moist

temperate ecozone (Roberts 1991) and consists of a range of forest

habitats, including coniferous, broadleaf and mixed coniferous-

broadleaf forests. These are characterised by the trees Cedrus

deodara , Pinus wallichiana , Abies pindrow, Picea smithiana , Taxus

wallichiana , Acer caecium , Bettila utilis, Berberis spp., Ouercus spp.,

Juniper us communis , Vibernum spp., Indigofera gerardiana,Juglans

regia and Aesculus indica. It has a mean annual rainfall of 1 25.7 cm,

with March and April being the wettest months, and is exposed to

heavy snowfall during the winter (Qureshi 1990).

We conducted a bird survey in SGR between May 2007 and

April 2008 using two methods: dawn and dusk call counts (Gaston

Forktail 28 (2012) An annotated checklist of birds and conservation issues in Salkhala Game Reserve, Pakistan

39

1980) for surveying Galliformes; and unlimited radius point counts

(Bibby etal. 2000) for surveying all other bird species. Twelve survey

points were positioned randomly and approximately 0.5 km apart

between 1,377 and 2,970 m elevation (Figure l), which was

representative of the altitudinal range and habitats covered by the

reserve. Ten points were located in coniferous forest (points 1-10

in Figure 1), and one each in mixed broadleaf-conifer forest and

scrub grassland. One point was surveyed during each dawn and dusk

survey, and each of the twelve points was surveyed twice per month,

once at dawn and once at dusk (total effort = 288 points).

Call counts of 60 minutes’ duration were conducted at 04h45-

05h45 (April-September) and 05h30-06h30 (October-March),

and 18hOO-19hOO (April-September) and 16h00-17h00

(October-March), with start time varying according to seasonal

differences in sunrise/sunset times. All calling Galliformes heard

were recorded and mapped. Point counts of 10 minutes’ duration

were carried out at the end of each dawn call count and start of each

dusk call count. All birds detected were identified and the number

of individuals recorded. If a bird group was only detected by call,

then a mean group size from visual contacts of that species was used

(Lee & Marsden 2008). A checklist for SGR was produced from

both sets of survey data. However, the survey methods employed

were not appropriate for effectively detecting birds of prey (Marsden

1998) and, consequently, these species are likely to be under¬

recorded in this study.

Species encounter rates were calculated based on the number of

individuals detected from all points surveyed, and presented as the

number of individuals per 100 point counts (± standard error).

Encounter rates were converted into ordinal categories of abundance:

<5 individuals per 100 point counts = ‘Rare’: 5.1-10 = ‘Uncommon’;

10.1-20 = ‘Frequent’; 20.1-40 = ‘Common’; and >40 = ‘Abundant’

(adapted from Lowen et al. 1996). These simple categories can be

used for future monitoring of the abundance of species within the

reserve (Robertson & Liley 1998). Mean encounter rates for each

species were calculated for each month (24 points/ month), and then

a standard error was derived from these sample means.

Bird survey data were supplemented by information gathered

from interviews with local staff of the AJ&K (Azad Jammu and

Kashmir) Wildlife Department (n = 15) and local community

members (n = 35); five community members were selected

randomly from each of the seven villages around SGR. Specifically,

this information was used to help confirm the presence/ absence of

Galliformes, and particularly Cheer Pheasant.

RESULTS

In total 1,959 bird records, comprising 101 species belonging to

38 families, were recorded in the survey. Of these, 45 were resident

species, 48 were summer migrants, six were winter migrants and

two were passage migrants. Two species were classified as

‘Abundant’, 16 as ‘Common’, 35 as ‘Frequent’, 44 as ‘Uncommon’

and four as ‘Rare’ (Appendix).

The survey recorded one of the three IBA trigger species, Western

Tragopan (29.1 ± 8.8 individuals/ 100 points; ‘Frequent’), and two

Near Threatened species, Pallid Harrier Circus macrourus (6.3 ± 4.2

individuals; ‘Uncommon’) and European Roller (6.9 ±4.9 individuals;

‘Uncommon’) (IUCN 2011), winter and summer migrants,

respectively. Cheer Pheasant and Kashmir Flycatcher, the two other

IBA trigger species, were not recorded during the survey. Interviews

with local communities did not provide any supporting evidence to

suggest that Cheer Pheasant is present within the reserve. In addition

to the tragopan, two additional restricted-range species (Stattersfield

et al. 1998), Kashmir Nuthatch Sitta cashmirensis (21.8 ± 5.7

Figure 1. Map showing the location and land cover types of Salkhala Game Reserve.

74°6‘0"E

i

73°54'0"E

73°55'0"E

73°56,0"E

73°57'0"E

Legend

© Survey Points

- Line of control

- Major Rivers

District Boundary

□ Salkhala GR

Sill Dense Conifer Forest

Medium Conifer Forest

Mixed Broadleaved / Conifer Forest

feS-Lj Grasses / Shrubs

: Agriculture / Cultivated Areas

Bare Land (Soil / Rocks)

Snow / Glacier

N

A

74°6'0"E

40

MUHAMMAD NAEEM AWAN, HASSAN ALI & DAVID C. LEE

Forktail 28 (2012)

individuals; ‘Frequent’) and Spectacled Finch Callacantbis burtoni

(12.5 ± 4.0 individuals; ‘Uncommon’), were also recorded. A

complete annotated checklist is given in the Appendix.

Interviews with local villagers revealed that a number of birds,

mainly Galliformes, are hunted to varying degrees within the

reserve. Hunting of Galliformes is probably higher in the reserve

during the winter months when birds move down to lower altitudes.

Western Tragopan is hunted for meat and feathers, and some skins

for taxidermy were for sale in local houses. A number of stuffed

Himalayan Monal Lopbopborus impejanus were also seen in many

homes. Kalij Pheasant Lopbura leucomelanos is hunted locally for

food, especially in the winter when pheasants migrate to lower

elevations. Koklass Pheasant Pucrasia macrolopha and Chukar

Partridge Alectoris chukar are also trapped and hunted for food by

local communities. Local villagers occasionally shoot Pallid Harriers

because they prey on their domestic chickens. All four species of

columbids recorded in the reserve are hunted, primarily by teenagers

and younger men, for food.

DISCUSSION

Salkhala Game Reserve is designated as an IBA due to the presence

of Western Tragopan, Cheer Pheasant and Kashmir Flycatcher. This

survey recorded 101 species, but included records for only one of

the IBA trigger species, Western Tragopan, lor which SGR is an

important site, along with Pallid Harrier, European Roller, and

Kashmir Nuthatch and Spectacled Finch, two restricted-ranges

species of the Western Himalayas EBA (Stattersfield et al. 1998).

Kashmir Flycatcher is an irregular and sparse summer migrant

to the area, so it is as feasible that it was present but undetected as

that it was absent in the reserve during the survey. Ol greater

conservation concern is the failure to detect Cheer Pheasant, with

its apparent absence from the reserve corroborated in local

interviews. For a species with a small and fragmented population

(BirdLife International 2011b), this loss from a protected site is a

worrying development.

The possible local extirpation of Cheer Pheasant from SGR is

indicative of a growing human population and an increasing

demand on natural resources affecting the conservation status of

species and habitats in what is a comparatively small protected area

(Awan 2010). Rising human activities are increasing the

conservation importance of the reserve in a landscape already

heavily impacted, raising concerns about site isolation and the

viability of populations of key species. The recent construction of

a road within the reserve, and its use for extracting trees that have

fallen due to heavy snow or landslides, has now made access to

wildlife relatively easy. Conservation threats within SGR include

habitat degradation and loss, through the collection of timber,

firewood and wild vegetables, hunting and overgrazing.

Hunting pressure is particularly high for Galliformes in the

reserve, with hunting for food, skins or recreation conducted by

local and non-local professional (trophy-hunting) and non¬

professional hunters alike. There is a seasonal shift in the type of

hunting pressure within the reserve. During the warmer months of

May-September, people from adjacent villages travel with their

cattle to higher grazing areas (above 2,400 m) and stay in their

summer homes in and around the reserve. At this time, people take

the opportunity to collect medicinal plants, vegetables and eggs

from pheasant nests, and to hunt wildlife (Qureshi 1990). Owing

to difficult terrain in the reserve, dogs are often used to flush birds,

especially pheasants, while traps may also be laid (Awan 2010).

During the winter months, people and their livestock move back

to lower elevations, and any hunting at this time tends to be

recreational rather than functional. In addition to the Galliformes,

Pallid Harrier, which is a rare winter visitor to SGR, experiences

some degree of hunting pressure as local villagers shoot it to protect

their chickens from predation.

Although commercial tree cutting is prohibited in all protected

areas in Pakistan, there is unlawful felling in SGR, especially in the

gullies in the north and south of the reserve (MNA pers. obs. 2008).

These areas tend to be at lower elevations, but logging activities

affect not only the species that occupy those elevations throughout

the year but also those that undergo seasonal migration during the

winter months. Of these, pheasants are most likely to be affected

since they are also hunted for food and trophies. To reduce the

impacts of harvesting forest resources, including hunting, Awan

(2010) recommended that the reserve be extended south-west to

the Gail Nullah area and east to the line of control and,

consequently, be better conserved under the protected area system.

Adding some form of mixed-use or buffer zone to try to shift

pressure away from core areas within what is a small reserve,

especially during the summer months when more people are

accessing and utilising the reserve, would seem likely to benefit the

reserve generally and the Western Tragopan in particular.

Man-made forest fires remain a threat to the conservation of

wildlife in the reserve, with large areas of forest affected by fires

every year (Qureshi 1990). These fires are especially prevalent

during the drier summer months, when people spend more time in

the forest and make fires for warmth at night and to help bring

down standing timber. From 1989 to 2003, cross-border firing

between India and Pakistan destroyed areas of natural forest growth

in and adjacent to the reserve.

A recent community-based awareness campaign was undertaken

to support the conservation of key bird and mammal species in the

reserve (Awan 2010). This included working with communities, in

schools, directly with hunters, and training local wildlife staff.

However, there remains a general lack of understanding of the

biodiversity importance of the reserve in those communities in and

around SGR. Consequently, it is vital that the efforts of this initial

programme are built on in a collaborative and constructive manner

to help improve the conservation status of SGR and the species

within it, while maintaining and supporting local livelihoods.

Now that the reserve is more accessible, it would benefit from

regular species monitoring to track general trends in species abundance

and habitat alteration, which, in turn, will help support effective

management of the site. In part, this could be included within the

planned surveys for Galliformes in the Western Himalayas of Pakistan,

coordinated by the World Pheasant Association-Pakistan and WWF-

Pakistan. At a basic level, using the same survey points as this study

may be a first step to establishing a bird monitoring scheme in SGR.

Although subjective and taking no account of detectability differences

between species, the ordinal categories of relative abundance that we

have used here may also provide a simple baseline to monitor and

detect any large-scale changes in the abundance of individual species

within SGR in the future.

ACKNOWLEDGEMENTS

We are very thankful to Prof. Z. B. Mirza for reviewing an earlier draft oi the

manuscript. We are also grateful to Mir Saleem, Supervisor Wildlife,

Muzaffrabad, and the wildlife staff of Salkhala Game Reserve for their support

during the field study. Thanks are also due to Baseer Qureshi who helped

with collecting data in SGR.

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Appendix

Annotated checklist of birds recorded in Salkhala Game Reserve

IUCN Red List status (IUCN 2011) follows the species name: VU = Vulnerable; NT = Near Threatened. RR after the species name indicates a

restricted-range species (Stattersfield et al. 1998). Species encounter rates are per 100 point counts ± standard error (SE). Abundance (ordinal

scale), with numbers of individuals encountered in parentheses: A = Abundant; C = Common; F = Frequent; U = Uncommon; R = Rare. Status,

with months observed in parentheses: R = Resident; S = Summer migrant; W = Winter migrant; P = Passage migrant; L = Local movement.

42

MUHAMMAD NAEEM AWAN, HASSAN ALI & DAVID C. LEE

Forktail 28 (2012)

Forktail 28 (2012) An annotated checklist of birds and conservation issues in Salkhala Game Reserve, Pakistan

43

FORKTAIL 28 (2012): 44-48

A review of the status of

Collared Laughingthrush Garrulax yersini

and Grey-crowned Crocias Crocias langbianis

SIMON P. MAHOOD & JONATHAN C. EAMES

A survey of the globally Endangered Collared Laughingthrush Garrulax yersini and Grey-crowned Crocias Crocias langbianis was conducted in

May 2009 on the Da Lat Plateau, southern Vietnam. Collared Laughingthrush proved to be common throughout broadleaved evergreen forest

habitat within its limited altitudinal distribution while the distribution of Grey-crowned Crocias was extended to the south. These results are

discussed in the context of the conservation status of these species and it is recommended that both species are retained as Endangered.

INTRODUCTION

For many species distributed in the tropics, a paucity of basic

abundance and distribution data makes the accurate assessment of

threat status difficult. There is still much to be learned about the

status and distribution of relatively well-known taxa, such as birds,

in countries like Vietnam. Vietnam supports over 800 resident bird

species (Robson 2008) and, owing to its complex topography, up to

six areas which are designated (or would qualify) as Endemic Bird

Areas (StattersfieldeTu/. 1998). In recent years the discovery of new

species and the revision of species limits has dramatically increased

the number of species endemic to Vietnam (or nearly so), revealed a

more detailed pattern of endemism, and reinforced the conservation

importance of the country (Eames etal. 1999a, Eames etal. 1999b,

Eames &: Eames 2001, Eames 2002, Collar 2006).

The Da Lat Plateau Endemic Bird Area (EBA 145) is located

on an isolated massif in southern Vietnam beyond the southern

tip of the Annamite mountains (BirdLife International 2011). The

natural vegetation cover is broadleaved evergreen forest with

emergent Pinus. The Da Lat Plateau Endemic Bird Area supports

more restricted range bird species than any other EBA in mainland

South-East Asia. Recent taxonomic revisions have reinforced the

status of the Da Lat Plateau EBA as the most important centre of

avian endemism in Vietnam. Following taxonomy used in the

BirdLife checklist version 4, it supports at least 14 species that

qualify as restricted-range species, of which five occur only in this

EBA, namely Orange-breasted Laughingthrush Garrulax

annamensis , Collared Laughingthrush Garrulax yersini. Grey-

crowned Crocias Crocias langbianis , Black-headed Parrotbill

Paradoxornis margaritae and Vietnam Greenfinch Carduelis

monguilloti. Approximately 40 subspecies of bird are also restricted

in range to this EBA (Eames 1995). Of the five species that are

restricted to the Da Lat Plateau EBA two are considered globally

threatened: Collared Laughingthrush and Grey-crowned Crocias.

Both are considered Endangered (BirdLife International 2011).

Accurately understanding the distribution and status of species

is of utmost importance in assessing their threat status. In Vietnam,

there have been few recent field surveys aimed at better

understanding species distribution and abundance. In this study we

conducted surveys to gather data on the abundance and distribution

of the two Endangered passerines that are endemic to the Da Lat

Plateau: Collared Laughingthrush and Grey-crowned Crocias.

Collared Laughingthrush Garrulax yersini

Collared Laughingthrush is part of a species complex that also

includes Chestnut-crowned G. erythrocephalus, Assam G.

chrysopterus, Silver-eared G. melanostigma, Golden-winged G.

ngoclinhensis and Malayan Laughingthrush G. peninsulae. Its

habitat, habits and vocalisations are similar to those species,

although unlike most of them, it is regarded by some as localised;

Robson (2008) describes it as ‘locally common’. Collared

Laughingthrush is an understorey bird restricted to high-elevation

broadleaved evergreen forest (>1,500 m elevation) on the Da Lat

Plateau. Its extent of occurrence (EOO) is only 720 km2 (BirdLife

International 2011) and it is known from only five sites: Chu Yang

Sin National Park in Dak Lak province, the contiguous Bi Duop

Nui Ba National Park and Dam Rong State Forest Enterprise (part

of which lies within Cong Troi Important Bird Area), Mt Lang

Bian in Dan Kia Suoi Vang Tourism Area, and Nam Ban Police

Forest Management Board (part of Tuyen Lam Important Bird

Area) (Tordoff 2002). Within these areas populations are

fragmented owing to the patchy distribution of suitable habitat,

which is often embedded in a matrix of unsuitable pine Pinus forest.

Collared Laughingthrush lives in pairs or small parties that forage

quietly and unobtrusively in dense understorey and on the forest

floor. It is usually detected by vocalisations, which are given

infrequently. However, it responds strongly to playback of pre¬

recorded vocalisations.

Grey-crowned Crocias Crocias langbianis

The Grey-crowned Crocias is known from only four specimens

collected at two localities. For a passerine with a continental

distribution this paucity of specimen records renders it most

unusual. After much intensive effort it was rediscovered in 1994,

in Chu Yang Sin National Park (Eames et al. 1995). It has since

been found in three parts of Tuyen Lam Important Bird Area, a

place known popularly as Ta Nung valley (and near Cam Ly, one

of the historical collecting localities) located within Lam Vien

Landscape Protection Forest, at Ho Tuyen Lam Tourism Area and

in the adjacent Nam Ban Police Forest Management Board (Pilgrim

etal. 2007), these constituting two discrete areas of the same forest;

at this site the populations may be small and the habitat fragmented.

Of the historical collecting localities little is known. At one site,

Bjorkegren’s ‘Lang Bian peaks’, which is presumably Mount Lang

Bian in Dan Kia Suoi Vang Tourism Area (Eames & Ericson 1996),

no suitable habitat within its altitudinal range remains. The location

of the other site is imprecisely known but is thought to be within

D'Ran Watershed Protection Forest (formerly known as Don

Duong Police Forest Management Board) (Pilgrim et al. 2007).

There are also recent reports from Bi Doup Nui Ba National Park,

including a photograph of an individual of this species purportedly

taken there in 2006 by the Vice-Director of the national park, Mr

Do Manh Hung (Le Trong Trai pers. comm. 2008). The location

at which this photograph was taken has been described variously

as ‘the slopes of Mount Lang Bian’ (Pilgrim et al. 2007) and ‘in

forest adjacent, along and close to road 723 at 1,450 m and 1,600

m elevation’ (Do Manh Hung pers. comm. 2009). Both locations

would place the record within Bi Doup Nui Ba National Park.

Forktail 28 (201 2) Status of Collared Laughingthrush Garrulax yersini and Grey-crowned Crocias Crocias langbianis

45

Plate 3. Hydro-electric dam construction in the Da Nhim Watershed

Protection Forest. A male Grey-crowned Crocias could be heard singing

from a tree top behind the photographer. (Jonathan C. Eames)

Plate 5. Road construction within the altitude range of Grey-crowned

Crocias in the Da Nhim Watershed Protection Forest. Note the gully

erosion and the tipping of soil over the side of the road. Note also the

clearance of pine forest for coffee. (Jonathan C. Eames)

The elevational range of Grey-crowned Crocias is poorly

understood, because although there are historical records from up

to 1,700 m elevation, recent records (excluding those claimed from

Bi Doup Nui Ba National Park and the 1 994 record from Chu Yang

Sin National Park, which was at c. 1,615 m) are from a narrowband

Plate 2. Habitat of Grey-crowned Crocias in the Da Nhim Watershed

Protection Forest. (Jonathan C. Eames)

Plate 4. Many parts of the Da Lat Plateau now support only an

anthropogenic landscape. Evergreen forest degraded by logging and

shifting cultivation can be seen in the far distance. A ridge covered by

Pinus kesiya plantation in the middle foreground and irrigated rice and

market gardening in the foreground. (Jonathan C. Eames)

between 910 and 1,130 m. It forages in the canopy and mid-storey

of broadleaved evergreen forest, sometimes utilising scrub between

forest patches, in pairs or small groups, in monospecific and mixed-

species flocks.

METHODS

Surveys were undertaken in four Forest Management Units with

suitable habitat for the target species located within the Da Lat

Plateau EBA, as part of a project to locate and map forest of high

conservation value (Mahoodef^/. 2009). We focused on sites where

the distribution of the target species was poorly known. The Forest

Management Units surveyed were Bi Doup Nui Ba National Park,

Da Nhim Watershed Protection Forest, D’Ran Watershed

Protection Forest and Don Duong Forest Company. Survey

locations are defined as Forest Compartments within Forest

Management Units. All of these Forest Management Units

receive some level of legal protection from illegal logging

and encroachment, although in the case of Forest Companies

(state owned logging concessions) and especially

Watershed Protection Forests legislation is rarely enforced. Bird

surveys were undertaken by three teams consisting of an

experienced ornithologist and a relevant member of staff from the

Plate 1. Expansion of coffee cultivation in the altitudinal range of Grey-

crowned Crocias within the Da Nhim Watershed Protection Forest.

(Jonathan C. Eames)

46

SIMON P. MAHOOD & JONATHAN C. EAMES

Forktail 28 (2012)

Plate 6. Grey-crowned Crocias in the Don Duong Forest Company at the southern limit of its range. (Jonathan C. Eames)

Forest Management Unit (local guides were also used where

required). Surveys were undertaken in the first two weeks of May

2009. Data were collected throughout the day, starting from dawn.

If there was moderate or heavy rain, or moderate or strong wind,

then the survey was abandoned.

The survey teams actively searched for indicator species on

foot, using existing roads and trails. Trails were walked slowly (1 —

1 . 5 km/hour) . The survey teams concentrated their effort in habitat

at elevations believed to be most suitable for target species. The

most suitable habitat for Collared Laughingthrush is believed to

be broadleaved evergreen forest with dense undergrowth above

1,500 m, and for Grey-crowned Crocias broadleaved evergreen

forest at 900-1,200 m, although this species was also searched for

in forest above 1,200 m. In suitable habitat, the survey team

broadcast prerecorded songs of the target bird species at 100 m

intervals and waited at each broadcast station for three minutes

afterwards to detect responding birds.

RESULTS

Collared Laughingthrush

Collared Laughingthrush was recorded at 93% (13 of 14) survey

locations where broadleaved evergreen forest reached over

1,500 m elevation. Our records in Da Nhim Watershed Protection

Forest are the first for that site, and they fill a gap in the species’s

range between contiguous forest in Bi Doup Nui Ba National Park

and Dam Rong State Forest Enterprise.

Grey-crowned Crocias

Grey-crowned Crocias was recorded at 57% (four of seven) survey

locations where contiguous forest reached below 1,200 m and 8%

(one of 13) survey locations where forest was only found above

1,200 m. Two sites where it was found were in areas where it was

previously unrecorded: pairs were seen at two locations at just over

1,500 m along a dirt road in Forest Compartment 62, Da Nhim

Watershed Protection Forest and a further seven pairs were found

along a 25 km transect in Forest Compartments 326 and 331, Don

Duong Forest Company (Plate 6). We also obtained the first

modern records of the species in D’Ran Watershed Protection

Forest. Three pairs were located in Forest Compartments 314 and

315 in a mosaic of broadleaved evergreen and coniferous forest.

Although the survey team spent four person-days surveying suitable

habitat for Grey-crowned Crocias in Bi Doup Nui Ba National Park

none was recorded.

DISCUSSION

The survey gathered new data for two globally threatened bird

species. These data have allowed us to expand the range of one

species (Grey-crowned Crocias) and re-evaluate the abundance of

another (Collared Laughingthrush). Based on these data we reassess

the global threat status of both of these species and recommend

that they be retained as Endangered. Because their altitudinal

distributions encompass almost the full elevational range of all of

the other restricted-range species that characterise the Da Lat

Plateau EBA and because they are reliant on broadleaved evergreen

forest, Grey-crowned Crocias and Collared Laughingthrush are

effective analogue species for the Da Lat Plateau EBA.

Collared Laughingthrush

Owing to the distribution of suitable habitat we only found this

species in Bi Doup Nui Ba National Park and the south-east corner

Forktail 28 (201 2) Status of Collared Laughingthrush Garrulax yersini and Grey-crowned Crocias Crocias langbianis

47

of the north-west sector of Da Nhim Watershed Protection Forest.

However, our records suggest that within its small range Collared

Laughingthrush is common wherever suitable habitat is found. The

population estimate given in BirdLife International (201 1) is

precautionarily placed in the band 2,500-10,000 based on the

observation that this species is less abundant than Black-hooded

Laughingthrush Garrulx milleti , which has been estimated to occur

at a density of 70 individuals/km2 in Chu Yang Sin National Park,

where it was recorded on 1 6 occasions compared to four encounters

with Collared Laughingthrush (Hill et al. 2001). Unlike Black-

hooded Laughingthrush, which occurs in demonstrative flocks that

vocalise often, Collared Laughingthrush is usually silent and

skulking unless provoked. These two species are therefore quite

different in habits and detectability. Pre-recorded vocalisations

were not used by Hill et al. (2001) to detect birds. We assert that

Collared Laughingthrush is more abundant than formerly believed

and that population estimates based on the encounter rate in Hill

et al. (2001) probably underestimate the population size; the

population might number at least 10,000 individuals. In addition,

we recorded Collared Laughingthrush at slightly lower than the

minimum recorded elevation, at 1,450 m (Robson 2008).

The high-elevation forests occupied by Collared

Laughingthrush are more secure than low- and mid-elevation forest

on the Da Lat Plateau, in part because they are mainly distributed

in protected areas which may confer some degree of security to the

species, at least in the short to medium term. Outside of the two

protected areas (Chu Yang Sin and Bi Doup Nui Ba National

Parks) there is relatively little broadleaved evergreen forest above

1.500 m, but because the overall area of habitat within this species

range is low, even these areas are of high significance in a

conservation context. In Da Nhim Watershed Protection Forest

clearance of high-elevation broadleaved evergreen forest, primarily

for coffee cultivation, is widespread. This cash crop is growing in

popularity with farmers, although much of the coffee produced is

robusta, a relatively low-value crop widely used for the production

of instant coffee; thus a relatively large area of coffee cultivation is

needed in order for farmers to turn a good profit. In 201 1 Nescafe

announced plans to build a large instant coffee factory in nearby

Dong Nai province, which is likely to increase the attractiveness of

coffee as a cash crop on the Da Lat Plateau.

Although habitat loss within its elevational range is less severe

than for Grey-crowned Crocias, and despite our belief that it is

more abundant than previously thought. Collared Laughingthrush

must have a relatively small global population, limited as it is to

broadleaved evergreen forest on the Da Lat Plateau. We

recommend it be retained as Endangered under criterion Bla+b

(i,ii,iii,iv,v).

Grey-crowned Crocias

Our records of Grey-crowned Crocias slightly expand its known

range to the south, to the southern limit of the Da Lat Plateau,

and bring the number of sites where it has been recorded to seven.

Our data indicate that it is uncommon and localised between 900

and 1,200 m and rare above those elevations. Its occurrence in forest

at the higher end of this elevational range may be dependent on

the persistence of contiguous forest lower down. The sites at which

we found the species contain small areas of forest at a suitable

elevation, and therefore do not significantly increase the global

population estimate. We concur with BirdLife International

(2011 ) that the population of this species is likely to be between

2.500 and 10,000 individuals. Don Duong Forest Company, where

we found seven pairs along a 25 km transect in logged broadleaved

evergreen forest (Plate 6) probably supports one of the largest

populations of the species. We have not been able to confirm its

occurrence in Bi Doup Nui Ba National Park, where it must be

rare and highly localised at best because although there is much

apparently suitable habitat it is almost entirely at the higher end of

the species’s altitudinal range. We suggest that records from this

location be treated as provisional until they are confirmed by an

independent survey.

Throughout its range most ol the forest at elevations suitable

lor Grey-crowned Crocias has already been converted to agriculture

or burned and replaced by coniferous forest (Plate 4). This process

is ongoing. At the location where we found the species in Da Nhim

Watershed Protection Forest, forest was being cleared for a

hydropower dam and coffee cultivation (Plates 1 and 3).

Throughout the Da Lat Plateau it is likely that forest conversion

activities will increase in line with ongoing road construction as

new areas are opened up to human colonisation. Because roads are

typically built at lower elevations, associated forest clearance

inherently affects lower- and mid-elevation forest before it impacts

higher-elevation forest. People have already settled alongside a road

that was built through Chu Yang Sin National Park in 2010;

this road connects to route 722 along which we found Grey-

crowned Crocias in DaNhim Watershed Protection Forest (Plates

2 and 5).

Suitable habitat for Grey-crowned Crocias is highly

fragmented, although its ability to tolerate and even possibly prefer

secondary habitats means that, with appropriate land management

(which is a long way off in Vietnam), it could potentially recolonise

much of its former range. Although it has recently been found in

Kon Turn province (Anon. 2012), its distribution there and in

intervening forest is extremely poorly known, and it is likely that,

overall, it has a very small global distribution within which all

suitable forest is rapidly declining in extent and quality, both inside

and outside protected areas. Our survey does not indicate a lower

level of threat than was previously assumed, and we recommend

that Grey-crowned Crocias be retained as Endangered under criteria

Bla+b (i,ii,iii,iv,v). It should be noted that although it has the same

threat category as Collared Laughingthrush, it is perhaps inherently

more vulnerable to extinction owing to its elevational preferences.

