'IqS 5-^0( &

Journal of Asian Ornithology

m

Forktail 29(2013)

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Cover picture: Cambodian Tailorbird Orthotomus chaktomuk, Highway 61, near Prek Kdam, Kandal province, Cambodia, 21 November 2012

by James A. Eaton

ISSN 0950-1746

© Oriental Bird Club 201 3

The Oriental Bird Club is a Registered Charity No. 297242

FORKTAIL

Number 29, 201 3

CONTENTS

S. P. MAHOOD, A. J. I. JOHN, J. C. EAMES, C. H. OLIVEROS, R. G. MOYLE, HONG CHAMNAN, C. M. POOLE, H. NIELSEN & F. H. SHELDON

A new species of lowland tailorbird (Passeriformes: Cisticolidae: Orthotomus) from the Mekong floodplain of Cambodia . 1

N.J. COLLAR &S. van BALEN

Notes for the conservation of the Rufous-fronted Laughingthrush Garrulax rufifrons . 1 5

N. J. COLLAR, J. A. EATON & R. O. HUTCHINSON

Species limits in the Golden Bulbul Alophoixus ( Thapsinillas ) affinis complex . 19

VLADIMIR YU. ARKHIPOV, TOM NOAH, STEFFEN KOSCHKAR & FYODOR A. KONDRASHOV

Birds of Mys Shmidta, north Chukotka, Russia . . . 25

PAUL J. LEADER, GEOFF J. CAREY & PAUL I. HOLT

Species limits within Rhopophilus pekinensis . 31

PAVEL KVARTALNOV, ABDULNAZAR ABDULNAZAROV, VERONIKA SAMOTSKAYA, JULIA POZNYAKOVA, IRINA ILYINA,

ANNA BANNIKOVA & EUGENIA SOLOVYEVA

Nesting of the Large-billed Reed Warbler Acrocephalus orinus: a preliminary report . 37

FRANCESCO GERMI, AGUS SALIM & ANDREA MINGANTI

First records of Chinese Sparrowhawk Acc/p/ferso/oens/s wintering in Papua (Indonesian New Guinea) . 43

SHIAO-YU HONG, YUAN-HSUN SUN, HSIN-JU WU & CHAO-CHIEH CHEN

Spatial distribution of the Tawny Fish Owl Ketupa flavipes shaped by natural and man-made factors in Taiwan . 48

M. MONIRUL H. KHAN

Population, breeding and threats to the White-rumped Vulture Gyps bengalensis in Bangladesh . 52

JOHN D. FARRINGTON, ZHANG XIULEI & ZHANG MIN

The birds of the Longbao National Nature Reserve and surrounding basin, Yushu county, Qinghai, China . 57

LI-HU XIONG & JIAN-JIAN LU

Habitat specialisation in the Reed Parrotbill Paradoxornis heudei — evidence from its distribution and habitat use . 64

MIYUKI MASHIKO& YUKIHIKOTOQUENAGA

Increasing variation in population size and species composition ratio in mixed-species heron colonies in Japan . 71

P. C. RASMUSSEN & N. J. COLLAR

Phenotypic evidence for the specific and generic validity of Heteroglaux blewitti . 78

H. EDEN W. COTTEE-JONES, JOHN C. MITTERMEIER & DAVID W. REDDING

The Moluccan Woodcock Scolopax rochussenii on Obi Island, North Moluccas, Indonesia: a 'lost' species is less

endangered than expected . 88

PER ALSTROM, GANG SONG, RUIYING ZHANG, XUEBIN GAO, PAUL I. HOLT, URBAN OLSSON & FUMIN LEI

Taxonomic status of Blackthroat Calliope obscura and Firethroat C. pectardens . 94

COLIN R.TRAINOR, STEPHEN J. S. DEBUS, JERRY OLSEN, JANETTE A. NORMAN & LES CHRISTIDIS

Bonelli's Eagle Aquila fasciata renschi in the Lesser Sundas, Wallacea: distribution, taxonomic status, likely origins

and conservation status . 100

EARL OF CRANBROOK, GOH WEI LIM, LIM CHAN KOON & MUSTAFA ABDUL RAHMAN

The species of white-nest swiftlets (Apodidae, Collocaliini) of Malaysia and the origins of house-farm birds:

morphometric and genetic evidence . 107

SOMSAK BUATIP, WANCHAMAI KARNTANUT & CORNELIS SWENNEN

Nesting period and breeding success of the Little Egret Egretta garzetta in Pattani province, Thailand . 1 20

SIMON P. MAHOOD, SEBASTIEN DELONGLEE, FLORIAN KLINGEL, FALK WICKER & RICHARD CRAIK

The status of Brown-chested Jungle Flycatcher Rhinomyias brunneatus in Vietnam . 124

JOHN C. MITTERMEIER, H. EDEN W. COTTEE-JONES, ENDANG CHRISTINE PURBA, NOVA MAULIDINA ASHURI,

EKA HESDIANTI & JATNA SUPRIATNA

A survey of the avifauna of Obi island, North Moluccas, Indonesia . 1 28

NATURAL HISTORY

MUSEUM LIBRARY

26 NOV 2013

PURCHASED

Forktail 29 (2013)

Short Notes

ASHOKA JAYARATHNA, SARATH W. KOTAGAMA & EBEN GOODALE

The seasonality of mixed-species bird flocks in a Sri Lankan rainforest in relation to the breeding of the nuclear species.

Orange-billed Babbler Turdoides rufescens . . . 138

T.M. BRAILE&K. WINKER

New distributional records of Philippine birds from Bohol, Mactan, Olango, Busuanga and Luzon islands . 140

MERWYN FERNANDES & JAN WILLEM DEN BESTEN

Some interesting breeding records for Pong Dam Wildlife Sanctuary, Himachal Pradesh, India . 141

TARIQ MAHMOOD, SYED M. USMAN-UL-HASSAN, MUHAMMAD S. NADEEM & AMJAD R. KAYANI

Population and diet of migratory Common Starlings Sturnus vulgaris wintering in agricultural areas of Sialkot

district, Pakistan . 143

EUN-MI KIM, CHANG-YONG CHOI & CHANG-WAN KANG

Causes of injury and mortality of Fairy Pitta Pitta nympha on Jeju Island, Republic of Korea . 1 45

A. MOHAMED SAMSOOR ALI, S. RAMESH KUMAR & P. R. ARUN

House Crow Corvus splendens nesting on pylons, Kutch district, Gujarat, India . 148

MARKR. BEZUIJEN

New waterbird count data from the Heihe river in Gansu province, western China . . 1 50

SHAOBIN LI, WEIJUN PENG, CHENG GUO & XIN LU

Breeding biology of the Small Snowfinch Pyrgilauda davidiana on the Tibetan plateau . 1 55

NATARAJAN EZHILARASI & LALITHA VIJAYAN

Nest, eggs and nest sites of the Andaman Crake Rallina canningi . 1 58

TERESA M. PEGAN, JACK P. HRUSKA & JUSTIN M. HITE

A newly described call and mechanical noise produced by the Black-and-crimson Pitta Pitta ussheri . 160

HUGH L. WRIGHT, SOK KO, NET NORIN & SUM PHEARUN

White-shouldered Ibis Pseudibis davisoni population size and the impending threat of habitat conversion . 162

Errata . 165

Editorial notes . 166

Guidelines for contributors

inside back cover

FORKTAIL 29 (2013): 1-14

A new species of lowland tailorbird (Passeriformes:

Cisticolidae: Orthotomus) from the Mekong floodplain

of Cambodia

S. P. MAHOOD, A. J. I. JOHN, J. C. EAMES, C. H. OUVEROS, R. G. MOYLE, HONG CHAMNAN, C. M. POOLE,

H. NIELSEN &F. H. SHELDON

Based on distinctive morphological and vocal characters we describe a new species of lowland tailorbird Orthotomus from dense humid

lowland scrub in the floodplain of the Mekong, Tonle Sap and Bassac rivers of Cambodia. Genetic data place it in the 0. atrogularis-O.

ruficeps-O. sepium clade. All data suggest that the new species is most closely related to O. atrogularis, from which genetic differences are

apparently of a level usually associated with subspecies. However the two taxa behave as biological species, existing locally in sympatry

and even exceptionally in syntopy, without apparent hybridisation. The species is known so far from a small area within which its habitat is

declining in area and quality. However, although birds are found in a number of small habitat fragments (including within the city limits of

Phnom Penh), most individuals probably occupy one large contiguous area of habitat in the Tonle Sap floodplain. We therefore recommend

it is classified as Near Threatened on the IUCN Red List. The new species is abundant in suitable habitat within its small range. Further work

is required to understand more clearly the distribution and ecology of this species and in particular its evolutionary relationship with O.

atrogularis.

INTRODUCTION

After a hiatus of over half a century owing to the intense human

conflicts in the area, the last two decades have witnessed the

discovery of a flush of novel bird taxa in Indochina. These recent

discoveries have been facilitated by better sampling of remote micro¬

habitats and to a much lesser degree the greater use of non-

morphological characters in delimiting species. Most of these

discoveries concerned babblers (Timaliidae) from isolated montane

areas in Vietnam (Eames et al. 1994,Eames et al. 1999a, b, Eames&

Eames 2001, Eames 2002). A smaller wave of discoveries involving

a diverse range of taxa took place in forested limestone karst in Lao

PDR, Vietnam and adjacent areas of China (Zhou Fang & Jiang

Aiwu 2008, Woxvold et al. 2009, Alstrom et al. 2010). Only one

new species, Mekong Wagtail Motacilla samveasnae, was named

from Cambodian specimens, but it also occurs in Lao PDR,

Thailand and Vietnam in ‘channel mosaic’ habitat on the Mekong

and its major tributaries (Duckworth et al. 2001, Le TrongTrai &

Craik 2008). Here we describe a new species of lowland tailorbird

Orthotomus , confined to low elevation humid evergreen scrub in

the floodplain of the Mekong and associated large rivers, in

Cambodia.

THE NEWTAILORBIRD

During routine sampling of birds for avian influenza in 2009, four

individual tailorbirds Orthotomus sp. were mist-netted and

photographed in the hand: one on 28 and another on 29 January

2009 in a patch of scrub near a pond at Kraing Check, Kandal

province (l l°4l'53.36"N 104°46'38.93,,E) (J. Reside per F. Goes

in litt. 2012), one on 24 February 2009 and another on 12 March

2009 near to Phnom Tamao Zoo, Takeo province (lCH'Sh^N

104°50'22"E), in 3-5 m high scrub near rice fields (HN, A. Yang

103°30'Q"E 104°0'0"E 104°30'0"E 105°0,0,,E 1O5°30'O"E 106°0,0"E 106°30'0"E 107°0,0"E

Figure 1. Distribution of records of

Orthotomus chaktomuk and O.

atrogularis within and close to the

floodplain of the Mekong, Tonle Sap

and Bassac rivers.

2

S. P. MAHOOD etal.

Forktail 29 (2013)

and P. Joyner in litt. 2012). Based on photographs and inferences

drawn from incorrect location data (which misleadingly indicated

that the birds had been caught close to the coast) these individuals

were incorrectly identified as Ashy Tailorbird O. ruficeps-, the

possibility of their representing aberrant Dark-necked Tailorbird

O. atrogularis was considered and rejected based on general plumage

similarity to O. ruficeps (F. Goes verbally 2012).

On 29 January 2012, HN found a similar bird at PrekKsach, in

a partially flooded construction site c.15 km from Phnom Penh

(1 1°4T37.14"N 104°53'43.79"E) and, owing to similarity to the

2009 birds, assigned it to O. ruficeps. In earlyjune 20 1 2, photographs

by AJIJ of an Orthotomus sp. from this site raised the interest of

SPM. Subsequent field observations by SPM, AJIJ and T. D. Evans

and discussion with J. W. Dqckworth, P. D. Round, CMP and C.

Robson indicated to SPM that these birds might not be 0. ruficeps,

but could perhaps represent an undescribed taxon. On 23 June 2012,

SPM, HN and AJIJ searched for additional individuals at Prek Ksach

and located five single males and two pairs. Between 23 June 2012

and 20 April 2013, intensive searches revealed at least 1 00 individuals

at nine additional locations (Figure 1, Table SOM 1 [supplementary

online material — see page 14]). Seven morphologically typical O.

atrogularis comprising six males and one female were seen at five

floodplain locations at or within 10 km of locations where birds of

the new taxon were found (Figure 1, Table SOM 1).

From photographs SPM re-identified all of the individuals mist-

netted in 2009 as conforming to the new taxon, thus temporarily

removing O. ruficeps from the list of birds recorded in Cambodia.

Typical O. ruficeps has since been recorded in mangrove forest in

coastal Cambodia close to the border with Vietnam (Mahood &

Martin 2013).

In August 2012 two adult males, one immature male and two

immature females (aged by plumage, sexed internally) were collected

for formal description (below). SPM was later able to compare these

specimens directly with the Orthotomus material held at the Natural

History Museum, Tring, UK (NHMUK) including a syntype

(NHMUK 1 886. 10.1. 1830) of O. atrogularis nitidus (the subspecies

in Cambodia) and the holotype of Olive-backed Tailorbird O.

sepium (NHMUK1 880. 1.1. 4473), and also examined specimens at

Naturalis Biodiversity Centre, Feiden, Netherlands (RMNH)

including the holotype of O. ruficeps cineraceus (RMNH 137559).

(The disjunct population of 0. ruficeps in coastal southern Vietnam

and Cambodia has not been assigned to a subspecies; however, O. r.

cineraceus is the subspecies recorded in mainland Asia: Madge 2006).

All other Orthotomus taxa differed so extensively from the new form

01 23456789 10

Plate 1. Dorsal, ventral and lateral views of the holotype of 0.

chaktomuk.

that detailed comparison would be superfluous. A detailed list of

all specimens examined is provided in Table SOM 2.

The new taxon shows significant morphological differences

from its close relatives and is sympatric with two lowland tailorbird

species, O. atrogularis and Common Tailorbird (). sutorius , without

signs of intergradation. We therefore consider that it represents a

new species, which we name:

Orthotomus chaktomuk, sp. nov.

Cambodian Tailorbird

http://zoobank.Org/urn:lsid:zoobank.org:act:23E9A09C-AD9C'

4346-A594-F 1 87DAFB60 1 3

Holotype and paratypes

Study skins deposited in NHMUK (Table 1, Plate 1, Plate 2a-c)

were collected by JCE and SPM at Bateay District, Kompong Cham

Table 1 . Mensural and other relevant data of holotype and paratypes (in mm, except mass in g).

M, Male; F, Female; ad, adult; lyr, first calendar year. State of wing moult lists feather tracts recently replaced; P, primaries; R, right wing; L, left wing.

HARRY TAYLOR © NATURAL HISTORY MUSEUM, LONDON

HARRY TAYLOR © NATURAL HISTORY MUSEUM, LONDON

Forktail 29 (2013) A new species of lowland tailorbird ( Orthotomus ) from the Mekong floodplain of Cambodia

3

(b)

V '* _ 2

,KIT 3 H i

V.lWotv TD ■-

SfrAi-i Toe j

0123456789 10

Plate 2. Specimens of 0. chaktomuk: (a) dorsal, (b) ventral and (c) lateral

views of the holotype and four paratypes (from left to right NHMUK

2012.9.1, 2012.9.2, 2012.9.3, 2012.9.4, 2012.9.5).

province, Cambodia (1 1°56'53.94,'N 104°56'50.94"E), c.43 km

north of Phnom Penh at c. 1 5 m elevation on 8 and 9 August 2012,

and prepared by JCE. Tissue samples from the same individuals

were deposited in Louisiana State University Museum of Natural

Science, Baton Rouge, Louisiana, USA (Table 1). Holotype

(NHMUK 2012.9.1): adult male; in active wing moult; one

large testis (left testis 4.3 mm length, right testis 2 mm length).

Paratypes aged by plumage: one adult male (NHMUK 2012.9.2);

one immature male (NHMUK 2012.9.3) with unmoulted

rectrices olive-green fringed; one immature female (NHMUK

2012.9.4) with some retained greater coverts fringed olive-green

indicating immaturity, all other rectrices as adult; and one

immature female (NHMUK 2012.9.5) with unmoulted rectrices

as immature male.

Diagnosis of species

Head: in male entirely rich cinnamon-rufous crown and

contrasting white cheeks, very similar to O. atrogularis , differing

from 0. ruficeps and O. sepium in cheek colour (Table 2, Plate 3a-

c). Rufous of crown less extensive in female. UpperpartS and

wings: mid-grey in adult, superficially similar to those of 0. sepium

but lacking olive tones, strikingly different from O. atrogularis

which is yellowish-green (Table 2, Plate 3a, c); tail with dark grey

subterminal band and whitish tips when fresh. UnderpartS: pale

Table 2. Qualitative summary of plumage of Orthotomus chaktomuk and closely related species (all adult male).

HARRY TAYLOR © NATURAL HISTORY MUSEUM, LONDON

4

S. P. MAHOODefa/.

Forktail 29 (2013)

grey ground colour with profuse blackish throat-streaking in males

(largely absent in females) with white drop-shaped marks, extensive

mid-grey on flanks, and white vent; underparts of both sexes

superficially similar to those of respective sexes of O. atrogularis

owing to throat-streaking, but greyer on flanks and vent white, O.

ruficeps and O. sepium lacking throat-streaking in both sexes (Table

2, Plate 3b, c); further distinguished from other members of the

genus by whitish-cinnamon thighs. Vocalisations : loud, lengthy,

complex and highly varied. Very similar to O. atrogularis.

Compared to O. atrogularis , phrases are given at a quicker pace

and the gaps between phrases are shorter. Subjectively, these

characteristics mean that the vocalisations of O. cbaktomuk sound

faster and more complicated than those of O. atrogularis.

Sexing and ageing

Based on field observations (Plate 4a-k, Media Files SOM 1-3)

and specimens (Plate 2a-c), female O. cbaktomuk can be

distinguished from males by paler cinnamon-rufous on crown,

which is restricted to forecrown and sides of mid-crown (in lateral

view this appears as a short cinnamon-rufous supercilium), paler

grey upperparts and wings and whitish underparts with usually faint

dark streaking. The latter is usually evident only at the edges of the

throat/upper breast, although some (possibly older birds) show

stronger and more extensive streaking on throat and breast. Even

in these extreme individuals, the degree of female streaking does

not approach that in males (Plate 4a-g). All three immature

paratypes show shorter tails than adults (Table 1). Immature birds

possess bright yellowish-olive fringing to the wing-feathers (Plate

4h), which are moulted during August and replaced with grey adult-

type feathers (Plate 4i-j). Immatures are browner (slightly olive)

above and paler below, with reduced streaking (Plate 4h— j). Wing-

feathers of subadults appear as in adults, except sometimes they

retain yellowish-olive-fringed greater coverts (Plate 4k). Overall,

subadults resemble adults, but are paler and less heavily marked

below. In adults, there is individual variation in colour tone of grey

feathering above and below, and intensity of throat-streaking (e.g.

Media Files SOM 1-3). It is unknown if this is age-related.

Description of species

The detailed description below was completed in the NHMUK

based primarily on the prepared specimens (Plates 1-2),

supplemented by information from individuals observed and

photographed in the field (Plate 4). It refers to the holotype unless

otherwise stated. Although moult of body feathers was almost

complete when specimens were collected, all adult specimens

retained a few head, throat or breast feathers in pin. The holotype

and paratypes were in wing moult. Moult of wing feathers is

complete by late August and followed immediately by moult and

replacement of tail feathers, which were very worn in all specimens.

Subjective colour assessments of plumage are, where possible,

followed by a formal colour classification taken from Smithe (1975).

Head and face

Crown from forehead to nape, lores, and feathers on orbital ring

and just behind eye rich cinnimon-rufous (136 Raw Sienna)

(slightly richer-coloured in the adult male paratype); hindcrown

slightly darker and more brownish (23 Raw Umber). Crown

feathers in moult with newer feathers slightly richer rufous. On

the immature female paratypes the crown is less richly coloured

than that of the holotype (240 Kingfisher Rufous) and the

cinnamon-rufous lores and feathering on the orbital ring and

immediately behind the eye are replaced by rufous-buff( 1 18 Warm

Buff). The rufous crown feathering extends from the bill only as

far back as the anterior of the mid-crown where dark-grey feathers

predominate, imparting an overall greyish-brown colour (129 Dark

Brownish Olive) to the hindcrown.

Plate 3. Specimens of the haiotype of Orthotomus chaktomuk and

closely related species (all males); (a) dorsal, (b) ventral and (c) lateral

views (from left to right of 0. chaktomuk, O. sepium sepium, 0. ruficeps

cineraceus and 0. atrogularis nitidus).