ACKNOWLEDGEMENTS

We thank the Trust Fund for Forests for financially supporting this survey

and the management boards of the special-use forests where we conducted

fieldwork for their cooperation. Le Trong Trai of BirdLife in Indochina

collected invaluable data for this publication. Pham Xuan Nguyen and other

staff from Bi Doup Nui Ba National Park were invaluable in arranging logistics

and were good company in the field, and Nguyen Cong Mao our driver took

us to places we would otherwise have not been able to reach. Craig Robson

and an anonymous referee provided comments on a draft of this manuscript.

REFERENCES

Anon. (2012) Grey-crowned Crocias crops-up in KonTum province, Vietnam.

Downloaded from http://birdlifeindochina.org/ on 28 March 201 2.

BirdLife International (201 1) Endemic Bird Area factsheet: Da Lat Plateau.

Downloaded from http://www.birdlife.org on 08/08/201 1.

BirdLife International (201 1) IUCN Red List for birds. Downloaded from http:/

/www.birdlife.org on 08/08/201 1.

Collar, N. J. (2006) A partial revision of the Asian babblers (Timalidae).

Forktail 22: 85-112.

Eames, J. C. (1995) Endemic birds and protected area development on the

Da Lat Plateau (Vietnam) Bird Conserv. tnternatn. 5: 593-425.

Eames, J. C. (2002) Eleven new subspecies of babbler (Timaliinae) from Kon

Turn Province, Vietnam. Bull. Brit. Orn. Club 122: 109-141.

Eames, J. C. & Eames, C. (2001) A new species of laughingthrush Garrulax

(Passeriformes: Sylviinae: Garrulacinae) from the Central Highlands of

Vietnam. Bull. Brit. Orn. Club 121:1 0—23.

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SIMON P. MAHOOD & JONATHAN C. EAMES

Forktail 28 (2012)

Eames, J. C. & Ericson, P. G. P. (1996) The Bjorkegren expedition to French

Indochina: a collection of birds from Vietnam and Cambodia. Nat. Hist.

Bull. Siam Soc. 44: 75-1 1 1 .

Eames, J. C., LeTrong Trai & Nguyen Cu (1995) The rediscovery of the Grey-

crowned Crocias Crocias langbianis. Bird Conserv. Internatn. 5: 527-537.

Eames, J. C., Le Trong Trai & Nguyen Cu (1999) A new species of

laughingthrush Garrulax (Passeriformes: Sylviinae: Garrulacinae) from

the Western Highlands of Vietnam. Bull. Brit. Orn. Club 1 19: 4-15.

Eames, J. C., Le Trong Trai, Nguyen Cu & Eve, R. (1999) New species of

barwing Actinodura (Passeriformes: Sylviinae: Timaliini) from the

Western Highlands of Vietnam. Ibis 141 : 1 — 1 0.

Hill, M„ Eames, J. C., Le Trong Trai & Nguyen Cu (2001) Population sizes,

status and habitat associations of forest birds in Chu Yang Sin Nature

Reserve, Dak Lak Province, Vietnam. Bird Conserv. Internatn. 1 1 : 49-70.

Inskipp, T„ Lindsey, N. & Duckworth, W. (1996) An annotated checklist of the

birds of the Oriental region. Sandy, UK: Oriental Bird Club.

Mahood, S. P., Le Trong Trai, Tran Van Hung & Le Anh Hung (2009)

Identification, planning and management of high-value forests:

final consultancy report. Hanoi: B i rd Life International Vietnam

Programme.

Pilgrim, J. D„ Nguyen Xuan Vinh, Nguyen Xuan Dang, Polet, G.,ThaiTruyen,

Tordoff, A. W„ Tran Huy Manh & Peters, J., eds. (2007) Biological

Assessment of the Dong Nai River Basin Conservation Landscape,

Vietnam. Final report, 1 9 October 2007.

Robson, C. R. (2008) A field guide to the birds of South-East Asia. London:

New Holland.

Stattersfield, A. J., Crosby, M. J., Long, A. J.& Wege, D. C. (1998) EndemicBird

Areas of the world: priorities for biodiversity conservation. Cambridge, UK:

BirdLife International (Bird Conservation Series No. 7).

Tordoff, A. W., ed. (2002) Directory of Important Bird Areas in Vietnam. Hanoi:

BirdLife International in Indochina and the Institute of Ecology and

Biological Resources.

Simon P. MAHOOD, Wildlife Conservation Society Cambodia

Programme, House 2 1, Street 21, Sangkat Tonle Bassac, Phnom

Penh, Cambodia. Email: simonmahood@gmail.com

Jonathan C. EAMES OBE, BirdLife International Cambodia

Programme, #9, Street 29 Tonle Basac, Chamkarmon, P.O. Box

2686, Phnom Penh, Cambodia. Email: eames@birdiife.org.vn

FORKTAIL 28 (2012): 49-52

Seasonal changes in plumage coloration of

Orange Bullfinches Pyrrhula aurantiaca

TILLTOPFER

Males of the sexually dimorphic Orange Bullfinch Pyrrhula aurantiaca have previously been thought to have a particular subadult body

plumage, whereas there is no such plumage stage known in all other Pyrrhula taxa.This particular plumage is characterised by light yellow

ventral colour compared to the intensive orange of adult males. It has alternatively been explained as representing geographic variation. In

contrast, reconstruction of the hitherto unknown moult cycle using museum specimens shows that birds are coloured richly orange from

late autumn to spring and become increasingly yellowish during the summer. The different coloration is therefore most likely a consequence

of colour fading because of carotenoid degradation (i.e. photobleaching) during the breeding season.

INTRODUCTION

The Pyrrhula bullfinch taxa are traditionally grouped on grounds

of distinctive plumage coloration, a pattern that is also supported

by molecular data (Bianchi 1907, Voous 1949,Vaurie 1956, Topfer

etal. 2011). Of the six (Dickinson 2003) or seven (Clement 2010)

currently recognised species, four are sexually dimorphic, with the

males being the brighter-coloured sex. For the first few months of

their life, juveniles of all Pyrrhula taxa are easily distinguished from

adults by an overall brownish and comparatively dull body plumage

(e.g. see plate 28 in Clement 1993). This plumage is replaced by

the adult plumage during the first autumn (Newton 1966, Jenni &

Winkler 1994).

In contrast, males of the sexually dimorphic Orange Bullfinch

P. aurantiaca have been suspected of having a distinctive subadult

body plumage, although there is no phenotypically distinct

subadult plumage known in any other Pyrrhula bullfinch. This

plumage is said to be characterised by a light yellowish ventral colour

compared to the intensive orange of adult males (Sharpe 1888,

Oates 1890, Roberts 1992, Rasmussen & Anderton 2005).

However, the different plumage colours are not always treated as a

sign of age (Grimmett et al. 1999), and have alternatively been

assumed to represent geographic variation (Bates & Lowther 1952,

Roberts 1992). Given the variety of interpretations and the lack of

comprehensive field data, I carried out a thorough re-examination

of study skins in order to identify the true nature of the light yellow

plumage and to understand the moult cycle of Orange Bullfinches.

METHODS

I examined 47 study skins (32 males and 15 females) of Orange

Bullfinches housed in the ornithological collection of the Natural

History Museum in Tring, UK (BMNH; Table 1), including the

male syntype of P. aurantiaca Gould, 1858 (BMNH 1858.9.7.5).

One male is a juvenile, with the characteristic brownish plumage.

The birds were collected at 14 localites in the Kashmir area

(Pakistan andlndia) during 1857-1937 (Table 1). These specimens

represent the most extensive set of Orange Bullfinch skins available

in natural history museums (very few specimens exist in other major

collections) and clearly outnumber the published records of

individuals from the wild. Most of the specimen-based descriptions

in the literature probably refer to these particular skins. The series

contains five from the Meinertzhagen collection that is notorious

for fraudulent data (Rasmussen & Prys-Jones 2003). However,

according to their preparation style, these individuals do seem to

be reliably labelled (R. Prys-Jones verbally 201 1), so were included

in the analyses. Exclusion of these specimens does not change the

results reported here.

I visually determined the colour of the ventral body plumage

of every male, distinguishing five colour classes: orange, yellowish-

orange, orange-yellow, yellow, and light yellow. I also compared

these with the best colour match in two colour guides (Smithe 1 975,

Anon. 2000). I documented the condition of the plumage in each

individual regarding signs of wear, feather gloss and abrasion of

feather tips, particularly of flight feathers and coverts, distinguishing

the stages: fresh, worn and heavily worn. In addition, I recorded

several morphometric parameters: wing length; wing tip; tail length

and graduation; bill length, depth and width; tarsus length; and

sagittal and lateral tarsus diameter. I analysed the measurements of

males according to body plumage colour, comparing orange (those

classified as orange or yellowish-orange) with yellow (orange -yellow,

yellow and light yellow), using unpaired t-tests (with a = 5%). All

altitudes of collected specimens were converted from feet to metres

in Table 1.

RESULTS

The occurrence of orange- and yellow-coloured male Orange

Bullfinches was confirmed. However, there are individual

differences, with some showing intermediate orange/yellowish

coloration. Taking into account this individual variation, there is

an obvious relationship between plumage colour and season (Figure

1, Table 1). A rich orange fresh plumage appears to be obtained in

October-November and remains bright until May, but gradually

fades to yellowish-orange and finally to yellow during summer

(Figure 2). This is well illustrated by a moulting bird from October

(BMNH 1897.12.10.244) that is growing some new feathers on

the belly and back that are contrastingly bright orange compared

with the other yellowish-orange feathers, while a male from mid-

November (BMNH 1858.9.7.5) shows a consistently rich orange

body plumage.

According to their labels, at least three males from the beginning

of July and one male from mid-August (BMNH 1949.25.3775,

3776, 3778 and 1949.Whi. 1.8743) were in full reproductive

condition. All four show a yellow or light yellow coloration.

Additionally, the labels of two females from the beginning of July

and from mid-August (BMNH 1949.25.3779 and 1969.41. 223)

confirm this time as the breeding season, since information on the

full development of the reproductive tract as well as data on nesting

material and nest construction are given.

No significant morphological differences were found between

orange and yellow individuals: in all ten morphometric dimensions

the two groups are statistically indistinguishable (Table 2). There

were no signs of moult limits in the greater wing-coverts of any of

the specimens and no distinctive differences in size or coloration

that allow ageing of first-year and adult Eurasian Bullfinches

50

TILLTOPFER

Forktail 28 (2012)

Table 1 . List of male Orange Bullfinch specimens examined, indicating locality and elevation, date, ventral body plumage colour as scored by

eye and compared with colour guide codes (Smithe 1 975, Anon. 2000), and plumage condition ('h.' = heavily). Specimens are listed chronologically

by day and month.

P. pyrrhula (Jenni & Winkler 1994, Winkler & Jenni 2007). For

example, in all birds the tips of the coverts are whitish-brown on

the outer vane and yellowish-brown on the inner vane. This pattern

does not differ between the juvenile specimen (BMNH

1897.12.10.245) and the other individuals, and is identical on the

growing coverts of one specimen from October (BMNH

1897.12.10.244).

The correlation between body feather colour and season is also

supported by examination of the overall plumage condition,

although with greater individual variation. Specimens from February

to mid-May show fresh and glossy remiges that subsequently abrade,

becoming heavily worn by August-September (Table 1). In some

individuals with only slightly worn flight feathers, the uppertail-

coverts show small white edges or tips, while the white edging is

Figure 1. Seasonal changes in

body plumage coloration of male

Orange Bullfinches illustrated by

selected specimens (ventral

view). The rich orange plumage of

birds in spring gradually fades to

yellow during summer, returning

to orange again after moult in late

autumn. From left to right: 1 965-

M-17967 (March), 1887.6.2.1522

(May), 1897.12.10.247 (July),

1887.6.1.1520 (September),

1858.9.7.5 (November).

© The Natural History Museum.

Forktail 28 (2012)

Seasonal changes in plumage coloration of Orange Bullfinches Pyrrhula aurantiaca

51

Table 2. Morphological comparisons of yellow and orange individuals.

absent from birds with heavy wear. This pattern holds true also for

female Orange Bullfinches that are not subject to such an obvious

change in body plumage coloration as males.

All of the specimens were taken from three adjacent regions that

today belong to the state Jammu and Kashmir in India and the

province Punjab and the special territory Gilgit-Baltistan of Pakistan.

The sampling localities are spread over an area of about 300 x 250 km

(see Wunderlich 1992), with two main concentrations of male

specimens: around Srinagar (Jammu and Kashmir, India; n=19) and

in the Gilgit area (Gilgit-Baltistan, Pakistan; n=10). Orange and

yellowish individuals occur among the birds from each of these

regions. From this geographic sampling, there is no evidence for a

geographic separation of orange and yellow plumage colours.

DISCUSSION

Differences in coloration of Orange Bullfinches appear to be

unrelated to age or geography, but instead linked to season and

caused by fading. The yellowish and orange colours are generated

by the carotenoid pigments canary xanthophyll A and B that are

known to be subject to fading over time (McGraw 2006). Because

carotenoids are usually deposited in feathers in their free, unesterified

form, they are probably more prone to light-induced degradation

than esterified pigments (through ‘photobleaching’: McGraw 2006).

Taking into account the higher levels of ultraviolet radiation in the

Orange Bullfinches’ high- altitude breeding grounds compared to

that in their much lower wintering habitats (2,400-3,900 m vs

1,550-2,330 m: Ali & Ripley 1974, Clement 1993, 2010,

Rasmussen & Anderton 2005), it appears likely that photobleaching

may be more extensive during summer. This could also explain the

apparent lack of change in the orange coloration of males during

winter (Table 1). Seasonal light-induced fading of carotenoid

plumage has also been demonstrated in House Finches Carpodacus

mexicanus (McGraw & Hill 2004) and Great Tits Parus major

(Figuerola & Senar 2005). Although different sets of carotenoids in

congeners may be differently susceptible to light-induced

degradation, a similar phenomenon to that reported here might be

expected for two closely related Himalayan species: Grey-headed

Bullfinch P. erythaca and Red-headed Bullfinch P. erythrocephala.

Since feather pigments are metabolised from dietary precursors

(McGraw 2006), the consumption of carotenoid-poor food during

or prior to feather growth could result in a dull yellow instead of

rich orange plumage colour. However, there is no evidence for this

in the skins studied: all the specimens dating nearest to the time of

moult (i.e. in winter) show bright orange plumage.

The variability of coloration is most probably not an artefact

of long-term colour fading in museum skins (as suggested by

Rasmussen & Anderton 2005): as early as 1888, Sharpe, referring

to specimens collected within less than 20 years, already remarked

that ‘every shade between yellow and deep orange is exhibited by

the series in the Museum’. Moreover, Doucet & Hill (2009) found

that reflectance spectra of museum specimens were very similar to

those from wild birds, including in the UV range. Notably, the

variation in reflectance spectra of wild populations might even

exceed the variation in museum specimens obtained from different

localities and over a century-long period (Doucet & Hill 2009).

Although some influence of specimen age on wavelength

reflectance cannot be denied (e.g. Armenata etal. 2008, Doucet &

Hill 2009), the results presented here are more likely to be

connected to season than to specimen age. The specimen BMNH

1897.12.10.244 (collected in 1880) which has some fresh, bright

orange feathers within its yellowish-orange body plumage further

indicates that proper century-long storage does not necessarily lead

to artificial colour fading.

It is very likely that seasonal pigment degradation has a

significant impact on plumage UV reflectance. Although not tested

here, it is possible that the UV reflection spectrum of the orange

plumage acts as a signal of sexual maturity, and that there is no need

for such recognition during the late stages of the breeding season.

Further work using UV reflectance spectrometry would be

informative, but is beyond the scope of the present study.

In general, the moult cycle of Orange Bullfinches roughly

corresponds to that of the European Bullfinch, in which adults have

a complete post-breeding moult, while juveniles undergo a partial

moult during which they renew the body feathers, but retain the flight

feathers until their second summer (Newton 1966, Busse 1984,

Roselaar 1994, Jenni & Winkler 1994, Glutz von Blotzheim 1997,

Winkler & Jenni 2007). Clement’s statements (1993, 2010)

suggesting that first-summer Orange Bullfinch males are deeper

orange above and below than females supports the conclusion that

males acquire their first orange body plumage during a post-juvenile

moult (Fig. 2). Interestingly, Bates & Lowther (1952) reported that

male Orange Bullfinches appear ‘clear yellow’ in the field, and

mentioned a specimen from March being much ‘deeper and duller

in tone’ than three other males from August that are ‘far brighter’, i.e.

more yellowish-orange. This observation precisely fits the findings

of this work. Unfortunately, other field studies are still lacking. Thus

the change in body plumage coloration from orange to yellow occurs

in summer without a moult, but appears loosely associated with the

timing of abrasion of flight feathers and uppertail-coverts in both

sexes. As there is no evidence for a distinct subadult plumage in the

Orange Bullfinch, there is also no indication for sexual plumage

heterochrony (see Bjorklund 1991. Badyaev & Hill 2003).

Feather abrasion, which leads to seasonal changes in plumage

colour and patterns in many bird species by revealing previously

hidden colours or patterns, can be excluded as a cause for the colour

Figure 2. Moult cycle of the Orange Bullfinch, reconstructed from study

skins. The colour change in male body plumage is indicated by the

colour gradient of the circle.

52

TILL TOPFER

Forktail 28 (2012)

changes in Orange Bullfinches. Individual body feathers are

bicoloured, having a black base and a yellowish to orange outer

portion covering about two-thirds of the feather. Thus, even though

heavy wear (particularly of flight feathers) can be associated with

orange -yellow or yellow plumages (Table l), extensive abrasion of

body feathers would not lead to fading but to darkening of the orange

coloration because of the underlying black feather bases. However,

such extensive abrasion of body feathers was not found in any of the

specimens examined (or in specimens of congeners).

The lack of statistical differences in measurements between

orange and yellow individuals does not necessarily mean that there

are not two age classes, because no significant difference exists in

wing length between first-year and adult European Bullfinches

(Glutz von Blotzheim 1997). However, while morphometric data

alone might not permit identification of age classes in Orange

Bullfinches, it is obvious from label annotations that yellow males

were in active breeding condition and were not immature. This is

further supported by the fact that most specimens from the

breeding season (late May to late August; Ward 1908, Ali & Ripley

1974, Roberts 1992, Clement 2010) are yellowish or yellowish-

orange instead of rich orange. As the pair-bonds form with the

break-up of winter flocks in early May (Roberts 1992, Clement

2010), a rich orange coloration may be important for mate choice,

although no field data are available to test this.

There is also no evidence for a geographic explanation for the

different plumage colours. While Roberts (1992) mentions males

from the Gilgit area being distinctively ‘much paler saffron yellow’

than those from the Hazara district (today Khyber Pakhtunkhwa,

Pakistan), there is no correlation between colour and locality in

the specimens I studied. In both of the areas from which specimens

were taken (the Srinagar area and Gilgit), there are both orange

and yellowish birds, with identical seasonal patterns of colour

change. Thus, there is no evidence for a potential subspecific

differentiation as suggested by Roberts (1992).

ACKNOWLEDGEMENTS

I cordially thank Robert Prys-Jones and Mark P. Adams for providing access

to the bird collection of the Natural History Museum at Tring and for their

helpful discussion of various aspects of this work. My sincere thanks go also

to Raffael Winkler and Mats Bjorklund for their useful remarks on the subject

and to Harry Taylor for taking the photograph. I also thank the anonymous

referees for their thorough review of the manuscript. This work was supported

by the Biodiversity and Climate Research Centre (BiK-F), Frankfurt/Main.

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Doucet, S. M. & Hill, G. E. (2009) Do museum specimens accurately represent

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McGraw, K. J.& Hill, G. E. (2004) Plumage color as a dynamic trait: carotenoid

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&W. M. Loskot, eds. Atlas der Verbreitung palaarktischer Vogel, 1 8. Berlin:

Akademie Verlag.

Till TOPFER, Senckenberg Natural History Collections Dresden,

Museum of Zoology, Konigsbrucker Landstrasse 159, 01109

Dresden, Germany; and Biodiversity and Climate Research

Centre & Senckenberg Gesellschaft fur Naturforschung,

Senckenberganlage 25, 60325 Frankfurt/M. Germany. Email:

TiU.Toepfer@senckenberg.de

FORKTAIL 28 (2012): 53-56

The use of tree-fall gaps by a forest interior avian frugivore

in a tropical evergreen forest

D. KHAMCHA & G. A. GALE

Previous studies suggest that forest birds commonly use tree-fall gaps owing to the relatively higher availability of fruits and arthropods.

However, others have argued that gaps may not be important for food resources but may offer other services, such as increased protection

from predators owing to denser vegetation associated with gaps. We examined the use of tree-fall gaps by Puff-throated Bulbul Alophoixus

pallidus, an abundant generalist frugivore found in tropical evergreen forests of East Asia. Bulbuls (family Pycnonotidae) are known to be

the most important small frugivores in this region (an area with the highest rate of deforestation in the world), and likely to be integral to

regional forest maintenance as well as regeneration. This bulbul did not preferentially use small gap interiors (average 1 63 ± 43 SE m2) and

probably avoided them, but it appeared to use areas immediately surrounding gaps more than expected particularly during the breeding

season. Such areas probably provide increased security for roosting adults and for young fledglings during the post-fledging period. This

first detailed study of the movement of forest bulbuls in tropical East Asia suggests that the use of gaps and gap edges by these frugivores

is highly variable in space and time (with equally complex reasons for their use or avoidance) but is likely to impact their role in forest

maintenance and restoration.

INTRODUCTION

Differences in age, size or location of tree-fall gaps — defined by

Brokaw (1982) as vertical holes in the forest extending from canopy

level down through a height of 2 m above ground — can greatly

impact the distribution and abundance of animals by influencing

the availability of resources (Fogden 1972). Here we focus on gaps

created through tree-fall or large branch-fall owing to wind storms

or other natural processes, excluding fire, rather than those caused

by human factors. Several initial studies suggested that frugivorous

birds foraged more frequently in gaps relative to forest interior sites,

presumably because gaps provided greater fruit (as well as arthropod)

resources. Additional light levels in tree-fall gaps have been suggested

to cause understorey shrubs and lianas to produce more fruits; gaps

also provide more foliage in the understorey for herbivorous

invertebrates and higher near-ground temperatures, which can also

be more suitable for a variety of forest arthropods (Blake Sc Hoppes

1986, Levey 1988, 1990, Malmborg& Willson 1988). These studies

also suggested that birds may change their foraging behaviour to

increase fruit and arthropod foraging in gaps depending on the

availability of resources (Blake & Hoppes 1986). However, more

recent detailed studies of gaps and food resources suggest that there

is a complex relationship between gap age and use by birds, and that

many birds do not appear to track resources tightly in gaps at least

at the scales measured (Restrepo et al. 1999). Nevertheless,

regenerating gaps may provide other services for birds, particularly

protection from predators owing to the density of vegetation

(Anders et al. 1998, Bowen et al. 2007).

The aim of this study was to investigate whether a common

frugivorous bird would preferentially use canopy gaps during the

course of a year and whether such preferential use was related to

food resources or other resources, particularly cover. We focused on

a forest bulbul because of the importance of bulbuls as seed dispersers

in East Asian forests (Corlett 1998) and because little is known

about the movements of forest bulbuls. Furthermore, South-East

Asia has the highest deforestation rate in the world (Sodhi & Brook

2006), so understanding the movement patterns of potential seed

dispersers such as these will be fundamental in maximising the

restoration potential of the region’s forests. We assessed ( 1 ) whether

Puff-throated Bulbuls Alophoixus pallidus , a locally abundant forest

species, would spend proportionately more time foraging in natural

tree-fall gaps or areas immediately surrounding gaps (referred to here

as ‘buffer’ areas) compared to areas away from gaps, and (2) whether

they would utilise gaps or buffer areas for resources other than food.

METHODS

The study was conducted in the 30-ha Mo-singto Long-term

Biodiversity Research Plot ( 14°26'N 101°22'E), Khao Yai National

Park (~2000 km2) in north-eastern Thailand. The plot consists of

a series of ridges and valleys with an elevation range of 730-860 m

asl. The vegetation is mostly mature evergreen forest dominated

by fleshy-fruited trees witli a ~0.25 ha section of secondary forest

at the north edge of the plot (Brockelman 1998). The plot is laid

out on a 20 m-square grid. All trees with diameter at breast height

(DBH) >1 cm have been identified, labelled with unique ID

numbers, and mapped using GIS with a precision ±1 m

(Brockelman 1998). When the birds are seen, the tree numbers were

used to reference their locations precisely.

One hundred and sixty-nine species of bird are known to occur

on the plot (Round et al. 2011), with Puff-throated Bulbul having

the highest density (>3 individuals/ha) (Gale et al. 2009). Puff-

throated Bulbul is a common resident of evergreen forest of

northern and north-eastern Thailand (Lekagul & Round 1991),

and studies from Khao Yai suggest that it is also the commonest

avian frugivore (Kitamuraet^/. 2002, Sankamethawee et al. 2011)

and consumes the widest variety of f ruits, based on faecal samples

(80 plants species from 185 samples) and direct observations of

fruiting plants (>100 species between 2003 and 2008)

(Sankamethawee et al. 2011). It forages in the lower to middle

canopy with an average foraging height of 9.4 m (range 1-15 m)

(P. D. Round et al. unpubl. data), in pairs or in groups of 3-7 birds

(Sankamethawee et al. 2009), with an average home range of

c.2.2 ha (Tanasarnpaiboon 2008). Birds also occasionally join

mixed-species bird flocks feeding on insects (McClure 1974,

Nimnuan et al. 2004). Their breeding season extends from

February to July (Pierce et al. 2004). Colour-banding was initiated

in January 2003 such that at the time of this study all family groups

(3 1-32 groups) and the majority of individuals (~300 individuals)

on the Mo-singto plot could be individually identified. As far as

we are aware no quantitative habitat selection data are available

for Puff-throated Bulbul, and while generally considered a forest

species (Lekagul & Round 1991) it has been observed foraging

along forest/ non-forest edges (Chaikuad 2000). Furthermore,

Bowen et al. (2007) found that many species generally classified as

forest-interior species were frequently observed in forest gaps, often

depending on season.

Vegetation was classified into three categories: tree-fall gaps,

gap buffer areas (0-10 m from the gap edge) and forest interior.

54

D. KHAMCHA & G. A. GALE

Forktail 28 (2012)

All the gaps in our study plot were created by natural tree-falls. We

used a somewhat narrower definition of a tree-fall gap than Brokaw

(1982): following Wongs riphuek (2008), who mapped gaps on the

Mo-singto plot just prior to the start of this study, we defined gaps

as open areas larger than 25 m2 and covered by vegetation shorter

than 5 m. Gap edges were marked with plastic stakes at 1 m intervals.

Tagged trees on the edge of gaps were used to georeference the gaps.

The coordinates were mapped in a GIS (ArcView 3.2a) to create

polygons for each gap and to estimate their area. The 1 0-m buffers

around these gaps were drawn in ArcView 3.2a using the Buffer

Selected Features command in the XTools extension.

We selected ten groups of Puff-throated Bulbul which occupied

c.20 ha of the core of the 30 ha Mo-singto plot, and precisely mapped

their movements using the georeferenced trees. All activities of the

ten selected groups were recorded to determine if they spent

proportionately more time in gaps, near gaps or away from gaps in

the surrounding forest. The ten selected groups were followed two

hours twice per month from May 2007 to April 2008 (for logistical

reasons no data were collected in September 2007). The

observations started when at least one individual in a group was

located. The behavioural data to examine the response to gaps were

taken from the second hour of each of the two-hour observations

to avoid possible biases relating to greater initial detection in gaps.

Observations on a particular group were distributed evenly

throughout the day to avoid potential biases associated with time

of day. Behavioural data were grouped into two seasons: breeding

(February-July) and non-breeding (August-January), which

included feeding (on both fruit and arthropods), breeding (nest

building, incubating, brooding and feeding offspring) and roosting

(perching, calling, preening and resting). To test the frequency of

gap and non-gap use, as well the frequency of the different

behaviours between breeding and non-breeding seasons, Fisher’s

exact tests were carried out using R version 2.1 1.0. Total use areas

were based on minimum convex polygons drawn around each of

the ten groups using the Minimum Convex Polygon command in

ArcView 3.2a for estimating the expected use. We used the ratios

of observed vs expected observations per group per habitat to

examine the overall trends within and among habitat types. To

generate the expected values we used the areal proportions of each

habitat type for each group mulriplied by the number of observations

of each group in each of the three habitat types.

RESULTS

There were 28 gaps (average 163±43 SE m2) within the home

ranges of the ten selected groups of Puff-throated Bulbul (mean

2.8±0.57 SE gaps/group), totalling 0.5 ha. The percentage areas

covered by gaps, 10-m buffers and interior forest were 2.09%,

10.05% and 87.86% for the total area occupied by the ten groups.

Based on the Mo-singto tree database, as expected most of the older

gaps were covered by a dense layer of herbaceous seedlings and

saplings, but had few trees with DBH >1 cm (0.1 trees/m2). The

10-m buffer areas were more densely vegetated (0.4 trees/m2) and

tree crowns were typically covered with a dense tangle of lianas. A

study of lianas on the Mo-singto plot also indicated that the number

of lianas was higher in gap areas (gap interiors and gap-edges) than

the primary forest (Wongsriphuek 2008). The database also

indicated that the forest interior had the highest tree density

(0.6 trees/m2).

Overall, 84.56% of the observations occurred in the forest

interior (n = 2,120), while the proportion of observations of bulbuls

in tree-fall gaps was 1.08% (n =27) and in the 10-m buffers 14.36%

(n = 360). The use of gaps and 10-m buffer areas tended to vary

with time. Of the observations in tree-fall gaps, the largest

percentage occurred in June (30%) (n = 7) and none in October-

Figure 1. The precentage of observations of Puff-throated Bulbul

activities in tree-fall gaps (n = 27), 10-m buffers surounding gaps (n =

360) and the forest interior (n = 2,120) as a percentage of the total

number of observations in each particular habitat based on the total

number of observations often different groups in each particular

habitat over an 1 1 -month period (May 2007— Apri I 2008).

December (Fig. 1). The number of observations in tree-fall gaps

was more frequent during the breeding season (February-July).

The highest percentage of observations in the buffer areas was

during March (17%) (n = 61), the lowest in August (3%) (n = 9)

(Figure 1). The number of observations in the forest interior

fluctuated relatively little during the study period (Figure 1).

Overall, there were 1,579 observations obtained during the

breeding season and 928 during the non-breeding season. For all

groups combined. Puff-throated Bulbuls used the gaps, 10-m

buffers and forest habitat differently between breeding and non¬

breeding seasons (Fisher’s exact test, p< 0.001) (Table 1 andbelow).

When analysed separately by group, this difference was significant

between seasons for only two groups; however, the other groups

seem to have similar patterns (Table 1).

Approximately 6 5.75%, 32.01% and 2.24% of the behaviours

were associated with roosting, feeding and breeding respectively.

There were significant differences in behaviour among habitats for

all groups combined (Fisher’s exact test, p < 0.001) (see below).

When analysed separately by group, behaviours were also different

among habitats for four of the ten groups (Fisher’s exact test, p =

0.01-0.03). Nine out of ten groups appeared to use gaps less than

expected (ratios were less than one), with three groups having no

detections in the gaps (Table 1). There was only one group that

appeared to use gaps more than expected (ratio was greater than

one), especially during the breeding season. The remaining groups

appeared in gaps more often during the breeding season but still

lower than expected overall (Table 1). In contrast, seven out of ten

groups used the 10-m buffer areas more than expected (ratios were

greater than 1), mostly for roosting and feeding, while two groups

appeared to use the buffer areas less than expected. Puff-throated

Bulbul was detected more often in the buffer areas during the

breeding season than non-breeding season (252 vs 108

observations). Although there were more hours of observation

during the breeding than the non-breeding season (12 vs 10 hours

per group), with an expected ratio of 6:5 for total observations,

there were 5.8 and 2.3 times more observations in the gaps and

buffer areas during the breeding season than the non-breeding

season respectively (Table 1). Fdowever, the magnitude of the overall

seasonal difference in gap use was somewhat skewed owing to the

relatively frequent use by one group during the breeding season.

There was no particular behaviour associated with the use of buffer

areas.

For the forest interior there were no trends among the different

behaviours or between seasons. There was only one group that

occurred in the forest interior less than expected (Table 1).

Forktail 28 (2012)

The use of tree-fall gaps by a forest interior avian frugivore in a tropical evergreen forest

55

Table 1. Use of tree-fall gaps, 10-m buffer zones surrounding gaps and forest interior by Puff-throated Bulbul from May 2007 to April 2008. The

breeding season (BR) is during February-July and the non-breeding (NB) August-January. The ratio refers to the ratio of observations per group

per habitat to expected observations. Expected observations were generated from the areal proportions of each habitat type for each group

multiplied by the total number of observations of each group in all habitats. The total expected observations were generated from the proportions

of each habitat type across all home ranges multiplied by the total number of observations of all groups in all habitats. Based on the total number

of hours of observation during each season, the expected ratio of breeding to non-breeding season observations was approximately 6:5.

Gap Buffer Forest

Season Season Season

'Groups which used habitat types differently between breeding and non-breeding seasons (Fisher's exact test, p< 0.05)

Although breeding behaviours did not occur significantly more

often in gaps or 1 0-m buffer areas, of the 56 observations associated

with breeding, 26 (46%) occurred during the post-fledging period.

Eight percent occurred in gaps, 19% in the 10-m buffer areas and

73% in the forest interior. Thus, while more than 25% of the

breeding-associated observations were related to caring for

fledglings in or adjacent to gaps, the proportion was not

significantly different from expected (%2 = 3.7, p = 0.057), although

suggestive of greater use of gaps and buffers for fledgling care. Of

the seven groups that successfully produced fledglings, three were

observed utilising gaps or 10-m buffer areas with their fledglings.

In the forest interior the number of observations during the post-

fledging period was not significantly different from expected (%2 =

0.37, p = 0.54).

DISCUSSION

Overall, there were only 27 observations of Puff-throated Bulbuls

using gaps out of >2,500 total observations, suggesting there was

no preference for gaps, and perhaps gap avoidance, with the

frequency of gap use being significantly lower than expected

(%2 = 12.02, p < 0.001; Table 1). During the non-breeding season,

Puff-throated Bulbuls were detected in gaps much less than

expected. Most of the detections in the gaps occurred during the

breeding season. Overall only one group out of ten appeared in the

gaps more than expected, while the other nine groups used them

less than expected, including three which were never observed to

use gaps. These results are in agreement with Bowen et al. (2007),

who suggested that forest bird use of gaps was seasonal and,

depending on species, more or less common than expected. For

these bulbuls open gaps may present a greater risk from predators

(Willson et al. 1982, Belisle & Desrochers 2002). Although we did

not determine the exact ages of the gaps, Puff-throated Bulbuls

tended to care for fledglings in older gaps where there was more

vegetation cover.

Overall, the 10-m buffer areas appeared to be particularly

important locations for roosting and feeding year-round,

presumably owing to the dense cover and density of food resources.

Regenerating gaps and areas immediately surrounding gaps were

covered by dense vegetation, especially lianas, at the gap edges rather

than in gap interiors. Pull-throated Bulbul prelerred to roost during

the day in areas adjacent to gaps where there was probably greater

cover and where they seemed to spend more time feeding fledglings

as well, presumably to reduce the risk from predators, especially

raptors (Anders et al. 1998, Belisle & Desrochers 2002, Bowen et

al. 2007).

While it is not possible to generalise from one species, this paper

and Khamcha et al. (2012) represent the first detailed studies of

the movement of forest bulbuls in tropical East Asia, and suggest

that the use of gaps and gap edges by bulbuls is probably highly

variable in space and time and the reasons for their use or avoidance

equally complex and probably dependent on the type of resource

that the gap and the surrounding habitat provides. Based on our

observations, the Puff-throated Bulbul probably contributes little

to forest gap regeneration, but rather to the overall maintenance of

forest interiors. Other forest bulbuls are worth investigation, as at

least some are highly likely to prefer gaps and therefore may be

significant contributors to the regeneration of fragmented forests

in the region.

ACKNOWLEDGEMENTS

We thank the Department of National Parks, Wildlife and Plant Conservation

and Khao Yai National Park for granting permission to conduct this study.

DK thanks W. Sankamethawee, N. Sukumal, M. Pliusungnoen, J.

Khunwongsa, P. Wongsorn, Y. Rattanapongsai, T. Ong-in, the Khao Yai

research team and the Conservation Ecology Program students for invaluable

assistance in the field. We also thank T. Savini, V. Chimchome and W. Y.

Brockelman for advice on the development on this project and manuscript.

We especially acknowledge the late David H. Reed for his advice on statistical

analysis; we will miss him greatly. The study was funded in part by the Thai

Biodiversity Research and Training Program (BRT T_350009).

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Chaikuad, K. (2000) Ecological niches of some bulbuls (family Pycnonotidae)

in KhaoYai Nation Park. MSc. dissertation, Kasetsart University, Bangkok.

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equatorial forest birds in Sarawak. Ibis 1 14: 307-343.

Gale, G. A., Round, P. D., Pierce, A. J., Nimnuan, S., Pattanavibool, A. &

Brockelman, W. Y. (2009) A field test of distance sampling methods for

a tropical forest bird community. Auk 1 26: 439-448.

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Influence of food availability and distribution on the movement

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pallidus).J. Trop. Ecol. 28: 1-9.

Kitamura, K., Yumoto,T., Poonswad, P., Chualua, P., Plongmai, K., Maruhashi,

T. & Noma, N. (2002) Interaction between fleshy fruits and frugivores

in a tropical seasonal forest in Thailand. Oecologia 1 33: 559-572.

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mixed-species insectivorous bird flocks in Khao Yai National Park,

Thailand. Nat. Hist. Bull. Siam. Soc. 52: 71-79.

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pallidus). Condor 111: 675-683.

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Daphawan KHAMCHA and George A. GALE, Conservation

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Bangkhuntien, Bangkok, Thailand. 10150. E-mails:

daphawan@hotmall.com and george.and@kmutt.ac.th

FORKTAIL 28 (2012): 57-61

Status and distribution of the endangered

Baer's Pochard Aythya baeri in Bangladesh

SAYAM U. CHOWDHURY, ALEXANDER C. LEES & PAUL M.THOMPSON

Continued declines in the now endangered Baer's Pochard Aythya baeri imply that this species may well be entering a downward spiral

towards global extinction. Knowledge of the drivers of this decline remain fairly anecdotal and even basic data on population trends are

hard to come by given the species's similarity to other diving ducks. We conducted a review of past records of Baer's Pochard in Bangladesh,

given concerns over the reliability of some past counts, to determine the true magnitude of recent declines, and also present counts from

targeted surveys in Bangladesh in the 2010-201 1 winter in addition to some basic autecological data. We conclude that several of the high

counts from the period 1 990-2005 may have been in error, but even accounting for these, the population has declined nationally by 99% in

just two decades.

INTRODUCTION

Baer’s Pochard Aythya baeri is a globally threatened diving duck

which breeds in Russia and north-eastern China and winters mainly

in eastern and southern mainland China, India, Bangladesh and

Myanmar. It was uplisted from Vulnerable to Endangered on the

IUCN Red List in 2008 in response to evidence of a drastic decline

in numbers at known wintering grounds. The principal drivers of

this decline are suspected to be habitat loss and degradation plus

over-hunting; the most recent global population estimate totalled

just 5,000 individuals. It is now absent or rare over much of its

former wintering grounds. For example, Thailand held as many as

426 individuals in the late 1980s but now only plays host to single¬

figure counts each winter (BirdLife International 201 1). BirdLife

International (2011) estimated the wintering population of Baer’s

Pochard in Bangladesh and India to be of the order of 1 ,000-2,000

individuals; however, recent counts from both the countries suggest

that this is a significant overestimate of at least one order of

magnitude, with a maximum of only c.100 individuals in the last

five years. The earlier estimate of 1,000-2,000 is probably based

on counts from the 1990s and early 2000s, some of which are of

questionable reliability (Li et al. 2009). Given the apparent

significance of Bangladesh for this species and a sharp population

decline in its wintering areas, we reviewed all past records and

counts of the species. In addition, SUC conducted a survey of the

wintering population of Baer’s Pochard in Bangladesh in 2010-

2011 that spans a wider spatial and temporal range than previous

surveys and complements recent data from Asian Waterbird

Censuses (Li et al. 2009).

METHODS

Review of records

In addition to compiling and reviewing past published records of

Baer’s Pochards, five senior birdwatchers in Bangladesh were

contacted for unpublished records of this species, with counts also

obtained from monitoring of a key site — Hakaluki Haor — and

information obtained on the reliability of past Asian Waterbird

Census (AWC) records. We compiled all records of Baer’s Pochard

from Bangladesh based on both published literature and

unpublished observations. It is important to note that Baer’s

Pochard is often difficult to separate from the sympatrically

wintering Ferruginous Duck Aythya nyroca and Tufted Duck

A. fuligula (Grimmett et al. 1998, BirdLife International 2001),

depending on expertise of the observer, viewing conditions and

quality of optical equipment. In order to assess the reliability of

old records, previous surveyors were contacted for Baer’s Pochard

survey methodology, field descriptions and in rare cases

photographic documentation. We treated older records as reliable

if the observers were known to be experienced surveyors with

previous experience of identifying this species. Some AWC records

were treated as unreliable if experienced observers, listed as present,

were subsequently found not to have participated in the counts.

Waterfowl surveys

SUC undertook waterfowl surveys in two haors, Hakaluki and

Tanguar, which are large depressions in north-east Bangladesh that

are deeply flooded in the wet season and retain water in several

beels (shallow waterbodies) in the dry season; historically the

majority of Baer’s Pochard sightings in Bangladesh have been

located at these haors. A total of 1 8 beels were thoroughly surveyed

for Baer’s Pochard and other globally threatened or Near

Threatened species such as Ferruginous Duck, Falcated Duck Anas

Jalcata and Baikal Teal A. formosa between December 2010 and

February 2011. Additional sites which had previously hosted Baer’s

Pochards such as Muhuri Dam, Hail Haor and some parts of the

Jamuna River were also surveyed. At least two observers conducted

surveys together at each site to minimise counting and identification

errors. Surveys were conducted both on foot and by boat. Counts

were repeated twice on all occasions and the maximum numbers

are presented here. Observations were made using 10x42 binoculars

and 25-50x spotting scopes.

Behavioural observations

SUC selected three male Baer’s Pochards to investigate their diurnal

time-activity budget, since males were easier to separate while

foraging within mixed flocks than females. These three individuals

were observed for c.180 minutes between 08h00 and 14h00 over

two days. Behaviour categories were established for four main

activities: foraging (diving, up-ending), resting (floating motionless

on the water with eyes open or sleeping), preening and swimming

(moving from one place to another). In each flock-scan, all birds in

the flock were checked from a rowing-boat for Baer’s Pochard,

working systematically from one side of the flock to the other using

a spotting scope. In addition, a GPS unit was used to measure

distance between the foraging site and the lakeshore. Water depths

were determined using a measuring tape attached to a bamboo stick.

RESULTS

Globally threatened waterfowl surveys in the

2010-2011 winter

During the winter 2010-2011 a minimum total of five Baer’s

Pochards were recorded by this survey, comprising four individuals

58

SAYAM U. CHOWDHURY, ALEXANDER C. LEES & PAUL M. THOMPSON

Forktail 28 (2012)

Table 1. Abundance of internationally significant waterfowl counted between December 2010 and February 201 1 during a search for Baer's

Pochard in selected key wetlands.

(three males and one female) at Hatirgata Beel and one female at

Rowa Beel, both in Tanguar Haor (Table 1). The four individuals

(2% of the total number of waterfowl counted in the flock) at

Hatirgata Beel were c.200 m from the shoreline at 1.88 m water

depth, within a mixed flock of 166 Red-crested Pochard Netta

rufrna (83%), ten Gadwall Anas strepera (5%), eight Ferruginous

Duck Aythya nyroca (4%), six Common Teal Anas crecca (3%) and

six Common Pochard Aythya ferina (3%). The single individual at

Rowa Beel was seen c.25 m from the shoreline at 1.26 m water depth

and was not accompanied by other waterfowl. In addition, in this

winter four individuals were recorded at Pashua Beel during the

regular mid-January waterbird census (E. Haquepers. comm. 2011),

and in Hakaluki Haor seven were seen at Haor Khal and one male

in Nagoa-Loribari during a census of the haor in mid-February,

giving a total of 17 Baer’s Pochard for the country (it is unlikely

that there was duplication in the counts given the close dates and

distances between sites). Three other globally threatened waterfowl

species (Table 1) were recorded including the third national record

of Baikal Teal and 17 Falcated Duck, which is the highest count

since 1993 in Bangladesh (Thompson et al. 1993, Thompson &

Johnson 2003).

Baer's Pochard diurnal time-activity budget

The activity budget data indicated (Figure 1) that Baer’s Pochard

spent most of its time between 08h00 and 14h00 resting (58%)

and swimming (32%), and considerably less time in foraging (7.4%)

and preening (2.6%). Baer’s Pochards were actively foraging and

swimming during the early hours of the day, mostly prior to 1 OhOO

and primarily stationary (either resting or preening) after 10h30.

Certainly, this information is insufficient to draw a firm conclusion

on the overall diurnal activity pattern of the Baer’s Pochard, and

further detailed observations are needed.

Figure 1 . Diurnal time-activity budgets of Baer's Pochard at Tanguar

Haor, Bangladesh, observed for c.180 minutes between 08h00 and

0% 20% 40% 60% 80% 100%

Percentage of time

Review of records since 1990

We were able to trace 59 reports of Baer s Pochards from Bangladesh

in the period 1990-2011, of which 40 records are previously

unpublished in detail (including observations from 2011), although

aggregate Asian Waterbird Census (AWC) counts have been used

in analyses of threat status of this species. Table 2 lists all records

known to us. Of these we consider 50 of the 59 records (85%) to

be reliable using the criteria outlined above. We consider that the

AWC published counts highlighted below and in Table 2 should

be treated as inconclusive, and the current national AWC

coordinator is of the same opinion (E. Haque pers. comm. 2011).

The counts deemed unproven comprise: 510 individuals from

Hakaluki Haor and Pasua Beel in 1995; 3,010 individuals mostly

from Pashua, Tanguar and Hakaluki Haors in 1996; and 3,580

individuals from Taguar Haor, Pashua Beel and Khana Muair Beel

of Sylhet division (Fi et al. 2009) in 2001. These AWC records (D.

Fi in lift. 2007) also included names of experienced but non-

participatory observers, so there was potentially a dearth of

experience within the survey team in 1996 and 2001; hence some

of the highest counts of this species which might have played a key

role in the current global population estimate (5,000 mature

individuals) (BirdFife International 2011) are considered

potentially unreliable.

Based on Table 2 the peak numbers recorded in a winter were

about 700 in January-February 1992 and about 1,700 injanuary-

February 1993 (the high estimates at three sites in 1993 are

considered reasonable, as very large numbers of duck were present

in the haors that winter), and just under 900 in 1994. Between 1995

and 2000 the true status is confused by the unreliable counts and

some gaps in coverage of key wetlands. After the last larger count

of 70 at Tanguar Haor in 2001 the most important site has been

Hakaluki Haor, where an average of about 10 birds has been

recorded in each of the last four winters. Table 3 summarises the

annual totals and peak counts each year for the two wetland systems

most regularly used by Baer’s Pochard — Tanguar and Hakaluki

Haors — and also indicates those years for which data are unavailable

or unreliable. The average annual count was about 660 during

1991-1995, is unknown during 1996-2000, was 99 during 2001-

2005 and has been only 6 during 2006-2010.

DISCUSSION

Population decline

Records of just five individuals during the targeted survey, an

additional 1 2 individuals during regular surveys in the winter 20 1 0-

2011, plus our review of past records, indicate that the wintering

population of Baer’s Pochard in Bangladesh has undergone a decline

of about 90% in the last ten years, and of 99% since the early 1 990s

even after discounting the highest counts that are considered

Forktail 28 (201 2) Status and distribution of the endangered Baer's Pochard Aythya baeri in Bangladesh 59

60

SAYAM U. CHOWDHURY, ALEXANDER C. LEES & PAUL M. THOMPSON

Forktail 28 (2012)

Table 3. Total and highest annual counts of Baer's Pochard in

Bangladesh and two main sites.

1 NS - Not Surveyed, NA - Not Accepted.

2 Year with square brackets indicate no count or no acceptable count from either of main sites.

unreliable (Table 2). Without historical data we cannot refute the

potential hypothesis that these early high counts may have been a

temporary phenomenon involving displacement of birds from other

wintering areas, but we consider this unlikely given that other

potential wetland sites were also covered by the AWC, at least

within Bangladesh. Moreover, there are few if any unsurveyed sites

to which birds might have relocated. Globally, between the period

1998 and 2008, a sharp decline (50-70%) has occurred, leaving an

estimated current global population of just 5,000 mature

individuals. Considering no double-counting between sites, an

estimated c. 1 ,000 Baer’s Pochards were recorded between late 2010

and early 201 1 in the wintering grounds, comprising 760 individuals

at Wuchang Hu, Anhui province, eastern China, which appears to

be the largest known current population of this species (M. Barter

inlitt. 2010); 152 in four sites of Hubei province, central China (J.

Lie in litt. 2011) and 1 in Deobali Jalah, Assam, India (P. Baruah

pers. comm. 201 1).

Current threats in Bangladesh

The lowlands of north-east Bangladesh formerly provided a wide

variety of habitats for waterbirds; shallow wetlands, seasonally

flooded grasslands and swamp forest were ubiquitous just a century

ago, but substantial areas have now been converted to agricultural

use (BirdLife International 2004). Illegal hunting of waterfowl using

poisoned baits (Alpha-Chloralose in boiled rice) in the two major

wetland complexes has proved to be a significant problem,

particularly at Hakaluki Haor with c.500 ducks poisoned in one

recent incident {The Daily Star 2010). The Northern Anas

acuta is the commonest victim of the poisoned baits, but

Ferruginous Ducks were also reported to be killed ( The Daily Star

2009). Although no evidence was found of trapping or poisoning

of Baer’s Pochard in recent years, hunting is still likely to be a

significant threat. Changes in attitudes and practices amongst

people who presently poison or trap ducks, and the enforcement

of laws, including an end to all use of poisoned baits, are urgently

needed for future protection of all waterfowl, globally threatened

or otherwise, in Bangladesh.

Human presence may also cause indirect effects through

disturbance. At Tanguar Haor fishing boats regularly pass through

most parts of the wetland. As these boats approach closer the ducks

abandon where they are feeding or loafing due to the interruption

(Muzaffar 2004). This might reduce the foraging efficiency of

migratory ducks, considering that the ducks are tolerant of boats

only if they are over 30 m from the toraging or roosting spot

(Muzalfar 2004). Recent management changes at Tanguar Haor (a

Ramsar site) have set rules to permit lishing by local people, but

have yet to zone the wetland effectively to provide disturbance-

free areas for wildlife. The resultant widespread small-scale fishing

may cause more disturbance than the traditional previous practice

of mass fishing of large areas once every three years. Moreover,

unsustainable fishing practices pose a huge threat to the haor

ecosystem and biodiversity as a whole (BirdLife International 2004,

Muzaffar 2004).

Time-activity budgets have been used extensively elsewhere to

provide valuable information on duck habitat use and wintering

strategies. However, knowledge of the behavioural ecology of diving

ducks (Aythyini) is lagging far behind that of dabbling ducks

(Anatini) (Houhamdi & Samraou 2008). Our observations suggest

that Baer’s Pochards actively forage between 08h00 and lOhOO at

1.26-1.88 m water depth; recently arrived individuals on the

breeding grounds were reported as foraging actively between

08h00-09h00 and diving to depths of 0. 5-2.0 m for c.40 seconds

at Xianghai Nature Reserve in China (Gao Jihong et al. 1992b).

Further studies on the behavioural ecology of wintering diving

ducks including the Baer’s Pochard are important for sound

evidence-based conservation planning decisions regarding

sustainable wetland management practices in Bangladesh.

CONCLUSION

This consolidation of both past and recent surveys confirms a

dramatic reduction in the wintering population of Baer’s Pochards

in Bangladesh, and indicates that the global population may be

substantially lower than previously thought. A thorough remote¬

sensing analysis is required to identify potential suitable wetland

habitats to be targeted by future surveys for Baer’s Pochard in north¬

east India and Bangladesh. Further studies are needed to determine

the potential negative effects on wintering waterfowl of current

fishing practices, synthetic agrochemical run-off, and disturbance

caused by intensive human activities. Urgent measures must be

taken to reduce the pressure of hunting, especially at key wintering

sites. Both Hakaluki and Tanguar Haors are recognised as

Important Bird Areas and are both considered nationally

Ecologically Critical Areas, and the latter also a Ramsar site

(BirdLife International 2004). Although both haors have been the

subject of recent projects aimed at establishing conservation and

wise use through co-management involving local communities, this

has not been reflected in recovery of wintering waterfowl

populations. Additionally, it is reported that wetland management

plans have not been properly implemented in either of these

internationally important sites (Muzaffar 2004). In order to

maintain a healthy wetland ecosystem to support both human use

and biodiversity, appropriate conservation-based wetland

management plans and better governance are required.

ACKNOWLEDGEMENTS

SUC is grateful to the British Ecological Society for funding this survey

(Application number: 2776 / 3471). ACL would like to thank CAPES -

Coordena^ao de Aperfeiqoamento de Pessoal de Nivel Superior for funding

support. SUC is thanklul to Ronald Haider and Farhad A. Pavel for their

support in the field. We thank Enam U1 Haque for information on past

waterbird counts under the AWC.

Forktail 28 (2012)

Status and distribution of the endangered Baer's Pochard Aythya baeri in Bangladesh

61

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The Daily Star (2009) Guest birds in danger: poachers active in Plakaluki

Haor. Downloaded from http://www.thedailystar.net/newDesign/

news-details.php?nid=75525 on 28.04.201 1.

The Daily Star (2010) Poachers poison to death hundreds of Plakaluki birds.

Downloaded from http://www.thedailystar.net/newDesign/news-

details.php?nid= 165072 on 28.04.201 1.

Thompson, P. M. & Johnson, D. L. (2003) Further notable bird records from

Bangladesh. Forktail 1 9: 85-1 02.

Thompson, P. M„ Plarvey, W. G„ Johnson, D. L., Millin, D. J., Rashid, S. M. A.,

Scott, D. A., Stanford, C. & Woolner, J. D. (1993) Recent notable bird

records from Bangladesh. Forktail 9: 1 2-44.

Sayam U. CHOWDHURY, Department of Environmental

Science and Management, North South University, Plot 15,

Block-B, Bashundhara, Dhaka 1229, Bangladesh. Email:

sayam_uc@yahoo.com

Alexander C. LEES, Dept, of Zoology, Museu Paraense Emilio

Coeldi, Caixa Postal 399, CEP 66040-170, Belem - Para, Brazil.

Email: alexanderlees@btopenworld.com

Paul M. THOMPSON, House 32, Road 10, Banani, Dhaka,

Bangladesh. Email: paul@agni.com

FORKTAIL 28 (20 12): 62-66

Breeding biology of Asian House Martin Delichon dasypus

in a high-elevation area

ZHIXIN ZHOU, YUE SUN, LU DONG, CANWEI XIA, HUW LLOYD & YANYUN ZHANG

We present data on the breeding biology of the largest known colony of Asian House Martin Delichon dasypus, located in the Jiangxi Wuyishan

Nature Reserve at 2,1 58 m in the Huanggang Mountains, China. Nest surveys conducted in abandoned buildings in a subalpine meadow

during March-August 2007 and 2008 yielded 163 and 132 clutches, from 84 and 82 nests, respectively. Breeding pairs also laid multiple

broods and replacement clutches. Average clutch size was 3.0 and 2.6 eggs for first and second broods respectively. Synchronous hatching

was detected in 79% of clutches. The proportion of eggs hatching was 0.7 and 0.6 for first and second broods respectively, and the proportion

fledging was 0.5 and 0.4 respectively. Nests situated inside buildings were more successful than those situated outside owing to greater

protection from severe weather, which was the major cause of breeding failure. Nest losses caused by severe weather were more pronounced

later in the breeding season.

INTRODUCTION

Many bird species raise only one brood per year because of a narrow

period of suitable environmental conditions which prohibits

multiple breeding attempts (Evans-Ogden & Stutchbury 1996).

Others raise multiple broods per breeding season (Verhulst et al.