PHOTOGRAPHS BY J. EATON (a,b,f), J. C. EAMES (c,d,j,k), A. J. I. JOHN (e,g,h,i)

Forktail 29 (201 3) A new species of lowland tailorbird (Orthotomus) from the Mekong floodplain of Cambodia

5

Plate 4. Orthotomus chaktomuk (a-b) adult males in fresh plumage, 21 November 2012; (c-d) adult male in active moult (holotype),

8 August 2012 (NHMUK 2012.9.1); (e) adult male in worn plumage, 29 July 2012; (f) adult female in fresh plumage, 21 November 2012;

(g) adult female in worn plumage, 29 July 2012; (h) immature male pre-moult, 16 July 2012; (i) immature male in active moult, 29 July

2012; (j) immature female in active moult, 8 August 2012 (NHMUK 201 2.9.5); (k) sub-adult female in active moult, 9 August 201 2 (NHMUK

2012.9.4;).

6

S. P. MAHOODeto/.

Forktail 29 (2013)

Five blackish rictal bristles per side, anterior two c.3 mm, twice

the length of posterior three. Ear-coverts, cheeks and moustachial

stripe almost white contrasting strongly with crown and

underparts; however, feathers have buff (124 Buff) tips imparting

an off-white wash. Feathers of submoustachial stripe and malar

stripe white with very dark grey (82 Blackish Neutral Gray) bases

and sometimes tips and fringes; white predominates, giving an

impression of white speckling on a blackish base and contrasting

strongly with the whitish cheeks. The malar stripe on the immature

paratypes is quite different to that of the adult male specimens. It

is made up of white feathers with pale grey central portions (85

Light Neutral Gray) and therefore contrasts little with the cheeks.

Upperparts

Boundary between hindcrown and upper neck abrupt. Upper neck,

mantle and rump concolorous mid-grey (84 Medium Neutral

Gray), slightly blue-toned approaching 78 Plumbeous (all feathers

fresh and body moult apparently completed). Feathers on mantle

and particularly rump relatively long and filamentous.

Wings

Wings of all prepared specimens in active moult (Table 1). On all

specimens, fresh adult feathers are slightly darker grey (83 Dark

Neutral Gray) than mantle, tinged very slightly brownish with mid¬

grey (84 Medium Neutral Gray) fringing (slightly broader on outer

webs). Fresh primaries with off-white inner webs; worn adult

rectrices buffy-brown (239 Ground Cinnamon) lacking fringing

or pale webs. Underside of remiges dull silver-grey (84 Medium

Neutral Gray). Underwing-coverts paler grey (85 Light Neutral

Gray). Alula and axillaries contrastingwhite. Unmoulted rectrices

of the immature male paratype and immature female paratypes

differ strikingly from those of adult male specimens in being fringed

bright olive-green (50 Yellowish Olive-Green).

Tail

Slightly rounded, outermost pair of rectrices 7 mm shorter than

central pair. T ail of holotype very worn, buffy-brown (239 Ground

Cinnamon), dorsal side slightly darker than ventral but heavily

worn. Whitish-buff terminal tips just visible on all but central

rectrices. Tail of immature female paratype (NHMUK 2012.9.4)

less worn than that of the holotype (and other paratypes) and is

dark greyish-brown (21 Fuscous) with broader whitish tips than

those shown by other specimens. Field observations indicate that

fresh tail feathers are mid-grey (similar in colouration to fresh wing

feathers and therefore probably 83 Dark Neutral Gray or 84

Medium Neutral Gray) with a blackish-grey subterminal band (c. 1

cm wide) and whitish tips.

Underparts

The holotype shows white chin feathers with very dark grey (82

Blackish Neutral Gray) bases, tips and fringes, therefore darker

overall than feathers on malar stripe, the latter overhanging those

on throat. In the holotype, feathers of throat in an advanced stage

of moult, some feathers in pin visible. Throat similar in colouration

to chin although with much less white; feathers almost entirely

solid dark grey (82 Blackish Neutral Gray) gradually becoming

darker towards the breast (83 Dark Neutral Gray) with some white

tips throughout. On the breast some dark grey feathers (83 Dark

Neutral Gray) possess contrasting white rachis and base of barbs

on distal two-thirds of feather, creating a pattern of whitish drops

on a mid-grey background. On the edges of the breast, solid mid¬

grey (84 Medium Neutral Gray) feathers predominate. On the

adult male paratype (NHMUK 20 12.9.2), the whitish drop-shaped

marks on the breast are better developed than on the holotype and

extend onto the throat, perhaps because the darker fringes are more

worn. Field observations indicate that there is variation in the

extent and intensity of dark throat-streaking in males (Media tiles

SOM 1-2). The boundary between breast and belly is gradual;

feathers tend towards lighter grey on belly (86 Pale Neutral Grey)

and flanks (85 Light Neutral Gray). Flank feathers are relatively

long. Vent greyish-white (paler than 86 Pale Neutral Gray). Thighs

whitish-cinnamon (6 Salmon).

The underparts of the immature male paratype (NHMUK

2012.9.3) differ from those of the adult male holotype in being

paler with reduced dark grey on the throat and upper breast. There

is an almost complete lack of dark tones on the throat, and the

very dark grey (82 Blackish Neutral Gray) area on the throat is

much smaller and barely extends onto the breast. On the throat

and breast, feathers with white shafts and distal portions are more

abundant than on the holotype, giving the throat a more speckled

appearance. On the breast, solid white and pale grey feathers

predominate such that the overall colour is whitish-grey (86 Pale

Neutral Gray) rapidly grading to off-white on the belly. Flank

feathers of the immature male are slightly whiter than those of the

holotype. The underparts of the immature female paratypes are

even paler than those of the immature male and almost completely

lack dark tones. In those two specimens the chin and throat are

white. Although there is a small area of mid-grey (84 Medium

Neutral Gray) on the sides of the upper breast and the flank feathers

are pale grey (86 Pale Neutral Gray or 85 Light Neutral Gray), the

underparts are otherwise off-white.

Bare parts

Upper mandible dark horn, lower mandible pink horn, paler and

pinker at base (more extensively pink on adult male paratype). Bill

slender. Culmen decurved close to tip, not strongly carinated, tip

very slightly hooked. Gonys convex. Tarsus and toes pinkish

(slightly darker in adult male paratype, paler in immature male

paratype); soles of the feet pale pink. Claws pale brownish pink,

becoming paler towards tips. On female paratypes tarsi, feet, soles

and lower mandibles are paler than those of the holotype. Iris

orange-brown. Inside of mouth pale pink.

Description of vocalisations

For clarity we use the following terminology to describe

vocalisations: note - a single song element; strophe - a continuous

flow of notes, separated from other strophes by silent pauses; phrase

- one or more strophes given in quick succession; and song - one

or more phrases given in quick succession; strophe pace - number

of notes per strophe/strophe length; phrase pace (for phrases with

more than one strophe) - phrase duration/strophes per phrase.

Note that recordings varied in length and quality, so only those

with good quality strophes were analysed.

Male O. chaktomuk songs are lengthy, often lasting more than

one minute (Figure Slo-s, Slu, Media Files SOM 1-6). They

consist of multiple phrases repeated at intervals of 0.42-4.30

seconds, typically much shorter than the maximum interval (mean:

1.7 seconds). Phrases are made up of 2-5 strophes, which are given

at 0.12-0.95 second intervals. Males occasionally switch to a

different strophe type mid-way through a song, although not within

the same phrase. Strophes are also sometimes given singly. Strophes

are trilled, consist of 3-18 notes and typically last 0. 17-0.49 seconds

(Table 3). Twelve distinct male strophe types are known, ranging

from up, down or ‘overslurred’ (the latter referring to sequences of

notes that rise and then fall) trills (often with a louder initial or

terminal note) to a mix of trilled notes and upslurs, downslurs or

‘overslurs’ (Table 3, Figure S la-1, Media Files SOM 1-6). Within

strophe type, number of notes varies slightly (Table 4).

Female O. chaktomuk vocalisations are typically emitted whilst

the male is vocalising, but are sometimes given between male

vocalisations (Figure Slo-s, Slu, Media File SI -6). Females give a

stereotyped trill at a higher frequency than male vocalisations

Forktail 29 (2013) A new species of lowland tailorbird ( Orthotomus ) from the Mekong floodplain of Cambodia

7

Table 3. Transliterations and univariate summary statistics of measurements of strophe characteristics of Orthotomus chaktomuk.

Analyses based on 12 recordings of male vocalisations from six pairs of Orthotomus chaktomuk in Kandal province, Cambodia, obtained as follows: (1) four pairs from c.40 km south-east of Phnom Penh (at or very

close to 1 1°19'45.77"N 105°11'48.41"E);(2) one pair from c.15 km north of Phnom Penh (11°41'37.14"N 104°53'43.79"E); and (3) one pair from c.30 km north of Phnom Penh (1 1°50'18.24"N 104°49'26.55”E).

Measurements taken in Raven Pro 1 .4 (Raven 2012). Values given are: minimum-maximum (mean; sd; sample size). In vocal transcriptions, notation follows Rasmussen & Anderton (2005).

(typically 5-16 notes lasting 0.24-0.84 seconds; Table 3, Figure

Sim, Media File S 1 — 6) . Females, and exceptionally males,

sometimes produce a nasal squeak consisting of a single note with

harmonics (Figure Sin, Media File S6). This vocalisation is usually

given singly (Figure Sip), but occasionally more than one is

repeated in quick succession; when many squeaks are given in

sequence the first is usually longer than others (Figure Sit, Media

File S6).

Etymology

The specific epithet ‘ chaktomuk ’ is a Khmer word meaning ‘four

faces’. It is used in reference to the low-lying area at which the T onle

Sap, Bassac and Mekong rivers come together to form an ‘X’ centred

on Phnom Penh, itself historically known as ‘Krong Chaktomuk’

(literally ‘City of Four Faces’). Based on current knowledge, the

global distribution of the new species is restricted to scrub within

the dynamic floodplain created by the confluence of these waters.

We use chaktomuk as a noun in apposition to the genus name, and

it is thus invariable.

Nomenclatural acts

The electronic edition of this article conforms to the requirements

of the amended International Code of Zoological Nomenclature

(International Commission on Zoological Nomenclature 2012),

and hence the new name contained herein is available under that

Code from the electronic edition of this article. This published

work and the nomenclatural act it contains have been registered in

ZooBank, the online registration system for the International

Commission of Zoological Nomenclature. The ZooBank Life

Science Identifiers (LSIDs) can be resolved and the associated

information viewed through any standard web browser by

appending the LSID to the prefix “http://zoobank.org/”. The

LSID for this publication is: urn:lsid:zoobank.org:pub:F177849T

B6EE-4225-95B2-2843B32CBA08. The electronic edition of this

work was published in a journal with an ISSN, and has been

archived and is available from the digital repository BioTaxa

(http://biotaxa.org).

ECOLOGY AND BEHAVIOUR

Habitat

All observations of O. chaktomuk were made on level ground in

very dense humid evergreen scrub (multi-stemmed woody

plants, 2-6 m tall), sometimes admixed with long grasses or trees

(Plate 5), at elevations of 3-25 m above sea level. Trees occur

exceptionally; where present they are typically scarce, the scrub

forming a dense layer with occasional tree canopies emerging from

8

S. P. MAHOODefa/.

Forktail 29 (2013)

')

Plate 5. Habitat at the type locality of Orthotomus chaktomuk.

it. Orthotomus chaktomuk has not been seen in forest (defined as a

habitat where trees predominate) and is therefore assumed to be

absent from it. At all locations where birds have been found, the

scrub is located within a floodplain and experiences seasonal or

permanent (artificial) flooding. The presence of seasonally flooded

scrub in any location is probably typically transitory, since in the

absence of disturbance by people, large ungulates or hydrological

processes it would presumably revert to seasonally flooded forest.

Orthotomus atrogularis is sometimes found in seasonally

flooded scrub occupied by 0. chaktomuk (Figure 1, Table SOM

1). Where the two species are syn topic, O. atrogularis is much the

rarer. Typically O. atrogularis is found in the edge and canopy of

taller forest habitats, showing a preference for disturbed and

secondary forest because these offer an abundance of vines (Madge

2006, Wells 2007). In some parts of the Tonle Sap floodplain where

0. chaktomuk is absent, O. atrogularis is common in seasonally

flooded forest and scrub, presumably because this habitat also offers

an abundance of edge surfaces. At the other end of the habitat

continuum, O. sutorius replaces O. chaktomuk in open scrub and

gardens, although at some locations the two species are syntopic,

even vocalising from the same individual plants (SPM pers. obs.).

Orthotomus chaktomuk possibly occupies a habitat intermediate

between those of O. atrogularis and 0. sutorius. However, as in

other geographic areas where more than one lowland tailorbird

species is present, habitat niches are difficult to define and

distinguish. Ecological interactions and habitat associations of O.

chaktomuk and other lowland tailorbirds are worthy of further

research.

Birds sharing the habitat of O. chaktomuk include widespread

species often associated with gardens, e.g. Yellow-vented Bulbul

Pycnonotus goiavier. Pied Fantail Rhipidura javanica, Oriental

Magpie Robin Copsychus saularis, sometimes O. sutorius , and species

usually associated with dense lowland humid evergreen scrub,

including Striped Tit Babbler Macronous gularis. Yellow-bellied

Prinia Prinia flaviventris, Plain Prinia Prinia inornata , Olive-

backed Sunbird Cinnyris jugularis and O. atrogularis. From

October to April, Palearctic migrants (e.g. Dusky Warbler

Phylloscopus fuscatus and Siberian Rubythroat Luscinia calliope ) are

abundant in this habitat. In locations where it occurs, O. chaktomuk

often appears to be one of the most abundant bird species.

Behaviour

Owing to the structural characteristics of its habitat, O. chaktomuk

is rarely seen without the aid of playback of vocalisations, and thus

data on ‘normal’ behaviour are few. Almost all encounters have been

with what appear to be adult male-female pairs, or adult male-

female pairs with one subadult. Prior to moult, immature birds were

seen singly, in male-female pairs of exclusively immature birds or

in male-female pairs consisting of one immature and one adult bird.

Birds usually stay within dense vegetation, where they glean and

sally-glean from live and dead leaves of multi-stemmed bushes and

occasionally vines, from ground-level to canopy. Orthotomus

chaktomuk has not been observed foraging in trees. When vegetation

is flooded, birds typically forage below the crown of the bush, on

hanging branches just above the water. One individual that was lured

out of dense scrub with the aid of playback foraged on long grass-

stems, gleaning leaves of a vine that was growing amongst the grass.

Individuals have been observed taking the following prey (once

each): a small fly Diptera, a small spider Araneae, a small caterpillar

Lepidoptera and a small katydid Tettigoniidae; all were consumed

immediately.

In response to playback, birds that have approached the observer

have been seen to sing, usually in a duet, while perched (usually on

or near the top of vegetation, including trees; Media F iles SOM 1 -

3) and occasionally in song flight. Singing is sometimes accompanied

by rapid downwards tail-wagging. Sometimes, while singing in duet,

perched birds droop and shiver their wings. Immature males gave a

simpler, less developed song than adults.

During March and April only males responded strongly to

playback of vocalisations; females typically did not respond, or did

so only briefly. Because this was in stark contrast to behaviour at

other times of year it is thought to indicate that females were on the

nest. Although there are no data on the timing of breeding of

lowland tailorbirds in Cambodia, in Thailand O. atrogularis pairs

with dependent young have been recorded from July to early

September (Round 2008). The nest and eggs of O. chaktomuk

remain to be described.

Distribution

The distribution of O. chaktomuk is incompletely known. It is

apparently constrained by the distribution of seasonally flooded

dense scrub within the floodplain of the Tonle Sap, Mekong and

Bassac rivers in Cambodia (Figure 1). However, based on current

data it is absent from part of this floodplain. Searches at various

locations in apparently suitable habitat in the Tonle Sap floodplain

have thus lar only found the species in the south-east (see Table

SOM 1 for a list of all locations in the floodplain of the Mekong,

Tonle Sap and Bassac rivers where searches for O. chaktomuk have

been conducted). In the north of the Tonle Sap floodplain (where

we have searched for and not found O. chaktomuk ), 0. atrogularis

is abundant in habitat that is superficially structurally similar to

habitat in the south-east, and it is unclear how far north and west

along the lakeshore the distribution of O. chaktomuk extends. There

is no biogeographic reason why O. chaktomuk should be absent from

parts of the Tonle Sap floodplain, and the causes of its absence are

unknown; O. atrogularis is scarce or absent at sites where O.

chaktomuk was recorded (Table SOM 1).

Orthotomus chaktomuk was not found in seemingly appropriate

small seasonally flooded scrub patches at the northern limit of the

Mekong floodplain (12°36'27.52"N 106° 01'36.06"E) in Kratie

province (Table SOM 1, J. A. Eaton verbally 2012). Satellite data

indicate that there is little, if any, suitable habitat for O. chaktomuk

in the Mekong floodplain in Vietnam and it is currently unrecorded

there (although no specific searches have been conducted). As might

be expected, we have located only O. atrogularis in scrub habitats

outside of the Mekong, Tonle Sap and Bassac floodplain (where

these records were within 1 0 km of superficially suitable habitat for

O. chaktomuk they are mapped on Figure 1). Based on current

knowledge of its range, the distribution of O. chaktomuk covers less

than c. 10,000 km2 (Figure 1); it therefore can be considered a

restricted-range species (sensu Stattersfield etal. 1998).

Conservation

Orthotomus chaktomuk is restricted in distribution. Suitable habitat

is patchy outside of the Tonle Sap floodplain and in the latter its

Forktail 29 (2013) A new species of lowland tailorbird ( Orthotomus ) from the Mekong floodplain of Cambodia

9

distribution is poorly understood. Trends in loss, degradation and

fragmentation of floodplain scrub are poorly documented and

subject to considerable local variation (e.g. Packman et al. 2013).

However, most floodplain scrub in Cambodia occupies land

suitable for rice cultivation and could be further threatened by

changes in ongoing burning, fuel-wood collection, cattle grazing

(all of which potentially have a dual role because they also serve to

slow succession) and the spread of the invasive plant Mimosa pigra.

Ironically, O. chaktomuk might now be dependent on human

activity to keep suitable scrubby habitat from becoming forest, since

other anthropogenic impacts — eradication of wild ungulates,

replacement of domestic animals by machines, water flow/level

control, and changes in agricultural practices such as fallows and

cyclical abandonment — have greatly curtailed processes that

maintained the scrub. The species occurs in one protected area,

Baray Bengal Florican Conservation Area, although at that site

habitat is managed to maximise the area of grassland. It has already

been lost from one site (Kraing Check) where birds were netted in

2009: visits in late 2012 found no birds and all suitable habitat had

been converted to aquaculture ponds.

We believe that 0. chaktomuk should be classified as Near

Threatened on the IUCN Red List because it approaches the

thresholds for Vulnerable under criteria Bla+bi,ii,iii,iv (IUCN

2001). Its Extent of Occurrence is 9,385 km2 and thus below the

threshold for Vulnerable (<20,000 km2; criterion Bl). Although

most locations where it occurs are small and isolated it has been

found in the Tonle Sap floodplain where there is a large area of

apparently suitable habitat (although it apparently does not occupy

all of it). Because of this, its habitat cannot be considered severely

fragmented (subcriterion a). Nonetheless it is inferred to be

undergoing a continuing decline (subcriterion b) in (i) extent of

occurrence, (ii) area of occupancy, (iii) area, extent and/or quality

ofhabitat, and (iv) number of locations or subpopulations. Its Area

of Occupancy has not yet been evaluated owing to uncertainty

regarding both the distribution of suitable habitat and its

distribution within apparently suitable habitat. Notwithstanding

this assessment, if the species is found to be more widely distributed

in the Tonle Sap floodplain, then it would warrant downlisting to

Least Concern.

Ongoing habitat loss is likely to be exacerbated by the impacts

of hydropower development on the Mekong and its tributaries.

Models of the effects of hydropower dams predict changes in the

duration and size of the annual flood-pulse that will lead to a

reduction in the extent of seasonally flooded habitats (Arias et al.

2012). Dam construction will also reduce fish populations (the

primary protein source in rural Cambodia), cause changes in flood

regime and lead to water shortages in the floodplain (Orr et al. 2012).

These changes will probably lead to additional loss of floodplain

scrub owing to expansion of agricultural land for rice production,

fish ponds and grazing land for cattle. Construction has started on

one mainstream lower Mekong dam (Xayaburi, in northern Lao

PDR) and numerous tributary dams and ‘pre-construction’ works

are thought to have begun on another (Don Sahong) in the far south

of Lao PDR (International Rivers 2012); nine more mainstream

dams are planned (Mekong River Commission 2011).