1997) and the total seasonal fecundity for these species depends

on the number of breeding attempts (Geupel & DeSante 1990).

One of the principal determinants of multiple broods is the

duration of the breeding season, which is often directly related to

the abundance and availability of food (Holmes et al. 1992,

Morrison 1998).

At high elevations, climatic conditions are more severe and

unpredictable, with greater daily variation (Lu 2005). Such variable

weather may significantly increase the risk of nest failure and may

influence parental investment in each clutch and offspring (Bulmer

1984). Consequently, bird species that occupy montane habitats

may adjust their reproductive strategy by breeding over shorter

periods and producing fewer broods or smaller-sized clutches than

species at lower elevations (Sandercock et al. 2005, Bears et al.

2008).

We examined the reproductive ecology of the Asian House

Martin Delichon dasypus in the high-elevation montane

environment of the Huanggang Mountains in south-eastern

China. This widespread migrant species is known to occupy a

wide elevational gradient spanning both lowland and montane

habitats from 800 to 3,000 m (Turner & Rose 1989). In the

Huanggang Mountains, birds visit during March-August each

year to breed at elevations above 2,000 m, where weather

conditions are unpredictable, with severe rainstorms and

typhoons. Our aim was to evaluate the reproductive biology and

breeding success of this population under such environmental

conditions.

METHODS

Study site

Fieldwork was conducted in the Jiangxi Wuyishan Nature Reserve,

in the Huanggang Mountains (27°51,38"N 1 17°47/0l"E), Jiangxi

province, south-eastern China. Nest surveys were conducted from

March to August in 2007 and 2008. The study site represents the

highest peak of Wuyishan Cordillera, at an elevation of 2,158 m,

with a mean daily temperature of 14.3°C during May and August,

and mean annual precipitation of 1,500 mm. From June to August,

rainstorms become more severe and weather conditions are

extremely poor when typhoons occur.

Nest surveys

The 3-ha study area is predominately subalpine meadow habitat in

which are situated more than 30 abandoned buildings and garages

that provide suitable nesting substrate for the breeding martins. The

nest of Asian House Martin is a closed cup typical of hirundines,

built with tiny mud pellets and then lined with dried moss, sear’

grasses and feathers, leaving only a small entrance hole. We classified

two different nest locations: (a) under the eaves outside the

buildings, or (b) inside the buildings. Where possible for each nest

we recorded the overall size (width x height x depth) and the

dimensions of the nest entrance (widest x smallest diameter).

Systematic surveys of nests and their contents began on 1 March

in each year. Weekly visits were made to determine the arrival dates

and record pre-breeding behaviour. Mist-nets (6-8 m long, with 1

x 1 cm mesh) were set 1.8 m from the ground to capture adult

birds. For each individual caught we measured mass (g), total body

length, wing length, tail length, bill length and tarsus length (all in

mm). We banded the birds with coloured plastic rings and also

marked each bird (using indelible markers) with a unique pattern

on the throat and undertail-coverts to enable individual

identification from a distance using binoculars following Massoni

etal. (2007).

Once nest-building commenced, we recorded details daily. Four

observers visited each nest every day to determine the exact timing

of egg-laying. All eggs were numbered (in laying sequence) at the

pointed end using a waterproof ink pen (Gosler et al. 2005). Egg

length and width were measured to the nearest 0.01 mm using

plastic vernier calipers, and egg mass was recorded to the nearest

0.1 g using an electronic balance. Egg volume was calculated as

length x width2 x 0.000507, following Manning (1979). Clutch

volume was calculated as the sum of the volume of each egg laid in

a clutch. Clutch size was defined as the number of eggs in the clutch

at the onset of incubation.

Hatching date was defined as the date on which the majority

of eggs in each clutch hatched. Synchronous hatching was defined

as occurring when all eggs in the clutch hatched on the same day.

Incubation duration was calculated as the time elapsed from the

beginning of incubation to the hatching date. During the hatching

period, nests were checked every day to monitor the incubation

progress and obtain hatching dates. After hatching, we checked

nests carefully every day to monitor the growth of nestlings and to

determine the exact date they fledged. On each day, nestlings were

marked with indelible markers on their legs, and. weighed to the

nearest 0.1 g with an electronic balance. The nestling daily mass

gain and growth rate were calculated following Ricklefs (1968).

Nestlings were not weighed after 18 days to reduce the possibility

of premature fledging. Fledging date was defined as the date when

Forktail 28 (2012)

Breeding biology of Asian House Martin Delichon dasypus in a high-elevation area

63

all nestlings left the nest, and the nestling period was calculated as

the duration between the hatching and fledging dates. We

determined the length of the breeding season for each year by

calculating the number of days between the date when nest-building

for the first breeding attempt commenced and the fledging date

for the last clutch of the year.

Statistical analysis

Data were examined for normality using Kolmogorov-Smirnov

tests, and we pooled the data of two years for subsequent statistical

analysis. Comparison of breeding parameters across successive

broods (clutch size, fledging morphology, number of nestlings, etc.)

was conducted using non-parametric Mann-Whitney 17-tests.

Variation in the duration of incubation and other periods of the

reproductive cycle across successive broods was examined using

non-parametric Kruskal-Wallis tests. Spearman’s rank correlation

was used to detect the relationship between clutch size and the

proportion of eggs hatching. Logistic regression was used to model

nestling growth. Values are reported as means ± SE with sample

size («) and range. Differences were considered significant at P <

0.05. Data were analysed using SPSS 16.0 (SPSS Inc. 2008).

RESULTS

Nest construction and breeding

Asian House Martins were first observed in the study area on 13

March in 2007 and on 10 March in 2008, with nest-site selection

and territorial defence by paired birds first observed in early April

each year. Asian House Martins apparently preferred re-using

existing nest sites: in 2008, all 195 nests remaining from the

previous breeding season were re-occupied by the earliest-arriving

individuals. Individuals that arrived later constructed new nests.

The average external dimensions of nests (»= 37) were 121.9 ± 4.5

mm (70-181.1 mm) wide, 109.3 ± 2.8 mm (73.9-130.9 mm) high,

and 101.9 ± 2.6 mm (80-171 mm) deep. The average dimensions

of entrance holes (#=3 6) was 37.3 ±1.4 mm (18.3-60 mm) by

22.9 ± 0.6 mm (13.9-30.4 mm).

The maximum number of nests on any one day during the

season was 628 in 2007 and 675 in 2008. The number of nests

increased rapidly during April and May and peaked during June

(Figure 1; results for 2007 were similar). The number situated

outside buildings subsequently fluctuated and then declined after

the end of August, whereas the number of nests situated inside

buildings remained relatively stable (Figure 1).

Figure 1. Number of Asian House Martin nests located inside and

outside buildings each month in 2008. Shaded blocks indicate the

timing of severe weather events.

400

The first eggwas laid on 20 April in 2007 and 27 April in 2008,

when the majority of pairs had only just begun to construct nests.

The last breeding attempt was recorded on 1 August in 2007 and 3

August in 2008. In both seasons, the Asian House Martin

population had completely left the area by late August.

Clutch size, incubation, fledging and

nesting success

Pooling both breeding seasons, we monitored 295 clutches totalling

834 eggs from 166 nests. Of these, it was possible to measure and

collect data from 571 eggs. Clutch size ranged from 1 to 4 eggs

(25, 52, 167 and 51 clutches contained 1, 2, 3 and 4 eggs,

respectively), averaging 2.8 ±0.1 eggs. No replacement eggs were

laid if any eggs were lost. Eggs were coloured pinkish-white with

blood-red spots just after being laid, but gradually faded to a pale

white colour during incubation. On average, eggs were 17.9 ±0.1

mm (15.5-21.3 mm, zz = 5 5 8 ) long, 12.9 ± 0.2 mm (1 1 .6- 14.6 mm,

72=558) wide, and 1.5 ± 0.1 g (0.4-2. 2 g, n= 571) at laying. Mean

egg volume was 1.5 ± 0.1 cm3 (1.2-2. 2 cm3, 72=558), and clutch

volume averaged 4.2 ± 0.1 cm3 (1.3-6. 8 cm3, tz=195 clutches).

Incubation by both parents (distinguished by colour rings) lasted

15.3 ± 0.2 days (7-25 days, 72=195 clutches, excluding data from

four nests with only one egg since we were unable to determine the

exact date of incubation). There was no significant difference in

the duration of incubation between clutches of different size

(Kruskal-Wallis test, 72=191, /22=5.46, P= 0.07).

The hatching proportion among all eggs in both two years was

0.66 (72=834 eggs). At least one nestling hatched in 75% (222) of

all clutches recorded, and the mean proportion of eggs hatched in

each clutch was 0.62 ± 0.02 (72=295 clutches); this was positively

correlated with clutch size (Spearman r= 0.185, P=0.001).

Synchronous hatching was detected in 175 clutches (79%),

asynchronous hatching in 47 clutches, with the majority of these

(87%, 72 = 41) hatching within 48 hours (1-5 days). The number

of nestlings in a brood averaged 2.5 ± 0.1 (1-4, 72=222 clutches).

Among clutches which led to the fledging of at least one chick

(72=157), the nestling period lasted on average 28.2 ± 0.2 days (22-

35 days) and increased with the number of nestlings in a brood

(Spearman r=0.213, 72=153, 73=0.008). For these 157 successful

clutches, the mean number of fledglings was 2.5 ±0.1 (1-4). The

mean proportion of fledged nestlings in the clutches which had at

least one nestling averaged 0.69 ± 0.03. Of the 834 monitored eggs,

47% fledged. The mean proportion of fledglings per egg in a clutch

was 0.44 ± 0.03 (0-1, 72= 295).

The survival rate (to fledging) of nestlings inside buildings

was 0.84 ± 0.04 (n=73), which was significantly higher (Mann-

Whitney U test, 72=166, P=0.006) than the survival rate of

nestlings outside (0.65 ± 0.04, 72=93). Severe weather conditions

(rain and wind) caused by a typhoon in June 2008 resulted in the

loss of 50 nests inside buildings and 112 nests outside. The total

number of broods lost during two breeding seasons was 141 (48%,

72=295). Of these, 72 were lost during incubation, comprising 46

(64%) lost owing to nest damage caused by severe weather

conditions, and 26 (36%) owing to egg ejection by parental or other

adult individuals for unknown reasons. During the nestling stage,

69 broods failed owing to severe weather (49 clutches, 71%),

nestling starvation (10, 14%), eggs falling accidently from the nest

(9, 1 3%), and predation (presumed by a small rodent based on bite

marks; 1 clutch, 1%).

Nestling growth

We measured 381 individuals in 2007 and 2008 (199 adults and

1 82 fledglings; Table 1 ). There were significant differences between

adults and fledglings in mass (Mann-Whitney U test, 72=333,

P<0.0001) and tarsus length (Mann-Whitney U test, 72 = 353,

P=0.000l) (Table 1). Mean mass of nestlings at hatching was 1.4

64

ZHIXIN ZHOU etal.

Forktail 28 (2012)

Table 1. Morphological characteristics of Asian House Martin adults

and fledglings in study area (means ± SE, with range in parentheses).

" Significant difference between adults and fledglings, Mann-Whitney U test.

clutch (Table 2), except for the duration of both the incubation

(Mann-Whitney U test, «=169, P=0.054) and nestling period

(Mann-Whitney U test, « = 136, P= 0.93). Of the pairs that failed

to rear their first clutch successfully, 39 also laid a replacement

clutch. Of the 79 nests which had successfully fledged nestlings 44

clutches failed, for which 1 1 pairs laid further replacement clutches,

but only three of these hatched nestlings and none fledged

successfully. Replacement clutches for first broods also had

significantly lower breeding parameters than the first breeding

attempt (Table 2).

DISCUSSION

Figure 2. Nestling mass (means ± SE) as a function of age. Day 0

corresponds to the hatching day. The horizontal line indicates the

mean adult mass.

21 r

18

15 -

12 -

6 L

3 -

1 1 1

0 3 6 9 12 15 18 21

Age (d)

± 0.1 g (1.2-1. 5 g, n = 5). Nestlings reached a peak mass of

approximately 18.6 ± 0.8 g at 14 days of age, but then showed a

marked decrease to 16.7 ± 0.7 g at 18 days (Figure 2). Fitting the

mass as a function of nestling age to a logistic growth curve gave a

growth rate constant (if) of 0.210.

Multiple clutches

After successful fledging of nestlings, a second breeding attempt

was recorded for 79 nests (48%, 72 = 1 66). The majority of breeding

parameters from the second brood, in particular clutch size,

hatching proportion, fledging success and nesting success, were

significantly reduced when compared to the first successful

Our detailed description of the breeding parameters for Asian

House Martins nesting at high elevation (2,158 m) reveals that they

lay apparently smaller clutches (2.8 ± 0. 1 vs 3.4 eggs) but with larger

eggs (1.5 ± 0.1 vs 1.1 g) compared to colonies at lower elevations

(data from Yu etal. 1998 and Mi etal. 2007, at 500 m and 1,290 m,

respectively), presumably as a local adaptation to boost reproductive

success by investing more in fewer offspring (Lu 2005), although

we were unable to test this. The Jiangxi Wuyishan Nature Reserve

breeding colony, with over 600 pairs, is the largest described to date.

Typically, hirundines breed in smaller colonies (Snapp 1976), and

smaller colonies of Asian House Martins are known at a number of

lower-elevation sites (<1,800 m) in the Wuyishan mountains. The

large colony size may be a consequence of a local shortage of nest

sites (Snapp 1976, Shields & Crook 1987), but we were unable to

test this.

We found that Asian House Martins at the Jiangxi Wuyishan

Nature Reserve breed over a five-month period, from April until

August, the beginning of which is apparently at least 30 days earlier

than the equivalent date for a population at l,290m(Mi etal. 2007)

and compared with Northern House Martins D. urbicum breeding

at a latitude of 39°N (Saygili & Yigit 2007). We speculate that this

earlier initiation of breeding may be an adaptation to the increased

risk of nest failure later in the season when extreme weather

conditions are more likely.

Selection of nest sites offering greater protection from severe

weather conditions was of critical importance to the breeding

success in this population. We found that fewer pairs used more

exposed nest sites outside buildings, and that such nests were subject

to more pronounced fluctuations in nesting success than those

located inside, owing to nest damage from heavy rain and strong

winds. Such differences have not previously been recorded in

hirundines, to our knowledge.

Asian House Martins produced multiple broods and

replacement clutches in an attempt to maximise the number of

Table 2. Breeding parameters for successive clutches.

4 = significant difference between first and second clutches (Mann-Whitney U test, P<0.05)

b = significant difference between first and first replacement clutches (Mann-Whitney U test, P< 0.05)

Forktail 28 (2012)

Breeding biology of Asian House Martin Delichon dasypus in a high-elevation area

65

offspring they produce per season. However, mean clutch size,

hatching proportion, fledgling success and nesting success declined

significantly across successive broods. Declines in mean clutch size

and hatching success through the breeding season have also been

reported in other hirundine species (Zhao 1981, Sakraoui et al.

2005) and other migrant species (Dhondt et al. 2002). Multiple

broods and replacement clutches have been reported for several

hirundine species including Northern House Martin (Bryant 1979,

Lahlah et al. 2006), Barn Swallow Hirundo rustica (Jones 1987,

Sakraoui etal. 2005), Tree Swallow Tachycineta bicolor (Clapp 1 997,

Monroe et al. 2008), White-rumped Swallow T. leucorrhoa

(Massoni etal. 2007) and Purple Martin Progne subis (Allen & Nice

1952). In one experimental egg-removal study, 41% of female Tree

Swallows laid replacement clutches (Rooneem & Robertson 1 997),

which is lower than the 63% of female Asian House Martins at the

Jiangxi Wuyishan Nature Reserve. The capacity for laying multiple

broods and replacement clutches has been shown in other species

to be dependent on a number of factors including female age and

quality, timing of first clutch, size of first clutch, and local resource

abundance (De Laet & Dhondt 1989, Smith & Marquiss 1995,

Tinbergen & Verhulst 2000).

Significant declines in breeding success across successive

broods are often related to parental energy budgets and how

many young both parents are able to feed (Lack 1968, Bryant &

Westerterp 1980). Whether or not the declines observed for

Asian House Martins are related to seasonal variation in prey

abundance, or other environmental constraints imposed on the

parents, remains to be seen. The seasonal declines in breeding

success may simply reflect reduced foraging efficiency by parent

birds caused by poorer weather during the later stages of the

breeding season, as recorded in Barn Swallow (Pilastro & Magnani

1997), Tree Swallow (Nooker et al. 2005), Brown-chested Martin

P. tapera (Turner 1984) and Crag Martin H. rupestris (Acquarone

etal. 2003).

ACKNOWLEDGEMENTS

We thank the staff of Jiangxi Wuyishan Nature Reserve for their help in

fieldwork and data collection, and an anonymous reviewer for helpful

comments. This study was supported by Ministry of Education Key

Laboratory for Biodiversity Science and Ecological Engineering, Beijing

Normal University.

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Zhixin ZHOU, Ministry of Education Key Laboratory for

Biodiversity and Ecological Engineering, College of Life Sciences,

Beijing Normal University, Beijing, 100875, China, and Key Lab

of Animal Ecology and Conservation Biology, Institute of

Zoology, Chinese Academy of Sciences, 1 00 101, China. Email:

zhixinzhou@gmail.com

Yue SUN, Ministry of Education Key Laboratory for Biodiversity

and Ecological Engineering, College of Life Sciences, Beijing

Normal University, Beijing, 100875, China, and State Oceanic

Administration Beijing, 100860, China. Email:

sunyuebird@ 1 63.com

Lu DONG & Canwei XI A, Ministry of Education Key Laboratory

for Biodiversity and Ecological Engineering, College of Life

Sciences, Beijing Normal University, Beijing, 100875, China.

Emails: tiantai_bird@yahoo.cn & xiacanwei@126.com

Hu w LLOYD, School of Science and the Environment,

Manchester Metropolitan University, Chester Street,

Manchester, UK, Ml 5GD. Email: H.Lloyd@mmu.ac.uk

Yanyun ZHANG, Ministry of Education Key Laboratory for

Biodiversity and Ecological Engineering, College of Life Sciences,

Beijing Normal University, Beijing, 100875, China. Email:

zhangyy@bnu.edu.cn (author for correspondence)

FORKTAIL 28 (201 2): 67-70

Sex bias in a wintering population of Dunlin

Calidris alpina in central Taiwan

DZUNG-YUN YANG, CHUNGYU CHIANG & YU-CHENG HSU

The sexes of many migratory shorebird species differ in timing and distance of their migrations, but this phenomenon has been little

studied for birds in the East Asian-Australasian Fly way (EAAF). Using molecular sexing techniques (but also taking morphometric data), we

compared the sex ratio of Dunlin Calidris alpina sampled on the west coast of Taiwan between September 2007 and May 2008. We found an

overall sex bias (59.2%) toward females, including juveniles. A discriminant function deduced from three morphological measurements

correctly identified 89.0% of Dunlin, but performed better in identifying males than females. Since Taiwan is at the southern end of the

Dunlin's East Asian winter range, our results support the 'arrival-time' hypothesis whereby the sex that arrives earlier on the breeding

grounds winters closer to them. To maintain stable populations, conservation must take account of this differential use of wintering grounds

by the two sexes.

INTRODUCTION

In many migratory bird species, individuals of different sex or age

differ in their routes and timing of annual migration (Cristol et al.

1999). Several hypotheses have been proposed to explain these

differential migrations, focusing on differences in territorial

behaviour, body size and behavioural dominance. The arrival time

hypothesis proposes that certain individuals winter closer to the

breeding grounds because competition for mates or territory

favours early arrival to the breeding grounds (see Morbey &

Ydenberg 2001). The body size hypothesis proposes that the

larger-bodied sex can winter at higher latitudes due to larger fuel

reserves and better thermal efficiency (see Catry et al. 2005). The

dominance hypothesis suggests that the dominant sex will out-

compete the less dominant one for resources, forcing the less

dominant sex to migrate longer distances or winter in inferior

habitats (see Marra 2000). Testing these hypotheses is difficult

because they often are not mutually exclusive. An early test of these

hypotheses by Myers (1981), using data from several shorebird

species, found that the sex that arrived earlier at the breeding

grounds was the sex that wintered at higher latitudes. He proposed

that, regardless of size and behavioural dominance, difference in

arrival time alone is sufficient to predict a latitudinal cline in

wintering shorebirds.

Dunlin Calidris alpina is a long-distance migratory shorebird,

breeding along the Arctic rim in Europe, Iceland, Scandinavia,

Siberia, North America and Greenland (Piersma et al. 1996) and

wintering in temperate and tropical regions throughout the

Northern Hemisphere (Wenink & Baker 1996). Studies on

wintering Dunlin in Hong Kong have found two distinct

mtDNA haplotypes, Siberian and Alaskan (Wenink & Baker

1996). Significant differences exist in culmen length and

mtDNA haplotypes among Dunlin of different breeding

populations (Wenink & Baker 1996, Wenink et al. 1996). Dunlin

ringed in Taiwan have been recaptured on breeding grounds in

both Alaska and Siberia, and in Taiwan we have re-sighted Dunlin

which were ringed in Alaska and Sakhalin Island (Lanctot et al.

2009, Fernandez et al. 2010, CC pers. obs.). During the non¬

breeding season, significant numbers migrate from breeding

grounds to winter in eastern China, Japan, Korea, Hong Kong and

Taiwan (Bamford et al. 2008). Around 950,000 Dunlin use this

flyway every year, making the East Asian-Australasian Flyway

(EAAF) one of the five major flyways for this species (Bamford

etal. 2008).

Female Dunlin are larger than males, but it is the males that

arrive earlier at their breeding grounds and defend territories each

spring (Holmes 1966). According to the arrival time hypothesis,

males should winter nearer the breeding grounds than females,

whereas the dominance and body size hypotheses predict the

opposite.

The sex of wintering Dunlin is difficult to identify in the field,

even in the hand (Prater etal. 1977). In many studies, sex is assigned

by difference in bill length (e.g. Chao et al. 1991, Warnock et al.

1997, Shepherd et al. 2001, Shepherd 8c Fank 2004). However,

bill length overlaps to some extent among the sexes (Brennan etal.

1984, Shepherd et al. 2001, Meissner 2005) and geographically

distinct groups of Dunlin also vary in bill length (Wenink 8c Baker

1996). Instead, by amplifying fragments of sex-specific DNA

sequences, the sex of all individuals of monomorphic bird species

like Dunlin can be accurately determined (e.g. Baker 8c Piersma

1999, Fridolfsson 8c Ellegren 1999).

Differential migration of Dunlin has been reported in North

America (Shepherd et al. 2001), but their migration patterns have

not been explored in the EAAF. In this study, we present data on

the sex ratio of Dunlin wintering in central Taiwan, one of their

southernmost wintering areas (Wei et al. 2009). In addition, using

the results of molecular sexing, we derive a discriminant function

and examine its accuracy in distinguishing the sex of Dunlin in

Taiwan.

METHODS

Field methods

Our study area is Han Pao wetland, Changhua County, Taiwan

(23-95°N 1 20. 36°E). Each year, more than 10,000 Dunlin migrate

through or winter here (Hu 2005). This area is one of the largest

intertidal mudflats in Taiwan (c.38 km2). It has a long history of

shellfish and oyster farming, and shorebirds often rest and feed in

these habitats (Hu 2005).

We mist-netted Dunlin from September 2007 to May 2008,

three to four times every month, at night from evening to high tide.

Most ringing sites were on agricultural lands with some shallow

water. We defined birds netted from September to November 2007

as the autumn population, December 2007 to February 2008 as

the winter population, and March to May 2008 as the spring

population. We ringed and weighed each Dunlin captured and, for

most, measured (to the nearest mm): maximum wing length, tarsus

length, total head length and bill length. We then extracted 20- 40

yl of blood by venipuncture from the brachial vein and placed the

blood in SET buffer (0.15 M NaCl, 0.05 M Tris, 0.001 M EDTA,

pH = 8.0). We aged the Dunlin based on the presence of either

buffy-edged inner median coverts (first-years) or white-edged inner

median coverts (after first year) (Choi et al. 2010). We could

68

DZUNG-YUN YANG, CHUNGYU CHIANG & YU-CHENG HSU

Forktail 28 (2012)

identify birds as first-years before February, but coverts became

worn and buffy fringes disappeared from March.

Molecular sexing

We extracted DNA from blood using standard methods (Gemmell

& Akiyama 1996). Fragments of the chromo-helicase-DNA

binding protein (CFID) gene from the sex chromosomes were

amplified by polymerase chain reaction (PCR), using the primers

2550F and 2718R (Fridolfsson & Ellegren 1999). The reaction

volume was 20 pi and contained c.10 ng DNA, 0.8 pi dNTP

(2.5 mM of each nucleotide), 0.4 pi 2550F primer (10 mM), 0.4 pi

2718R primer (10 mM), 2.0 pi 10 x PCR buffer, 15.92 pi dH,0

and 0.4 U pi Taq DNA polymerase (Protech). The PCR included

3 minutes at 95°C, followed by 35 cycles of 30 seconds at 95°C, 40

seconds at 46°C and 50 seconds at 72°C, and a final extension

reaction at 72°C for 5 minutes. The PCR products were

electrophoresised in 1% agarose gel, using a Bio- 100™ DNA

Ladder (Protech) as size reference. We then dyed the gels in

ethidium bromide and examined them under ultra-violet light. We

identified the sex of birds by the presence or absence of the female-

specific DNA fragment of the CHD-W gene.

Statistical analysis

We used a chi-square goodness-of-fit test to determine if the sex

ratio significantly deviated from 1:1 for both adult and juvenile

Dunlin. We also compared the difference of sex ratios within each

season with the same test. Mean bill length, total head length,

maximum wing length and tarsus length were compared between

sexes using the Student’s /-test. We used three measurements

(maximum wing length, tarsus length and bill length) to derive a

discriminant function using SPSS 14.0 to assign sex.

RESULTS

From September 2007 to May 2008, we caught and sexed 748

Dunlin in the Han Pao wetland (Figure 1). Excluding recaptured

individuals, 693 birds were ringed (Table 1). The sex ratio was

significantly different from 1:1 (xr= 23.27, P < 0.001): 410

individuals were female (59.2%) and 283 individuals were male

(40.8%). We identified 138 Dunlin as juvenile (<20%), but failed

to age some individuals because the buffy-edged inner median

Table 1 . Number of Dunlin ringed from September 2007 to May 2008

at Han Pao Wetland, Taiwan.

coverts were unclear after midwinter. Among juveniles, 94 were

female (68.1%) and 44 were male (31.9%), significantly female-

biased (-),12= 18.1, P < 0.001). Combining data from all captures,

the sex ratio did not significantly deviate from 1:1 in autumn (73

females, 61 males, zl2 = 1.07, P = 0.3). However, in winter and

spring, the sex ratio was significantly female-biased (winter: 238

females, 163 males, %i2 = 14.03, P < 0.001, spring: 137 females, 76

males, xl2 = 17.47, P < 0.001 respectively).

Of Dunlin that were both sexed molecularly and measured

(n = 565 individuals), females were significantly larger than males.

However, these measurements showed overlap between the sexes

(Table 2). For discriminant analysis, we excluded total head length

in the analysis because it was not independent from bill length and

both were highly correlated (Pearson’s correlation coefficient

r =0.92, P< 0.001).

The best discriminant function included bill length (BL), tarsus

length (TL) and wing length (WL). The resulting function was:

140

Figure 1. Number of female and

male Dunlin captured from

September 2007 to May 2008 at Han

Pao Wetland, Taiwan.

Month

Forktail 28 (2012)

Sex bias in a wintering population of Dunlin Catidris alpina in central Taiwan

69

Figure 2. Distribution ofthe D values

of the discriminant function in male

and female Dunlin captured from

September 2007 to May 2008 at Han

Pao Wetland, Taiwan.

D = 0.515 x BL + 0.05 xTL + 0 .105 x WL - 33.018 (Wilks’

lambda = 0.413).

The value of D < 0 was identified as male and D > 0 was

identified as female. This function correctly identified the sex of

89.03% of Dunlin, with 92.9% of males and 86.2% of females

respectively (Figure 2). This function performed better in

identifying males than in females (%2 = 6.46, P = 0.01 1).

DISCUSSION

The wintering sites of Dunlin in East Asia range from the eastern

coast of Liaoning province, China (45°N), Japan, South Korea to

Hong Kong (22°N). Few Dunlin are found wintering farther south

than Hong Kong (Bamford et al. 2008). As our study site (Han Pao

wetland) is located at 23.95°N, we assume it is one of the southernmost

wintering sites. Numbers of Dunlin reached their peak from

December to February, indicating that most Dunlin used the Han

Pao wetland as a wintering ground, instead of as a stopover site.

Within the E AAF, at least three species of shorebirds show sex-

biased migration: Curlew Sandpiper Calidris ferruginea males

winter farther south; and Eastern Curlew Numenius

madagascariensis and Bar-tailed Godwit Limosa lapponica males

winter farther north (Nebel 2007). We found that the overall sex

ratio of Dunlin in central Taiwan is significantly female-biased. Both

the body size hypothesis and the dominance hypothesis predict that

females, which are larger and therefore presumably more dominant

than males, would winter at northern sites. Our data thus support

the arrival time hypothesis, which states that the sex that arrives

later at the breeding grounds will winter farther south (Morbey &

Ydenberg 2001). The arrival time hypothesis also predicts more

male Dunlin in the northern wintering grounds, such as Japan,

Korea or northern China. With further studies from these regions,

this hypothesis can be further tested.

In North America, Shepherd et al. (2001) found male-biased

populations of Dunlin at higher latitude wintering areas and

suggested that there is a latitudinal cline in the sex ratio ofwintering

Dunlin populations. Our data are consistent with this. Another

possibility for the female-biased sex ratio in our study site comes

from local differences in habitat preference between sexes. Shepherd

& Lank (2004) reported that male Dunlin preferred agricultural

land and females preferred mudflats on wintering grounds in British

Columbia. All Dunlin we sampled were caught on agricultural land,

but we found more females, suggesting that the findings of

Shepherd & Lank (2004) are not generally applicable. Our

sampling, however, was conducted during high tides when all

Dunlin came into agricultural lands, thus minimising such habitat-

based sex bias.

We used the results of molecular sexing to develop a

discriminant function to identify Dunlin sex by morphological

measurements. For sex-size dimorphic species, discriminant function

analysis of morphometric data is a common technique to sex birds

that are monomorphic in plumage (Brennan et al. 1984, Iko et al.

2004). Previous discriminant function and maximum likelihood

function analyses have correctly identified the sex of over 90% of

non-breeding Dunlin (Brennan et al. 1984, Shepherd et al. 2001,

Meissner 2005), but were all conducted on a single subspecies. In

Taiwan, wintering Dunlin come from several geographically distinct

subspecies (Lanctot et al. 2009), which differ in body size (Wenink

& Baker 1996, Bamford et al. 2008). The mix of subspecies in our

study site may result in the lower discriminant power of the function,

or might be a result of differing morphological measurements among

ringers in our study (Barrett et al. 1989).

CONCLUSION

We found a significant female -biased wintering Dunlin population

in central Taiwan. This is the first study to reveal a differential sex

ratio in this species in the EAAF. East Asia has one of the highest

human population densities in the world and the fastest population

and economic growth, especially in coastal areas (Liu 2009).

Shorebird habitats are therefore being rapidly altered (Ge et al.

2007). The loss of Dunlin habitats used heavily by either sex could

result in disproportionate population declines (Steifetten & Dale

2006). Identification of sites that are preferred by members of one

sex is crucial for conservation (Nebel 2007). Our data alone are

not sufficient to test hypotheses for the differential migration of

Dunlin in this region. More long-term data from different latitudes

in the EAAF, such as eastern China, Korea, Japan and other parts

of Taiwan, will help the understanding of the migration ecology of

this species.

70

DZUNG-YUN YANG, CHUNGYU CHIANG & YU-CHENG HSU

Forktail 28 (2012)

ACKNOWLEDGEMENTS

We thank J. Pilgrim, N. Warnock and K. Mathot for their insightful comments

on the manuscript, Taiwan Bird Banding Center for providing bird rings and

the Changhua County Government for issuing a ringing permit. The ringing

work could not have succeeded without the involvement of the wader-catching

crew of the Taiwan Wader Study Group and the students of Department of

Environmental Science and Engineering, Tunghai University. We also thank

many volunteers for their hard work with us at night.

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Dzung-Yan YANG, Institute of Natural Resources, National

Dong Hwa University, No. I, Sec. 2, Da-Hsueh Rd. Shou-Feng,

Hualien, 97401, Taiwan. Email: dzungyun@yahoo.com.tw

Chungyu CHIANG, Department of Life Science, Tunghai

University, Taichung, 40704, Taiwan. Email:

dec.chiang@gmail.com

Yu-Cheng HSU, Department of Natural Resources and

Environmental Studies, National Dong Hwa University, No. 1,

Sec. 2, Da-Hsueh Rd. Shou-Feng, Hualien, 97401, Taiwan. Email:

ycsheu@maiindhu.edu.tw (corresponding author)

FORKTAIL 28 (2012): 71-76

Habitat associations of the Manchurian Reed Warbler

Acrocephalus tangorum wintering on the Tonle Sap

floodplain and an evaluation of its conservation status

JEREMY P. BIRD, BERRY MULLIGAN, ROURS VANN, PHILIP D. ROUND & JAMES J. GILROY

Sixty tape playback trials and 17 net rides were used to investigate the habitat associations of Manchurian Reed Warbler Acrocephalus

tangorum at what is potentially its most significant wintering site, the Tonle Sap floodplain in Cambodia. Fieldwork in March 2006 concentrated

at three sites within the inundation zone during the dry season when floodwaters are at their lowest. This corresponds with the non-breeding

season, when Palearctic migrant Acrocephalus warblers visit. We analysed cover of different habitat types at two scales: close to and broadly

surrounding playback locations. Detections within broad habitat types differed significantly from random (c2 = 32.8, d.f = 5, P <0.001 ) with

an apparent bias towards grass habitats represented in the study area by tall (> 1 .0 m) grassland. A principal component analysis (PCA) of

the proportionate abundance of different habitats within 10 m of playback locations generated just two PCA axes, correlating strongly with

the abundance of grassland and wet habitat features (PCA1 ) and woodland and scrub (PCA2). Logistic regression with both axes as predictor

variables revealed a significant effect of PCA1 (z = -2.566, P = 0.010), but no significant effect of PCA2 (z = 0.088, P = 0.419). All sites with

detections had a low loading on PCA1 , suggesting a strong association with grasslands. Capture rates were extremely low compared with

one wintering location in Thailand, so while our study suggests theTonle Sap is of global importance for the species, we cannot find sufficient

evidence to warrant revising the species's IUCN Red List status from Vulnerable to a lower category of threat.

INTRODUCTION

The Manchurian Reed Warbler Acrocephalus tangorum is a

migratory passerine listed as Vulnerable on the IUCN Red List

(BirdLife International 2012) because it is inferred to have a small

and declining population. The species is only known to breed at a

few sites in extreme south-east Russia and north-east China. It is

considered rare or threatened in north-east Asia by Gluschenko

(1989) and of the three sites in the Chinese breeding range

(excluding Inner Mongolia), Alstrom et al. (1991) reported that

two did not appear to have much suitable habitat remaining and

they ‘doubt that significant numbers of tangorum remain’, but it

remained ‘locally common’ at Zhalong.

The species’s status in, and even the extent of, its wintering range

remains particularly poorly understood. The wintering grounds

were unknown until a population was found in Phragmites

reedswamp atKhaoSam RoiYot, Thailand, in 1981 (Round 1993,

Round & Rumsey 2003). Subsequently, it has been found wintering

in sedge beds within deciduous dipterocarp forest at a number of

sites: Champasak province, southern Lao PDR (Round 1998),

Preah Vihear province, northern Cambodia (Clements et al. 2005)

and southern Mondulkiri province, eastern Cambodia (Bird et al.

2007). In 2000 ‘at least 20’ Manchurian Reed Warblers were

recorded at Krous Kraoum, KompongThom province, on the Tonle

Sap floodplain in Cambodia (Robson 2000, Davidson 2001), and

more records have since followed in Kompong Thom and Siem

Reap provinces, Cambodia. These observations originated from a

variety of habitats, particularly tall grass stands (away from water),

sedge beds (both wet and dry), scrub-fringed lotus swamps, and

heterogeneous scrub/grass mixes away from water (BirdLife

International 2012).

To date, the only study examining winter habitat use by the

species was conducted within the least disturbed part of a c.50 km2

marsh in Khao Sam Roi Yot National Park, Thailand (Round &

Rumsey 2003). This demonstrated a strong positive association of

Manchurian Reed Warbler with stands of mature Phragmites.

Because Phragmites is rare elsewhere in Thailand, and because a

moderate increase in mist-netting in South-East Asia and an

improved knowledge of the species’s identification failed (at least

before 1997) to produce many records, the Red Data Book asserts

that Manchurian Reed Warbler should be considered truly scarce

and locally distributed with a small global population (BirdLife

International 2001). However, more recent records at wintering; sites

away from Khao Sam Roi Yot demonstrate that other wetland

vegetation is widely used in the absence o l Phragmites. and it remains

undetermined whether this is truly suboptimal habitat (Round &

Rumsey 2003). The extent of potentially suitable habitat in the

Tonle Sap Lake inundation zone, South-East Asia’s largest freshwater

wetland, is vast (P. Davidson in litt .) and therefore, even if

population densities in floodplain habitats other than Phragmites

should prove to be lower, Manchurian Reed Warbler may be more

widespread and numerous than previously considered.

At Khao Sam Roi Yot, the best documented (and formerly only

known) wintering site for the species, the extent of Phragmites is

declining owing to illegal encroachment with plantations of

casuarinas, eucalyptus and coconut palms established around the

margins of the area, while major prawn farms have been established

in the centre, pumping salt or brackish water into previously

freshwater areas (BirdLife International 2001). There is now very

little Phragmites habitat in Thailand and wetlands in the country

are imminently threatened by reclamation and urbanisation

(Round 1993). The Tonle Sap Lake floodplain supports large areas

of natural and semi-natural seasonally inundated grassland (Evans

et al. 2005) that are currently under intense pressure from land-

use change, with grassland cover declining since the late 1990s

owing to scrub encroachment and, since 2004, an extraordinary

intensification of rice cultivation (Gray 2006, Gray et al. 2007).

Gray et al. (2009) documented losses of 27% of grassland cover on

the Tonle Sap floodplain within 10 grassland blocks between

January 2005 and March 2007, primarily as a result of land

conversion for intensive dry season rice cultivation. Bengal Florican

Houbaropsis bengalensis was uplisted to Critically Endangered in

2007 as a direct result of these land-use changes in the Tonle Sap

floodplain (BirdLife International 2012). The impacts of land-use

change on Manchurian Reed Warbler remain unknown but, based

on the above, a population decline has been inferred in assessing

its current IUCN Red List status (BirdLife International 2012).

We used tape playback and mist-netting to investigate the

habitat associations of Manchurian Reed Warbler on the Tonle Sap

floodplain. Our results are discussed in the context of the species’s

likely abundance and trends, in order to review its assessed

extinction risk.

72

JEREMY P. BIRD ero/.

Forktail 28 (2012)

METHODS

Study area

The Tonle Sap Lake in Cambodia is the largest freshwater lake in

South-East Asia (Gray et al. 2009). The lake floodplain system is

characterised by notable seasonal differences in water level and

surface area (Kummu & Sarkkula 2008 in Vastila et al. 2010). The

lake expands from some 2,500 km2 in the dry season (December to

June) to between 12,500 km2 and 15,000 km2 during the wet season

(Campbell et al. 2006). The floodplain landscape is a complex

mosaic of flooded forests, wet and dry scrub, cropped grasslands,

tall grasslands structurally similar to Phmgmites reedbeds, and areas

of rice cultivation radiating out from the lake in roughly concentric

rings. During March 2006 we conducted surveys on the northern

boundary of the Tonle Sap Lake inundation zone around the Siem

Reap and Kompong Thom provincial border (Figure 1 ) in natural

and semi-natural habitats and areas of dry season rice production

(headponds for irrigation and rice paddy). The study area

experiences annual flooding up to 4 m depth for 4-7 months a year

(Gray 2007). Three relatively distinct sites were studied during the

dry season (Figure 1): (1) an area ofseasonally inundated grassland

and short tree scrubland near Stoung (dry at the time of field visits),

(2) natural and semi-natural seasonally inundated grassland at

Kruos Kraom (fields were dry at the time of fieldwork but

headponds remain inundated year-round for irrigation) and (3)

permanently flooded forest and seasonally flooded bankside scrub

at Boeung Tonle Chhmar, one of the core areas of the Tonle Sap

Biosphere Reserve.

Tape playback

Manchurian Reed Warbler has previously been shown to respond

to playback (Alstrom et al. 1991 ). To investigate habitat preferences

of the species, relative abundance was compared in seven broad

habitat types using 60 separate playback trials. A series of random

playback positions was generated and a representative sample

chosen for each habitat type. At each playback position, five minutes

of continuous vocalisations by Manchurian Reed Warbler was

played on a portable Sony CD player through a pair of Philips

SBA220 speakers at equivalent volume to reed warbler vocalisations

(judged by ear). The vocalisations included song and calls

alternating for a minute each for the total five minutes. During the

following ten minutes, two observers standing back-to-back

recorded all Manchurian Reed Warblers seen and heard from the

position of the speakers. An estimation of the total number of

individuals was made.

Parameters recorded at each playback position were site number,

date, time, cloud cover and precipitation. Broad habitat type was

subjectively assigned by eye to dry scrub (which is inundated by

shallow water during the wet season), permanently flooded forest,

wet/flooded scrub (potentially submerged in the wet season),

scrub-island within paddy (inundated in the wet season), grass-

island within paddy (inundated within the wet season), tall dry

grass (>1.0 m average height, inundated in the wet season), or tall

wet / flooded grass ( > 1.0 m average height, artificial headponds and

natural). Percentage cover within a 10 m radius (measured with a

measuring tape) of playback positions of sedge, floating vegetation,

cultivated rice, Mimosa sp., tall grass (>1.0 m), scrub, flooded forest,

water and mud was estimated by eye. A radius of 10 m was selected

by default: no comparable studies were found to indicate the

attractant radius of playback on individual reed warblers; to

maximise the number of playbacks completed, a manageable radius

was selected; and because isolated habitat islands were part of the

trial and these pockets were small, a 10 m radius was optimal.

Surveys were carried out from 06h00 to lOhOO and again from

14h00 to 18h00, periods of maximum bird activity, between 4 and

24 March 2006.

Using R version 2.13 (R Core Development Team 2011), we

applied principal component analysis (PCA) to data on the

proportionate cover of habitat features within 10 m of each

playback location in order to isolate key components of habitat

variation. Using ordination plots classified according to the presence

or absence of Manchurian Reed Warbler detections, we then

performed a visual assessment of the relationship between habitat

variation and warbler detection. We used logistic regressions with

presence/absence as a response variable to test for significant

relationships with extracted values from habitat principal

Kampong Svay

Stueng Saen

lantuk

Prasat Balangk

Siem Reap

Province

Chi Kraeng

Stoung

Kampong Thom

Province

TONLE SAP LAKE

V S'

r \ Preah/Mhear

i Scale

- Kampong Chhnang

Province

Prasat Sambour

Figure 1. Study sites where

playback was performed around

the Tonle Sap, Cambodia.

Forktail 28 (2012)

Manchurian Reed Warbler Acrocephalus tangorum wintering on the Tonle Sap floodplain

73

components, as well as overall habitat classes. We also compared

detection rates across the broad habitat types at playback sessions

using a Chi-square test.

Mist-netting

Birds were caught during 14 mornings of netting and ringing

sessions on 17 separate net rides, each of which was classified under

a broad habitat type. Only where there was relatively extensive

homogeneous habitat of the same type were net rides established.

Nets were operated for up to four hours, mostly between 06h00

and lOhOO. All birds caught were identified, ringed, measured and

examined for moult and feather wear. Birds were weighed to the

nearest O.lg using Pesola spring balances before being released.

Capture rates allow comparison of relative abundance of

Manchurian Reed Warbler between sites and with other species.

The number of Manchurian Reed Warbler caught per metre-hour

(mh) of mist-netting was calculated for different habitat types.

Owing to the difficulties of mist-netting in flooded scrub and

forest, the survey of Manchurian Reed Warbler was limited to

playback in the maze of channels and streams linking Boeung

Chhmar and the Tonle Sap Lake.

RESULTS

An estimated 35 Manchurian Reed Warblers were recorded (based

on both calls and direct sightings) during 60 playback sessions.

Overall densities of Manchurian Reed Warbler appeared to be

relatively low, with detections being made at just 16 playback

locations (27% of total). Detection rate across different habitat

types was significantly different from random (Table 1, c2 = 32.8,

d.f=5, P<0.001), with most detections occurring in grass habitats

(Table l). Tall grassland habitats, both wet and dry, appeared to be

the main vegetation types supporting Manchurian Reed Warblers

within the study area. Multiple encounters also occurred in man¬

made habitats, mostly in tall grasses surrounding artificial wet

headponds (used for dry season rice irrigation — an estimated 12

individuals encountered in 6 playbacks), as well as from small

‘islands’ of tall grass habitat in and along the edges of rice fields (an

estimated 7 individuals encountered in 10 playbacks). Detection

rates in forest and scrub habitats, both flooded and dry, were very

low, and no detections were made in extensive areas of permanently

inundated scrub and flooded forest in the Boeung Chhmar/Moat

Khla IBA. However, logistic regression with habitat type as an

explanatory variable revealed no significant differences between

habitat types, potentially owing to the small sample sizes in each

habitat.

Table 1 . Broad habitat classifications of playback locations, together

with the numbers of Manchurian Reed Warblers recorded in each

habitat type and numbers that would be expected if habitat selection

was random. The distribution of records was significantly different from

Table 2. Factor loadings for each variable in a principal component

analysis of habitat variation across playback locations, showing the

first two principal components (proportionate eigen values: PCA1

69.2%, PCA2 23.7%).

Fig ure 2. Ordination plot showing playback locations, classified

according to the presence or absence of Manchurian Reed Warbler

in relation to principal component axes explaining variation in

habitat characteristics. Axes PCA1 and PCA2, collectively, explain

93% of variation in the habitat characteristics shown. NB 'snaow' is

Sesbania spp.

PC A of the proportionate abundance of different habitats

within 10 m of playback locations revealed that habitat variation

was rather simple (Table 2), with 93% ofvariation being explained

by just two PCA axes, correlating strongly with the abundance of

grassland and wet habitat features (PCA1) andwoodland andscrub

(PCA2). Logistic regression with both PCA1 and PCA2 as

predictor variables revealed a significant effect of PCA1 (z =

-2.566, P = 0.010), but no significant effect of PCA2 (z = 0.088,

P = 0.419). Figure 2 shows an ordination plot of playback sample

locations against both PCA axes, with sites classified according to

the presence or absence of Manchurian Reed Warblers. All sites at

which the species was recorded have a low loading on PCA1,

suggesting a strong association with locations dominated by

grasslands. Manchurian Reed Warblers were absent from all

locations which received a high loading for PCA1, suggesting

avoidance of habitats associated with wet features such as floating

vegetation, open water, mud and sedges. Figure 2 also shows that

locations with a high or low loading on PCA2 did not produce

detections, suggesting avoidance of wooded or scrub-rich sites. This

analysis underlines the importance of tall grassland for Manchurian

Reed Warblers within the study area. Grasslands, structurally

similar in height and density to Phragmites reedbed, also tended to

be the dominant habitat at playback locations at which detection

rates were high. Tall grass comprised 76% habitat cover within

10 m of playback locations where Manchurian Reed Warbler was

detected versus just 26% at locations where none was detected.

The sample of Manchurian Reed Warblers from mist-netting

was too small to test statistically, but their distribution was heavily

74

JEREMY P. BIRD era/.

Forktail 28 (2012)

Table 3. Habitat types and capture rate for Manchurian Reed Warblers

(MRW) netted in Kompong Thom and Siem Reap provinces.

*Data from captures at Khao Sam Roi Yot, Thailand, from Round & Rumsey (2003)

skewed towards grassland. All eight of the net rides where

individuals were caught were dominated by grass (Table 3). Capture

rates (n/ mh netted) for the species were very low in comparison to

those recorded by Round & Rumsey (2003) in pure Phragmites

karka stands at Khao Sam Roi Yot (Table 3). During 6,643 mh of

mist-netting, captures of all Acrocephalus warblers involved 9

Manchurian Reed Warbler, 73 Black-browed Reed Warbler A.

bistrigiceps, 3 Blunt-winged Warbler A. concinens , 118 Oriental

Reed Warbler A. orientalis and 1 Thick-billed Warbler A. aedon.

DISCUSSION

Habitat association and abundance

A strong positive association of Manchurian Reed Warbler with

grassland was demonstrated using playback. The species has been

observed in a wide variety of wetland habitats in Cambodia, but

within the Tonle Sap inundation zone areas of tall grass (possibly

a Saccharum sp.), similar in appearance to Phragmites reedbed,

appear to be strongly favoured over alternative habitats. It remains

unclear whether the species is wintering in suboptimal habitat on

the Tonle Sap floodplain, as no Phragmites reedswamp could be

located for direct comparison with earlier studies (e.g. Round &

Rumsey 2003).

Densities of Manchurian Reed Warbler on the floodplain were

low relative to other Acrocephalus species and locations. Despite

6,643 mh mist-netting effort, just nine individuals were captured

at a rate of 0.0013 individuals per metre hour, whereas Round &

Rumsey (2003) recorded a capture rate of 0.022 in stands of

mature Phragmites at Khao Sam Roi Yot, capturing 60 different

individuals during April 1995. The high levels at Khao Sam Roi

Yot might partly be explained by influxes of migrant individuals

and a concentration effect in limited habitat area, but this is

conjecture.

Inundated grasslands within the Tonle Sap floodplain are of

exceptional importance for conservation (Gray et al. 2009). It has

been suggested that Manchurian Reed Warbler might have a large

wintering population in Cambodia given the extensive available

habitat in the Tonle Sap floodplain (Round & Rumsey 2003, P.

Davidson pers. comm. 2005). Our findings support this assertion:

although densities throughout the floodplain are likely to be low

given the habitat associations and densities we observed, the available

area of tall grass stands remain vast (but needs quantifying).

Furthermore, with a spate of new wintering records in the late 1990s

and 2000s redefining the species’s non-breeding range to include

much of Cambodia to southern Lao PDR and the Vietnamese

border, the availability of tall grass within this range is unlikely to

be a limiting factor considering its prevalence in human modified

landscapes on the floodplain. Although Manchurian Reed Warbler

is now known to be more widespread in the non-breeding season,

the Tonle Sap lies at the core of its wintering range, and as South-

East Asia’s largest permanent freshwater wetland it probably

represents a stronghold for the species. Continuing destruction of

wetland habitats in Thailand may further increase the global

significance of this population (BirdLife International 2001, Round

& Rumsey 2003).

Potential threats

Alluvial grasslands have been cleared preferentially for agriculture

in most parts of East and South-East Asia, so the large areas

remaining around the Tonle Sap Lake are of outstanding regional

importance (Evans et al. 2005). Drainage and conversion of natural

wetlands and grasslands, primarily for rice agriculture, have resulted

in a significant decline in the area of available habitat for

conservation priority species in Cambodia (Seng et al. 2003) and

expansion of dry season rice is probably the most serious threat to

the biodiversity of the Tonle Sap grasslands (Evans et al. 2005). It

is possible, however, that Manchurian Reed Warbler has been less

heavily impacted than other species. The major land-use changes

occurring around the floodplain, particularly the shift from

traditional wet season rice cultivation to dry season rice cultivation

(Chan et al. 2004, Evans et al. 2005), may influence habitat

availability for the winteringpopulation. Manchurian Reed Warbler

appears to persist around headponds created for dry season rice

irrigation, as well as in small blocks of tall grass within paddyfields

themselves. This implies that the influence of these land-use changes

will not necessarily be negative. It is also likely that traditional

management systems are beneficial to the species, as regular grazing

and burning help maintain grasslands (both tall and short swards)

by preventing successional reversion to scrub and forest (BirdLife

International 200 6).

Despite the availability of apparently suitable habitat within

Cambodia, it is important to emphasise that population densities

are likely to be low in most areas. The continuing degradation and

inadequate habitat protection at Khao Sam Roi Yot, the only other

relatively large area of known habitat to support this species in

South-East Asia, highlights the need for conservationists to ensure

the maintenance of suitable habitat for this species in Cambodia.

Management measures such as retaining tall grass margins may be

relatively simple and cost-effective to implement, and should be

considered in mid- to long-term management objectives for

inundated grassland areas around the floodplain.

Another potential threat to the species, not reported in the

IUCN Red List account, is trapping. Trapping is widespread in

South-East Asia for food and for merit release (the practice of

releasing caged wild-caught birds for religious purposes). Trapping

has been implicated in rapid declines in Yellow-breasted Bunting

Emberiza aureola populations: roosting flocks in reedbeds are

disturbed and then caught in mistnets, and are cooked and sold as

‘sparrows’ or ‘rice-birds’ (BirdLife International 2012). This practice

has now become widespread and, although it is illegal in some areas,

over a million individuals are reportedly killed annually to be sold

as snacks (BirdLife International 2012). While Manchurian Reed

Warbler is unlikely to be a focus of trapping effort it may well be

caught as a bycatch species. We observed passerine trapping while

conducting fieldwork. Merit release is now also widespread at

tourist sites. JPB observed a tail-less individual of either^.

bistrigiceps or A. tangorum in central Phnom Penh that was

presumably a merit-released bird.

Red List implications

The IUCN Red List criterion that has been triggered to qualify this

species as Vulnerable, C2a(ii), states that the population is estimated

to number fewer than 10,000 mature individuals and is undergoing

a continuing decline, with all mature individuals in one

subpopulation (IUCN 2001). The population estimate for

Manchurian Reed Warbler of 2,500-9,999 mature individuals

Forktail 28 (2012)

Manchurian Reed Warbler Acrocephalus tangorum wintering on theTonle Sap floodplain

75

(BirdLife International 2012) is derived from the original Red Data

Book assessment that it ‘must be presumed to have a fairly small

world population’ owing to the scarcity of wintering records and

its restricted wintering range (BirdLife International 2001). We

recommend revising the text that accompanies the Red List account

to reflect that the wintering range is now known to be much larger

than when the Red Data Book account was compiled. Based on

our findings it seems unlikely that wintering habitat is population-

limiting and it should not be used as the basis for the current

population estimate.

However, it would be premature to revise the population

estimate upwards and downlist the species to Near Threatened or

Least Concern as a consequence, for two reasons. First, although

the species has a broad non-breeding range we cannot definitively

say its population is larger than the current estimate. It is possibly

noteworthy that increased mist-netting in Phragmites in central

Thailand (Bung Boraphet), during which several thousand

Acrocephalus warblers were ringed in 2007-2012, yielded fewer

than ten Manchurian Reed Warblers (PDR unpubl. data), and very

few have been caught on migration in Hong Kong, suggesting the

global population is indeed relatively small. Second, the Red Data

Book provides sufficient justification for a cautious approach:

various problems exist at the three best known breeding grounds

of the species, Khanka Lake in Russia and Zhalong and Dalai Hu

National Nature Reserves in China, while the two other reported

breeding sites in China (excluding Inner Mongolia) are regarded

as no longer suitable for the species (Alstrom et al. 1991, BirdLife

International 2001). We are unaware whether updated

information exists from these sites in the Russian or Chinese

literature. If it does, it could provide valuable insights into the

conservation status of the species. If not, gathering this information

should be a priority for further research. It is also clear from the

Red Data Book account that assessing the species’s status in Inner

Mongolia is important.

After small population size, the second assertion of the current

IUCN Red List assessment is that the species’s population is

declining. Despite conversion and degradation of habitat in some

wintering sites (notably Khao Sam Roi Yot), the Manchurian Reed

Warblers ability to exploit a variety of habitats throughout its non¬

breeding range suggests that habitat conversion may not be causing

declines on the wintering grounds, and this should be reflected in

the Red List text account. However, again we feel it is premature

to recommend changing the Red List status owing to uncertainties

on the breeding grounds, where habitat conversion may well be

causing the population to decline.

While the population consequences of documented threats

remain unquantified, there is insufficient evidence to infer a rapid

population decline (>30% over 10 years or three generations) that

could qualify the species as Vulnerable under the A criterion of the

IUCN Red List (IUCN 2001). Furthermore, now that the

wintering range has been increased substantially through improved

knowledge, the species’s estimated extent of occurrence, 490,000

km2 (BirdLife International 2012), is much larger than the

threshold triggering the B criterion. Overall we recommend that

the status quo, listing the species as Vulnerable under criterion

C2a(ii), be maintained.

CONCLUSION

This study is the first to investigate the wintering habitat

associations of Manchurian Reed Warbler in Cambodia since

records revealed the country encompasses the majority of its known

non-breeding range. While the findings deserve further scrutiny

and ideally more prolonged research, they go some way towards

outlining Cambodia’s importance for this threatened species.