TAXONOMIC CONSIDERATIONS

Higher-level systematics

Assignment of the new species to the genus Orthotomus is

straightforward based on its overall structure, plumage and habits,

which typify this genus. Genetic analysis provide additional support

for this arrangement, as DNA sequence comparisons (detailed

below) included specimens of the type species for the genus: O.

sepium Horsfield, 1821. Orthotomus was previously placed in an

expanded Sylviidae (Sibley & Ahlquist 1990) until that family was

shown to be paraphyletic and the genus was transferred to the

Cisticolidae, along with the cisticolas ( Cisticola ), prinias ( Prinia )

and a number of other genera (Alstrom et al. 2006). Within the

Cisticolidae, Orthotomus occupies a clade with the genera Heliolais ,

Prinia and Urorhipis (Olsson et al. 2013). Orthotomus remained

intact until molecular evidence led to the removal of four

superficially similar species: the two African tailorbird species were

transferred to the resurrected genus Scepomycter (Nguembock et al.

2007) and the Asian mountain tailorbirds, Mountain Tailorbird

Phyllergates cucullatus and Rufous-headed Tailorbird P.

heterolaemus, were shown to be not particularly closely related to

‘lowland tailorbirds’ — the most appropriate English group name

for the remaining Orthotomus species (Alstrom et al. 2006, 2011).

The Orthotomus comparisons by Sheldon et al. (2012), based on

one mitochondrial and two nuclear DNA markers, suggested that

O. sutorius is sister to the rest of the lowland tailorbirds, which

comprise four relatively divergent clades: (l) Rufous-fronted

Tailorbird O. frontalis, (2) Rufous-tailed Tailorbird 0. sericeus, (3)

O. atrogularis-O. ruficeps-O. sepium, and (4) the rest of the

Philippine endemics.

Taxonomic implications from morphology

Lowland tailorbirds appear to be relatively conservative in the

evolution of distinctive plumage. For example O. atrogularis , the

widespread O. sutorius and the Philippine endemics Philippine

Tailorbird O. castaneiceps, 0. frontalis and Grey-backed Tailorbird

O. derhianus share the same basic plumage pattern and colouration,

but are not particularly closely related (Sheldon etal. 2012). Owing

to this morphological congruence, various Philippine taxa were long

considered part of O. atrogularis (Delacour & Mayr 1946). These

were later split from that species and grouped under O. castaneiceps

and 0. derhianus (Dickinson et al. 1991, Kennedy et al. 2000).

Collar (2011) considered that morphological data alone were

insufficient to afford O. casteneiceps chloronotus species status to

resolve the geographically vexing situation created by the specific

recognition of O. derhianus. Equally, treatment of O. frontalis as a

species (proposed by Madge 2006) was deemed untenable using

morphological information alone (Collar 2011).

The plumage of 0. chaktomuk is typical of the O. atrogularis-

0. ruficeps-O. sepium clade. Within this grouping, all species are

characterised by a rulous crown; white or rufous cheeks; grey, olive-

grey or bright olive-green upperparts; and grey or whitish-grey

underparts with or without heavy blackish throat-streaking (Table

2, Plate 3). Superficially, the head and underparts pattern and

colouration of O. chaktomuk are similar to O. atrogularis, while

colouration of the upperparts is more similar to 0. ruficeps.

However, 0. chaktomuk shows a suite of plumage features that in

combination are unique, and there are various additional subtle

plumage differences between it and closely related species (Table 2,

Plates 3,4). Immatures of all species within the clade possess yellow-

olive fringing on the wing-feathers; adult O. atrogularis exhibit the

same colouration on the wings, tail and much of the upperparts.

Examination of specimens suggests that this colouration is not as

vivid in immature O. ruficeps and 0. sepium as in 0. atrogularis and

O. chaktomuk.

In common with other lowland tailorbird species, those in the

O. atrogularis-O. ruficeps-O. sepium clade show relatively minor

geographic variation in morphology. All three species within the

clade are polytypic. Although a detailed examination of

morphological variation within these species was not completed,

examples (including some type material) of multiple subspecies were

examined (Table S2). Within each species, all examined specimens

were superficially very similar. Morphological variation within 0.

atrogularis is most marked along the Sabah-Sarawak border area,

in common with other species that share a similar distribution (e.g.

10

S. P. MAHOODeta/.

Forktail 29 (2013)

White-rumped Shama Copsycbus malabaricus). The most

morphologically divergent taxon within O. ruficeps , O. r.

cagayanensis of Cagayan Sula, Philippines, which is apparently

extensively brown or rufous-washed above with pale eyes (Madge

2006), was not examined. Within this context of limited geographic

variation within species, any suggestion that O. chaktomuk should

be considered a highly distinctive localised subspecies of O.

atrogularis is untenable. Equally, there is no evidence that any

lowland tailorbird species possesses regularly occurring colour

morphs.

Biometrics ol O. chaktomuk are similar to other species within

the O. atrogularis- O. ruficeps-O. sepium clade (Table SOM 3).

Using bill length/wing length as a proxy lor size, there is an

indication that 0. chaktomuk is smaller than closely related species

(Figure 2). This could not be confirmed statistically, because the

sample size of O. chaktomuk is too small (Table SOM 3). In the

field, O. chaktomuk appeared to have a shorter tail than local O.

atrogularis. However, this could not be confirmed from the

specimens collected of O. chaktomuk , because their tails are very

worn.

The description of a new species provides an opportunity to

test the quantitative criteria for species delineation proposed by

Tobias et al. (2010). These criteria use a scoring system for

morphological, vocal and ecological features to assess taxonomic

rank. Even when applied only to morphological features, O.

chaktomuk exceeds the threshold score (7) for species status when

compared with O. atrogularis, 0. ruficeps and 0. sepium. It scores 8

against O. ruficeps-. cheeks white rather than orange-rufous (3),

throat and breast very dark grey with white speckling rather than

unmarked pale grey (3), thighs whitish-cinnamon rather than

orange-rufous (2). Its scores against O. sepium are similar. It scores

8 against O. atrogularis-. upperparts mid-grey rather than bright

yellowish-olive (3), vent white rather than yellow (3), thighs whitish-

cinnamon rather than yellowish-orange (2).

Taxonomic implications from vocalisations

Vocal data reaffirm the close relationship between O. chaktomuk

and 0. ruficeps, O. sepium and O. atrogularis, in particular the last.

Orthotomus vocalisations are difficult to define. Males and females

often duet, or if three birds are present, all will vocalise

simultaneously. Vocalisations of O. chaktomuk are typically lengthy,

and those of males are extremely varied (Media File SOM 4-6).

We do not think that we have documented the full vocal range of

the species. In addition, comparisons with closely related species

were hampered because it is doubtful that the full vocal repertoire

of such species has been documented.

The vocalisations of species in the O. chaktomuk-O.

atrogularis-O. ruficeps-O. sepium clade fall into two distinct types.

Those of 0. ruficeps and O. sepium are largely short, pure-tone

whistles, while those of O. chaktomuk and O. atrogularis are

restricted to short trills. To quantify the distinctiveness in songs

of O. chaktomuk in relation to other species within the clade, we

conducted a discriminant analysis (DA) on male vocalisations using

XLStat (Addinsofc 2013). We randomly selected one song (defined

as above) from each individual of O. chaktomuk from which we

had obtained recordings (total five individuals) and randomly

selected a similar number of songs from five individuals each of O.

a. nitidus (the geographically closest subspecies of O. atrogularis ),

0. ruficeps and O. sepium. These included three recordings of O.

atrogularis from Cambodia (including one from the floodplain of

the Tonle Sap) and one recording from Vietnam made within 100

km of the border with Cambodia. (For a full list of recordings used

in analyses see Table SOM 4.) Recordings were downloaded from

www.xeno-canto.org and http://avocet.zoology.msu.edu/. From

each recording we calculated mean values of the following variables:

notes/strophe, length (in seconds) of longest note (one per

0.4

0.38

0.36

0.34

0.32

0.3

0 28

0.26

0.24

0.22

0.2

(a)

O. chaktomuk

O. a. nindus

O. r onereceaus

O. s. sepium

0.4

0.38

0.36

0.34

0.32

0.3

0.28

0.26

0.24

0.22

0.2

(b)

O. chaktomuk O. a. nitidus O. r. dnereceaus O. s. sepium

0.4

0 38

036

0.34

0.32

03

0 28

0.26

024

0.22

0.2

(c)

O chaktomuk O. a. nitidus O. r onereceaus O. s. sepium

Figure 2. Box plots of bill-length (to skull) divided by wing-length of

(a) males of Orthotomus chaktomuk (n = 3), 0. atrogularis nitidus (n =

1 5), 0. ruficeps cineraceus (n = 14) and 0. sepium sepium (n = 20); (b)

females of 0. chaktomuk (n = 2), 0. a. nitidus (n = 10), 0. r. cineraceus (n

= 6) and 0. s. sepium (n = 8); (c) males and females of 0. chaktomuk (n

= 5), 0. o. nitidus (n = 25), 0. r. cineraceus (n = 20) and O. s. sepium (n =

28). Small squares represent the median; box indicates 50% of samples;

bars indicate maximum and minimum.

strophe), strophe length (seconds), maximum and minimum

fundamental frequencies (one each per strophe), number ol

strophes per phrase, length ol interval between phrases (in seconds)

and, for phrases with more than one strophe, interval between

strophes and length ol phrase (in seconds); we also calculated

bandwidth (maximum minus minimum fundamental trequency

within a given strophe), strophe pace (number ol notes per strophe/

strophe length) and, for phrases with more than one strophe, phrase

pace (phrase length/strophes per phrase).

Forktail 29 (2013) A new species of lowland tailorbird ( Orthotomus ) from the Mekong floodplain of Cambodia

11

-10 -SO S 10

FI (94.65 \)

Figure 3. Multivariate vocal space of lowland tailorbirds in the 0.

chaktomuk-O. atrogularis-O. ruficeps-O. sepium clade from

discriminant analyses based on twelve song traits. Scatter plot of the

first two canonical functions that discriminated songs of 0. chaktomuk,

O. atrogularis, O. ruficeps and 0. sepium. • represent 0. chaktomuk ;

@1, 0. atrogularis-, A, 0. ruficeps; +, 0. sepium.

Table 4. Results of discriminant analyses of songs of species in the 0.

chaktomuk-O. atrogularis-O. ruficeps-O. sepium clade, based on 1 2

acoustic variables, showing first three (of four) canonical functions.

Canonical function

Most of the analysed songs of O. chaktomuk and O. atrogularis

clustered separately in multivariate vocal space and could be

discriminated from each other and from the songs of O. ruficeps

and O. sepium (Wilks’s X= 0.005, F = 2.222, P = 0.047; Figure 3).

Overall, the analysis assigned songs to the correct species with 60%

accuracy. Songs of O. chaktomuk were classified with 60% accuracy

and songs of O. atrogularis were classified with 40% accuracy. Songs

of O. ruficeps and O. sepium were classified with 40% and 100%

accuracy, respectively. The discriminant analysis was mainly

influenced by the length of the longest note, strophe pace and the

number of notes per strophe (Table 4).

In accordance with their acoustic similarities, O. chaktomuk and

O. atrogularis respond to playback of each other’s vocalisations,

indicating that inter-specific territoriality is common. There is

individual variation in the magnitude of response, but this is poorly

understood at present. Interspecific territoriality is a common trait

in avian sister species whose ranges come into contact (e.g. Orians

& Willson 1964, Murray 1971, Murray 1976). At locations where

O. chaktomuk is sympatric with O. sutorius , the latter sometimes

also responds to broadcast of vocalisations of the former by

ascending the vegetation and singing.

Taxonomic implications from ecology

The apparently restricted distribution of O. chaktomuk differs from

those of other Orthotomus species on the Asian mainland, which

are typically wide (Madge 2006). Lowland tailorbirds are thought

to have originated in southern Asia or possibly Sundaland. They

rapidly spread widely in Sundaland and the Philippines and, more

recently, additional species have evolved in both island groups

(Sheldon et al. 2012). The O. chaktomuk-O. atrogularis-O.

ruficeps-O. sepium clade emerged relatively recently, and, in

contrast to the clade containing Philippine endemics, exhibits lower

species richness (four versus six species), presumably owing to the

lack of opportunity for speciation in lowland populations in a

continental mainland versus an oceanic island setting. Lowland

passerines in mainland Asia typically have large distributions. Those

with smaller distributions are largely confined to successional

habitats in the floodplains of large rivers, such as Black-breasted

Parrotbill Paradoxornis flavirostris and Marsh Babbler Pellorneum

palustre (Rasmussen & Anderton 2005).

T ailorbird habitats are notoriously difficult to define (e.g. Mitra

& Sheldon 1993), although lowland tailorbirds within the 0.

chaktomuk-O. atrogularis-O. ruficeps-O. sepium clade have slightly

clearer habitat preferences. In this clade, greater specialisation is

thought to have helped these younger species avoid competition

with the pre-existing generalist species O. sutorius and O. sericeus

(Sheldon et al. 2012). The habitat preferences of O. chaktomuk are

thought to be somewhat intermediate between O. atrogularis and

O. sutorius. In addition, O. chaktomuk is apparently confined to

successional floodplain habitat. Vegetational patterns in what is

now its distribution have been shaped by cyclical ice-ages and

interglacials over the past two million years. During glacial maxima,

the sea-level was much lower than today and southern Vietnam

and Cambodia were connected by land to what is now Peninsular

Malaysia by the now submerged Sunda Shelf (Sathiamurthy &

Voris 2005). It is thought that there were four large river basins on

the Sunda Shelf and these great river systems connected the

freshwater riverine faunas of many of today’s rivers (such as the

Mekong) that are now restricted to Indochina, the Malay Peninsula

or one of the greater Sunda Islands (Voris 2000). Floodplain

habitats were therefore probably much more extensive during

glacial maxima and it is possible that O. chaktomuk evolved on the

Sunda Shelf. Orthotomus chaktomuk is now restricted to a much

smaller area, constrained by the reduced availability of suitable

habitat during an interglacial.

The known distribution of O. chaktomuk lies within the

distribution of O. atrogularis. No parts of the distribution of O.

chaktomuk are more than a few tens of kilometres from locations

where O. atrogularis is found. The two species are locally syntopic

(Figure 1, Table SOM 1); for instance, c.200 m from the type

locality a pair of O. atrogularis was observed in the same bush as a

single O. chaktomuk (SPM, AJIJ, HC pers. obs.; photographed).

Although nearly 50 O. chaktomuk-O. chaktomuk male-female

associations have been observed, neither mixed pairs nor birds that

are phenotypically identifiable as hybrids have been detected.

Taxonomic implications from molecular analyses

Within the genus Orthotomus , superficial morphological and vocal

similarities between taxa have frequently clouded their taxonomic

status. In this context, molecular techniques can provide a useful

tool to infer relationships between taxa. T o determine the position

of O. chaktomuk in the molecular phylogeny of Orthotomus we

compared DNA sequences of mitochondrial ND2 and nuclear

MUSK and TGF(32 genes of the type specimens with all other

species of lowland tailorbird (Sheldon etal. 2012) (see Table SOM

5 for details of all specimens used in the genetic analyses). Tissues

from the five O. chaktomuk specimens (Table 1) were preserved in

95% ethanol and stored in the University of Kansas Natural

12

S. P. MAHOODefo/.

Forktail 29 (2013)

History Museum (KUNHM) and Louisiana State University

Museum of Natural Science (LSUMNS) tissue collections. DNA

was extracted and sequenced following the protocol described in

Sheldon et al. (2012), and the sequences deposited in GenBank:

accession numbers KF015230-KF015247. The total number of

DNA nucleotides was 1,041 of ND2, 614 of MUSK, and 613 of

TGF(32. Separate and concatenated Bayesian phylogenetic analyses

of these sequences using MrBayes ver. 3.2.1 (Ronquist et al. 2012)

as in Sheldon et al. (2012) placed the novel taxon in a clade with

0. atrogiilaris , O. ruftceps and (). sepium ; all trees except TGF[32

placed 0. atrogiilaris and O. chaktomuk as sisters (Figure 4). This

arrangement concurs with morphological and vocal analyses.

The ND2 sequences of the four taxa had 90 variable and 78

parsimony informative sites. Between the one sample of O.

atrogularis and the five samples of O. chaktomuk were 22 variable

sites of which 12 consistently differed. The ND2 p-distance from

O. chaktomuk to O. atrogularis averaged 1.3% (range 1 . 1 %— 1 .4%),

to O. ruficeps 5.0% (4.6%-5.2%), and to O. sepium 6.5% (6.3%-

6.8%). Variation in the MUSK and TGFa2 sequences between O.

chaktomuk and O. atrogularis was negligible (three and six sites,

respectively). These genetic data are insufficient to resolve the

relationship between O. chaktomuk and O. atrogularis owing to

the small number of samples of 0. atrogularis compared (one) and

because that sample was not of the subspecies sympatric with O.

chaktomuk. Instead, the O. atrogularis sample was of the nominate

subspecies collected in Sarawak, which is restricted to the Sundaic

region (except Sabah, north Borneo). The genetic divergence

between O. chaktomuk and the 0. atrogularis sample (1.3%) is small

and broadly comparable to that between other lowland taxon-pairs

on Borneo and mainland Asia that are considered subspecies,

although there is considerable inter-species variation in genetic

distances (e.g. Lim et al. 2010, Sheldon et al. 2012). A

phylogeographic study including samples from all subspecies and

biogeographically relevant populations of O. ruficeps , O. atrogularis

and O. chaktomuk is required to clarify their evolutionary

relationships.

Relationship of Orthotomus chaktomuk to species

within Orthotomus

Orthotomus chaktomuk is locally syntopic (Figure 1, Table SOM

1) with the only species from which it shows apparently relatively

low genetic divergence, O atrogularis. We have found no evidence

of hybridisation, and the taxa satisfy the precepts of the biological

species concept because they behave like separate species when they

come into contact (e.g. Mayr 1963, 1999). Classification of the

new taxon as a subspecies or highly localised colour morph of O.

atrogularis is therefore untenable. Because O. chaktomuk is on a

distinct evolutionary trajectory it also satisfies the phylogenetic

species concept (Cracraft 1989).

Although reported genetic divergence among sister species is

typically lower in temperate regions than in the tropics this may be

an artefact of incomplete sampling and incorrect taxonomy (Tobias

etal. 2008, Sangster 2009). Recent studies are overturning the trend

for lumping distinctive taxa into polytypic species and revealing

cryptic diversity in widespread species (e.g. Collar 2006, 2011,

Rheindt & Eaton 2010, Leader 2011, Moltesen et al. 2012,

Rasmussen et al. 2012). By taking an integrated approach to

taxonomy (as here), sister species are being recognised in tropical

regions that differ in sampled regions of nuclear DNA by levels

that are in line with those used in temperate regions (e.g. Irestedt

etal. 2013).

There are a number of plausible explanations for the apparently

low genetic divergence between O. chaktomuk and O. atrogularis.

The molecular phytogenies suggest that O. chaktomuk might be a

relatively young lineage. Its diagnostic phenotypic traits that

apparently prevent modern hybridisation might be encoded by a

100

100

100

J-L

V:

100

100

q

100

- O. atrogularis 57037

O. chaktomuk 201293

O. chaktomuk 201294

O. chaktomuk 201295

O. chaktomuk 201291

O. chaktomuk 201292

O. ruficeps 47143

O. ruficeps 17284

O. ruficeps J 1 1 56

r O. sepium 219589

O. sepium 269052

O. sepium 56705

O. sepium 220247

O. sepium 703336

f

0.02

Figure 4. The Orthotomus chaktomuk, O. atrogularis, O. ruficeps, and

0. sepium clade extracted from the entire Orthotomus phylogeny,

which was constructed from concatenated DNA sequences of ND2,

MUSK, and TGFp2 via Bayesian phylogenetic inference as described in

Sheldon et al. (2012). Numbers along branches indicate Bayesian

posterior probabilities. The topology is the same as ND2 and MUSK

trees by themselves.

small number of genes that evolved rapidly under sexual selection

(cf. Uy et al. 2009). This process might have occurred too rapidly

for significant additional genetic differences to accumulate in parts

of the genome not under intense selection or which are selected

for other purposes (as are mitochondrial genes). If 0. chaktomuk is

indeed a Sunda Shelf species, now confined to relict habitat in

Indochina, then it might have been derived from Sundaic O.

atrogularis rather than mainland populations. If this were the case

then it might be expected to show greater genetic divergence from

mainland O. atrogularis to which it is now locally syntopic than to

the Sundaic nominate used here in the genetic analyses. An

alternative explanation is that the apparent low genetic divergence

between the species is a result of genetic introgression sometime

during the last two million years (Rheindt & Edwards 2011).