Preliminary evidence suggests Cambodia, and the Tonle Sap Lake

floodplain in particular, probably supports a substantial proportion

of the global population. Establishing more reliably what that

population is likely to be requires considerably more study, and is

probably best achieved by concentrating on the breeding sites.

Currently there is little evidence that the species is unduly

threatened by habitat conversion in Cambodia but we must stress

that this conclusion requires confirmation. Additionally, the

unregulated trapping of small birds (including reed warblers)

through the use of nets is widespread within the species’s non¬

breeding range and its impacts at the population level warrant

further investigation.

ACKNOWLEDGEMENTS

We thank Pete Davidson for helpful advice and information before

undertaking fieldwork, Tom Evans and colleagues at the Wildlife

Conservation Society in Cambodia for providing essential help and support

throughout, and the many donors for sponsoring the research: the Astor of

Hever Trust, British Ecological Society, British Ornithologists’ Union, Chris

and Liz Shepley, IdeaWild, Opticron, Oriental Bird Club, People’s Trust for

Endangered Species, Robert and Elaine Dawson, Royal Geographical Society,

Sevenoaks RSPB members’ Group, Sir Phillip Reckitt Educational Trust,

University of East Anglia Travel and Expeditions Committee, The Wetland

Trust and Wexas International. Also, thanks to Santhosh Kurian for his

assistance with the map. Finally we thank two anonymous referees for helpful

comments that greatly improved the manuscript.

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FORKTAIL 28 (201 2): 77-92

The avifauna of Alor and Pantar, Lesser Sundas, Indonesia

COLIN R.TRAINOR, PHILIPPE VERBELEN & RON E. JOHNSTONE

The status of birds on Alor and Pantar is reviewed based on the literature plus surveys in 1991, 2002, 2009 and 2010 totalling about five

weeks and two weeks respectively. A total of 116 birds (86 resident landbirds) were recorded on Alor including 37 new island records

(including 27 resident landbirds) and 97 birds were recorded on Pantar including 75 new island records (including 55 resident landbirds).

The most significant records were the discoveries on Alor of a population of Timor Bush Warbler Locustella timoriensis which may represent

an undescribed subspecies, an undescribed species or subspecies of honeyeater resembling Crimson Myzomela Myzomela kuehnii, and an

undescribed subspecies of Timor Stubtail Urosphena subulata. The endemic subspecies of Southern Boobook Ninox boobook plesseni collected

historically from Alor was found to be vocally distinct and may be a distinct species; it was recorded on both Alor and (a new population)

Pantar. Vocalisations of a presumed scops owl Otus on Pantar are distinct from Moluccan Scops Owl Otus magicus and may also be an

undescribed taxon. Alor appears to have an important population of the Critically Endangered Flores Hawk Eagle Nisaetus floris but

populations of the heavily traded Critically Endangered Yellow-crested Cockatoo Cacatua sulphured have declined greatly since 1991. The

Vulnerable Flores Green Pigeon Treron floris was recorded regularly on Alor, which may be a stronghold. Other notable records are significant

range extensions of Mees's Nightjar Caprimulgus meesi to both islands, new island records for Little Cuckoo Dove Macropygia orientalis and

Ruddy Cuckoo Dove M. emiliana, and improved knowledge of the montane avifauna of Alor including the first records of Mountain White-

eye Zosterops montanus. Our fieldwork shows that the avifauna of these islands contains more globally threatened species than previously

thought, and a stronger representation of birds originating from Timor. Alor and Pantar are therefore of greater global conservation

significance than previously highlighted.

INTRODUCTION

The Flores island chain spans more than 1,000 km from Lombok

in the west through to Alor in the east: these islands delineate the

Northern Nusa Tenggara Endemic Bird Area (EBA) (Stattersfield

et al. 1998). Most islands in this chain would have coalesced

(joined) or have been briefly isolated during ice-ages and associated

sea-level changes during the Pleistocene period (10,000 yrs BP and

earlier) (Voris 2000). This global biological hotspot hosts at least

29 restricted-range bird species, including 17 endemic to the EBA

(Stattersfield et al. 1998). These numbers are even greater when

the recently described Mees’s Nightjar Caprimulgus meesi (Sangster

& Rozendaal 2004) and several taxonomic splits including the

Critically Endangered Flores Fdawk Eagle Nisaetus floris (Gjershaug

et al. 2004) are included. Most of the avian endemics of the Flores

island chain are found on the large islands of Flores (at least four

island endemic birds and 24 restricted-range birds; 13,540 km2)

and Sumbawa (22 restricted-range birds; 15,448 km2), and

consequently most historical and modern-day fieldwork has

concentrated on these islands (White & Bruce 1986, Butchart

et al. 1996, Johnstone et al. 1996, Pilgrim et al. 2000, Hutchinson

etal. 2007).

Despite recent efforts (Trainor 2002a, 2003, Schellekens et al.

2011), most other islands in the group have been neglected by

ornithologists, with little published on the avifauna of Pantar

(728 km2) since the nineteenth century (Hartert 1898,

Johnstone 1994). Alor Island (2,800 km2, 1,825 m) is rhe fourth

largest island in the EBA but has been visited rarely, and

consequently its avifauna has also remained poorly known.

A. H. Everert visited both islands in March-April 1897,

collecting 64 bird species on Alor and 22 species on Pantar (Hartert

1898). The collecting on Alor ‘cannot be regarded as at all an

exhaustive one’ because a lack of water and food hampered the visit,

causing Everett to become feverish and then injure a leg which

required hospitalisation on Sulawesi (Hartert 1898). The collection

was enough to determine that Alor has a ‘Flores ornis’ (Hartert

1898) although a few species typical of Timor Island, such as Olive¬

headed Lorikeet Tricboglossus euteles , Bar-necked Cuckoo Dove

Macropygia magna and the Alor-endemic subspecies of Southern

Boobook Ninox boobook plesseni were collected on Alor, with the

latter taxon being described as late as 1929 (Stresemann 1929,

White & Bruce 1986). V. von Plessen visited Alor in July- August

1927, collecting 68 bird specimens of 31 bird species (Rensch

1929).

A major study ( 1987- 1 993) of the avifauna of the Lesser Sundas

and the islands of Maluku by the Western Australian Museum

(WAM) and the Indonesian Institute of Sciences (LIPI) included

a visit to Pantar and Alor in April-May 1991. This resulted in new

information on the shorebirds and seabirds of Pantar and Alor

(Johnstone 1994), the discovery and description of a new form of

Sunda Bush Warbler Horornis [ Cettia\ vulcanius kolichisi

(Johnstone & Darnell 1997a) on Alor, and a redescription of the

Alor subspecies of Southern Boobook (Johnstone & Darnell

1997b). Substantial additional information on the landbirds of Alor

and Pantar, including many new island records, has remained

unpublished. The first published record of a montane bird on Alor

(Lesser Shortwing Bracbypteryx leucophrys) by Holmes (1995) was

notable. Recent visitors to Alor include limited ornithological

observations by Mason (1991, 1993) and Holmes (1995), and a

status survey of rhe Critically Endangered Yellow-crested Cockatoo

(Setiawan et al. 2000).

Following a failed attempt to reach Wetar Island by CRT, birds

on Alor were surveyed at several locations over three weeks in April-

May 2002, with an account of the birds at one location in the

vicinity of Mt Koya Koya and Tanglapui Timur published (Trainor

2005a). A visit to Alor by PV and Veerle Dossche (VD) in June-

July 2009, and to Pantar and Alor in September 2009 (PV), resulted

in the discovery of a population of Timor Bush Warbler Locustella

timorensis on Alor (Trainor et al. 2012), prompting further

fieldwork by CRT on Alor and Pantar in December 2009 and

January 2010.

The aim of this paper is to document the new general

ornithological observations of REJ, PV and CRT, particularly

significant new island records, and records of globally threatened

and restricted-range bird species. Many of the supposed new island

records mentioned in Holmes (1995) and Trainor (2005a) had

actually been recorded during the WAM-LIPI expeditions, so we

take the opportunity to correct the record. Recent discoveries by

PV also prompted brief visits to Alor and Pantar by Peter Collaerts

(PC) and a BirdTour Asia tour led by James Eaton (JE) and Rob

Hutchinson (RH), with significant records from these

ornithologists also noted in this paper.

78

COLIN R. TRAINOR, PHILIPPE VERBELEN & RON E. JOHNSTONE

Forktail 28 (2012)

METHODS

We independently visited seven sites on Alor over about 40 field

days, including visits to three islets (Kepa, Ternate and Sika off

Alor), and five sites on Pantar Island over about 14 days. Details of

the sites, survey dates and habitat are provided in Table 1 and Figure

1 . The main sites were in the far east near Tanglapui Timur and Mt

Koya Koya, and above Apui (near Subo village and Manmas

hamlet): both these locations were surveyed over a total of about

14 field days.

To support bird identifications, and descriptions, CRT and PV

took photographs using respectively a Canon 7D and a Canon 40D

digital camera with a 100-400 mm lens. Sound recordings were made

with an Olympus LS-10 recorder (CRT) and an Edirol R09 HR,

and Olympus WS 331 M digital recorder and a Sony Minidisk HI-

MD MZ-RH 1 (PV) — all in combination with an ME-66 Sennheiser

directional microphone, and playback was done with a RadioShack

mini amplilier-speaker. A set of recordings will be uploaded to web

databases such as xeno -canto and the Avian Vocalization Centre

(AVoCet). REJ visited three sites on Alor and two sites on Pantar as

part of the WAM-LIPI survey of the Nusa Tenggara islands. Birds

were collected by trapping with mistnets and general observations.

Specimens are housed at the WAM and LIPI.

Taxonomy and nomenclature follows Gill & Donsker (2012)

except that we use Common Golden Whistler Pachycephala

pectoralis instead of Rusty-breasted Whistler P. fulvotincta and

Brown Honeyeater Lichmera indistincta instead of Indonesian

Honeyeater L. limbata.

ANNOTATED LIST

We recorded a total of 116 bird species on Alor including 37 new

island records (27 resident landbirds) and a total of 97 bird species

on Pantar including75 new island records (55 resident landbirds).

Only eight bird species, including one Australian migrant and

several visiting waterbirds, appear to have not been recorded on

Alor since the historical collections (Australasian Grebe

Tachybaptus novaehollandiae , Pacific Baza Aviceda subcristata. Red-

backed Buttonquail Turnix maculosa , White-headed Stilt

Himantopus leucocephalus, Swinhoe’s Snipe Gallinago megala,

Channel-billed Cuckoo Scytbrops novaehollandiae , Savanna

Nightjar Caprimulgus affinis and Thick-billed Flowerpecker

Dicaeum agile). On Pantar, only two species recorded historically

(Australian Hobby Falco longipennis and Sunda Cuckoo Cuculus

lepidus ) went unrecorded in the current surveys. The following

annotated list focuses on significant resident landbirds, but includes

information on a few migrants. Many of the numerous new island

records are unsurprising and these are denoted clearly, with the

tabulated summary of all records in the appendix.

Table 1 . Study site details, survey dates and habitat. Refer also to Figure 1 .

Apui-Subo-Manmas/Alor Selatan (700-1,250) 8°17'39"S 124°43'17"E

Tanglapui Timur (including Kunggwera, Lipa,

Kobra, Mt Koya Koya/Alor Timur (400-1,200)

8°14'56"S 124°59'30"E

22-25 April 1991 (REJ); 1 July 2009 (PV &VD),

31 August 2009 & 9 September 2009 (PV),

4-6 December 2009 (CRT), 14 January 2010 (CRT)

2-7 & 1 1-16 May 2002 (CRT);21-27 June 2009 (PV & VD),

16— 18 January 2010

Mixed tree crops; and regrowth forest; E urophylla woodland

Regrowth forest; E. alba woodland

Figure 1. Map of Pantar and Alor

showing survey sites visited or

localities mentioned in the text.

Sites: 1 = Puntaru village; 2 = Mt

Wasbila; 3 = Baranusa 4 = Mt

Sirung 5 = Kabir; 6 = Treweg

Island; 7= Batu village; 8= Pura

Island, 9= Ternate Island; 10=

Buaya Island; 1 1 = Kepa Island, 1 2=

Kalabahi; 13= Wahwah; 14= Mali;

1 5= Mainang; 1 6= Sika Island; 1 7=

10 km north of Apui; 18= Apui-

Subo-Manmas; 19= Kunggwera;

20= Mount Koya Koya; and, 21 =

Tanglapui Timur (Lipa, Kobra

hamlets). Source is Google Earth,

10 March 2012.

Forktail 28 (2012)

The avifauna of Alor and Pantar, Lesser Sundas, Indonesia

79

Yellow Bittern Ixobrychus sinensis

Alor: An adult male (WAM A24495) was collected in rice fields at

Apui-Subo on 24 April 1991 (REJ). The Yellow Bittern is poorly

known in Wallacea. It was thought to be largely migratory, except

on Sulawesi and Flores (Coates & Bishop 1997), but there appear

to be no confirmed breeding records on Flores (Verheijen 1964,

Mees 2006). Dated specimens range from September to April

(White 1976). A nest with one chick was discovered on Sumbawa

in May 198-8 ( Johnstone et al. 199 6). The number of April, May,

June and July records for East Timor, and a recent July record on

Sumba (Trainor 2011), suggest that this bird either breeds widely

in Nusa Tenggara or that some individuals occasionally oversummer

in the region.

Eastern Osprey Pandion cristatus

Pantar: The first island record was of a single bird observed in flight

along the coast at Baranusa village on 10 December 2009 (CRT).

Eastern Osprey appears to be uncommon to rare in the central Lesser

Sundas (White & Bruce 1986). Everett observed one bird on Alor

during April or May (Hartert 1898), and one bird was observed in

the hills ofAlor during December 1994(Holmes 1995). During about

300 field-days in Timor-Leste, CRT has only observed a single bird

(11 October 2004). These presumably refer to resident birds.

Migratory Western Osprey/? haliaetus has been recorded in northern

Wallacea (Sangihe, Talaud and Sulawesi) but none has been noted

entering Wallacea from Bali (White & Bruce 1986, Germi 2005,

2006) , suggesting that the migratory species is either very rare or does

not enter Wallacea along the Continental Flyway (Germi 2009).

Short-toed Snake Eagle Circaetus gallicus gallicus

Alor: One was observed and sketched over Apui-Subo on 24 April

1991 (REJ). One bird was observed above Apui-Subo on 1 July 2009

(PV and VD); a pair were seen at the same site on 9 September 2009

(PV), and one bird photographed at Mainang on 9 January 2010

(CRT). Pantar: One bird was observed by WAM staff near Kabir on

21 April 1991 (REJ). Singleswere observed daily in September 2009,

including one bird photographed, on the slopes of Mt Wasbila (PV) .

The resident Lesser Sundas population (Lombok to Timor/Roti) of

Short-toed Snake Eagle is isolated by at least 2,500 km from the nearest

resident populations in India and Nepal ( Robson 2000). Migrant birds

reach Bali at least (Coates & Bishop 1997). Despite the isolation of

this population, there are few apparent morphological differences from

the nominate subspecies (Mees 2006).

Bonelli's Eagle Aquiia faseiata renschi

Alor: One bird was observed about 12 km north-east of Mt Koya

Koya (on the road to Tanglapui Timur) on 10 May 2002 soaring

over tropical forest; a pair was seen on the slopes of Koya Koya on

13 May 2002, and a single bird was observed south of Lipa hamlet

on 14 May 2002 (CRT). Two pairs and a single were observed near

Mainang on 9 January 2010 flying over Eucalyptus alba woodland

and secondary forest, with one pair photographed (CRT). A pair

was photographed near Baifui village (5 km south-west of Apui-

Subo) on 1 1 January 2009 (CRT). Pantar: One adult was giving a

display flight on the slopes of Mt Wasbila on 3 September (PV).

The isolated resident Lesser Sunda population of Bonelli’s Eagle

ranges from Lombok to the Tanimbar Islands (Debus et al. in press)

with the nearest resident populations in north-west Thailand and

northern Laos (Robson 2005). The photographs of birds near

Mainang confirm the presence of this eagle on Alor. However,

records of this eagle from Alor in 2002, and Pantar in 2009, may

have been confused with Flores Hawk Eagle Nisaeetusfloris.

Flores Hawk Eagle Nisaetus floris

Alor: A pair was seen daily giving a territorial display over evergreen

forest at Kunggwera (23-25 June 2009) and two birds were

photographed in the hills above Apui-Subo on 9 September 2009

(PV). A pair and two singles were subsequently photographed at

Apui-Subo on 6 December 2009 with a pair on 14 January 2010

(CRT). One was photographed in flight with a large bird (probably

a rail) in its talons (CRT). A pair was photographed at Mainang at

800 m on 9January 2010 (CRT) displaying over secondary tropical

forest. One bird was seen in highly degraded forest in the hills near

Kalabahi on 3 July 2011 (Collaerts et al. 201 1). This critically

endangered species was known from Lombok through to Flores

(including Satonda, Sumbawa and Rinca islands) with a population

estimated at fewer than 100 pairs (Gjershaug et al. 2004). Flores

Hawk Eagle has not been recorded on Komodo Island

(Raharjaningtrah & Rahman 2004). However, it was recently

recorded in mangrove habitat on an unnamed islet 800 m off

Komodo, apparently hunting amongst a fruit-bat colony (probably

Large Flying Fox Pteropus vampyrus ) (M. van Buuren pers. comm.

2001). Our Alor records are therefore an important range extension.

The use of matrix habitats on Alor including secondary tropical

forest (about 10-20% cover) on valleys and slopes, mixed with more

extensive Eucalyptus woodland and agricultural land, suggests that

the species can persist in highly fragmented forest estates. The only

extensive (>20 km2) area of tropical forest on Alor appears to be

on the slopes of Mt Koya Koya, but tropical forest remnants are

frequent throughout the island’s mountains, especially in gullies

and on slopes. The frequency of records on Alor suggests that a

good population is likely to present, although repeat observations

above Apui-Subo are likely to refer to the same pair. If the home

range of birds on Alor is similar to the 38.5 km2 estimated for Flores

(Raharjaningtrah & Rahman 2004) then a maximum of 73 pairs

could be present on the island (assuming that they can use all

available habitat, which is unlikely). Flores Hawk Eagle is likely to

be present on Pantar (where similar habitat was surveyed but the

species went unrecorded), and perhaps Wetar, Lembata and

Adonara. Although the Flores Hawk Eagle is a relatively large and

distinctive raptor, in the field we (CRT, PV) initially assumed that

these birds were Bonelli’s Eagles, and relied on species identification

from photos.

Crested Honey Buzzard Pernis ptiiorhynchus

Alor: One bird was photographed above Apui-Subo on 6 December

at 900 m, and another near Mainang on 1 1 and 1 3 January 20 1 0 at

700 m (CRT). Pantar: One bird was photographed in flight below

Mt Wasbila over agricultural land on 10 December 2009 (CRT).

Despite the lack of historical specimens (White 1976, White &

Bruce 1986) this bird is a common migrant to Nusa Tenggara during

the northern winter, with more than 1,000 birds sighted crossing

the Lombok Strait during October-November 2004 (Germi 2005)

and 7,717 birds sighted in September-November 2005 (Germi

2006). It should be expected in small numbers throughout Nusa

Tenggara, and there are Christmas Island and recent Australian

mainland records (Gregory 2007).

Black-winged Kite Eianus caeruleus hypoleucos

Alor: A single bird was observed in Eucalyptus alba woodland in

Lipa hamlet on 2 May 2002, and a pair of birds was seen on 7 May

2002, hovering over woodland and perched in an E. alba tree in

the same area (CRT). One bird was also seen on 21 June 2009 in

coastal/:, alba woodland about 15 km north-west of Lipa (PV and

VD). Three birds were also observed by Widodo (2009): one at

Tanglapui and two at Tuti Adagae Nature Recreation Park. These

appear to be the first Alor records since Everett’s observation of

one bird (Hartert 1898). Pantar: One adult perched in a

Tamarindus indicus tree at the edge of mangroves at Kabir on 20

April 1991 (REJ). In Nusa Tenggara this species appears to be ‘a

local and sparsely distributed resident’ (Coates & Bishop 1997)

perhaps with the exception of the grassland and woodland habitats

80

COLIN R. TRAINOR, PHILIPPE VERBELEN & RON E. JOHNSTONE

Forktail 28 (2012)

on Sumba (Olsen & Trost 2007) and Sumbawa, where seven birds

were seen including one sitting on a nest on 7 July 2000 (Trainor

2002b). A nestling has been recorded on Flores during July (Mees

2006). The only recent Lombok records appear to be the four birds

seen in 1997 (Myers & Bishop 2005), and it appears to be

uncommon and local on Flores (Butchart etal. 1994, Pilgrim 2000,

Mees 2006).

Peregrine Falcon Falco peregrinus ernesti

Alor: The first island records were of a pair displaying over

agricultural land at Kunggwera on 5 May 2002 (CRT).

Subsequently one bird was photographed above Apui-Subo on 9

September 2009 (PV), one was seen on a tower in Apui-Subo village

on 14 January 2010 (CRT) and one bird was photographed in a

roadside tree about 4 km north of Mainang on 13 January 2010

(CRT). Pantar: The first island record was of one observed in

Baranusa village on 10 December 2009 (CRT). The photographs

confirm that these birds are the resident subspecies F p. ernesti.

Migrant F p. calidus appears to migrate through Nusa Tenggara in

low numbers, with seven birds noted on Bali flying towards Lombok

(Germi 2006).

Red-legged Crake Rallina fasciata

Alor: The trill-like call (grebe-like chatter, see http://www.xeno-

canto.org/xc33425) of Red-legged Crake was heard and sound-

recorded from tall grass above Apui-Subo on 4 December 2009

(CRT). The first Alor record was of a bird killed by a catapult near

Kalabahi on 9 December 1994 (Pdolmes 1995). Eggs were collected

from a nest in central Flores on 25 February 1959, and specimens

of adults were collected in December (Mees 2006). The eggs had

similar dimensions to birds breeding on Java (Mees 2006). On

neighbouring Timor, the first records in February and March

included observations of breeding (Dymond 201 1, Trainor 2011).

These records suggest that Red-legged Crake migrates into the

Lesser Sundas during the northern winter (wet season) to breed.

Red-legged Crake has also been recorded recently from the east

Lesser Sunda islands of Sermata and Romang (CRT unpubl. data).

Flores Green Pigeon Treron fioris

Vulnerable. Alor: During 2002 there were about 1 0 records mostly

of 1-2 birds in tropical forest, or feeding in figs at 580-780 m, and

one flock of 40-50 birds feeding in figs in forest-lined ravines at

860 m on the slopes of Mt Koya koya (CRT). In July 2009, Flores

Green Pigeon was quite common in groups of up to 15 in fruiting

trees about Kunggwera and calling from the forest canopy in ravines

(PV and VD). A small party of 5-10 birds were observed feeding

in a roadside fruiting tree near Mainang, at 700 m (CRT); one bird

called from secondary forest at 1,050 m above Apui-Subo on 14

January 2010 (CRT); and two birds called from a gully 2 km west

of Lipa at 720 m on 17 January 2010 (CRT), two birds were seen

in tropical forest at about 230 m on the road to Apui on 9

September 2009 (PV). Pantar: The only records are two

observations of single birds in flight on 3 September 2009 over

tropical forest at c.400 m on Mt Wasbila (PV).

Alor appears to be a stronghold for the Flores Green Pigeon,

which seems to be common around the sparsely populated

Kunggwera village and on the slopes of Mt Koya Koya, which

retains some of the most extensive tropical forest on the island. In

western Alor our records were less frequent, and populations of

the Flores Green Pigeon might be limited by intensive hunting.

However, throughout inland Alor there are also many extensive

steep forest-lined ravines that are relatively remote from villages.

There is little recent information on the status of Flores Green

Pigeon from Flores and Sumbawa apart from bird tour reports

(Eaton 2009), and the last published Lombok record was from

about 1909 (Myers & Bishop 2005).

Pied imperial Pigeon Ducula bicolor

Alor: A flock of eight birds feeding in a fruiting Garuga tree

and later a flock of four in a teak Tectona plantation near

Kalabahi on 10 April 1991 (REJ) are the first Alor records. The

Pied Imperial Pigeon is a small-island species that possibly visits

islands nomadically (Coates & Bishop 1997) to access fruit

resources. There are few Nusa Tenggara records, but most of these

have come from Komodo, Rinca and West Flores, associated with

extensive mangrove landscapes (apparently used for feeding and

roosting) during July-November and January and March

(Schellekens et al. 2009). It is frequent on the large island of

Yamdena in the Tanimbar islands (Bishop & Brickie 1998). Pied

Imperial Pigeon is clearly not resident on Alor, but the likely source

of these birds is unclear. It typically breeds on small islets, but the

islands around Alor (Treweg, Kepa, Pura, Ternate and Buaya) are

relatively heavily populated and are unlikely breeding habitat. It

might be expected to occur about the extensive mangrove stands

on Pantar Island.

Green / Pink-headed Imperial Pigeon

Ducula rosacea laenea polia

Alor: Pink-headed Imperial Pigeon D. rosacea was recorded in

small flocks of up to five birds in coastal forest near Kalabahi, with

Green Imperial Pigeon D. aenea also moderately common in flocks

of up to 18 birds in the same area in April 1991 (REJ). Imperial

pigeons were common in forest in Kunggwera at 600-900 m with

up to 1 5 Green present in a fruiting fig, and occasional on the slopes

of Mt Koya Koya (CRT). Green Imperial Pigeons were also

recorded frequently in the forests around Kunggwera in June

2009 (PV). A few imperial pigeons, probably Pink-headed, were

recorded a couple of times on Kepa Island and Ternate Island

(CRT). Pantar: Green Imperial Pigeons were recorded in small

flocks in forest and mangrove at Batu and Kabir in April 1991

(REJ). Pink-headed Imperial Pigeons were moderately common in

small flocks up to nine at Batu and in mangrove at Kabir in

April 1991 (REJ). Green were recorded relatively commonly in

tropical forest on Mt Wasbila in September 2009 (PV) but were

heard only a few times in December 2009 over two days at the same

site (CRT). Small numbers of imperial pigeons (2-4), most

probably Pink-headed, were observed and heard calling on

5 September 2009 in a stand of mature mangrove forests some 5 km

from Baranusa (PV). Imperial pigeons appear to have been severely

impacted by hunting on Alor: they were relatively common in

remote forested areas in the east, but none was recorded at Apui-

Subo or Mainang in west Alor, with only a few birds heard on two

islets. A total of 13, 9 and 17 Green Imperial Pigeons were recorded

at three Alor sites and seven Pink-headed were recorded at one site

by Widodo (2009). Imperial pigeons may be more abundant in

lowland coastal forests on Alor, especially those dominated by

Canarium trees, although this habitat has not been covered in

recent surveys.

Bar-necked Cuckoo Dove Macropygia magna magna

Alor: Th is dove was frequently seen and heard in tropical forest

and sometimes adjacent Eucalyptus woodland at Kunggwera in

2002 and 2009 (CRT, PV and VD), and up to five birds were

heard calling (and sound-recorded) in tropical forest at 250 m

about 10 km north of Apui-Subo, and above Apui-Subo on 1 July

2009 and 9 September 2009 (PV). A few birds were also observed

in degraded forest in the hills near Kalabahi on 3 July 2011

(P. Collaerts in litt. 2012). Pantar: Relatively commonly heard

and seen (and sound-recorded) in tropical forest on Mt Wasbila

(PV) with up to 20 birds calling at dusk from a small area

there on 10 December 2009 (CRT). One bird was calling in

beach forest at Puntaru (CRT). Bar-necked Cuckoo Dove occurs

on the Flores Sea islands of Tanahjampea and Kalaotoa, and in

Forktail 28 (2012)

The avifauna of Alor and Pantar, Lesser Sundas, Indonesia

81

the Lesser Sundas from Pantar through to Tanimbar (White &

Bruce 1 986). Both Alor and Pantar probably host large populations

of this dove but these may be impacted by hunting in heavily

populated areas.

The vocalisations of the different races are distinct, suggesting

that at least three species are involved (Eaton & Hutchinson 2011).

On Pantar, Alor, Wetar, Atauro, Timor and associated islands At.

m. magna gives a two-note call, while At. m. timorlaoensis of the

Tanimbar archipelago gives a three-note call (Coates & Bishop

1997, www.xeno-canto.org) and subspecies At. m. longa of the

Flores Sea also gives a distinctive call (Eaton & Hutchinson 201 1 ).