Periods of peak sea-level might be the most plausible time for

genetic introgression to have occurred. A higher sea-level might

have constrained suitable habitat for O. chaktomuk into a narrow

band between the distributions of O. ruficeps and O. atrogularis.

This process perhaps drove 0. chaktomuk through a population

bottleneck and increased the chances of hybridisation with 0.

atrogularis , leading to genetic introgression. If a comprehensive

study of relationships within the O. chaktomuk-O. atrogularis-O.

rufiiceps-O. sepium clade reveals that O. chaktomuk is more closely

related to O. a. nitidus than to Sundaic taxa, then this is perhaps a

more likely explanation for the low genetic divergence.

Final remarks

The modern discovery of an undescribed bird species close to sea-

level within the limits of a large city in a populous country is

extraordinary, but not unprecedented (cf. Stymphalornis sp. nov.,

an as-yet undescribed taxon restricted to marshes close to Sao Paulo,

Brazil, discovered in 2005: Reinert et al. 2007). At least three

interacting factors probably account for O. chaktomuk having gone

unnoticed for so long. It inhabits a very small geographic range, and

within this it is restricted to a very specific habitat type: dense

floodplain scrub. This habitat is of little interest to birdwatchers

and ornithologists because the other species that it supports are some

of the most widespread and abundant birds in tropical South-East

Asia. Even if its habitat were to attract more attention, the denseness

of the habitat and the species’s skulking habits would more often

than not render it invisible to the casual would-be observer.

Vocalisations of O. chaktomuk are similar to those of O.

atrogularis , which has a perplexing array of vocalisations with which

birdwatchers rarely attempt to familiarise themselves fully.

Orthotomus atrogularis is a common species within suitable habitat

across a fairly broad range, and therefore there is little a priori reason

Forktail 29 (2013) A new species of lowland tailorbird ( Orthotomus ) from the Mekong floodplain of Cambodia

13

for a birdwatcher or ornithologist to invest effort in trying to see a

hidden, vocalising tailorbird in dense scrub in mainland South-East

Asia. Moreover, collecting effort in Cambodia has been low: we have

been able to trace only two O. atrogularis and one O. sutorius

specimens (NHMUK 1928.6.26.1210, Eames & Ericson 1996) (the

identification of all these specimens has been verified by the primary

author, either first-hand or using photographs). Modern

birdwatching effort in Cambodia is also limited and very localised.

The factors discussed above also help explain why the first four

individuals known were all mist-net captures. Their

misidentification can be accounted for by the species’s superficial

similarity to other species, observer inexperience and the sheer

unlikelihood of alternative options (cf. Woxvold et al. 2009). The

discovery of 0. chaktomuk indicates that new species of bird may

still be found in familiar and unexpected locations.

ACKNOWLEDGEMENTS

We are most grateful to Keo Omaliss, Director of the Department of Wildlife

and Biodiversity of the Forestry Administration of the Ministry of Agriculture,

Forestry and Fisheries, Kingdom of Cambodia, for permission to collect and

export scientific specimens; Mark Adams, Natural History Museum, Tring,

and Steven van der Mije, Naturalis Biodiversity Centre, Leiden, for access to

and high-quality images of scientific specimens; Will Duckworth for unwavering

support and good advice; Philip Round and Craig Robson for their support for

our initial identification; James Eaton for good advice and the use of his

photographs and sound recordings; Robert Martin for the use his sound

recordings; Martin Kennewell for his videos as supplementary online material;

Tom Evans for his early role in the species’s discovery; Sarah Brook, Nigel Collar

and John Pilgrim for their counsel; Phien Sayon for making the map; Harry

Taylor for photographs of the type material at NHMUK; Angela Yang for

information on the 2009 records; Andy Symes and Stuart Butchart at BirdLife

International for advice on the conservation section; Normand David and

Edward Dickinson for advice on nomenclature; Mauricio Arias for information

on floodplain processes; Alexander Lees and Sidnei Dantas for help with

sonagrams; Lucy Keatts, Chantha Yuthea and Tan Setha for help obtaining

permits; Ulf Johansson for photographs of Orthotomus specimens from the

Swedish Museum ofNatural History; Ann Mahood for assistance at NHMUK;

Robert van Zalinge and Frederic Goes for field observations; Son Virak for

assistance at Baray Bengal Florican Conservation Area; Tom Clements at WCS

Cambodia for overlooking SPM’s absences in the field or museum, and for

logistic support; and Frank Rheindt, Alexander Lees and five anonymous

reviewers for comments that substantially improved the manuscript.

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Wells, D. R. (2007) The birds of the Thai-Malay Peninsula, 2. London:

Christopher Helm.

Woxvold, I. A., Duckworth, J. W. & Timmins, R. J. (2009) An unusual new

bulbul (Passeriformes: Pycnontidae) from the limestone karst of Lao

PDR. Forktail 25: 1-12.

Zhou Fang & Jiang Aiwu (2008) A new species of babbler (Timaliidae:

Stachyris) from the Sino-Vietnamese border region of China. Auk 1 25:

420-424.

SUPPLEMENTARY ONLINE MATERIAL

Available on Oriental Bird Club website, links at http://

www.orientalbirdclub.org/ publications/ forktail29

Tables SOM 1 to SOM 5

Figure SOM 1

Media Files SOM 1 to SOM 6

Simon P. MAHOOD, Wildlife Conservation Society Cambodia

Program, PO Box 1620, Phnom Penh, Cambodia. Email:

smahood@wcs.org

Ashish Joshia Ingty JOHN, Wildlife Conservation Society

Cambodia Program, PO Box 1620, Phnom Penh, Cambodia.

Email: ajohn@wcs.org

Jonathan C. EAMES, BirdLife International Cambodia

Programme, PO Box 2686, Phnom Penh, Cambodia. Email:

Jonathan.Eames@birdlife.org

Carl H. OLIVEROS, Biodiversity Institute and Department of

Ecology and Evolutionary Biology, University of Kansas,

Lawrence, KS 66045, USA. Email: oliveros@ku.edu

Robert G. MOYLE, Biodiversity Institute and Department of

Ecology and Evolutionary Biology, University of Kansas,

Lawrence, KS 66045, USA. Email: moyle@ku.edu

HONG CHAMNAN, Forestry Administration, Ministry of

Agriculture, Forestry and Fisheries, Cambodia and Wildlife

Conservation Society Cambodia Program, PO Box 1 620, Phnom

Penh, Cambodia. Email: chong@wcs.org

Colin M. POOLE, Wildlife Conservation Society Singapore, 352

Tanglin Road, Strathmore Block #01-08, Tanglin International

Centre, Singapore 247671. Email: cpoole@wcs.org

Howie NIELSEN, 193 Hollywood Blvd, Whitefield, Maine, USA.

Email: birderhowie@gmail.com

Frederick H. SHELDON, Museum of Natural Science and

Department of Biological Sciences, Louisiana State University,

Baton Rouge, LA 70803, USA. Email: fsheld@lsu.edu

FORKTAIL 29 (2013): 15-18

Notes for the conservation of the Rufous-fronted

Laughingthrush Garrulax rufifrons

N. J. COLLAR &S. van BALEN

The Rufous-fronted Laughingthrush Garrulax rufifrons, endemic to Java, has been recorded from a total of 15 montane sites, 14 in West Java

(nominotypical rufifrons) and one in Central Java (subspecies slamatensis). It occupies montane forest generally in the range 1 ,000-2,000 m,

although this may vary with site, and occurs in monospecific parties of birds but also in bird waves, and has or had an association with Javan

Green Magpie Cissa thalassina. Breeding appears to be extended through the year, but lack of records in January-February and July-

August may reflect real breaks in the cycle. A lack of recent records from bird markets and a recent hike in prices of captive birds supports

other concerns that the Javan bird trade may have affected the species, which in the past 20 years appears only to have been observed at

Gunung Gede-Pangrango. Surveys of known sites and of several montane forest reserves are needed before a heavy investment in captive

breeding is made.

INTRODUCTION

Of all the species bearing the English name ‘laughingthrush’, now

proposed as components of a large subfamily of babblers named

Leothrichinae (Moyle etal. 2012), Rufous-fronted Laughingthrush

Garrulax rufifrons — called Red-fronted in Andrew (1985) and

MacKinnon (1988) and Plain-brown in Hellebrekers &

Hoogerwerf (1967) — is the southernmost, being confined to the

island of Java, Indonesia (Collar & Robson 2007). This fact,

combined with its restriction to montane forest (Stattersfield etal.

1998), suggests a relictual distribution, and Berlioz (1930), in

considering it ‘truly aberrant’, attributed this in part to its

geographical isolation.

The species has received virtually no attention from biologists,

ecologists and scientific ornithologists, and there are no studies of

it in the wild, but because, by contrast, it has received considerable

attention from bird trappers, it has been treated all this century as

Near Threatened (Stattersfield & Capper 2000, BirdLife

International 2001). Recent anecdotal evidence suggests that its

conservation status may have declined further since the 1990s

(Collar et al. 2012 and below). This paper is therefore an attempt

to assemble basic information relevant to its long-term conservation

and make some appropriate preliminary recommendations.

DISTRIBUTION

Mees (1996) listed and mapped 1 1 localities for the species (treating

Gn [=Gunung] Endut and Gn Salak separately) and on this basis

remarked that it ‘may be assumed to occur throughout the highlands

of West Java’. Our further collation of records (initials of museums

are glossed in the Acknowledgements) suggests that this prediction

was correct. Since its description in 1831 the Rufous-fronted

Laughingthrush has been recorded at the following localities (listed

as far as possible from west to east), increasing the total to 1 5 (but

treating Gn Endut as part of Gn Salak):

• Gn Karang above Ciomas and Ujungtebu, 1-13 April 1920

(Robinson & Kloss 1924: 285) and at an unspecified locality

in April 1991 (D. A. Holmes in litt. 1991);

• Gn Halimun, August and September 1922 (2 specimens in

Naturalis), July 1982 (K. D. Bishop in litt. 2013);

• Gn Salak (type locality designated by Deignan 1964), on the

south-east slope, October 1882 (Vorderman 1886), at Gn

Endut, 10 June 1897 (Bartels 1902, 1906, Mees 1996; 1

specimen in Naturalis), at Cianten, April-June 1932 (2

specimens inMZB),at Pasirreungit, 12-15 August 1981 (SvB),

at Warungloa, heard once, 15 July 1981 (SvB), on the south¬

west slope at Awibengkok, 10 records of 1-3 birds, 3-9

September 1988 (SvB);

• Gn Gede-Pangrango, May 1889 (1 specimen in Naturalis;

Vorderman 1892), 1900-1926 (34 specimens and 3 clutches in

Naturalis, 4 specimens in MZB), 1943-1947 (8 clutches in

Naturalis, 2 specimens in MZB; also Hoogerwerf 1948),

specifically at Puncak, 1970s (W. G. Harvey in a list supplied by

thelateD. A. Holmes to SvB), Telaga Warna, 1979-1981 (SvB),

Cibodas, October 1896 (1 specimen in Naturalis) and 21

September 1918 (Spennemann 1923), Cibodas and

Kandangbadak, February-March 1916 (Robinson & Kloss

1924, Delsman 1927) and April 1941 (1 clutch in Naturalis),

with many encounters 1979-1989 (Andrew 1985, SvB)

including one at Cimungkat, July 1987 (SvB), sight records

through the 1 990s and 2000s (J. Chance inlitt. 1991, J. A. Eaton,

C. R. Robson in litt. 2013) and audio recordings in June-July

2009 (XC30475-76 by B. Cox, XC40473-74 by D. Edwards);

• Cianjur, Cibeber, in the period 1946-1949 (G. F. Mees

notebooks seen by SvB);

• Gn Patuha, Koleberes, 1927-1929 (Bartels 1931: 336; hence

Hoogerwerf 1948);

• Situ Lembang, 15 March 1984, 8 birds (P. Andrew in litt.

2013);

• Gn Tangkubanprahu (Mees 1996), July and October 1926,

December 1955 and December 1957 (8 specimens in

Naturalis);

• Gn Malabar at Tirtasari, 12 May 1910 (Mees 1996; 1 specimen

in Naturalis);

• Gn Wajang, Cibitung (Mees 1996), April and May 1910 (3

specimens in Naturalis);

• Gn Papandayan, late 1920s (Stresemann 1930), 1941-1942

(2 specimens in MZB), with subsequent records specifically

at Kawahmanuk, 2 birds, 3 September 1987 (SvB); Gn

Kendang, flock of 10-15 birds tape-recorded, 6 September

1987 (SvB);

• Gn Rakutak, March 1900 (1 specimen in AMNH);

• Gn Guntur, Garut, October 1900 (2 specimens in AMNH),

including Kawah Kamojang, May 1923 (1 specimen in MZB),

and ‘near Garut’ (Siebers 1929);

• Gn Ciremay (Mees 1996), June 1930 (1 specimen in Naturalis);

and

• Gn Slamat (type and only locality for race slamatensis ) at

Kaligua, 1916-1917 (Siebers 1929; type specimen in Naturalis,

3 paratypes in MZB), and at Purwokerto, March 1925 (Voous

1948, Mees 1996; 2 specimens in Naturalis).

16

N. J. COLLAR & S. van BALEN

Forktail 29 (2013)

ELEVATIONS, ECOLOGY AND NATURAL HISTORY

The species is resident in and confined to ‘mixed original forest’ or

‘broadleaved evergreen forest’ at 900-2,500 m (Sody 1956, Collar

& Robson 2007), this being a minor shift from elevations of 1,000-

2,400 m (Stattersiield et al. 1998, BirdLite International 2001).

However, these limits represent extremes amalgamated from

individual sites, and may vary considerably at each known site

depending on ecological conditions, mountain height (the peaks

of several mountains listed above lie below 2,400 m), and levels of

deforestation. Moreover, nothing is known about the species’s

relative abundance at different elevations, although Hoogerwerf

(1950) indicated that on Gn Gede-Pangrango it was a common

bird from Cibodas up to near the tops of the mountains.

The site-specific elevations in Hoogerwerf (1948) — 1,500-

2,600 m on Gn Papandayan, 600-1,000 m at Ciomas on Gn

Karang, 800-1,200 m at Cimungkat on Gn Cede, 600-1,000 m

at Koleberes on Gn Patuha and 500-2,300 m on Gn Salak — are

not intended to indicate the limits between which the species was

certainly encountered; nor is there clear evidence to support

Hoogerwerf ’s (1948) characterisation of the species as one ‘in

certain areas probably living permanently between 2500 and

(above) 3000 m’. On Gns Endut and Pangrango, Bartels (1902)

gave its elevation as ‘3,000-3,500 feet’ (900-1,100 m), later

changing this to ‘3,000-6,000 feet’ (900-1,800 m) (Bartels 1906);

records from Cimungkat on Gn Gede were at 1,200 m (SvB). Some

specimens on Gn Tangkubanprahu were at 1,500 m (Naturalis label

data), as was the first record from Gn Papandayan (Stresemann

1930) , although subsequently birds were found in the latter locality

at 1,900 m (Kawahmanuk) and at 2,525 m (Gn Kendang) (SvB).

Records from Gn Salak are at 1,500 m (Vorderman 1886), and

specifically at Pasirreungit at 1,350-1,900 m and Awibengkok at

1,000-1,150 m (SvB). It therefore appears that only one record,

hitherto unpublished, pins the species to an elevation higher than

2,000 m; all other records traced come from below this altitude.

The record from Situ Lembang was at 850 m (P. Andrew in litt.

2013), and those on Gn Karangwere at 600-900 m (Robinson &

Kloss 1924), these apparently being the lowest elevations recorded

for the species.

The Rufous-fronted Laughingthrush occupies all strata of the

forest but chiefly the undergrowth, and is ‘very agile’ (Hoogerwerf

1950). It occurs in loose, sometimes large monospecific groups but

also participates in bird-waves (Hoogerwerf 1950, Andrew 1985),

these latter sometimes comprising up to 1 5 different species on Gn

Gede (van Balen 1992); in particular, it associates with the Javan

Green Magpie Cissa thalassina (Koningsberger 1901, Bartels 1915—

1931) , such that on Gn Halimun in 1982 the two species were

found together in a bundle of birds being carried by a poacher (K.

D. Bishop in litt. 20 1 3). Its presence is best determined by its noisy,

whinnying call, earning it the local name ‘horsebird’ (van Balen

1992) and placing it with the group oflaughingthrushes that possess

a laughing call (Collar & Robson 2007). Various authors have given

glancing accounts of the diet: ‘berries and insects, mostly beetles’

(MacKinnon 1988), beetles, snails and fruits of Melastoma

malabathricum (Sody 1989), and these plus mantids and caterpillars

(Collar & Robson 2007). Hoogerwerf ’s (1950) mention of small

hard seeds and Sody’s (1989) of Melastoma may well both refer

back to Vorderman’s (1886) account of stomachs ‘coloured black

by fruit pulp, and filled with small hard seeds’ (our translation).

The closest observer of the species described its diet as mainly and

sometimes exclusively various forest fruits, supplemented with

insects, mainly beetles including weevils, plus bugs, caterpillars,

locusts, spiders, ants and small vertebrates such as frogs and lizards

(Bartels 1915-1931; also Delsman 1927). Specimen labels in

Naturalis mention Anomala beetles, small beetles, a large weevil, a

phasmid, looper caterpillars and Ficus andLantana fruit as stomach

contents. In captivity, birds caught wild mice in their enclosure

(Pithart 2009).

The nest is a sturdy, relatively small cup placed on a horizontal

branch or in a fork usually fairly close (about 2 m) to the ground in

smaller trees at the edge of forest (more details in Hoogerwerf 1 950,

Hellebrekers & Hoogerwerf 1967). The usual clutch is three (blue-

green) eggs, but sometimes two; nests have been found in March,

April, May, June, September, November and December

(Hoogerwerf 1949, 1950, Hellebrekers & Hoogerwerf 1967).

Whether the gaps in breeding in January-February and July-

August represent real seasonal differences, random variation or

temporal patchiness in observer coverage is an open question.

However, breeding in Prague Zoo followed a roughly similar

schedule, with nests in April-June and August- October (Pithart

2009). Naturalis possesses birds marked as juveniles from January

(1), May (2), June (1) and August (1), but these are full size and it

is impossible to pin them to a likely month of birth; it also contains

four specimens labelled as having full-sized gonads in April (male

and female) and May (two males). In captivity the female was noted

to do almost all incubation, which lasted 14-15 days, while the

nestling period was 15-16 days; moult occurred slowly from

autumn (occasionally July) through to December (Pithart 2009).

Indeed, birds at the end and start of the year have been described as

‘gut im Gefieder’ (Bartels 1902), which presumably best translates

as ‘in fresh plumage’.

POPULATION TRENDS ANDTHREATS

There has been no systematic monitoring of populations of this or

any other forest bird species in Java, so a quantitative assessment of

population trends is impossible. However, various items of

qualitative information have accumulated to suggest that the

Rufous-fronted Laughingthrush may now be in a more serious

condition than has hitherto been realised, largely as a result of the

singular Javanese tradition of bird keeping.

‘I am afraid that aviculture is a major source of bird destruction

in Indonesia’, wrote Morrison (1980), having found Java to be ‘a

singularly birdless island’. This was over 30 years ago. At that time,

however, the Rufous-fronted Laughingthrush, being a bird of high,

remote forests, may still have been common. On Gn Gede-

Pangrango it was common in the 1940s (Hoogerwerf 1950) and in

the 1980s (Andrew 1985), and there is no reason to imagine that it

was less common at the other localities listed above in the 1980s,

although as ajavan endemic it was protected under Indonesian law

in 1979 (Noerdjito & Maryanto 2001). Only once in the early

documentation was there an indication of relative rarity: it was

scarce at Gn Patuha in the years 1927-1929 (Bartels 1931),

presumably for natural reasons (‘only in the northern forests’).

Extrapolation from experience at Gn Gede presumably lies behind

MacKinnon’s (1988) general description of the species as ‘Locally

not uncommon in montane forests’ and behind Mees’s (1996)

remark that ‘Where this species occurs it is common, noisy, and

conspicuous.’

Nevertheless, only two years after this comment, the species

was said to be ‘fairly heavily exploited as a cagebird, which has

rendered it uncommon in otherwise moderately secure habitat’ (D.

A. Holmes in litt. 1998 in BirdLife International 2001), leading to

its designation as a Near Threatened species, and in the mid-2000s

it was described as ‘formerly common in Gede-Pangrango National

Park... but now rare along main trail, reportedly owing to trapping’

(Collar & Robson 2007), although inquiries of leaders taking bird

tours to Gn Gede do not suggest that numbers have obviously

declined there (C. R. Robson in litt. 2013, J. A. Eaton in litt. 2013).

Other evidence, however, certainly tends to support the notion

that a real decline has been occurring for some years. Bird dealers

Forktail 29(2013)

Notes for the conservation of the Rufous-fronted Laughingthrush Garrulax rufifrons

17

in markets in Medan, Sumatra, recently reported that Rufous-

fronted Laughingthrush is ‘becoming increasingly rare or difficult

to find in economically viable numbers’ (Shepherd 201 1).

Independently, it has been reported to have Vanished from the bird

markets in Sumatra’ (P. Hospodarsky in Pithart 2009). Moreover,

on Java at the start of the century the species ‘could be found in

bird markets as a cheap local songster, selling for Rp 150,000 ($16)’,

but in the past few years the price has increased tenfold and in 20 1 2

no birds could be found in bird markets (R. Sozer in Collar et al.

2012 and in litt. 2013). This latter testimony was independently

supported by C. R. Shepherd [in litt. 2013):

Dealers in the Barito Market and the Pramuka Market [Jakarta,

Java] stated in 20 1 1 that this species was ‘difficult to find, or all

gone’ ( susah or sudah habis). These kinds of statements do

usually mean trappers are rarely bringing them in and are not

finding them in their usual trapping areas. In 2012,1 only carried

out one survey in Jakarta’s three largest bird markets ( June 2012)

and did not see any.

Moreover, there are parallels with declines and near-disappearances

in other species that have been attributed to the demands of Javan

bird-keeping, most notably that of the Javan Green Magpie (van

Balen et al. 2013; also Collar et al. 2012).

However, the current plight of the magpie, and the

laughingthrush’s reported association with it, opens up the plausible

if very remote possibility that the laughingthrush’s conservation

status may not be so desperate. Since the magpie is a much more

prized species in the Javan bird trade, it might conceivably be that

when targeting the magpie trappers took many laughingthrushes

simply as a ‘bycatch’, which could explain the latters’ low prices and

wide availability a few years ago. Moreover, now that trade has

reduced the numbers of magpies to near-zero (van Balen et al.

2013), trappers are perhaps no longer visiting areas where magpies

once occurred, in which case the sudden disappearance of

laughingthrushes from markets might simply reflect lack of

trapping effort rather than lack of birds. Nevertheless, the rather

high prices now commanded by the laughingthrush tend to suggest

that its rarity is real and, as V. Nijman (in litt. 2013) has commented,

there are high numbers of montane bird species still available for

sale in Java’s markets, and ‘not all of them are expensive’.

CONSERVATION NEEDS

If protected area status improves the chances of long-term

habitat conservation, then Gn Halimun-Salak, Gn Gede-

Pangrango and perhaps Gn Guntur are likely to be the best-

preserved of the sites at which the laughingthrush occurs (although

at Gn Gede in the past 1 0 years there has been ‘shocking clearance’

for vegetable plots, ‘apparently inside the protected area, probably

up to 2, 1 00 m in a c. 1 km belt above and east of the Cibodas Botanic

Garden’: F. R. Lambert in litt. 2013). The reserve at Kawah

Kamojan on Gn Guntur covers 8,000 ha at 1,400-2,250 m

(MacKinnon et al. 1982), but other sites at which the species has

been recorded have very small areas protected: the only reserves

larger than 100 ha are at Telaga Patengan on Gn Patuha (150 ha),

Gn Papandayan (844 ha) and Gn Tangkubanprahu (1,660 ha)

(MacKinnon et al. 1982), but it is not known if they encompass

laughingthrush habitat and viable populations. An area of

1 5,000 ha on Gn Slamat was long ago recommended for protection

(see Stattersfield et al. 1998) but only two reserves, both less than

20 ha, exist there (MacKinnon etal. 1982); since it is the sole locality

for the highly distinctive subspecies slamatensis (Siebers 1929,

Voous 1948, Mees 1996) of Garrulax rufifrons , formal protection

of the site is clearly highly desirable.

Last records of the species from all known sites are: Gn Karang

1991, Gn Halimun 1984, Gn Salak (where on Endut it was ‘not

rare’ at the start of the twentieth century: Bartels 1902, 1906) 1988,

Gn Gede-Pangrango 2012, Cianjur at least 1949, Gn Patuha before

1931, Situ Lembang 1984, Gn Tangkubanprahu 1957, Gn Malabar

1910, Gn Wajang 1910, Gn Papandayan 1987, Gn Rakutak 1900,

Gn Guntur 1923, Gn Ciremay 1 930 and Gn Slamat 1925. D. Liley

(in litt. 2013) spent 5-6 weeks at Cikuya, on the southern slopes

of Gn Halimun, mostly at 1,000-1,200 m, without seeing the

species, and K. D. Bishop (in litt. 2013) visited Gn Halimun in

August 2011 after a gap of 29 years and found no laughingthrushes;

however, it is fair to note that the one location that most

birdwatchers go to at Gn Halimun, Cikaniki, probably never had

the species (between 1996 and 2009 field teams never recorded it:

Prawiradilaga et al. 2003, Noske et al. 2011). Even so, it appears to

be at least 20 years since there was a record of the species away from

Gn Gede-Pangfango.

Naturally, therefore, these sites need urgent surveying to

determine the status of the forests and the continuing presence of

the species (for which, given its noisiness, playback techniques

would probably be highly effective). Other under-explored reserves

which might hold the species are: GnBurangrang (2,700 ha; 1,000-

2,000 m); Gn Tampomas (1,250 ha; 1,000-1,700 m); Gn Sawal

(5,400 ha; 600-1,764 m); Gn Simpang( 15,000 ha; 600-1,600 m)

and Gn Tilu (8,000 ha; 1,200-2,177 m). Preferably, however, such

a survey would involve line-transect or point-count work to

establish baseline densities at the sites, and would target other rare

species such as Javan Hawk Eagle Nisaetus bartelsi, Javan Trogon

Apalharpactes reinwardtii, Javan Cochoa Cochoa azurea and Javan

Green Magpie, along with (e.g.) certain primates.

Study of the culture and economy of bird-keeping in Indonesia

has led Jepson et al. (2011) to ‘argue that, in Indonesia at least,

conservationists need to move beyond the moralistic, animal rights

and protectionist logic that dominate \sic\ much wildlife trade

discourse and embrace the development logic of pro-poor growth

and more, better jobs’. Whatever one makes of this prescription it

predicates a time-scale that completely mismatches the short-term

needs of many species native to Java, and if acted on would merely

vaporise their chances of survival. If conservationists do not focus

on birds that are at greatest risk from trade activities on the island,

the only logic they are likely to embrace is the logic of extinction.

Captive breeding for conservation purposes (‘conservation

breeding’) may therefore now be a lifeline for the Rufous-fronted

Laughingthrush (Collar etal. 2012). However, the species has been

bred only with some difficulty and only, apparently, in two

European institutions, Tierpark Berlin (Kaiser 2006) and Prague

Zoo (Pithart 2009). It has proved an aggressive and problematic

species to keep, and endeavours to develop a significant captive stock

may only be worth making once the evidence is clearer about its

status in the wild.

ACKNOWLEDGEMENTS

We are most grateful to T. J. Trombone for locality data from specimens at

the American Museum of Natural History (AMNH), and to the authorities

at the Zoological Museum, Bogor (ZMB), and Naturalis, Leiden, for access

to their collections. K. D. Bishop, N. Brickie, J. A. Eaton, F. R. Lambert, D.

Liley, R. Sozer, C. R. Shepherd and A. Vaidl all greatly helped with our

inquiries. R Andrew and V. Nijman were most helpful referees.

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N. J. COLLAR, BirdLife International, Girton Road, Cambridge

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S. (BAS) van BALEN, Basilornis Consults, Muntendampad 15,

6835 BE Arnhem, Netherlands

FORKTAIL 29 (2013): 19-24

Species limits in the Golden Bulbul Alophoixus

(Thapsinillas) affinis complex

N. J. COLLAR, J. A. EATON & R. 0. HUTCHINSON

The Golden Bulbul Thapsinillas affinis of the Moluccan islands, Sula archipelago, Banggai islands, Togian islands and Sangihe, Indonesia,

was until recently treated in Alophoixus before being placed in the resurrected genus Thapsinillas and shortly afterwards split into Northern

and Southern Golden Bulbuls T. affinis and T. longirostris, but with a general consensus that a break-up into more species was required. We

used plumage and morphometric analysis of museum specimens, supplemented by vocal samples, to determine where new species limits

might be drawn. We found that the nine generally accepted subspecies break down into seven full species, five monotypic and two with

two subspecies each: T. chloris on Morotai, Halmahera and Bacan (small, featureless; undifferentiated olive-green lores and ear-coverts,

blackish base to submoustachial area; song reportedly a 'jumbled babbling'); T. lucasi on Obi (round yellow lores, yellow-tinged ear-coverts,

seemingly simple often squeaky-toy-like vocalisations); T. affinis on Seram with race flavicaudus on Ambon (larger than previous two, with

half-wedge yellow lores, broad yellow tips to tail, song a group of strong rich flat whistles); T. mysticalis on Buru (half-wedge yellow lores,

partial yellow eye-ring, olive-green underparts, olive-grey tail, whistled phrases recalling domestic canary); T. longirostris on Sula with race

harterti on Peleng and Banggai (longest-billed, large, undifferentiated olive-green lores, song a loud jumble); T. aurea on theTogian islands

(golden-yellow underparts, vague half-wedge yellow lores, blackish frontal supercilial line, yellow-tinged rump, song seemingly more

complex than in longirostris) and T. platenae on Sangihe (vivid yellow chin and submoustachial area to throat and breast, bright yellow

triangular lores, almost-complete yellow eye-ring, song seemingly simple and nasal). Comprehensive vocal sampling and molecular work

may shed light on the origins and colonisation routes of this geographically unusual cluster of species.

INTRODUCTION

The taxonomy of the Golden Bulbul Alophoixus ( Thapsinillas )

affinis complex of Wallacea, Indonesia, has long been considered

problematic, owing to the considerable variation in plumage pattern

and size shown by most of its subspecies (Hartert 1922, Delacour

1943, White & Bruce 1986). These subspecies possess an unusual

and indeed unique distribution for a species in the region, in the

geographic sequence given by White & Bruce (1986) as follows:

chloris (North Moluccas: Morotai, Halmahera, Bacan); lucasi (Obi);

affinis (Seram); flavicaudus (Ambon); mysticalis (Buru); longirostris

(Sula); harterti (Peleng, Banggai); aurea (Togian Islands) and

platenae (Sangihe).

It is perhaps a measure of the uncertainty surrounding this

complex that it has appeared in so many generic guises in the past

hundred years. Until at least 1922 it was largely treated in Criniger

(e.g. Wallace 1862a, b, 1863, Blasius 1888, Hartert 1903, 1922),

but Delacour (1943) placed it in Microscelis (subgenus Iole ), Rand

& Deignan (I960), Morony et al. (1975) and Andrew (1992) in

Hypsipetes , White & Bruce (1986) and Coates & Bishop (1997) in

Ixos, and Sibley & Monroe (1990) and Inskipp et al. (1996) in

Alophoixus. Finally Dickinson & Gregory (2002) resurrected the

genus Thapsinillas for the complex (a decision we follow hereafter),

citing as diagnostic characters ‘typically dark oily green [plumage] ,

relieved by areas of yellow in some forms; crown not crested and

feathers only slightly elongated ; bill much like Iole but perhaps more

hooked and with lower mandible deeper; rictal bristles fewer and

weaker’, but unaccountably omitting mention of the key criterion

in the original description, namely that ‘from all the related genera

with lengthened nostrils Thapsinillas may easily be distinguished...

by its very short tarsus, this being considerably less than the exposed

culmen’ (Oberholser 1905).

Continuing this theme of taxonomic hesitancy, both Dickinson

& Gregory (2002) and Dickinson & Dekker (2002) suspected that

the variation between the subspecies in this resurrected genus ‘will

justify subdivision into two to four species’. However, Delacour

( 1 943) bluntly cited ‘distribution’ as the reason to resist a split into

two species based on ‘size and tail pattern’ (larger taxa with

‘particolored tail, dark olive and bright yellow’, smaller ones

‘strangely similar to M. ictericus (= Yellow-browed Bulbul Iole indica

in Inskipp et al. [1996]). By contrast, Fishpool & Tobias (2005)

took what they regarded as ‘a preliminary measure’ by separating

the ‘Northern Golden Bulbul’ T. longirostris (with chloris, lucasi ,

harterti, aurea and platenae) from ‘Southern Golden Bulbul’ T.

affinis (with flavicaudus and mysticalis) on account of reported vocal

differences between these groups, thereby ‘drawing attention to the

broadest rift in the complex, and paving the way for appropriate

fieldwork and research into the song, morphology and genetics of

all taxa involved’. These authors, like Dickinson & Dekker (2002),

judged that ‘further subdivision’ would almost certainly be

required, ‘in view of significant differences between the various

island populations’. This was partially achieved by Rheindt &

Hutchinson (2007), who, without going into detail, considered

‘Southern Golden Bulbul’ to comprise two morphologically and

vocally distinct species, Buru Golden Bulbul T. mysticalis and Seram

Golden Bulbul T. affinis (including flavicaudus).

Steadily accumulating evidence on apparent differences in

vocalisations of most of the taxa in the Thapsinillas affinis complex

now prompts a more detailed review of their morphological and

morphometric characters in order to attempt to reach a further stage

in the revision of the Golden Bulbul complex. As Fishpool & Tobias

(2005) observed, this is important not least because ‘some island

races would prove to be very rare...’ such that ‘taxonomic review is

vital for the compilation of a realistic conservation strategy for

Wallacea, and must be made a priority’.

METHODS

We considered one line of hard evidence in this review, namely

plumage and mensural characters from museum material, and

supplemented it with morphological evidence from photographs

as well as recordings and reports of vocalisations.

Museum specimens of Golden Bulbuls were examined (NJC)

in the Natural History Museum, Tring, UK (NHMUK), Naturalis,

Leiden, Netherlands (Naturalis), Staatliches Museum fur Tierkunde,

Dresden, Germany (SMTD), Staatliches Naturhistorisches

Museum, Braunschweig, Germany (SNMB) and Zoologisches

20

N. J. COLLAR, J. A. EATON & R. O. HUTCHINSON

Forktail 29 (2013)

Museum (Museum fur Naturkunde), Berlin, Germany (ZMB).

Each specimen was measured (by NJC) for length of bill (skull to

tip), tarsus, wing (curved) and tail (tip to point of insertion), the

characters of each taxon were logged in a matrix, and representative

specimens were photographed. From these collections the numbers

of specimens by taxon and island were:

• chloris — North Moluccas: 39 specimens, 10 from Morotai, 16

from Halmahera, 13 from Bacan (11 males [m], 8 females [f],

20 unsexed M)

• lucasi — Obi: 13 (7 m, 5 f, 1 u)

• affinis — Seram: 12 (4 m, 3 1, 5 u)

• flavicaudus — Ambon: 8 (6 m, 1 f, 1 u)

• mysticalis — Buru: 21 (4 m, 10 1, 7 u)

• longirostris — Sula (Taliabu & Mangoli): 23 (7 m, 2 f, 14 u)

• barterti — Banggai (Banggai & Peleng): 13(1 m, 2 f, 10 u)

• aurea — Togian: 2 (1 m, 1 f)

• platenae — Sangihe: 3 (3 m)

The large number of unsexed specimens and an occasional

numerical bias in the sexed specimens prompted a comparison

of males only (Table 2), but the full figures and standard

deviations given in Table 1 are used in the analysis of character

difference below.

Photographs of live birds were assembled from our own

collections (JAE, ROH), from those of colleagues, contacts and

friends, and (with due care as to identification and provenance)

from the internet (notably Oriental Bird Images). Sound recordings

were likewise assembled from our own collections (JAE, ROH),

Table 1. Means and standard deviation (in brackets) of four

morphometric variables in all specimens of the Thapsinillas complex.

Notes: a = sample size reduced by 1; b = sample size reduced by 2; c =

sample size reduced by 6. These reductions were caused by damage

to the parts being measured or (in the case of tarsi) their inaccessibility

(being tucked tightly against the body).

Table 2. Means of four morphometric variables in male specimens of

the Thapsinillas complex. Note: 3 = sample size reduced by 1 .

those of others and the internet (AVoCet [AV],Xeno-Canto [XC]

and the Internet Bird Collection [IBC]). They were compared

qualitatively and informal descriptions and transcriptions of them

prepared. Use of capitals in the transcriptions indicates emphasis

(volume).

We measured the degree of phenotypic differentiation between

each taxon using a system in which 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) scores 3; a medium character

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

colour) scores 2; and a minor character (weak difference, e.g. a

change in shade) scores 1 ; a threshold score of 7 is set to allow species

status; species status cannot be triggered by minor characters alone,

and only three plumage characters, two vocal characters, two

biometric characters (assessed for effect size using Cohen’s d where

0.2-2 is minor, 2-5 medium, 5-10 major and >10 exceptional),

and one behavioural or ecological character may be counted (Tobias

et al. 2010). Where additional characters are apparent but under

these rules cannot be scored, the formula ‘ns [1]’ is used, signalling

‘not scored’ but giving in parenthesis the estimated value of the

difference in question.

RESULTS

We review each taxon in turn for its diagnostic morphological,

morphometric (Tables 1 and 2) and acoustic distinctiveness.

However, the acoustic component of the analysis remains

qualitative, because the vocalisations of each taxon appear to be

variable and complex, so that only tentative and general comments

on their diagnostic distinctiveness can be ventured from the limited

and fragmentary material available. From this evidence a shared

pattern of song nevertheless seems to exist between all taxa, which

involves a hesitant series of staccato nasal or guttural notes that

accelerate and switch abruptly either to a short jumble of babbled

and fluty notes on often widely differing pitches or to a short series

of fairly even whistles; but most taxa sound in varying degrees

different, and if these findings are replicated widely by other

recordings in future then they will add substantially to the case made

below for the redrawing of species limits based on morphology.

Photographs and museum label data indicate that there are no

significant differences in the bare -part colours of any of the taxa:

basically the bill is shiny black to plumbeous, reflecting light and

looking whitish at some angles or in some photographs; the legs

are brownish-grey; and the iris is reddish-brown to brown. There

are slight variations in how museum labels report iris colour: for

example, for the taxon mysticalis NHMUK 1969.29.203 gives ‘iris

brown’, 1923.9.15.91 iris dark crimson’ and 1923.9.15.92 ‘eye red’,

while the describer, Wallace (1863), also gives ‘iris red’, although

photographs repeatedly show reddish-brown irides. Hombron &

Jacquinot (1841) likewise gave 'iris rouge for their new species

affinis , but in photographs it is reddish-brown. Two of the three

known specimens of the very rare platenae are labelled by the

collectors as having 'iris: rot-braun.

Sample sizes of specimens of aurea and platenae were

respectively two and three; and recordings of all taxa were

inadequate in number, duration and representativeness. However,

no clinching evidence depends on data stemming from these limited

sources.

In the following account, the size and shape of (yellow) lores

are mentioned and require definition here. ‘Round’ (taxon lucasi )

lores means that the shape of the yellow patch is large and relatively

circular, and comes into contact with the leading edge of the eye.

‘Half-wedge’ (taxa affinis, flavicaudus, mysticalis and aurea )

indicates that the patch of yellow is compressed into a flat triangular

Forktail 29 (2013)

Species limits in the Golden Bulbul Alophoixus (Thapsinillas) affinis complex

21

bar close to the line of the upper mandible and separated from the

eye by an olive-green area. ‘Triangular’ (taxon platenae) describes a

fuller area of yellow than the wedge, extending to the eye.

Taxon chloris (Morotai, Halmahera, Bacan)

This form is characterised by its small size (it is the smallest of the

taxa in the complex) and its relatively featureless plumage; no

differences were apparent between the three island populations. It

differs from its geographically and morphologically closest relative,

lucasi of Obi, by its olive-green vs yellow lores (3), olive-green vs

olive -yellow ear-coverts (1), blackish base to submoustachial area

vs all olive-green (2) and slightly smaller size and distinctly shorter

wing (effect size -2.28) (2) — total score 8.

Originally described by Wallace (1862a) under the pre¬

occupied name simplex, this form was renamed and further

described by Finsch (1867), who pointed out that Wallace failed

to mention the blackish submoustachial line. Finsch found this a

very distinctive (‘ ganz besonders’) character, but in specimens

examined for this review it proved to be constant but somewhat

variable in strength.