Little Cuckoo Dove Macropygia ruficeps orientalis

Alor: Frequently heard and seen in tropical forest at Kunggwera

(PV) and secondary forest above Apui-Subo (500-1,000 m)

(PV.CRT), and a few birds heard at Mainangat 700 m (CRT), and

a single bird heard calling from secondary forest c.2 km east of Lipa

village (CRT). Pantar: A pair was photographed at c.400 m in

tropical forest on Mt Wasbila (PV), and heard calling there about

10 times over two days at 300-400 m in December 2009 (CRT).

Little Cuckoo Dove occurs widely in South-East Asia and the

Greater Sundas, with the endemic subspecies At. r. orientalis

reputedly present on the islands from Lombok to Flores, Lembata

and Timor, Sumbawa, Flores, Pantar (White & Bruce 1986) and

now Alor. It is generally poorly known in Nusa Tenggara, but is a

common bird in the mountains of Timor (CRT unpubl. data). The

call of birds on Timor, Pantar and Alor is unlike other members of

the species and unlike those of the endemic Lesser Sundas

subspecies At. r. orientalis , suggesting that it is an undescribed

species (Eaton & Hutchinson 2011). An analysis of calls is currently

underway (JE in lift. 2012).

Ruddy Cuckoo Dove Macropygia emiiiana

Alor: A sound recording of this bird was made in tropical forest

about 10 km north of Apui on 29 August 2009 (PV). On 23

October 2011 it was observed in flight at least twice in secondary

forest at Mainangat c.900-950 m (RH, JE) with afield note at the

time of a ‘reddish large cuckoo dove’ (JE in litt. 2012). Ruddy

Cuckoo Dove was known from Lombok, Sumbawa, Flores and

Paloe in Nusa Tenggara (White & Bruce 1986), with these records

the first for Alor. This bird calls rarely on Java and Flores ( JE in

litt. 20 1 2) and consequently it is relatively little known in Wallacea.

A recorded bird gave a downslurred double note whu-ivhu repeated

at about 5-second intervals, which is similar to calls described from

Flores (Coates & Bishop 1997). This bird is considered to be

generally uncommon on Flores (Verhoeye & Holmes 1999), but

locally common in the mountains of Flores (Pilgrim et al. 2000)

and rare in lowland forest, but common in upper montane forest

on Sumbawa (Johnstone et al. 1996).

Yellow-crested Cockatoo Cacatua sulphurea parvula

Alor: A loose flock comprising six, eight and four birds was

observed over forested gullies at c. 230 m, 1 0 km north of Apui, on

1 July 2009 (PV and VD). Local people said that cockatoos were

regularly seen in these river valleys (PV). In 2002, local people at

Kunggwera stated that flocks of up to 10 birds were occasionally

seen about the village, and that c.20 birds were present in a nearby

river valley, but none was seen there during 2002 or 2009 (CRT

PV). Pantar: In September 2009, a caged bird was seen in the port

of Baranusa (PV) ready to be shipped off the island, and another

caged bird, caught on the island, was seen and photographed in

the village of Anis Lilo (PV). This latter, a male, was on sale for

Rp. 400, 000 (c.$US40), but prices of up to Rp. 1,500,000

(c.$US150) are sought for ‘red-eyed’ female birds on Pantar (PV).

The Yellow-crested Cockatoo is currently a rare bird on both

Alor and Pantar as a result of intense historical and ongoing captures

for trade. On 9 April 1991 a total of 23 adults and 12 immature

cockatoos were seen in small cages at Kalabahi ready to be shipped

out (REJ). No birds were observed at a Kalabahi market on 9 May

2002 (CRT). No cockatoos were seen by Holmes (1995) during a

brief visit to sites in west Alor, but in the late 1990s a total of 80

birds were seen at seven Alor sites (4—30 birds per site) and 29 were

seen at two sites on Pantar (Setiawan et al. 1999; no survey dates

given in report). The total population was crudely estimated at

678-782 birds for Alor and 444-534 birds on Pantar (Setiawan et

al. 1999) based on the extent of preferred riverine forest. No

cockatoos were recorded at four Alor sites during a specific search

for this species (Widodo 2009; no survey dates given in article).

Riverine and coastal swamp forest dominated by Canarium are

probably the most important habitats for Yellow-crested Cockatoo

(and possibly imperial pigeons) on Alor (Setiawan et al. 1999,

Widodo 2009), as these large trees form hollows suitable for nesting

and the large nut of this tree is a preferred food item. These habitats

were poorly covered during our fieldwork (CRT, PV). No Yellow-

crested Cockatoo have been observed during recent visits to the

islands of Lembata, Adonara and Solor (Trainor 2002a, c,

Schellekens et al. 2011), and the species must now be close to

extinction on most Nusa Tenggara islands except Sumba and

Komodo, and possibly Timor-Leste (East Timor).

Little Bronze Cuckoo Chrysococcyx minutillus

Alor: One bird called from coastal dry forest near Kalabahi on 1 1

April 1991 (REJ). This cuckoo was recorded frequently about

Tanglapui Timor area in 2002, particularly in Eucalyptus woodland

above c. 1,000 m (CRT), but was unrecorded and apparently absent

from dense tropical forest at Kunggwera in 2002 and 2009 (CRT,

PV). Pantar: The first island records were of several birds

photographed and sound-recorded in Eucalyptus alba woodland

and coastal scrub in September 2009 (PV).

The taxonomic relationships within this species complex remain

unresolved, but most authorities prefer to re-unite Little Bronze

Cuckoo with Goulds Bronze Cuckoo C. russatus (Mees 2006, Gill

& Ddnsker 2012). In the Lesser Sundas this bird occurs on Flores,

Timor, Wetar and several Banda Sea islands (White & Bruce 1986).

Previously it was listed for Alor as Gould’s Bronze Cuckoo (Trainor

2005a). The lack of records of this bird on Alor during 2009 and

2010 (PV, CRT) was notable, but it may vocalise less duringperiods

of high rainfall such as December and January. The new record on

Pantar is expected, although surprisingly it is yet to be recorded for

Lembata or Adonara (Trainor 2003, Schellekens et al. 2011). The

host species on Flores is Golden-bellied Gerygone Gerygone

sulphurea (Mees 2006), and probably sunbirds Cinnyris (Coates &

Bishop 1997). It appears to be uncommon on Flores with the only

recent published records by Pilgrim et al. (2000). On Alor the

abundance of Little Bronze Cuckoo in high-elevation Eucalyptus

woodland matched the high abundance of its likely host in that

habitat, the Golden-bellied Gerygone.

Sunda/Oriental Cuckoo Cuculus lepidus/optatus

Alor: A single Cuculus cuckoo was observed in flight in forest above

Apui-Subo on 9 September 2009 (PV), and a single bird was

observed in Eucalyptus woodland south of Mainangat 1,050 m on

23 October 201 1 ( JE in litt. 2012). The distinctive calls of Sunda

Cuckoo have not been heard by any recent visitors to Alor (or

Pantar) although there is a specimen from Pantar (Hartert 1898)

and it is common on the adjacent islands of Atauro (Trainor &

Soares 2004, where mentioned as Oriental Cuckoo), Wetar

(Trainor et al. 2009) and Timor (Trainor et al. 2008). It would be

expected to occur on Alor. The brief observations cannot exclude

Oriental Cuckoo, which is one of the commonest non-passerine

migrants in Wallacea, with records from September to March

(White 1976).

82

COLIN R. TRAINOR, PHILIPPE VERBELEN & RON E. JOHNSTONE

Forktail 28 (2012)

Asian Koef/Pacific KoeS

Eudynamys scolopaceus/cyanocephala

Alor: One of the most frequently heard birds in tropical forest on

Mt Koya Koya in May 2002 at 600-1,200 m (CRT), heard on 31

August 2009 in highly degraded forest near Kalabahi (PV);

common above Apui-Subo in September and December 2009

(PV.CRT), at and below Mainang (350-900 m) and in mangroves

at the head of Muitara Bay (CRT). Pantar: Commonly heard in

forest on the slopes of Mt Wasbila during September 2009 (PV),

and a few birds heard calling on Mt Wasbila at 200-400 m on 12

December 2009 (CRT). A. H. Everett collected koels but reported

to Hartert (1898), who identified them as Asian Koel, that ‘it [is

not] easy to determine these birds from Alor’. The status of Pacific

Koel E. cycanocephala in Wallacea is unclear, with White & Bruce

(1986) suggesting that the Australian bird occurs mainly as a

migrant with some resident populations likely. On current

knowledge Pacific Koel reaches Ashmore Reef (140 km south of

Roti Island) as a rare austral migrant (M. Carter in litt. 2012). Alor

birds have been considered as Pacific Koel (White & Bruce 1986,

Coates 8c Bishop 1997), although December records on Alor were

considered by Holmes (1995) to be of resident Asian Koel. Our

records suggest that a large population presumably of Asian Koel

is present throughout the year, but we observed few birds and took

no photos to confirm the taxonomic status of this bird.

Southern (Aior) Boohook Ninox (boobook) plesseni

Alor: This bird called frequendy from Eucalyptus urophylla woodland

and tropical forests in gullies on Mt Koya Koya (700-1,200 m), with

up to four birds calling simultaneously and sound-recorded before

and at dusk; they also called throughout the night until dawn (CRT).

In June 2009, up to five birds were heard to call simultaneously (in

forest in the Kunggwera area), and 2-3 birds called, and were

photographed, at dusk in a forested gully about 2-3 km from Lipa

on 21 June (PV, VD). Pantar: Birds were photographed and sound-

recorded in coastal E. alba woodland, and called frequently from

mangrove forest adjacent to eucalypt woodland and from evergreen

forest on Mt Wasbila up to at least 600 m (PV), and several were

heard calling on Mt Wasbila at 600 m (CRT).

The subspecies N. b. plesseni was described from a single female

collected at c. 1,000 matTanglapoi [Tanglapui] (Stresemann 1929)

and was discussed by Mees (1964) and re-described by Johnstone

& Darnell (1997b), but there were no field observations of this

bird between 1927 and our fieldwork. Using morphological features

and colours to distinguish the many subspecies of Southern

Boobook has been of relatively limited value and it is likely that

several of these subspecies are specifically distinct (Johnstone &

Darnell 1997). However, vocalisations are often distinctive. Birds

on Aior and Pantar have the same low-pitched call which is repeated

rapidly 6- 14 times over c. 1 .5 seconds and which is highly distinctive

among all members of the complex, making it a prime candidate

for upgrading to species level. Most members of the Southern

Boobook complex have similar low-pitched calls, but vocalise with

bru-book or bru-brook double notes. We intend to publish further

details of the distinctiveness of this taxon elsewhere. Despite the

restricted global distribution of the ‘Alor Boobook’, it appears to

be under little threat of extinction because it can use both Eucalyptus

woodland and tropical forest. This appears to be the only species

endemic to Alor and Pantar, although this may change when the

distinctiveness of several passerines recently recorded on Alor is

resolved. We had no records from west and central Alor during

limited (c.4 nights) nocturnal observations (PV, CRT) but

presumably it is present throughout much of the island.

[Pantar scops owl?

Pantar: Calls presumed to belong to a scops owl Otus were heard

and sound-recorded after dusk and irregularly throughout the night

until at least 0 lhOO at the edge of mangrove forest some 5 km from

Baranusa in September 2009 (PV). These were again heard and

recorded at the same site on 3 July 201 1 (Collaerts etal. 2011) and

on 22 October 2011 during a Birdtour Asia visit (JE in litt. 2012).

The calls are reminiscent of those of Moluccan Scops Owl Otus

magicus of Flores and Lembata (www.xeno-canto.org, Schellekens

et al. 2011) but clearly differ in having a less harsh barking quality.

Sound recordings were posted on the Xeno-Canto website (http:/

/ www.xeno-canto.org/ asia/ discussion. php ?snd_nr= 1 7 04). When

an attempt was made by PC to get closer to the source of the call, it

retreated further away. Unfortunately there was not the typical Otus

response to playback of flying closer to the playback source. During

a visit to the Pantar mangroves in October 2011 there was again no

response to playback by the bird, which suggested that the calls

may not have originated from a bird (JE in litt. 2012). However,

future visitors to Pantar and Alor should remain vigilant for the

presence of a scops owl perhaps especially about mangrove habitat

and playback of these unidentified calls should be tried to confirm

the identity of the sources of the calls.]

Mees' s Nightjar Caprimuigus meesi

Alor: One bird was seen and sound-recorded after it called in

response to playback in Eucalyptus alba woodland near Lantoka

(7 km east of Tanglapui Timur) on 27 June 2009 (PV and VD).

Pantar: The species was targeted with playback on Pantar because

of the likelihood it was present on the island following its discovery

on Alor. In open woodland at 100 m about 3 km east of Puntaru

on the road to Sirung volcano on 4 September 2009 a nightjar

responded strongly to a recording of Mees’s Nightjar, by

immediately flying towards the recording and landing on the

ground less than 1 m from a motorbike. After it flew off a recording

of Savanna Nightjar C. afftnis was played, but the bird did not

respond (PV). The bird was not photographed or observed long

enough to confirm identification. Playback was used twice, with

the same result: no reaction to playback of Savanna Nightjar calls,

but immediate reaction to a recording of Mees’s Nightjar recording.

The lack of a vocal response makes confirmation impossible, but

its behaviour indicates that it was likely to be Mees’s Nightjar rather

than Savanna Nightjar. The recently described Mees’s Nightjar

(Sangster & Rozendaal 2004) was previously known from Flores

and Sumba. These two new island records are an important range

extension, and presumably it also occurs on the intervening islands

of Adonara, Lembata and possibly Solor, although little nocturnal

work has been conducted on these islands (Schellekens etal. 2011).

Stork-billed Kingfisher Pelargopsis capensis

Pantar: One bird was observed in mangroves near Desa Batu on

19 April 1991 (REJ). The Stork-billed Kingfisher is known from

Lombok, Sumbawa and Flores in the Lesser Sundas (Coates &

Bishop 1997), with several recent records from mangrove and

estuary habitat on Lembata (Trainor 2003). This bird is generally

uncommon to rare in Nusa Tenggara, but there are regular records

on Flores from mangroves and Lake Rana Mese at 1,200 m (Pilgrim

etal. 2000) and occasional records on Sumbawa from the coast to

hills up to 870 m (Butchart etal. 1994, Johnstone etal. 1996).

Oriental Dwarf Kingfisher Ceyx erythraca ssp.

Pantar: Three pairs of Oriental Dwarf Kingfisher were seen about

mangroves, coconut and shrub, 3 km north of Baranusa on 22

October 2011, including a pair apparently using a nest-hole on a

steep slope (JE in. litt. 2012). This bird occurs widely in South

and South-East Asia, and in Nusa Tenggara it occurs on Lombok,

Sumbawa, Flores and Sumba (White & Bruce 1986). On Alor, none

was noted during two visits (7 December 2009 and 10 January

2010) to extensive mangroves at the head of Mutiara (Kalabahi)

Bay (CRT). Recently, the Oriental Dwarf Kingfisher was found to

Forktail 28 (2012)

The avifauna of Alor and Pantar, Lesser Sundas, Indonesia

83

be locally common in evergreen swamp forest on neighbouring

Lembata Island (Schellekens et al. 201 1). There are relatively few

Nusa Tenggara records but it is a small and inconspicuous bird (with

a high-pitched and near-inaudible call) and is likely to be regularly

overlooked. It has been considered ‘rare’ on Flores (Butchart et al.

1994, Coates & Bishop 1997), there are no recent Lombok records

(Myers & Bishop 2005), but there have been records from riverine

forest on Sumbawa up to 650 m (Johnstone et al. 1996).

Sunda Pygmy Woodpecker

Dendrocopus moluccensis grandis

Alor: One adult male (WAM A2451 1, length 160 mm, weight 22 g)

was collected at Apui-Subo on 26 April 1991 (REJ). This bird was

recorded once or twice per day in tropical forest and Eucalyptus

woodland, usually as single birds, especially along gullies in the

Tanglapui Timur area (600-1,200 m) (CRT, PV), above Apui-Subo

(PV, CRT) and Mainang (CRT). Pantar: The only record was of

two birds on 4 September 2009 in Eucalyptus urophylla woodland

on the slopes of Mt Sirung at about 500 m (PV). In Wallacea the

Sunda Pygmy Woodpecker occurs from Lombok through to Alor

(White & Bruce 1986) and would have been expected to occur on

Pantar. It seems to prefer tropical forest on Alor, but also occurs in

woodland and plantations when in close proximity to tropical forest.

Specimens on Alor average larger than birds on Lombok, Sumbawa

and Flores, and were originally described as an endemic subspecies

D. m. excelsior (Hartert 1898), but differences were not considered

significant by White & Bruce (1986) and this subspecies has not

been recognised recently by authorities.

Elegant Pitta Pitta elegans concinna

Alor: Moderately common, calling with a single bird observed in

forest at Apui (650-740 m) and in tropical dry forest at Mali in

April 1991 (REJ), with three specimens collected (WAM A24458,

A24544 and A24488). Pittas called occasionally at dawn and dusk

from tropical forest at Kunggwera and the slopes of Mt Koya Koya

(600-950 m) during May 2002 (CRT), and called there frequently

in June 2009 (PV). At least three birds called at dusk in degraded

forest 15 km east of Kalabahi on 30 June 2009 (PV); two birds

called from forested gullies at Mainang on 9 January 20 10 (750 m),

and up to four birds called along the road to Wah Wah (4 km

north-east of Kalabahi at c.400 m) on 20 January 2010 (CRT).

Pantar: One bird was recorded from dense Acacia and shrubs at

the edge of mangroves on 19 April 1991 and one bird called from

dense stands of the weed Wedelia biflora near Ivabir on 20 April

1991 (REJ). A total of 5-7 birds called from tropical forest on

Mt Wasbila during 3 September 2009 (PV) and several birds

were sound-recorded in a dry thicket at 120 m and heard at 650-

800 m there on 10 December 2009 (CRT). Elegant Pitta occurs

widely in the Lesser Sundas with four subspecies present (White

& Bruce 1986). The subspecies P. e. concinna of Lombok-Alor

appears to be resident with records throughout the year, but on

the drier islands of Roti and Timor the subspecies P. e. elegans

(which has a three-note call) is migratory (White & Bruce 1986,

Trainor 2005b, Tebb et al. 2008, PV unpubl. data). Surprisingly,

there are no published records of Elegant Pitta from Timor-Leste

(East Timor), Wetar (Trainor et al. 2009) or Atauro (Trainor &

Soares 2004). Recordings of the two-note calls of P. e. concinna

from Alor, Pantar and Lembata have been uploaded to the

www.xeno-canto website.

Wallacean Cuckooshrike Cottacina petrsonata alfrediana

Alor: Moderately commonly observed in ones, twos and threes in

Eucalyptus woodland at Apui in April 1991 (REJ). Regularly

observed in small groups of 3-5 birds in Eucalyptus woodland on

Mt Koya Koya (600-1,100 m) and frequently observed, mostly as

singles, in Eucalyptus woodland and tropical forest at Kunggwera,

above Apui-Subo (PV, CRT) and Mainang (CRT). Pantar: Several

pairs were observed in dry tropical forest at the foot of Mt Wasbila

at 250 m (PV); several birds were observed in tropical forest and

Eucalyptus woodland at 300-600 m (CRT) and one pair was

observed in open eucalypt woodland en route to Mt Sirung (PV).

Wallacean Cuckooshrike occurs widely in the Lesser Sundas and

the Kai Islands (Maluku), with seven subspecies described (White

& Bruce 1986). On Alor, C. p. alfredianus was described from six

specimens which differed from C. p. floris (Sumbawa-Flores) by

being a ‘distinctly paler grey above and below, and the males do not

have a black throat and the sides of the head’ (Hartert 1898). A

specimen from Lembata was ‘a shade darker, thus pointing towards

[...]_ floris’ (Hartert 1898). The presence of Wallacean Cuckooshrike

on Pantar is expected: presumably this bird is included with C. p.

alfrediana. Based on distribution, the Lembata population is as

likely to be C. p. floris as C. p. alfrediana , but plumage characteristics

were not noted during recent field surveys (Trainor 2003).

Wallacean Cuckooshrike is part of a complex which also includes

Black-faced Cuckooshrike C. novaehollandiae. Some taxa within

this complex have already been split (e.g. Buru Cuckooshrike C.

fortis ) and Mason & McKean (1982) argued for species status for

the Timor group subspecies C. p. per sonata but without considering

other forms (Mees 2006). A thorough review of this group is

needed. There are substantial differences in vocalisations among

populations (e.g. Figure 2a, b), and these may help to set taxonomic

limits.

Figure 2. Song of the Wallacean

Cuckooshrike from Apui-Subo,

Alor, 5 December 2009 (a) and

Hatu Builico, East Timor, 28 April

2010 (b) (both recordings by C. R.

Trainor).

84

COLIN R. TRAINOR, PHILIPPE VERBELEN & RON E. JOHNSTONE

Forktail 28 (2012)

Common Cicadabird Coracina tenuirostris s$p.

Alor: A single female was initially heard calling (slow took notes)

and then seen in the subcanopy of a candlenut Aleurites moluccana

plantation above Apui-Subo village at 950 m on 4 December 2009.

The bird had obvious barring on the neck, breast and belly, but no

other notes were taken before it flew off (CRT). A single immature

male bird was collected on Lembata (Hartert 1896), but there have

been no subsequent records on that island (Trainor 2002c, 2003,

Schellekens etal. 2011). On nearby Atauro Island a single male bird

observed gave ruk ruk notes (Trainor & Soares 2004), and females

on Timor also give reasonably similar contact notes (http://

www.xeno-canto.org/32585). The taxonomic affinities of birds on

Lembata and Alor are unclear, but on biogeographic grounds (and

probably vocalisations) they may be close to C. t. timoriensis.

Common Cicadabird is an inconspicuous bird that is generally

uncommon throughout its range. Ir is most easily detected by its

distinctive vocalisations. There are at least 23 described subspecies

of Common Cicadabird (Gil & Donsker 2012) including many

distinctive forms (Rheindt et al. 2010). A review of these taxa is

needed.

Lesser Shortwing Brachypteryx leucophrys

Alor: Common in tropical forest along creeks near Apui (650-

800 m) in April 1991 with four specimens collected (REJ). The song

of this bird was frequently heard on the slopes of Mt Koya Koya

(PV and VD), above Apui-Subo (PV, CRT), particularly in dense

vegetation and along streams and gullies. At Mainang, and along

the 17 km walk to Apui-Subo, this bird was restricted to watered

gullies above about 850 m (CRT). In the village ofTanglapui Timur

it occurred in a gully, and was recorded down to 680 m about 2 km

west of the village (PV, CRT). This bird was noted also by Holmes

(1995) on 10 December 1994 on the road between Mainang and

Apui-Subo. Records of this bird given by Trainor (2005a) were

confused with Timor Stubtail Urosphena subulata (see below),

including the description of a call given which clearly refers to the

latter bird. Examination of recordings in 2002 have since confirmed

that Lesser Shortwing songs and high-pitched single notes were

sound- recorded. Field notes show that three Lesser Shortwings were

observed in watered gullies, but perhaps these were also misidentified

Timor Stubtail. This bird is patchily common throughout much of

Alor above 600-700 m, but is absent from the extensi Eucalyptus

woodlands. None was heard on the upper slopes of Mt Wasbila on

Pantar (PV, CRT). At least to the human ear, the song of birds on

Timor and Alor bird sound similar to birds on Java (www.xeno-

canto.com). Mimicry of the song of Timor Bush Warbler by Lesser

Shortwing was sound-recorded (www.xeno-canto.com/xcl03157).

The subspecies is presumably B. 1. leucophrys.

Timor Bush Warbler Loeustella timorensis ssp.

Alor: First discovered at Apui-Subo on 9 September 2009, when

up to five singing males were heard calling from tall grass and thickets

below secondary tropical forest at 1,050 m (PV). Subsequent visits

to the same location noted at least 6-8 singing males along a ridge

at 900-1,250 m, mostly calling from tall grassland beneath

Eucalyptus urophylla woodland (CRT). A second population was

located at Mainang with seven males heard singing in grassland below

E. alba at 859-930 m. A total of 1 1 birds were heard, with some

birds photographed and videographed near Mainang at 928-

1,070 m on 23 October 2011 ( JE and RH in litt. 2012). Details of

this discovery were recently published (Trainor et al. 2012) and

videos and photos are available online (http://ibc.lynxeds.com/).

Timor Bush Warbler is clearly patchily distributed on the island.

For example, none was heard singing in the vicinity of Mt Koya Koya

in 2002 (CRT) or 2009 (PV) or 2010 (CRT) despite the use of

playback (in 2010) in apparently suitable habitat, and none was

heard during a 17 km walk between Mainang and Apui-Subo

through seemingly suitable Eucalyptus woodland (CRT), indicating

that they are local and have specialised habitat requirements.

Differences in song characteristics (frequency) suggest that the

Alor birds may be subspecifically distinct from the Timor

population (Trainor et al. 2012). None was recorded after using

playback in apparently suitable habitat on Mt Wasbila at 800 m,

but birds may be present elsewhere on Pantar.

Chestnut-backed Thrush Geokichla dohertyi

Alor: Birds were observed and heard occasionally in forested ravines

on the slopes of Mt Koya Koya (CRT ) and one was seen and several

heard singing in a forested gully at Kunggwera in June 2009 (PV).

This bird was occasionally heard and recorded singing from

secondary forest above Apui-Subo village in December and January,

with one bird killed with a catapult by a local villager on 4 December

2009 (CRT).

Chestnut-backed Thrush occurs on the large Nusa Tenggara

islands of Lombok, Sumbawa, Flores, Surnba and Timor, mostly

in hill or montane forest above 500 m (White & Bruce 1986). A

‘ Zoothera' thrush was noted by A. H. Everett on Alor but no

specimens were collected (Hartert 1898). Its presence on Alor is

unsurprising, particularly given the extent of forested montane

habitat; by contrast, the absence of records from Pantar was

noteworthy (PV, CRT). Chestnut-backed Thrush is an

inconspicuous forest-dweller and is easy to overlook if not

vocalising. There have been no records from Lembata and Adonara

(Trainor 2002a, c, 2004, Schellekens etal. 201 1), Atauro (Trainor

& Soares 2004) or nearby Wetar Island (CRT unpubl. data) but

more effort is needed in the hills and mountains of these islands.

Kamchatka Leaf Warbler Phylloscopus examincmdus

Alor: Birds were occasionally observed on Ternate Island on El

April 2002, and on Sika Island on 29 April 2002; one bird was

observed in a gully in Lipa village at 700 m on 2 May 2002, and six

were heard or seen 2-4 km east of Lipa, at 440-620 m, on 1 7 January

2010 (CRT). Pantar: A single bird was heard in coastal vegetation

at Puntaru on 14 December 2009 (CRT). The Kamchatka Leaf

Warbler was recently split from Arctic Warbler (Alstrom et al.

2011). This bird breeds in south Kamchatka, Sakhalin and north¬

east Hokkaido (Alstrom etal. 2011) and winters throughout South-

East Asia with Alor at about the southern limit of its wintering range.

The ‘Arctic Warbler’ subspecies examinandus was described from

material collected on Flores (Mees 2006), and birds were collected

on Alor in April by A. H. Everett, but no details of subspecific

identity were noted (Hartert 1898). The 'Arctic Warbler’ was the

second most commonly collected passerine migrant to Wallacea

(White 1977). There are regular records of Kamchatka/Arctic

Warbler on Adonara (Trainor 2002a) and Lembata (Trainor 2002c,

Schellekens et al. 2011), but it is very uncommon on Timor

(R. Noske in litt. 2012) with no East Timor records (CRT unpubl.

data).

Sumda Bush Warbler Horornis vulcanius kolichisi

Alor: Noted as moderately common in weedy Wedelia biflora

shrubs and grass at c.700 m above Apui, with eight specimens

collected including the holotype on 24 April 1991 (REJ, Johnstone

& Darnell 1997a). In 2002, found to be frequent on the slopes of

Mt Koya Koya and Lipa village in weedy tall grassland gullies and

stream channels (CRT), and in 2009-2010 it was common in the

same habitat at Mainang (700-900 m, CRT), Apui-Subo (800-

1,200 m) and Tanglapui Timur area at 650-1,150 m (PV and VD,

CRT).

The Sunda Bush Warbler is a poorly known bird which occurs

on Sumatra, Borneo, Java and Bali, with Nusa Tenggara

populations on the mountain areas of Lombok, Sumbawa and

Timor (Coates & Bishop 1997). Like many bush warblers its

Forktail 28 (2012)

The avifauna of Alor and Pantar, Lesser Sundas, Indonesia

85

plumage is conservative and nondescript (Kennerley & Pearson

2010) but vocalisations are often distinctive. There are recent new

observational records for Sumbawa (Johnstone et al. 1996) and

Wetar (Trainor et al. 2009). The Alor subspecies H. v. kolichisi

differs from both H. v. vulcanius (of Bali, Lombok) and H. v.

everetti (of Timor and possibly Wetar) by its smaller size, duller

dark brown upperparts lacking the rusty tones of the nominotypical

form and the olive tones of H. v. everetti (Johnstone & Darnell

1997a). The bill is also proportionately longer and finer with a more

extensive dark zone on the lower mandible (Johnstone & Darnell

1997a). The songs ofNusaTenggara birds are relatively distinctive

from birds of the Greater Sundas (PV unpubl. data) and work

analysing calls has begun (G. Sangster/« litt. 2011). The Sumbawa

bird is presumably similar to those of Lombok (Johnstone &

Darnell 1997a) and the song of the Wetar bird sounds close to

birds on Timor (CRT, PV unpubl. data). Birds on Timor and

Wetar occur from sea-level to the mountains (Noske & Saleh 1996,

Trainor et al. 2009) but the Alor bird appears to occur in the hills

only (but more lowland surveys are needed on Alor). The Sunda

Bush Warbler appears to be absent from Pantar.

Timor Stuhtail Urosphena subulata ssp.

Alor: This bird was not identified during the 2002 survey, but its

high-pitched note was regularly sound-recorded from the slopes

of Mt Koya Koya, and these were later (following publication of

Trainor 2005a) identified as Timor Stubtail (CRT). Subsequently

this diminutive bird has been found to be relatively common and

vocal, especially in gullies at Kunggwera (PV and VD), Tanglapui

Timur (PV, CRT), above Apui-Subo (PV, CRT), Mainang (CRT)

and Wahwah (CRT) at elevations of 660- 1,200 m. During a 17 km

walk between Mainang and Apui-Subo it was recorded only in a

single well-vegetated gully near Baifui village, and was absent from

Eucalyptus alba woodland.

This is a significant and unexpected range extension for this

essentially ‘Timor-group’ bird. Only recently has the range of this

species become better known, with new island records for Wetar

(Coates & Bishop 1997), Atauro (Trainor & Soares 2004), Roti

(Trainor 2005b) and Romang (CRT unpubl. data). It is generally

common in tropical forest throughout its range, although suitable

habitat appears to be limited on Roti and Atauro. The closest

relatives of the Timor Stubtail are restricted to the mountains of

north-east Borneo (Bornean Stubtail U. wbiteheadi ), while Asian

Stubtail U. squameiceps is also confined to mountains when

wintering in South-East Asia (Kennerley & Pearson 2010). On

Timor this bird occurs from sea-level to the mountains, but there

is limited data on the elevational limits of this bird on Alor. It may

be restricted to the hills, but there has been limited lowland survey

effort on Alor. The song has a uniform structure on all islands — a

monotone whistled pseeee (Kennerley & Pearson 2010) which varies

substantially in length, and less so in frequency. The relationships

of these island forms are still unclear, but they are probably

distinctive at the subspecific level.

Little Pied Flycatcher Ficedula westermanni mayrii

Alor: One bird was observed in the forest canopy at Apui on 27

April 1991 (REJ). Noted as occasional in Casuarina junghuhniana

and tropical forest-lined gullies, and E. alba woodland (650-

1,120 m) around Tanglapui Timur in 2002 (CRT). A pair was seen

in a gully at Tanglapui Timor in 2010 (CRT), and birds were

frequent in secondary forest above Apui-Subo in 2009 and 2010

(PV,CRT), but the species was recorded only once on the walk from

Mainang to Apui-Subo (CRT ) . One male bird was observed singing

c.10 km north of Apui at 230 m (PV).

Little Pied Flycatcher is typically a montane bird (above about

800 m) throughout its broad South-East Asian range, but on Timor

(Noske & Saleh 1996), Atauro (Trainor & Soares 2004), Wetar

(Trainor et al. 2009) and now Alor it occurs from the lowlands to

the mountains. This unusual aspect of its habitat use in the Lesser

Sundas has been widely overlooked (White & Bruce 1986, Coates

& Bishop 1997, Mees 2006). The Little Pied Flycatcher is

morphologically conservative with relatively minor differences in

plumage throughout its range. Analysis of songs might help to

disentangle cryptic diversity within this complex. Three specimens

(all females) were collected on Alor by von Plessen (White & Bruce

1986) presumably at an elevation of about 1,200 m (which was given

as the Alor elevation range for this bird: Coates & Bishop 1997).

YeSlow-breasted Warbler Seircercus montis ssp.

Alor: Th is bird was local in well-forested gullies at 800-930 m on

the slopes of Mt Koya Koya, with only a few birds seen and recorded

during April 2002 (CRT). Several birds were photographed and

recorded in dense forest in the hills above Apui at c. 1 ,000-1 ,100m

on 9 September, 2009 (PV), but none was recorded there in

December or January (CRT).

In Nusa Tenggara this bird occurs on Flores (subspecies floris-.

compared to S. m. montis of Borneo it has less well developed dark

lateral crown-bands to nape) and Timor ( paulinae : compared to S.

m. montis it has a brighter rufous crown, brighter olive back,

yellower rump, and less developed crown-bands) in Wallacea

(White & Bruce 1986). Photos of the Alor bird indicate that it is

probably closer to S. m. paulinae with a bright orange crown which

appears to lack lateral crown-bands entirely. Although the

subspecies described for Nusa Tenggara have been considered to

have diverged little from S. m. montis of northern Borneo (White

& Bruce 1986) morphological differences shown by photos on

www.orientalbirdimages.org, and differences in vocalisations

(www.xeno-canto.org) suggest that species-level splits should be

expected in these taxa. Birds on Timor and Alor have a song

consisting ofa ‘high pitched piercing note’ (Coates & Bishop 1997)

similar to the song of birds in the Greater Sundas. This is similar to

songs of some Dicaeum flowerpeckers. However, contact notes

regularly given on Timor and Alor (a buzzing bresh note given

singly, doubly or often repeated) are unlike calls of birds from

Borneo and mainland Asian populations archived at http://

www.xeno-canto.org or http://macaulaylibrary.org/.

Asian Paradise-flycatcher Tersiphone paradisi floris

Alor: There were three observations of this bird in gallery forest at

580-820 m on the slopes of Mt Koya Koya (CRT), and birds were

observed daily and considered as frequent in tropical forest at the

same sites in 2009 (PV), and in forest above Apui-Subo (PV).

Pantar: One bird was observed in vine forest at Desa Batu on 17

April 1991 (REJ). Several birds were observed and sound-recorded

in lowland tropical forest and coastal woodland (PV) and one male

was observed and sound-recorded at 630 m on Mt Wasbila on 10

December 2009 (CRT).

Alor delimits the south-eastern limits of this widespread Asian

bird. The subspecies T. p. floris is known from Sumbawa, Flores,

Besar, Lembata and Alor, with our Pantar records filling an

important distributional gap. Asian Paradise-flycatcher is typically

one of the least common resident birds through its Nusa Tenggara

range (Coates & Bishop 1997, Mees 2006), although can be locally

common on Flores (Butchart et al. 1994, Pilgrim et al. 2000).

Females appear to be particularly inconspicuous, and we only

observed males on Alor and Pantar, matching historical collections

of males from Alor and Lembata (Hartert 1896). All males in Nusa

Tenggara are predominantly white above, whereas a high proportion

of males in South and South-East Asia are rufous (Owen 1964).

Broad-billed Flycatcher Myiagra ruficoilis ruficollis

Alor: Singles or pairs were recorded four times in E. alba woodland

usually next to streams or a dam at 625-750 m about Tanglapui

86

COLIN R. TRAINOR, PHILIPPE VERBELEN & RON E. JOHNSTONE

Forktail 28 (2012)

Timur (CRT), and occasional in mangroves near Kalabahi (CRT),

degraded lorest near Kalabahi (PV) and coastal strand forest on

Sika Island (CRT). Pantar: Regularly recorded in mangroves and

E. alba woodland in September 2009 (PV), and once heard from

secondary shrubland along the road to the Sirung volcano (CRT).

Broad-billed Flycatcher was known from Lembata and Alor in the

Flores island chain, with the Pantar record filling a gap in its

distribution. In the Lesser Sundas it occurs on Timor and most

associated islands, as well as Sumba (Coates & Bishop 1997). It

appears to be generally uncommon to rare on Alor except locally

near water and in mangroves, and on Pantar it appears to be

common in mangroves as on neighbouring Lembata (Trainor 2003)

and Timor (CRT unpubl. data).

Common Golden Whistler

Paeycephala pectoraiis fulvotincta

Alor: This bird was common in tropical dry forest, lantana thickets

and prickly shrubs at Kalabahi and Mali and in secondary forest

and plantations at Apui in April 1991, with eight specimens

collected (REJ). It was observed on several occasions in secondary

forest above Apui-Subo and in evergreen forest at Kunggwera (PV),

and was heard regularly from secondary forest and plantations at

400-900 m near Mainang, and in secondary forest above Apui-

Subo to 1,250 m (CRT). Pantar: Moderately common in vine

forest at Batu and in mangrove at Kabir in April 1991 with two

specimens collected (REJ). Birds were photographed and sound-

recorded in coastal woodland (PV), degraded forest patches in the

lowlands (<200 m) as well as on the slopes of Mt Wasbila and

frequently heard in tropical forest on Mt Wasbila from 100-800 m

and coastal shrubland about Puntaru (CRT).

The Common Golden/Mangrove Whistler P. pectoraiis/

mela.nu.ra complex, with 66 named populations, is one of the most

complex avian examples of geographic variation (Jonsson et al.

2008). The IOC currently treats Alor and Pantar populations as

part of a species called Rusty-breasted Whistler P. fulvotincta (of

Java, Lesser Sundas, Flores Sea islands: Gill & Donsker 2012) but

we prefer to wait until genetic and vocal analyses are published on

the complex to support this split. Common Golden Whistler is

typically one of the most vocal birds in secondary and primary forest

in the Lesser Sundas (Coates & Bishop 1997), although it is

inconspicuous and difficult to observe without the use of playback.

It appears to be mostly absent from Eucalyptus woodland on Alor

and Pantar, with a clear preference for tropical forest. For example,

it was only heard once while walking from Lipa to Lantoka village

(7 km), and heard once on the walk from Lipa to Naumang (CRT).

Cinereous Tit Parus cinereus

Alor: This bird was uncommon to moderately common in gardens

at' Apui in April 1991, when two specimens were collected (REJ).

In 2002, 2009 and 20 1 0 it was occasional to frequent at all sites, in

all wooded habitats, from sea-level to 1,250 m (PV, CRT). Pantar:

Observed in small numbers in tropical dry forest at 200 m (PV)

and common in coastal shrubland about Puntaru village on 14

December 2009, but unrecorded from Mt Wasbila (CRT). Alor

delimits the easternmost occurrence of this bird, which ranges from

Afghanistan, India, mainland South-East Asia, Greater Sundas and

the Lesser Sundas from Lombok along the Flores island chain to

Sumba (White & Bruce 1986). Typically this bird is considered a

common species along the Flores island chain (Butchart et al. 1 994,

Johnstone et al. 1996, Pilgrim et al. 2000, Trainor 2002a, c, Myers

<3c Bishop 2005).

Thick-billed Ftowerpecker Dieaeum agile

Pantar: One bird was collected at Batu (WAM A24474) and several

were observed in canopy of woodland on 19 April 1991 (REJ).

Thick-billed Flowerpecker is typically uncommon to rare (and

inconspicuous with low detectability) throughout its Lesser Sunda

range (Lombok, Sumbawa, Flores, Lembata, Alor and Timor)

(White & Bruce 1986, Johnstone et al. 1996, Coates & Bishop

1997), with only four specimens known for Flores (Mees 2006).

Although collected on Alor by A. FT. Everett (originally

thought to have been a specimen of Golden-rumped Flowerpecker

D. annae ), there have been no subsequent records on the island.

This is the first Pantar island record. Consistent with its general

rarity is the single observation of one bird during a recent seven-

week survey of Wetar (Trainor et al. 2009). This is an aberrant

flowerpecker with recent genetic analyses failing to resolve its

generic affinities, although it may actually be a Prionochilus (Nyari

etal. 2009).

Mountain White-eye Zosterops montanus

Alor: Common at c. 700-1, 271 m, particularly in Eucalyptus

woodland at Mainang, Apui-Subo and Tanglapui Timur (CRT).

Both Mountain White-eye and Ashy-bellied White-eye

Z. citrinellus were present in Tanglapui Timur village, but only

the former was heard during a 9 km walk from Lipa to Lantoka at

700-750 m. At Mainang this bird mostly occurred in Eucalyptus

woodland above 1,150 m, while Ashy-bellied White-eye was

more frequent in secondary tropical forest below c. 1,1 50 m. During

2002 only Ashy-bellied White-eye was discriminated (Trainor

2005a), with Mountain White-eye obviously overlooked.

Mountain White-eye was known from the large Nusa

Tenggara islands of Lombok, Sumbawa, Flores and Timor

(Coates & Bishop 1997) but has recently been recorded on

neighbouring Atauro Island above 800 m (CRT unpubl. data),

and in the mountains ofWetar (CRT unpubl. data). Elevation use

by Mountain and Ashy-bellied White-eye remains poorly

known on Alor, but the latter was widespread on Pantar from

sea-level to c.400 m (PV) and at Batu and Kabir (REJ), and was

recorded down to sea-level on Alor including the islets of

Kepa (REJ, CRT, PV), Ternate and Sika (CRT). About 20

specimens of Ashy-bellied White-eye were collected at Mali and

Apui in 1991 (REJ). Mountain White-eye would be expected to

occur on Pantar but none was definitively heard or seen in

apparently suitable habitat on Mt Wasbila up to 850 m (CRT).

Cryptic taxa are expected to occur within this complex but their

vocalisations are unlikely to help unravel the affinities of various

island forms (Rheindt & Hutchinson 2007). Photos of Alor birds

show they are typical of.Z. m. montanus (as described in Coates &

Bishop 1997) with forehead greenish to yellowish, abdomen

whitish, without a yellow median stripe, and the undertail-coverts

are yellow.

Lemon-bellied White-eye Zosterops ehloris/

Oriental White-eye Z. palpebrosus

Pantar: Lemon-bellied White-eye was observed and sound-

recorded in coastal scrub and secondary forest at 400 m on Mt

Wasbila (PV). In Nusa Tenggara this bird occurs on Lombok,

Sumbawa, Komodo, Rinca, Flores, Besar and Paloe mostly in the

lowlands (White & Bruce 1986). The identity of white-eyes

occurring on islands east of Flores remains poorly known. Records

of Oriental White-eye on Adonara (Trainor 2002a) were queried

by Mees (2006), who considered that Lemon-bellied was more

likely to occur on the island. We were unable to confirm Oriental

White-eye on Pantar, but it may also be present.

Short-tailed Starting Aplonis minor

Alor: Locally common in pairs and small flocks of up to 10 birds

in gallery forest and fruiting trees at Kunggwera and the slopes of

Mt Koya Koya in 2002 (570-1,1 50 m, CRT), with flocks of eight

and 12 birds seen in the same area in June 2009 (PV). An immature

bird with white below and black streaking on the chest was observed

Forktail 28 (2012)

The avifauna of Alor and Pantar, Lesser Sundas, Indonesia

87

near Tanglapui Timur on 3 May 2002 (CRT). This bird was

uncommon above Apui-Subo, with several voice records and flocks

of up to six birds seen (800-1,100 m), but was frequently observed

at Mainang in flocks of 2-8 birds (CRT). Pantar: The first island

records of Short-tailed Starling were of four and six birds observed

in flight near Sirung volcano (PV), and a single voice record from

evergreen forest on Mt Wasbila at 750 m (CRT). Short-tailed

Starling occurs widely as a typically uncommon forest bird in the

Lesser Sundas (Lombok, Suntbawa, Flores, Paloe, Sumba, Timor,

Wetar, Mo.a, Romang: Coates & Bishop 1997), and was recently

confirmed for Atauro Island (Trainor & Soares 2004). The

presence of this bird on Alor and Pantar would be expected,

although it has not yet been recorded from Lembata or Adonara

(Trainor 2002a,c, 2003, Schellekens et al. 2011).

Crimson-hooded (Alor) Myzomela Myzomela ( kuehni )

Alor: A single record of one bird feeding at the flowers of Eucalyptus

alba, at 900 m, on the slopes of Mt Koya Koya in April 2002

(CRT), which was thought to be a misidentification at the time,

with a brief notes crossed out in a notebook (CRT). In 2009 and

2010 there were occasional records of 1-4 birds feeding at flowers

or roosting in E. urophylla. Acacia, Casuarina junghuhniana and

the montane shrub Photinia integrifolia above Apui-Subo village

at 1,150-1,250 m (CRT, PV). Photographs show that the Alor

Myzomela is almost identical in appearance to the nominotypical

Wetar bird, except for the extent of red on the head, neck and chest.

The red hood on the Alor bird is much reduced and only reaches

midway on the hindcrown before sharply changing to grey on lower

hindcrown (nape), mantle, back and scapulars, whereas the red on

the Wetar bird reaches the back of head (c. 1 cm further than the

Alor bird). Alor birds have a red throat, but unlike Wetar birds

this does not extend down to the chest. Substantial differences in

contact notes and song of Alor birds compared to Wetar suggest

that the Alor birds have diverged significantly and represent an

undescribed taxon at either subspecies or species level. Alor birds

appear to be restricted to ‘upper’ montane habitat, although it was

apparently absent from forest at 850-1000 m above Apui where a

suite of other montane species were present, and was also

unrecorded from extensive £. alba woodlands at 700-1,000 m; by

contrast, Wetar birds are common from sea-level to the mountains

(CRT unpubl. data). None was heard or seen on Pantar, but no

habitat above 900 m was surveyed. It is intended to publish further

details on this bird elsewhere.

Common Hill Myrsa Gracula religiosa venenata

Alor: In April 1991 this bird was moderately common in ones and

twos in forest areas around Kalabahi and Mali and in forests at

Apui (REJ). It was locally common about Kunggwera village with

15 birds observed in a large fruiting fig tree on 3 May 2002, and

pairs and small parties were observed in the surrounding tropical

forest (CRT). One flock of six birds was observed in flight at

Kunggwera in June 2009 (PV and VD). One bird was heard calling

from secondary tropical forest about 2 km north of Mainang on 9

January 2010, and two birds were seen and photographed in the

same area on 13 January 2010 (CRT). Pantar: Three specimens

were collected from Batu village and one was collected from forest

at Kabir in April 1991 (REJ). One bird was sound-recorded on the

slopes of Mt Wasbila in September 2009 (PV), and four were

photographed on the slopes of Mt Wasbila and 10-11 (some

possibly double-counted) were heard on 10 December 2009

(CRT).

In Nusa Tenggara the Common Hill Myna occurs from

Sumbawa to Alor (White & Bruce 1986). It is uncommon on

Sumbawa, but moderately common on Moyo Island (Johnstone et

al. 1996), locally common on Flores (Butchart et al. 1994, Pilgrim

et al. 2000) and uncommon to rare on Adonara and Lembata

(Trainor 2003, Schellekens et al. 2011). Populations have

undoubtedly declined precipitously in Nusa Tenggara because of

capture for trade during the 1980s and 1 990s, but the residual birds

on the islands are still being caught. At a market in Kalabahi during

May 2002 nine chicks were available for sale, including two not

fully feathered, three fully feathered (selling for Rp 300,000 [c.

$US30] each) and four fledged subadults (selling for Rp 250,000

each). On Pantar in 2009, local people said that Common Hill

Myna was valued at Rp 200,000- 1 ,500,000 (PV). A total of three

individuals including one bird at Tanglapui (Alor) were recorded

during an undated survey by Widodo (2010). Despite ongoing

trade Alor may still retain some of the best populations of this bird

in Nusa Tenggara (Trainor 2005a).

Wallacean Drang© Biarurus densus bimaensis

Alor: Moderately common in pairs at Kalabahi (one specimen

collected), Mali and Apui mainly in forest but also in plantations

during April 1991 (REJ). One of the most vocal and frequently

observed birds in tropical forest at Kunggwera and Mt Koya Koya

at 570-1,250 m (CRT, PV); above Apui-Subo (900-1,250 m) (PV,

CRT) at Mainang (300-900 m); present in mangroves at the head

of Mutiara Bay, but less common in Eucalyptus woodland (CRT).

Pantar: One specimen was collected at Batu village on 17 April 1991

(REJ). Several observations on the slopes of Mt Wasbila at 400 m

(PV) and frequently recorded there to 800 m (CRT), and in

secondary forest and agricultural land about Puntaru village (CRT).

The Wallacean Drongo is typically one of the most vocal and

conspicuous birds in forest and secondary habitats in Nusa

Tenggara (Coates & Bishop 1997). However, it is worth

highlighting that there are substantial morphological and vocal

differences between the various Lesser Sunda subspecies, with this

subspecies informally named the ‘Lesser Wallacean Drongo’ D.

bimaensis and the bird on Timor and associated islands dubbed

the ‘Greater Wallacean Drongo’ D. densus (Eaton 2009): The

subspecies D. d. bimaensis is moderately common to abundant on

Sumbawa (Johnstone et al. 1996), Flores (Butchart et al. 1994,

Pilgrim et al. 2000), Adonara (Trainor 2002a) and Lembata

(Trainor 2002c), and was recently noted on Solor (Schellekens et

al. 2011). It is under no threat of extinction.

Eurasian Tree Sparrow Passer montanus

Alor: This introduced bird was very common in the large town of

Kalabahi during 2009 and 2010 (PV, CRT). Pantar: During

September 2009, it was observed and sound-recorded at Baranusa

(PV). At least three birds were noted at Puntaru village at the

church (CRT). This bird has become widely naturalised

throughout Nusa Tenggara over the last 30 years. None was noted

on Alor during 1991 (REJ) or 2002 (CRT) but birds are thought

to have been present on Flores since about 1955 (Mees 2006). They

were locally abundant on Adonara during December 2000 (T rainor

2002a), absent from Lembata in December 2000 (Trainor 2002c,

2003) and abundant there in 2009 (Schellekens et al. 2011).

DISCUSSION

Our records substantially improve knowledge of the avifaunas of

both Alor and Pantar. We expected to add many new island records

to the Pantar Island list because it was poorly surveyed historically,

with little information published on the island since the nineteenth

century, but the addition of 37 (including 27 resident terrestrial)

bird species to the Alor list was surprising. The continued

accumulation of new records for both islands by ornithologists in

2011 suggests that more species will be added in the future. We also

show that Alor and Pantar have greater levels of endemicity than

previously known, and the presence of the Critically Endangered

88

COLIN R. TRAINOR, PHILIPPE VERBELEN & RON E. JOHNSTONE

Forktail 28 (2012)

Flores Hawk Eagle and additional endemic and restricted-range

birds elevates the global conservation significance of these islands.

Both islands were assumed to be dominated by a ‘Flores ornis’

(Hartert 1898) or possess avifaunas ‘evidently derived from Flores’

(White & Bruce 1986), but our fieldwork shows that the avifaunal

origins of these islands is more complex. Alor demarcates the eastern

limits of a suite of well-known and ‘typical’ Asian bird species (e.g.

Ruddy Cuckoo Dove, Rusty-breasted Cuckoo Cacomantis

sepulcralis, Sunda Pygmy Woodpecker, Golden-bellied Gerygone,

Cinereous Tit and Asian Paradise-flycatcher), but many species

typical of the Timor-group islands were also added to the list (e.g.

‘Little Cuckoo Dove’, Common Cicadabird, Timor Bush Warbler

and Timor Stubtail) and other widespread Asian birds which appear

to be most closely related to Timor birds (Yellow-breasted Warbler,

Little Pied Llycatcher and Lesser Shortwing).

The discovery of Timor Bush Warbler on Alor is surprising on

biogeographical grounds. Some variation in song characteristics of

this species have been published (Trainor et al. 2012) but further

advances in determining the affinities of this bird in relation to other

members of the complex, including those from Timor, Bali and

Java, will rely on the collection of specimens and molecular analyses.

Timor Bush Warbler is currently considered Near Threatened, but

the Alor population appears under little threat. Only two

populations were discovered, highlighting that this bird is patchily

distributed in suitable montane habitat. The absence of records

during a 17 km walk between the two populations also highlights

the high degree of habitat specificity of the species (although

significantly this pattern was mirrored by a suite of other passerines

that are forest-dependent). Although it requires tall grass and low

shrub, this needs to be in close proximity to tropical forest because

it avoids extensive Eucalyptus woodland. Interestingly, the opposite

was the case on Timor, where the species was recorded from

landscapes dominated by Eucalyptus woodland with a grassy

understorey on the slopes of Mt Ramelau (Trainor et al. 2012).

The presence of an endemic subspecies of Sunda Bush Warbler,

and a likely endemic subspecies of Timor Stubtail also highlight the

importance of Timor as a source of Alor’s avifauna. The Timor

Stubtail is a diminutive bird which is most often seen walking on the

forest floor, but new island records for Alor, Roti, Atauro and Romang

attest to its strong powers of flight and capacity for island

colonisation. The song of this bird has a similar structure on all islands

(including Babar: PV and CRT unpubl. data) but these scattered

populations are presumably at least subspecifically distinctive.

The distinctiveness of endemic subspecies of Lesser Sunda

populations of Southern Boobook has been the subject of much

speculation (Johnstone & Darnell 1997b, Olsen et al. 2010,

Verbelen 2010) but only recently have sound recordings of

vocalisations started to help unravel the apparent hidden levels of

endemism in this complex. The calls of the Southern Boobook on

Alor and Pantar are among the most distinctive in the complex,

indicating that this form there should be considered a full species.

The ‘Alor Boobook’ occurs in tropical forest and Eucalyptus

woodland and probably is widely distributed on both islands, and

therefore under little threat of extinction. A study is underway on

the vocalisations of this complex including samples from most

Lesser Sunda islands (F. Rheindt and JE in litt. 2012). The Otus

owls have also harboured substantial hidden diversity in south-east

Asia (e.g. Lambert & Rasmussen 1998). We are unable to confirm

the presence of an Otus owl on Pantar. More fieldwork on Pantar

targeting this bird would be of value.

Further work is also needed to determine the affinities of the

Myzomela honeyeater recorded on Alor. In appearance it is

obviously close to Crimson-hooded Myzomela of Wetar, but its

calls and songs are different. This is of biogeographical interest

because it may be the only clear example of a direct relationship

between the avifaunas of Alor and Wetar.

Despite our efforts, the avifaunas of both Alor and Pantar

remain poorly known. The status of the Flores Hawk Eagle

particularly deserves attention. It would be valuable to produce a

population estimate, assess the degree to which it is restricted to

forested habitats and determine whether it is also present on islands

such as Lembata, Pan tar and Wetar. There has been little attention

given to the Yellow-crested Cockatoo in Nusa Tenggara since the

late 1990s, despite ongoing illegal bird trade which has probably

resulted in further population decline. Further effort on Alor,

Pantar and other islands would be of value to provide updated

information on its conservation status. A study of the status of the

Flores Green Pigeon on Alor, Pantar and other islands in its range

would be similarly valuable. The taxonomic status of several bird

species on Alor (e.g. ‘Little Cuckoo Dove’, Timor Bush Warbler,

‘Crimson-hooded Myzomela’ and perhaps several other montane

species) is poorly understood, with vocal or molecular studies now

needed to clarify their affinities, and further fieldwork needed to

define their distribution, habitat use and conservation status.

ACKNOWLEDGEMENTS

CRT thanks Heri Ndolu and Neny Herdianawati for logistical and/or field

assistance during the 2002 survey, Ian Cowie (Northern Territory Herbarium)

for identifying some plants, Mike Carter for information on the status of Pacific

Koel on Ashmore Reef and Richard Noske for information on the status of Arctic

Warbler in West Timor. PV thanks Veerle Dossche for companionship during

his first Alor visit. REJ thanks Dr Soetikno Wiroatmodjo of LIPI and Dr D. J.

Kitchener of WAM for facilitating the fieldwork on Alor and Pantar islands

and Mr N. Kolichis whose grants to WAM funded some of the field costs. We

also thank James Eaton, Rob Hutchinson and Peter Collaerts for their field

records and other information which confirmed the presence of several species

on the islands, and Bas van Balen for confirming the presence of Ruddy Cuckoo

Dove on Alor from a sound recording. Thanks to Nick Brickie and James Watson

for suggesting improvements during peer-review of the manuscript.

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