Fishpool & Tobias (2005) provided a description (‘a hurried,

cheery, jumbled babbling’) that conforms closely with the general

structure of Thapsinillas songs available to us. FFowever, brief

recordings by ROH of two consecutive song strophes consist (after

2-3 brief staccato introductory twis notes) of three or so simple

clear paired whistles, high-pitched at the start but each pair slightly

lower than the preceding, morphing subtly into a slightly more

drawn-out double-whistle with the stress on the first syllable, each

again slightly lower than the last: pi-pi, pi-pi, pi-pi, wiwi, wiwi, wiwi,

wiiwii, thus fairly closely resembling the falling-pitch song of

T. affinis (below). Otherwise the only recording we have found is

of a bird giving quiet thin sii calls in apparent mild alarm or for

contact (IBC video under T. longirostris, A bird softly calling from

a branch’).

Taxon lucasi (Obi)

Hartert (1922), while itemising Rothschild’s type specimens and

therefore not reviewing the Golden Bulbul complex in any detail,

remarked of lucasi , which he himself established as a full species

(Fdartert 1903), that ‘though differing by itsyellow lores and larger

size, [it] can hardly be anything but a subspecies of chloris', and

lumped it accordingly (albeit keeping chloris separate from affinis).

However, the morphological differences with chloris, as scored

above, gainsay this judgement.

The island of Obi is roughly equidistant from Seram, Buru and

Taliabu, where three further relatives of lucasi occur, respectively

affinis, mysticalis and longirostris. Of these, lucasi is closest in size

and general structure to mysticalis and remotest from longirostris,

but differs in turn from

• mysticalis by its shorter bill, tarsus and tail (effect size for bill

-3.22) (2); larger, much rounder yellow lores (2); lack of yellow

partial eye-ring (2); largely yellow chin to vent vs largely (yellow-

tinged) olive chin to vent (3); yellower ear-coverts (ns [1]) —

total score 9;

• affinis by its smaller size (effect size for bill -4.83) (2); larger,

rounder yellow lores (2); yellower ear-coverts and

submoustachial area (at least 1); paler and less extensive olive-

green on breast and flanks (ns [ 1 ] ) ; lack of yellow tips to

uppertail-coverts (ns [1]); olive-grey vs broadly yellow-tipped

and -edged rectrices with entire undertail bright yellow (3) —

total score 8;

• longirostris by its smaller size (effect size for bill -6.03) (3); large

round yellow vs olive-green lores (3); all-olive-grey vs bright

yellow-fringed (on inner webs) rectrices (3); narrow whitish vs

narrow yellow inner fringes to tertials ( 1); yellower ear-coverts

(ns [1]) — total score 10.

Recordings kindly sent by M. Thibault reveal only very simple

calls: (a) a flat nasal penetrating tuuu-tuuu-tuuu-tuuu (3-4 notes

separated by short pauses); (b) a high, thin, dropping-then-rising

TSIIiuuuuii, starting like a squeaky toy but ending more richly

whistled, this evidently the tweeuwip described by Linsley (1995)

and mentioned in Coates & Bishop (1997); and (c) an equally high

thin squeaky toy zu-WIIIT! zu-WIIIT! zu-iVIIIT! — these last

sounds not dissimilar to those recorded from platenae (see below)

but much thinner in tone, lacking the latter’s thrush-like richness.

Linsley (1995) also mentioned groups giving ‘raucous calls

reminiscent of Charmosyna placentis although without the harsh

or scratchy quality of that species’.

Taxon affinis (Seram)

Morphological differences from lucasi (and by extension chloris ),

aurea and platenae are scored above and below. It differs from

• chloris by its greater size (effect size for bill length 4.68) (2);

half- wedge yellow lores vs all olive-green lores (2); yellow tips

to uppertail-coverts (1); rectrices broadly tipped and edged

yellow (entire undertail bright yellow) vs olive-green (3) — total

score 8;

• mysticalis by its slightly larger size (effect size for bill length

1.99) (1); lack of partial yellow eye-ring (2); yellow vs olive-

green belly to vent (3); rectrices broadly tipped and edged yellow

(entire undertail bright yellow) vs olive-green (3); yellow tips

to uppertail-coverts (ns[ 1 ] ) — total score 9;

• longirostris by its rather smaller size and notably shorter tail

(effect size for latter -4.82) (2); half-wedge yellow vs olive-green

lores (2); darker and more extensive olive-green breast (2);

different tail pattern, with broad yellow tips and all-yellow

undersides vs broad yellow edges on both surfaces (3) — total

score 9.

A recording by F. R. Lambert (AV4805, XC67566) captures

a single song strophe which starts with some scratchy clucking

calls and then abruptly turns into a sequence of seven strong rich

flat whistles, each longer and perhaps a shade lower in pitch than

the previous, the last note most obviously lower: p’tupwupwud’p-

p’TI-WI-WII- Will- Will I- Will II- WUUUUU. Another, by

JAE, involves a very similar song but with the last two notes

rolled throatily. Rheindt & Hutchinson (2007) also describe this

song (‘a clean descending melodious whistle’) and present a

sonogram of it. Isherwood et al. (1997) found that at one of

their study sites (Wae Salas) ‘this species was found to possess a

distinct variety of the usual call’, and Coates & Bishop (1997)

independently mentioned two types of song (see ‘Conclusion and

conservation’).

Taxon flavicaudus (Ambon)

Bonaparte (1850) gave a nugatory diagnosis of this taxon

(translated from Latin: ‘olivaceous green, greenish-yellow

below; throat, undertail mostly strong yellow’), but his

scientific name nails the only discernible plumage difference from

affinis-. in the rather small sample in NHMUK the specimens

appear to have less olive markings in the rectrices than those of

affinis and hence seem more fully yellow-tailed. White & Bruce

(1986) suggested that flavicaudus males ‘tend to be lighter and

yellower dorsally and on the breast, with a deeper yellow throat’,

but admitted that ‘it is only a slightly differentiated form’.

Measurements suggest that flavicaudus is also marginally larger than

affinis (Tables 1 and 2). Consequently, always accepting that a larger

sample of flavicaudus may show all these slight differences to be

inconstant, flavicaudus is provisionally retained here as a valid taxon,

but it is clearly conspecific with affinis. Given the proximity and

biogeographical unity of Seram and Ambon, this is hardly

surprising.

Recordings of flavicaudus could not be found.

22

N. J. COLLAR, J. A. EATON & R. 0. HUTCHINSON

Forktail 29 (2013)

Taxon mysticalis (Burn)

Differences from lucasi (and by extension cbloris ) and ajflinis

(including flavicaudus) are scored above; chose from aurea and

platenae are given below.

Wallace (1863) gave this taxon the name mysticalis (not,

incidentally, mystacalis ), meaning moustached (Jobling 2010),

evidently because of its ‘remarkable half-yellow gape-bristles’. This

character (rictal bristles yellow basally, black distally) is not

particularly striking in specimens or photographs, nor is it unique

within the complex, being shared with platenae and to a lesser

degree with other taxa which show yellow lores; but olive-lored

member taxa have all-black rictal bristles). Unique to mysticalis ,

however, is the extent of olive-green on the undersides, with only

vague areas on the chin and vent being distinctly shaded yellow,

the rest having the merest yellow tinge (score 3). It further differs

from longirostris (including barterti) by its considerably smaller size

and notably shorter tail (effect size -3.62) (2); half-wedge yellow

vs olive-green lores (2); partial yellow eye-ring (ns [2]); dark olive-

grey rv bright yellow-fringed rectrices (3); narrow whitish vs narrow

yellow inner fringes to tertials (ns [2]) — total score 10.

A recording by F. R. Lambert (AV4147, XC 67565) consists of

single nervous low clucks, with occasional higher, very rapid

chatters, and three times a drawn-out, flat whistle with a very curt

downward inflection at the end, tweeeee(ub). These three calls also

feature in recordings by JAE, but with the drawn-out whistle

starting with a distinct short higher strangled tone,

tswiUUUUUU(uh). However, other recordings by JAE also capture

a series of song-phrases, starting with hesitant staccato accelerating

notes before breaking into longer, musical whistles on (sometimes

greatly) varying pitches and sometimes with glissandos, somewhat

reminiscent of a domestic canary: pip up... pip-up... pipup-pipupipu

WEE- WEE-WEE-puu -puu-puu- WEE-puii-PII- WEE- WEE-

WEE. Jepson (1993) reported: ‘Call comprised a descending “si-

si-seeow seeow seeow”, and typical bulbul chattering notes’.

Taxon longirostris (Sula)

As the name given it by Wallace (1862b) indicates, this form is the

longest-billed taxon in the complex, although flavicaudus runs it

close, and it is altogether the largest form, with the possible exception

of aurea. It differs from lucasi (and by extension cbloris ), ajflnis

(including_/7 avicaudus), mysticalis , aurea and platenae by the

characters scored under those taxa. It differs little from barterti (see

below).

Recordings of longirostris by ROH all contain a song that

consists of a throaty, rolling cb(a)rrrr, rapidly repeated several times

and accelerating before breaking into a loud jumble of short whistled

notes, some very clear: cbarrr... cbarrr... charrr-charrr-charrr-

d idly!) 0 OdidlyD 0 O dully I) 0 0 ; or cbarrr... cbarrr... charrr-charrr-

charrr-wididlyWAAbeDIbeDI, etc. However, the cbarrr component

may nor be obligate, given the evidence under barterti below.

Taxon harterti (Peleng, Banggai)

Stresemann (1912) separated this form from longirostris on account

of the darker olive coloration of the breast, less yellow upperparts

and narrower yellow edges to the outertail. Specimens in SMTD,

where 10 barterti are held alongside 7 longirostris , confirm this

diagnosis; but as Eck (1976) observed, barterti is ‘only subtly

differentiated’ (which is true also of its morphometries: see Table

1) and on morphological grounds it must remain a subspecies of

longirostris , as biogeography might predict.

Recordings by ROH reveal song-phrases similar or identical to

those of longirostris ; however, two by P. Verbelen (AV3344, 3345)

are of a singer that gives several clucks and only one very brief cbarrr

before launching into its song, suggesting that the cbarrr

component may be a separate call that is sometimes run together

with the song.

Taxon aurea (Togian)

While noting the morphological proximity of this lorm to

longirostris (which is indeed the closest taxon in plumage and size),

Walden (1872) diagnosed it on its smaller size, ‘much shorter bill’

and ‘bright golden colouring of its plumage’. However, while a

female specimen (ZMB 2000/26784) conforms in these respects,

the type of this taxon, a male, actually has wing and tail longer and

bill only 1.6 mm shorter than the mean for two male longirostris

(Table 2). Both specimens are distinguished by their notably more

golden-yellow underparts (2); much reduced yellow fringes to the

tips and inner vanes of the rectrices (2); vague half-wedge yellow

lores below a very narrow blackish-brown frontal supercilial line

and notably darker olive-green crown (2); rump a shade yellower,

less green (ns [1], well shown but perhaps a shade too obvious in

Fishpool & Tobias 2005: 236); and presumed shorter bill (allow

1) — total score 7.

Acoustically, aurea seems rather close to longirostris /barterti.

However, multiple recordings by ROH on different dates suggest

that (a) the homologous call in aurea to the 'cb(a)rrr call of

longirostris lacks the latter’s rolling throaty quality, and (b) the short

fluty babbling song is somewhat abrupt and simple in longirostris

whereas in aurea it can be more protracted and typically ends with a

set of very rich notes, slightly tailing off in pitch and volume, vaguely

recalling the yaffling cadence of a Green Woodpecker Picus viridis.

Taxon platenae (Sarsgihe)

This is the most isolated, most threatened and in some ways most

distinctive form in the Golden Bulbul complex. Blasius (1888),

working with two syntypes (illustrated, with a photograph of one

of them, in Hevers 2004), accurately characterised this bird as closest

to aurea and longirostris but distinguished by its shorter bill (this is

true for longirostris but not tor aurea). almost entirely uniform olive-

green upperparts, and vivid yellow colour of the chin, throat,

submoustachial area, eye-ring and inner vanes of all five outer

rectrices. Our own examination of the only three specimens in

existence (SNMB N 13945 and N43300, and RMNH [Naturalis]

84768) indicates that it is distinguished from all other taxa by its

bright yellow triangular lores (much fuller and brighter than the

yellow triangular lores of mysticalis against which it is here scored

on this feature) extending to and contiguous with the eye-ring (2);

bright yellow eye-ring, only broken by a narrow gap at the rear of

the eye (much more obvious and complete than in mysticalis , in

which it is confined to the ‘brow’ and a short arc on the lower rear

edge) (3); yellowish ear-coverts and yellow submoustachial area,

producing a broad yellow throat (ns [2]); and very broad yellow

fringes to the inner vanes of the rectrices extending the length of

the feathers, creating a different pattern from other taxa (2) — total

score 7.

A recording by P. Verbelen (AV3347) consists of a vigorously

delivered series of fairly short, simple strophes composed of little

groups of repeated thrush-like whistles. Recordings of this form by

ROH reveal a consistent pattern of song, comprising two short

abutting components, (a) four nasal but rich notes, each rising in

pitch but each lower than the previous, the last cutting to (b) usually

three high whistled notes, approximately: cui-cui-cui-cui-DEEP-

pDEEP-pDEEP! (As noted above, in structure these sounds

vaguely resemble those on a recording of T. lucasi , but are much

richer and less strangled in tone.)

CONCLUSION AND CONSERVATION

Fishpool & Tobias (2005) separated the Golden Bulbul into

Northern longirostris (with cbloris, lucasi , barterti, aurea and

platenae as races) and Southern ajflnis (with flavicaudus and

mysticalis as races) on account of their songs, the former lacking

Forktail 29 (2013)

Species limits in the Golden Bulbul Alophoixus ( Thapsinillas ) affinis complex

23

the ‘long sliding notes and descending cadence’ of the latter, affinis

and flavicaudus possessing ‘a distinctive mournful series of sweet

and minor-key notes, lasting 2-4 seconds, slightly erratic or

meandering in pace and note length, but essentially slow and

leisurely, sliding down scale almost throughout’, mysticalis ‘vaguely

similar but much more complex’ — and hence a reason why Rheindt

& Hutchinson (2007) recommended its separation Irom affinis.

However, while Coates & Bishop (1997) support the account of

the voice of affinis (‘main song... a lovely descending series of c. 15

short, clear, mellow whisdes... slightly slurred as the song dies away’)

they also mention a second song type, ‘a rapidly swelling series of

20-30 pure, high-pitched whistled notes that climbs to a notably

high pitch and ends abruptly’. Moreover, the clear resemblance of

songs of chloris and affinis tends to confound the notion of a north-

south divide in song types. This all suggests that the vocalisations

of the taxa in this complex may be considerably more varied but

also perhaps ultimately more homologous than we yet know, and

that the sample used in descriptions above should not be considered

anything more than partially representative.

Even so, from the very limited material available to us we derive

the impression that vocal differences largely support the seven-way

split of the Golden Bulbul complex which the morphological

evidence indicates, using the scoring system of Tobias et al. (20 1 0) :

Halmahera Golden Bulbul Thapsinillas chloris

Morotai, Halmahera, Bacan

Obi Golden Bulbul Thapsinillas lucasi

Obi

Seram Golden Bulbul Thapsinillas affinis

T. a. affinis Seram

T. a. flavicaudus Ambon

Buru Golden Bulbul Thapsinillas mysticalis

Buru

Sula Golden Bulbul Thapsinillas longirostris

T. 1. longirostris Sula

T. 1. harterti Peleng, Banggai

Togian Golden Bulbul Thapsinillas aurea

Togian Islands

Sangihe Golden Bulbul Thapsinillas platenae

Sangihe

The conservation status of these seven species will require

formal assessment against the IUCN Red List criteria, but a few

preliminary remarks may be made here. From evidence in Fishpool

& Tobias (2005), our own observations in the field (JAE and ROH)

and material cited below, the first six species in the list above are

relatively common in their various woodland/ forest habitats.

Poulsen & Lambert (2000) tabulated records of chloris

(Halmahera) indicating a high encounter rate, with birds found

(albeit less commonly) even in mangrove. Linsley (1995) saw lucasi

(Obi) in ‘small numbers (less than ten)... daily’, with two instances

of breeding evidence ‘in scrub on the edge of disturbed forest’.

Bowler & Taylor (1989) reported affinis (Seram) ‘common and

widespread... in forested areas’ from sea-level up to c.900 m, while

JAE saw them up to at least 1,300 m; Isherwood et al. (1997) also

found the species common. Jepson (1993) called mysticalis (Buru)

‘common and widespread... in all types of forest’ (confirmed in

Poulsen & Lambert 2000, and by JAE, ROH pers. obs.). Stones et

al. (1997) found longirostris (Sula, specifically Taliabu) ‘abundant

at each study site, in all habitat types surveyed, but most common

in primary forest, both lowland and montane’ (confirmed by JAE,

ROH pers. obs.), while Indrawan et al. (1997) reported harterti

(Peleng) as ‘commonly seen’ in groups of three to four birds... in

degraded forest at Monggias’ (confirmed by JAE, ROH pers. obs.).

Coates & Bishop (1997) were concerned that aurea (Togian

Islands) was ‘apparently rare and local’, but Indrawan et al. (2006)

documented records from three of the seven larger islands in the

group, finding it ‘relatively frequently’ on Togian itself and ‘relatively

common’ on Walea Bahi (confirmed by ROH pers. obs., and J. Riley

in litt. 2013).

The status of platenae (Sangihe) is, however, worrying.

Although Bishop (1992) observed it ‘commonly in secondary

woodland and mixed tree crop plantations’ during a visit over lb-

19 May 1986, others have not been able to repeat this finding (Riley

1 997a,b). A year before, on 30 May 1 985, a male specimen (RMNH

84768) was collected on Gunung(Gn) Sahendaruman in ‘primary

forest on eastern slope: 750 m: S of Liwung and SW of Kuma’

(Naturalis label data) by F. G. and C. M. Rozendaal, but it took

until November 1996 before the species was seen again, with records

of three birds twice and one bird once on three days, all evidently

in the same area-on Gn Sahengbalira (Riley 1997b). These records

were the only ones in four months’ fieldwork in 1995 and 1996,

when the only local people to recognise photographs of the species

(presumably from museum skins) were ‘in the village closest to the

forest on Gunung Sahengbalira’ (Riley 1997b). Further fieldwork

on Sangihe between August 1998 and March 1999 led Riley (2002)

to suggest that platenae ‘is one of the island’s most endangered

species’, being found only on Gn Sahendaruman with an estimated

population of 50-230 birds. However, he noted that it was missed

at one locality when not calling but found to be common there

when it became vocal (Riley 2002), thereby confirming an earlier

remark that ‘this can be a cryptic species, despite its bright

coloration’ (Riley 1997b). Even so, visits to its small fragment of

remaining habitat on Gn Sahengbalira in recent years have not

produced any evidence to revise the view that this species is in

trouble: JAE and ROH found four birds in August 2004, although

a subsequent visit over two days in 2012 by ROH failed to record

any. Of other observers visiting the area this century, R Verbelen

saw several in November 2008 but B. Demeulemeester, R Gregory,

J. Hornbuckle, C. Robson and M. Thibault ( in litt. or verbally to

JAE, ROH) all failed to find it. Consequently, we judge that the

Sangihe Golden Bulbul now requires urgent attention in order to

secure its future.

Clearly it would be valuable if this new arrangement of

Thapsinillas were to be tested and corroborated by molecular study.

Such work might also reveal the biogeographic history and

colonisation routes of the taxa across this unusual range (which no

other species or genus shares) . Moreover, a far more comprehensive

sampling of vocalisations would also be of great interest, in part

simply to determine the variation within individual taxa, in part to

assess more confidently the degree of difference between taxa, and

in part to test whether such differences correspond to the hoped-

for molecular evidence.

ACKNOWLEDGEMENTS

We thank Robert Prys-Jones (NHMUK), Stephen van der Mije and Cees

Roselaar (Naturalis), Michaela Forthuber (SNMB), Martin Packert (SMTD)

and Sylke Frahnert (ZMB) for access to the specimens in their care, Alison

Harding (NHMUK) for finding and assembling copies of the earlier literature

on the complex for review, and those mentioned in the text for their recordings

and records of members of the complex. We also thank L. D. C. Fishpool and

J. A. Tobias for their helpful comments as referees.

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FORKTAIL 29 (2013): 25-30

Birds of Mys Shmidta, north Chukotka, Russia

VLADIMIR YU. ARKHIPOV, TOM NOAH, STEFFEN KOSCHKAR & FYODOR A. KONDRASHOV

A survey of avifauna was carried out in the Mys Shmidta area, north Chukotka, Russia from 8 June to 1 2 July 2011 . A total of 90 species was

recorded in the area, which together with literature data made a final list of 1 04 species. For several species this area is beyond the northern,

north-eastern or north-western limits of their known distribution. We collected new data for 19 globally or locally threatened species. Tundra

Swan Cygnus columbianus, Emperor Goose Anser canagica, American Golden Plover Pluvialis dominica, Western Sandpiper Calidris mauri,

Semipalmated Sandpiper C.pusilla, Northern House Martin Delichon urbica and Barn Swallow Hirundo rustica were all confirmed to be breeding.

Breeding of Brent Goose Branta bernicla nigricans, Spectacled Eider Somateria fischeri and Steller's Eider Polysticta stelleri was judged to be

'very likely'. There was no evidence for breeding of Ross's Gull Rhodostethia rosea despite several records. Two Eurasian Dotterels Eudromias

morinellus were recorded displaying for the first time in the area, but the status of the species is unclear. The area is important for Snowy Owl

Nyctea scandiaca, and as moulting grounds for Emperor Goose. Canada Goose Branta canadensis, Baikal Teal Anas formosa, Bar-tailed Godwit

Limosa lapponica, Slaty-backed Gull Larus schistisagus, Thayer's Gull L. thayeri, Black-headed Gull L. ridibundus, White-tailed Eagle Haliaeetus

albicilla, Steller's Sea Eagle H. peiagicus, Osprey Pandion haliaetus, Arctic Warbler Phylloscopus borealis and House Sparrow Passer domesticus

are more likely to be rare vagrants or migrants. An observation of a Pine Siskin Carduelis pinus is the first record for Eurasia.

INTRODUCTION

Mys Shmidta or the Cape of Shmidt is a prominent headland on

the Arctic coast of Chukotka, Russia, ft is a remote place with a

very harsh climate. Information on birds from this area is difficult

to obtain but has inherent value because several globally or locally

threatened species breed here and human impact on the habitat has

been small. No systematic checklist of birds of the area has been

made until now.

Since Portenko (1972, 1973) worked on the birds of the

Chukchi Peninsula, few ornithological papers have been published

on the region and no avifauna surveys have been made. Existing

publications include a short note by Tomkovich et al. (1991) on

the area around Mys Shmidta airport, several notes by Stishov (1991,

1 992, 1 997) including one on the bird community of the Ekvyvatap

river, two papers by Stishov & Maryuhnich ( 1 99 1 a, b) on particular

species and short communications on brief visits by Andreev &

Kondratyev (1996), Dorogoi (1996, 1997, 1998), Dorogoi &

Beaman (1998) and Menyushina (2000).

In this paper the results of an avifauna survey carried out in the

Mys Shmidta area, north Chukotka, Russia, in summer 2011 are

presented. Together with data from the literature the final list of

birds for the area is 104 species. The purpose of the investigation

was to survey this area for potential breeding grounds and suitable

habitats for the Spoon-billed Sandpiper Eurynorhynchus pygmeus

and to document the avifauna of this remote and hard-to-reach

region, which had previously received little ornithological attention.

STUDY AREA AND METHODS

During an ornithological expedition to the Mys Shmidta area

between 8 June and 12 July 2011 to survey breeding areas of the

Spoon-billed Sandpiper, the other avifauna was also surveyed. Birds

were identified mainly using binoculars and telescopes, and the

species, number of individuals and habitat was noted in each case.

Overall, research was focused on the Akatan lagoon and the

Ekvyvatap river delta, plus as much of the Kosa Dvukh Pilotov Spit

(Spit of Two Pilots) as could be accessed on foot or by kayak, the

area immediately surrounding Mys Shmidta settlement and an area

to the north-west of Mys Shmidta around the Erokynmanky lagoon

(Table 1). There were four main habitat types in the area: gravel-

sand, tussock tundra, grassy tundra and polygonal tundra. The spits

running north-west to south-east were mainly gravel and sand, either

pure gravel-sand or only sparsely covered with lichens and grass, at

least on the main narrow sections running immediately parallel to

the ocean. About 60% of the spits visited were covered by this type

of vegetation. The higher ground on the spits and the areas around

the lagoons were generally cotton-grass tussock tundra, and grassy

tundra was found on lowlands surrounding the spits, which were

still flooded until almost the end of June, being frequented by

different species of geese. Dried polygonal tundra was found in

patches around the Ekvyvatap river delta. Small hills around the

settlement were covered by typical tussock tundra.

The temperature in the region is generally cold; the highest

temperature experienced during the day was 14°C and the typical

daytime temperature was 5-7°C. Storms were f requent, with a week-

long storm in the first week of July with strong winds and heavy

snow.

Table 1. Main survey sites in the Mys Shmidta, north Chukotka area,

2011.

Ninety species were recorded in the area which together with

literature data yielded a final list of 104 species (Appendix 1).

Details of the most interesting and significant are given below,

26

VLADIMIR YU. ARKHIPOV etal.

Forktail 29(2013)

including new records, new breeding records, regional rarities and

globally threatened species.

Whooper Swan Cygnus cygnus

Two records: one across the Akatan lagoon on 18 June was pursued

by a Tundra Swan that appeared to strike it on the head and neck.

Another Whooper Swan stayed near a tundra lake with a Tundra

Swan on 23 and 24 June, in the Ekvyvatap delta. Images were

obtained as these records are far to the north of known nesting

areas (Krechmar & Kondrat’ev 2006).

Tundra Swan Cygnus columbianus

A common breeding species around Mys Shmidta: six nesting pairs

within 25 km2 were recorded in the Ekvyvatap delta tundra. Some

non-breeding pairs were also recorded, but no large groups were

seen. A considerable number of birds had a mix of traits attributable

to Cygnus columbianus bewickii or C. c. columbianus ; some

individuals that could be reliably identified as either bewickii or

columbianus belonged to mixed pairs. Of note, in all such mixed

pairs the males appeared to possess columbianus traits. Possibly the

study area is located in the area of overlap of the two forms (Rees

2006). Overall three nests with eggs were found: a nest in the

Ekvyvatap River delta with one egg on 14 June and three eggs on

19 June; a nest in the same area found on 23 June but not checked

for eggs; and a nest with three eggs found on 7 July on the

Nutechikun spit.

Snow Goose Anser caerulescens

Breeding was not recorded. From 9 to 28 June Snow Geese were

seen regularly with a maximum of 64 birds in several flocks on 13

June in the mouth of the Ekvyvatap River and 60 birds in four

flocks on 14 June; all flocks were flying north-west. All observed

individuals belonged to the white morph. Residents of Mys

Shmidta reported that there is a pronounced autumn migration,

occasionally reaching thousands of birds per day. Three Snow Geese

rings were obtained from local hunters; two individuals, a male and

a female, had been ringed as adults on Wrangel Island and one male

as a juvenile in Alaska, 45 km east of Deadhorse.

Emperor Goose Anser canagica

Near Threatened. Common breeding species in the Ekvyvatap delta

and surrounding area. W e observed several pairs holding territories,

including one chasing away a Parasitic Skua Stercorarius parasiticus.

A nest with three eggs was found in the Ekvyvatap delta on 14

June and a second nest was found nearby on 26 June. A pronounced

migration of Emperor Geese to the north-west of Mys Shmidta

was observed from 17 to 29 June. A maximum of 1,670 flew north¬

west between lOhOO and I6h00 on 27 June. It appears that this

species is expanding its range westwards, as it was not observed in

the vicinity of Mys Shmidta in the early twentieth century, but by

1970 it was breeding to the west ofUkouge lagoon (Portenko 1972)

about 100 km south-east of the eastern part of the surveyed area.

Hunters handed over a ring from a male bird collected at nearby

Ryrkaypiy settlement; the bird had been ringed as an adult in Alaska

37 km south-east of Chevak.

Canada Goose Branta canadensis

A solitary Canada Goose was observed on 19 and 20 June in the

Ekvyvatap delta. It was flying with two Greater White-fronted Geese

Anser albifrons and was slightly larger than them. Judging by several

traits, such as the relatively short bill and neck, it may be th eparvipes

subspecies, common in northern Alaska. It appears that this is the

first record of this form in Russia, apart from the introduced

population in Eurasia (Koblik etal. 2006). Prior to this observation

there was only one record of a wild Canada Goose in East Asia, seen

on Hokkaido, Japan, in 2006, which was also presumed to be a

parvipes form (Brazil 2009). The form minima was also observed in

the vicinity of Anadyr airport in Chukotka. A solitary bird was seen

in a flock of Brent Geese on a lagoon near the airport from 3 to 5

June 2011. Images of both birds were obtained.

Brent Goose Branta bernicla

Observed daily from 9 to 27 June in flocks and pairs in the

Ekvyvatap delta and nearby on the Akatan lagoon with up to 143

birds on 17 June. On 1 1 June one pair was observed to be possibly

breeding, but the birds were not seen on subsequent days. However,

breeding is possible, as sporadic breeding pairs were observed in

the first half of the twentieth century in the vicinity of Mys Shmidta

(Portenko 1972). This species is commonly hunted by the local

population; in the late 1990s one female shot in the vicinity of

Polyarniy settlement had been ringed as a juvenile in Alaska,

20 km to the south of Chevak.

Baikal Teal Anas formosa

A single male was swimming on a lake near Mys Shmidta on 21

June. The area is outside the current breeding range (Krechmar &

Kondrat’ev 2006), although in the first half of the twentieth

century the species was observed in the area relatively frequently

(Portenko 1972).

Spectacled Eider Somateria fischeri

Uncommon, possibly breeding in the tundra near the sea. Three

pairs were observed on 26 June on lakes in the Ekvyvatap delta.

Overall it was observed on 12 days with a maximum daily count of

32; all were flying to the north-west on 1 6 June, and 30 birds, mostly

young males, were observed on the sea near Mys Shmidta on 10

July. Stishov (1992) found the species to be common on lakes and

bogs around the lower reaches of the Ekvyvatap River.

Steller's Eider Poiysticta stelleri

Vulnerable. Common to uncommon at the time of migration.

Possibly breeding in the tundra on coastal lakes. Four pairs were

holding territories on 14 June in Ekvyvatap delta. The species was

regularly seen in the vicinity of Mys Shmidta, with a maximum of 60

birds, including some pairs, on 9 June. Unfortunately the species,

which is Vulnerable, is hunted in the area; eight rings were collected

from individuals shot near Mys Shmidta; all had been ringed in Alaska.

Bar-tailed Godwit Limosa lapponica

Vagrant; only one record of a female feeding on tundra near the

Ekvyvatap delta on 26 June. It is also a rare vagrant to Wrangel

Island (Stishov etal. 1991).

Red Knot Calidris canutus

Seven observations were made, all on the sand spits or mudflats

nearby: flocks of nine and five birds on 1 1 and 14 June on mudflats

near the mouth of the Akatan lagoon; three flew north across the

Akatan lagoon in the direction of Wrangel Island on 18 June; two

were in tundra on the Spit ofTynkergynpil’gyn lagoon on 20 June

with two more 1 .5 km away; in the same area on 21 June a bird was

resting and also in the same area two birds were seen on 22 June;

finally on 27 June three birds were in tundra on the spit of Akatan

lagoon. Although most records were made on lichen tundra —

suitable breeding habitat — displays were not seen or heard. Three

individuals had colour flags and had been ringed in Australia —

two with yellow flags in north-west Australia, and one with a green

flag at Moreton Bay, near Brisbane, Queensland.

Western Sandpiper Calidris mauri

Late arrivals; during the first survey on 14 June in Ekvyvatap delta

no more than five displaying males were seen, and on the following

days, several other birds appeared in nearby areas. Two breeding

Forktail 29 (2013)

Birds of Mys Shmidta, north Chukotka, Russia

27

areas were found near Mys Shmidta. The first included 10

territories on the Ekvyvatap delta tundra, where a nest with four

eggs was found on 25 June. The second area was to the north-west

of Mys Shmidta on the Nutechikun spit, where a nest with four

eggs was found on 7 July.

Spoon-billed Sandpiper Eurynorhynchus pygmeus

Critically Endangered. During the surveys the species was not

recorded. However, the expedition found suitable habitat in the

region, although limited in extent, and obtained invaluable

information on the habitat in the area. The last record of this species

in the area was in mid-June 1990 when several displaying males

and at least two pairs were found in the Ekvyvatap delta (Stishov

& Maryuhnich 1991a). In the 1970s Kishchinski (1988) found it

to be a common breeder at the Ukouge lagoon about 100 km south¬

east of the eastern part of the area surveyed.

' Semipalmated Sandpiper Calidris pusilla

Near Threatened. A rare breeding species in Eurasia, it was found

nesting in areas with sparse vegetation on gravel and sandy patches

of the tundra near lagoons. Breeding was recorded in two areas:

Ekvyvatap delta (2-3 territories) and in the vicinity of Mys Shmidta,

where on 9-1 1 June up to 10 males were displaying. A brood with

four two-day-old chicks was observed on 25 June on the edge of the

town. On 28 and 29 June in the vicinity of Mys Shmidta three more

broods were observed and one pair was making alarm calls.

The first nest of this species in this area was found on 8 June

1993 near the Mys Shmidta airport (Andreev & Kondratyev 1 996).

In 1997 at least three pairs were breeding between the airport and

the town; a nest at an advanced stage of incubation found on 1

July 1997, and on 4 July 1997 the same nest already had chicks.

Two more nests with chicks were observed nearby on the same day,

and one pair was observed between the airport and the town on 6

July 1997 (Dorogoi & Beaman 1998). Thus, the population of the

Semipalmated Sandpiper in the vicinity of Mys Shmidta continues

to thrive and, possibly, is growing slightly.

American Golden Plover Pluvialis dominicus

One territorial pair near a nest with a clutch of four eggs was found

in the Ekvyvatap delta, with observations continuing between 23

June and 25 June. Images of adults and nest were obtained. This is

the first documented breeding record in the Palearctic with both

birds confirmed as American Golden Plovers. Prior to this

observation an unsuccessful breeding attempt by an American

Golden Ploverwith a Grey Plover Pluvialis squatarola was reported

(Taldenkov 2006). In 1987-1990 on the lower reaches of the

Ekvyvatap River about 30% of plovers had traits that resembled

American Golden Plovers (Stishov 1991) and the same author

mentioned breeding of the American Golden Plover in the same

area in 1990-1993 without further details (Stishov 2004).

Eurasian Dotterel Eudromias morinellus

Three records in 2011. A solitary bird was displaying over the

Ekvyvatap delta tundra on 14 June. On 15 June in the same area

two displaying birds were seen about 1 km from each other, but

during later surveys in this area the species was not seen. On a flat

hilltop near the Odnobokiy stream a flock of nine was seen on 2

July. This species is regarded as rare since prior to our observations

it was reported in Chukotka only once since 2000 (Tomkovich

2007a; P.S. Tomkovich in litt.). Records of Eurasian Dotterels near

Mys Shmidta are contained in reports on breeding tundra birds in

Russia (Dorogoi 1996, 1997).

Thayer's Gull Larus thayeri

On 17 June, one bird was resting on the ice of Akatan lagoon with

Vega Gulls Larus vegae. The bird was about a third smaller

than the Vega Gulls. This is the third record of Thayer’s Gull for

Russia.

Black-headed Gull Larus ridibundus

Vagrant. On 18 June a bird in second-summer plumage was seen

near the mouth of the Akatan lagoon and on 20 June two adult

birds were swimming in a small bay near Kozhevnikova Cliff.

Ross's Gull Rhodostethia rosea

A summer visitor and migrant. Over 10 days between 10 June and

25 June juvenile and adult Ross’s Gull were observed at the

confluence of the Akatan lagoon with up to eight birds on 1 1 June.

Common Murre Uria aalge or Thick-billed Murre U. lomvia

We observed seven murres flying north-west over the sea on 10

July, but unfortunately too far away for definitive identification to

species level. The predominant species on Wrangel Island is Thick¬

billed Murre, but several hundred Common Murre also breed there

(Stishov era/. 1991).

Osprey Pandion haliaetus

On 3 July an Osprey was seen attempting unsuccessfully to catch

fish on Erokynmanky lagoon. Images were obtained. This is the most

north-easterly record of the species in Eurasia and approximately

600 km from its known breeding range (Chereshnev 2008).

White-tailed Eagle Haliaeetus albieilla

The Mys Shmidta area is about 500 km from the known breeding

range of the White-tailed Eagle (Chereshnev 2008). In June 2011

there were three records of White-tailed Eagles: a subadult was

sitting on the sea ice in the mouth of Akatan lagoon on 13 June, an

adult was seen in the mouth of Tynkergynpil’gyn lagoon on 23

June, and a first-year was flying along Two Pilot Spit on 27 June.

Steller's Sea Eagle Haliaeetus pelagicus

Vulnerable. An adult was observed on 27 June on the sea ice in the

mouth of Akatan lagoon, and images were obtained; after about

10 minutes, the bird flew away to the north-west. This is the

northernmost record of Steller’s Sea Eagle in Asia (BirdLife

International 2001, Chereshnev 2008).

Bam Swallow Hirundo rustica

Two pairs of Barn Swallows were regularly observed in the

settlements of Mys Shmidta and Ryrkaypiy. The pair in Mys

Shmidta was nest building on 29 June inside an abandoned house.

After the severe weather in early July, the nest was empty and birds

were not seen again. One and two Barn Swallows were seen on 12

and 17 June respectively, on the spit near the mouth of the Atakan

lagoon. Most of these records were of white-bellied birds. However,

in Mys Shmidta settlement birds with bright red bellies were

recorded alongside the white-bellied form: one bird on 8 June and

two on 29 June, it is possible that these birds were the American

subspecies H. r. erythrogaster. U nfortunately all observations of red-

bellied birds were too brief for identification to subspecies to be

confirmed.

Northern House Martin Delichon urbica

Breeding species in Mys Shmidta settlement. Nine nests were found

on two four-storey buildings in June 2011. Following the severe

storm at the beginning of July only two nests were still occupied.

This record is the most north-easterly breeding record of this

species in Eurasia.

House Sparrow Passer domesticus

Vagrant. On 17 June, an adult male was seen on Kosa Dvukh

Pilotov Spit near an abandoned hut; an hour later it was seen again

28

VLADIMIR YU. ARKHIPOV etal.

Forktail 29 (2013)

near another abandoned hut about 3 km to the east. The closest

breeding location is Pevek where breeding was reported in previous

years (Tomkovich 2007b) and at least 10 lamilies with broods were

seen there on 13 July 2011.

Pine Siskin Carduelis pinus

On 9 and 10 June 2011 we found a Pine Siskin on a sandy gravel

spit near Mys Shmidta (68.872°N 179.358°W). The bird was not

shy and it was possible to approach to a few metres and images

were obtained. This is the first documented record for Russia and

Eurasia (Arkhipov etal. in press).

DISCUSSION

During the 2011 expedition to northern Chukotka 90 species were

recorded, 30 of which were first records for the wider region.

Overall, this confirms the paucity of previous studies of this area.

Including additional data Irom the literature, a total of 1 04 species

has been recorded for the region indicating a relatively complete

list in comparison with nearby areas. Near Pevek town 75 species

have been recorded (Tomkovich 2007b) and 152 species have been

found on Wrangel Island (Stishov et al. 1991). Nevertheless, the

list may be far from exhaustive and be expanded by subsequent

visitors, especially in respect of migrant birds. Northern House

Martin and Barn Swallow were recorded attempting to breed

around the settlements; these species had never previously been

recorded breeding this far to the north-east.

Breeding of several other species on the borders of their known

western or north-western breeding range was recorded: Emperor

Goose, American Golden Plover, Western Sandpiper and

Semipalmated Sandpiper. Several other rare or endangered species

were observed that were suspected to be breeding but without direct

evidence, including two displaying Eurasian Dotterels and many

Snowy Owls Nyctea scandiaca , with up to 13 individuals recorded

per day. Emperor Geese moult in the area and a maximum day

count of 1,670 was recorded. Finally, several species were observed

which may be migrants, including Canada Goose, Baikal Teal, Bar¬

tailed Godwit, Slaty-backed Gull Larus schistisagus, Thayer’s Gull,

Black-headed Gull, White-tailed Eagle, Steller’s Sea Eagle, Osprey,

Arctic Warbler Phylloscopus borealis and House Sparrow. An

observation of Pine Siskin is the first record for Eurasia.

Disappointingly, in 2011 neither the Critically Endangered

Spoon-billed Sandpiper nor the Near Threatened Yellow¬

billed Loon Gavia adamsii, both of which have been recorded in

the area in previous years, were seen.

ACKNOWLEDGEMENTS

We thank Natalya Kveten and Oksana Makarova, heads of administrations

of Mys Shmidta and Ryrkaypiy for hospitality and for help with organising

our excursions. Warm thanks too to Pavel Tomkovich for useful comments

on local birds and ornithological literature. We are very grateful to The David

and Lucile Packard Foundation for the support to Birds Russia’s Spoon-billed

Sandpiper conservation programme in 2011 and to Evgeny Syroechkovsky

Jr, the leader of the Spoon-billed Sandpiper conservation team in Russia.

REFERENCES

Andreev, A. V. & Kondratyev, A. V. (1996) A new case of breeding

Semipalmated Sandpiper in Chukotka. Information Materials of the

Working Group on Waders 9: 34 (In Russian.)

Arkhipov, V., Koshkar, S. & Noah, T. (in press) First record of Pine Siskin

Carduelis pinus in Eurasia. BirdingASIA.

B i rd Life International (2001) Threatened birds of Asia: the BirdLife

International Red Data Book. Cambridge UK: BirdLife International.

Brazil, M. (2009) Birds of East Asia. London: Helm.

Chereshnev, I.A. (2008) Red data book of the Chukchi autonomous district

Vol. 1 . Animals. Magadan: Dikiy Sever. (In Russian.)

Dorogoi, I.V. (1996) Breeding conditions report for Ryrkarpiy settlement,

Chukotka, Russia, 1996. In M. Soloviev & P.Tomkovich, eds .ARCTIC BIRDS:

an international breeding conditions survey. (Online database), http://

www.arcticbirds.ru/info96/ru60ru11596r.html.Updated 11 December

2008. Accessed 23 August 2013.

Dorogoi, I.V. (1 997) The fauna and distribution of waders in North-East Asia.

Pp.53-87 in A.V. Andreev, ed. Species diversity and population status of

waterside birds in North-East of Asia. (Series: Biological Problems of the

North.) Magadan: North-East Science Centre, Far East Department of

the Russian Academy of Sciences. (In Russian.)

Dorogoi, I.V. (1998) Breeding conditions for waders in Russian tundras in

1 997: Mys Shmidta area. Information Materials of the Working Group on

Waders 1 1 : 40. (In Russian.)

Dorogoi, I.V. & Beaman, M. (1998) New data on breeding of Semipalmated

Sandpiper in Eurasia. Information Materials of the Working Group on

Waders 1 1 : 48-49. (In Russian.)

Koblik, E. A., Redkin, Ya. A. & Arkhipov, V. Yu. (2006) Checklist of the birds of

Russian Federation. Moscow: KMK Scientific Press. (In Russian with

English introduction.)

Kishchinski A. A. (1988) Ornithofauna of North-East Asia. History and modern

state. Moscow: Nauka. (In Russian.)

Krechmar, A.V. & Kondrat'ev, A.V. (2006) Waterfowl of North-East Asia.

Magadan: North-East Science Centre, Far East Department of the

Russian Academy of Sciences. (In Russian.)

Menyushina, I.E. (2000). Breeding conditions report for Mys Shmidta

settlement, Chukotka, Russia, 2000. In M. Soloviev & P.Tomkovich, eds.

ARCTIC BIRDS: an international breeding conditions survey. ( Online

database). http://www.arcticbirds.net/info00%5Cru36ru1 1600.html.

Updated 1 1 December 2008. Accessed 23 August 2013.

Inskipp, T„ Lindsey, N. & Duckworth, W. (1996) Checklist of the birds of the

Oriental region. Sandy UK: Oriental Bird Club.

Portenko, L.A. (1972) Birds of Chukotski peninsula and Wrangel island. Part I.

Leningrad: Nauka. (In Russian.)

Portenko, L.A. (1973) Birds of Chukotski peninsula and Wrangel Island. Part

II. Leningrad: Nauka. (In Russian.)

Sibley, D. (2000) The North American bird guide. New York: Chanticleer Press.

Rees, E. (2006) Bewick's Swan. London: T & A. D. Poyser.

Stishov, M.S. (1991) New data on distribution of some bird species on

Chukotka Arctic coast. Ornithological problems of Siberia

conference. Available: https://www.google.ru/#fp=5085c1 fl e640d3bd

&newwindow=1 &psj = 1 &q=%220rnithological + problems+of

+Siberia%22 +Conference+abstracts (In Russian.)

Stishov, M.S. (1992) Bird community of the lower Ekvyvatap River (Vancarem

Lowland, Chukotka). Russian Journal of Ornithology 1 (2):245— 25 1 . (In

Russian.)

Stishov, M. S. (1997) Breeding conditions report for Mys Shmidta settlement,

Chukotka, Russia, 1997. In M. Soloviev & P.Tomkovich, eds .ARCTIC BIRDS:

an international breeding conditions survey. (Online database), http://

www.arcticbirds.ru/info97/ru30ru11697r.html. Updated 11 december

2008. Accessed 23 August 201 3.

Stishov, M. S. (2004) Wrangel Island - truly natural but a natural

anomaly. Yoshkar-Ola: Izdatelstvo Mariyskogo Poligrafkombinata. (In

Russian.)

Stishov, M. S. & Maryuhnich, P. V. (1991a) Spoon-billed Sandpiper in the

western Vankarem Lowlands. Pp.1 25-1 26 in V. Yu. Il'yashenko and L. N.

Mazin, compilers The study of rare animals in the R.S.F.S.R. Moscow:

Central Science Research Laboratory for Game Management and

Nature Reserves. (In Russian.)

Stishov, M. S. & Maryuhnich, P. V. (1991 b) Buff-breasted Sandpiper in the

Cape Yakan area and Ekvyvatap River valley (Arctic coast of Chukotka).

Pp.1 26-1 29 in V. Yu. Il'yashenko & L. N. Mazin, compilers The study of

Forktail 29 (2013)

Birds of Mys Shmidta, north Chukotka, Russia

29

rare animals in the R.S.F.S.R. Moscow: Central Science Research

Laboratory for Game Management and Nature Reserves. (In Russian.)

Stishov, M. S., Pridatko, V. I. & Baranyuk, V. V. (1991 ) Birds of Wrangel. Island.

Novosibirsk: Nauka. (In Russian.)

Taldenkov, I. A. (2006) Record of mixed breeding of American Plover and

Gray Plover at northern Chukchi peninsula. Information Materials of the

Working Group on Waders 1 9: 39-41 . (In Russian.)

Tomkovich, P. S. (1998) Breeding conditions for waders in Russian tundras

in 1994. International Wader Studies 10: 132-144.

Tomkovich, P. S. (2007a) Population dynamics of the Dotterel Eudromias

morinellus: alarming thoughts on the background of poor knowledge.

Information Materials of the Working Group on Waders 20: 43-45. (In

Russian.)

Tomkovich, P. S. (2007b) Annotated bird list for Pevek town vicinity,

Chukotka autonomous area, the Far East of Russia. Ornithologia 34 (2):

1 76-1 85. (In Russian.)

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Shmidta, Chukchi Sea. Ornithologia 25: 1 75-176. (In Russian.)

Vladimir Yu. ARKHIPOV, Institute of Theoretical and

Experimental Biophysics RAS, Pushchino, Moscow 142290,

Russia; State Nature Reserve Rdeysky, Kholm, Novgorod 1 75270,

Russia. Email: v.arkhipov@rambler.ru

Tom NOAH, Bergstr.14 D - 15910 Schlepzig, Germany. Email:

tomnoah@t-online.de

Steffen KOSCHKAR, LiebigstraBe 91, D-35392, D-GieBen,

Germany. Email: fasciolata@web.de

Fyodor A. KONDRASHOV, Bioinformatics and Genomics

Programme, Centre for Genomic Regulation, Dr. Aiguader 88,

Barcelona, 08003 Spain; Universitat Pompeu Fabra (UPF), 08003

Barcelona, Spain; Institucio Catalano de Recerca i Estudis

Avangats (ICREA), 23 Pg. Uuis Companys, 08010 Barcelona,

Spain. Email: Fyodor. Kondrashov@crg.eu

Appendix

Annotated checklist of birds recorded in Mys Shmidta area

Indicates a species not recorded during surveys and added from the literature.

Status: R=Resident, P=Passage, S=Summer visitor, B=Breeder, B? = Possible breeder, V=Vagrant, U=Unclear, (Lit) indicates status according to

30

VLADIMIR YU. ARKHIPOV et al.

Forktail 29(2013)

Previous records: 1 : Andreev & Kondratyev (1996), 2: Dorogoi (1996), 3: Dorogoi (1997), 4: Dorogoi (1998), 5: Dorogoi & Beaman (1998), 6: Menyushina (2000), 7: Portenko (1972), 8: Portenko (1973), 9: Stishov (1991),

10: Stishov (1992), 1 1 : Stishov (1997), 12: Stishov (2004), 13: Stishov & Maryuhnich (1991a), 14: Stishov & Maryuhnich (1991b), 15:Tomkovich (1998), 16: Tomkovich etal. (1991)

FORKTAIL 29 (2013): 31-36

Species limits within Rhopophilus pekinensis

PAUL J. LEADER, GEOFF J. CAREY & PAUL I. HOLT

Rhopophilus pekinensis is a passerine endemic to north-east Asia occurring primarily in China; two or three subspecies are variously

recognised. A review of museum material and fieldwork on the breeding grounds indicates that only two taxa ( R . p. pekinensis and R. p.

albosuperciliaris ) are valid, and using criteria that grade morphological and vocal differences between allopatric taxa (Tobias etai. 2010),

both achieve the threshold for species status. The English names Beijing Babbler and Tarim Babbler are proposed reflecting both the type

location of each and the recently elucidated taxonomic affinities of Rhopophilus.

INTRODUCTION

The White -browed Chinese Warbler Rhopophilus pekinensis is a

passerine endemic to north-east Asia, occurring from north-west

China to north-east China and North (and previously also South)

Korea (Cheng 1987, Dickinson 2003, Duckworth & Moores 2008,

Brazil 2009, Moores et al. 2009, BirdLife International 2013a).

Whilst placed in the family Cisticolidae (Dickinson 2003), it was

included in the Timaliidae, in a clade with Sylvia and Paradoxornis,

by Alstrom et al. (2006), based on myoglobin and cytochrome b

sequence data. In light of this, it was placed within the Timaliidae

by Collar & Robson (2007) and Gill & Donsker (2012), using the

English name Chinese Bush-dweller to reflect the fact that it was

no longer considered a warbler. Subsequently Gelang et al. (2009)

and Moyle et al. (2012) proposed treatment of the Sylviidae (which

includes Rhopophilus) as a family rather than a subfamily within

the Timaliidae.

Most authorities (Cheng 1987, Dickinson 2003, Zheng 2011,

Gill & Donsker 2012) recognise three subspecies: pekinensis

(eastern part of the range, type locality Beijing, China),

leptorhynchus (central part of the range, type locality Gansu, China)

and albosuperciliaris (western part of the range, type locality

Xinjiang, China). The validity of leptorhynchus was questioned by

Vaurie (1955), who suggested treatment as a synonym of pekinensis,

and this is followed, albeit tentatively, by Collar &c Robson (2007).

Two further taxa, ‘ beicki ’ (type locality north-west Nei Mongol,

China) and ‘ major (type locality Qinghai, China) are not currently

recognised and both have long been treated as synonyms of

albosuperciliaris (Vaurie 1955, 1959).

In this paper the relationship between pekinensis and

albosuperciliaris and the validity of leptorhynchus are reviewed

based upon an examination of museum material and fieldwork

conducted in China; the taxa ‘ beicki ’ and ‘ major are also

discussed.

METHODS

Museum specimens were examined at the Natural History Museum,

Tring, UK (NHMUK) and the Museum fur Naturkunde, Berlin,

Germany (ZMB). The type specimens of pekinensis,

albosuperciliaris (NHMUK), leptorhynchus and ‘ beicki ’ (ZMB)

were examined, as was material from the type locality of ‘ major

(NHMUK). In total 55 specimens were examined comprising 29

albosuperciliaris (including one ‘ beicki ’ and three ‘ major ), 15

pekinensis and 1 1 leptorhynchus. The following biometrics were

taken: wing (maximum chord), tail length (to base of tail measured

under the undertail-coverts) and bill length (to skull);

measurements taken accord with standard procedures (Redfern &

Clark 2001). All measurements were taken by PJL. No plumage

differences between males and females exist, but plumage

differences attributable to age and especially feather wear were

noted (juvenile birds were characterised by very fresh plumage and

loose contour feathering).

During fieldwork on the breeding grounds, sound recordings

were obtained from Beijing, Hebei, Qinghai and Xinjiang.

Recordings were made using Telinga Pro 5 or Pro 7 parabolic

microphones with either a Sound Devices 722 or an HHB

Portadisc MDP 500, and a Sony PCM-M10 with a Sennheiser

ME66. Spectrograms were produced and analysis of various

parameters carried out using Raven Pro 1.4 (Cornell Laboratory

of Ornithology 2003-11). Contrast was adjusted for each

recording to ensure all elements (defined as any continuous

line on a sonogram) were retained, while minimising reverberation.

Measurements were made using a spectrogram window size

of 512.

In all 122 strophes were analysed, comprising 67 from nine

pekinensis and 55 from eight albosuperciliaris. Analysis of parameters

of each strophe was based on those proposed by Tobias et al. (2010),

and comprised:

• start and finish times (from which duration was calculated);

• lowest and highest frequency (from which frequency range was

calculated);

• peak frequency (the frequency at which peak power occurs);

• pace (calculated by dividing strophe length by number of

elements).

For each individual, we calculated the mean of each

parameter; we then used the mean and standard deviation of

all individuals of each taxon to calculate Cohen’s d values (see

below). Due to their regular occurrence in flocks, the exact

number of different individuals recorded was not always certain,

although the figures provided are considered conservative

estimates.

In order to review species limits between taxa we applied the

quantitative scoring system proposed by Tobias et al. (2010) to

assess the degree of phenotypic difference between allopatric taxa.

These criteria were summarised by Collar (2011a, b) thus: an

exceptional difference (a radically different colouration or pattern)

scores 4; a major character (a pronounced and striking difference

in the colour or pattern of a body part, or in measurement or

vocalisation) 3; a medium character (clear difference reflected, e.g.

by a distinct hue rather than a different colour) 2; and a minor

character (a weak difference, e.g. a change in shade) 1. Tobias et al.

(2010) set a threshold score of 7 to allow for species status; species

status cannot be triggered by minor characters alone, and only three

plumage characters, two vocal characters (one spectral and one

temporal), two independent biometric characters and one

behavioural or ecological character may be counted. Vocal and

biometric characters were assessed for effect size using Cohen’s d

computed via the online calculator at http://www.uccs.edu/

~faculty/lbecker/, where 0.2-2 is minor, 2-5 medium, 5- 10 major

and >10 exceptional.

PAUL J. LEADER PAUL J. LEADER

32

PAUL J. LEADER, GEOFF J. CAREY & PAUL I. HOLT

Forktail 29 (201 3)

RESULTS

Morphological differences between taxa

As noted elsewhere ( Vaurie 1959, Collar & Robson 2007), there

are pronounced plumage differences between pekinensis and

albosuperciliaris. In general, albosuperciliaris is much paler and

more uniform than pekinensis ; the key differences between

the two are detailed in Table 1 and illustrated in Plates 1-4.

During fieldwork it became apparent that there is a

highly distinct difference in iris colour, with albosuperciliaris

having a dark brown iris and pekinensis a glaring pale yellow

iris.

Table 1. Plumage and bare part differences between adult Rhopophilus pekinensis pekinensis and R. p. albosuperciliaris.

Plate 1 . Adult male R. p. pekinensis, Shanxi, China, April 2012.

Plate 2. Adult male R. p. albosuperciliaris, Xinjiang, China, June 2012.

Plate 3. Adult male R. p. pekinensis, Shanxi, China, April 2012

Plate 4. Adult male R. p. albosuperciliaris, Xinjiang, China, June 201 2

PAUL J. LEADER PAUL J. LEADER

Forktail 29 (2013)

Species limits within Rhopophilus pekinensis

33

Consistent structural differences also exist with albosuperciliaris

being larger than pekinensis in terms of wing, tail and bill length

(Table 2), such that when wing and bill lengths are plotted there is

no overlap between the two (Figure 1).

Table 2. Average wing length, tail length and bill to skull (all

measurements in mm) and standard deviation (SD) of pekinensis (n =

26) and albosuperciliaris (n = 29).

Figure 1. Bill to skull (mm) and wing length (mm) of pekinensis and

albosuperciliaris.

Wing length (mm)

The validity of leptorhynchus and comments on 'major'

and 'beicki'

As noted above the treatment of leptorhynchus is inconsistent.

Vaurie (1955) recognised leptorhynchus but noted that it was poorly

differentiated from pekinensis and concluded that 'it is a matter of

opinion whether or not it should be recognised in the

nomenclature’. Specimens of leptorhynchus examined as part of this

study were on average slightly smaller than pekinensis (0.8 mm

shorter-winged, 0.9 mm shorter-tailed and 1.0 mm shorter-billed).

There was, however, extensive overlap in biometrics (Figure 2). In

addition there were no consistent plumage differences between the

two, and plumage of the type specimen fell within the range of

pekinensis sensu stricto. As such, we concur with Collar & Robson

(2007) and consider leptorhynchus a synonym of pekinensis.

Vaurie (1955) concluded that ‘ major was comparable to

albosuperciliaris and not larger and more densely streaked as noted

by Meise (1937) and that birds from the type locality of 'major

fell within the range of plumage variation and size of

albosuperciliaris from Xinjiang. An examination of specimens from

the Qaidam Basin, Qinghai (the type locality of ‘major), and of

birds in the field there provides nothing with which to contradict

Vaurie’s conclusion.

Meise (1937) described ‘beicki' from a single specimen collected

in north-west Nei Mongol, China (note: Vaurie [1995] correctly

mapped the type locality of ‘ beicki ’, but incorrectly labelled the

province as Ningxia), and considered it similar in colouration to

‘major but smaller in size. Vaurie (1955) regarded any differences

insufficient to establish the validity of' beicki and questioned the

wisdom of recognising it based on just a single specimen.

Examination of the type specimen as part of this study established

that in terms of plumage it falls within the range of variation of

albosuperciliaris. Differences in biometrics are limited to wing

length (62.0 mm), with values for tail (90.5 mm) and bill to

skull (16.2 mm) falling within the range of albosuperciliaris.

Examination of the type also suggested nothing unusual regarding

the condition or preparation of the specimen which may have

resulted in the smaller wing measurement and, whilst further

material may prove otherwise, there appears no reason at this stage

to recognise ‘beicki'.

Vocalisations

Both pekinensis and albosuperciliaris are garrulous and gregarious,

and are most often found in small foraging flocks, the members of

which frequently utter contact and other vocalisations. Both

taxa have a wide repertoire of vocalisations, comprehensive

comparative analysis of which would require a very large dataset of

recordings.

Both taxa appear to have more than one territorial song,

although we collected insufficient samples to clarify the situation.

However, in the samples taken for this study, a single common

vocalisation that appears to have the same territorial and/or

advertising function was identified, and both taxa were seen perched

prominently uttering it; based on this, we regard it as song. As a

result, it has been possible to carry out the analysis described above.

The relevant vocalisation is a short series of 2-5 very similar notes

transcribed as pyoo, each descending in pitch; typical examples for

each of the taxa are illustrated in Figures 3 and 4. The mean,

standard deviation and Cohen’s d. values of the various

Figure 2. Tail length (mm) and wing length (mm) of pekinensis and leptorhynchus.

Wing length (mm)

34

PAUL J. LEADER, GEOFF J. CAREY & PAUL I. HOLT

Forktail 29(2013)

Figure 3. Typical pyoo vocalisation of pekinensis, Miyun Reservoir,

Beijing, 4 November 2009. (Paul I. Holt)

4-

Figure 4. Typical pyoo vocalisation of albosuperciliaris, Aksu, Xinjiang,

10 August 2005. (Paul I. Holt)

measurements are presented in Table 3. Sample sizes of other

vocalisations were insufficient to allow comparison in the absence

of a thorough understanding of their function.

Table 3. Mean, standard deviation (SD) and Cohen's d values of

parameters (see text) selected for analysis of pekinensis and

albosuperciliaris.

low frequency, high frequency and peak frequency the mean values

are higher in albosuperciliaris than pekinensis, with little overlap

between the two taxa; these differences are clearly audible in

recordings.

Habitat differences

A bird of dense secondary shrubland, pekinensis ranges from sea

level (where generally rare) to at least 1,200 m and is found in

degraded hill slopes, forest edge and forest clearings (Plate 5).

Species regularly recorded in the same habitat include Vinous-

throated Parrotbill Paradoxornis webbianus , Godlewski’s Bunting

Ember iza godlewskii and Meadow Bunting A. cioides. On the other

Plate 5. Typical habitat of R. p. pekinensis, Shanxi, China, April 201 2.

Plate 6. Typical habitat of R. p. albosuperciliaris, Xinjiang, China, June

2012.

hand, albosuperciliaris is a desert species occurring in areas of mature

tamarisk and dense desert shrubland (Plate 6) particularly in areas

where Phragmites are mixed with Chinese Date Ziziphus jujuba or

‘Shazhao’ — a central Asian xerophyte. It prefers low-lying, arid,

sandy and often, but not always, well-drained areas and occurs from

780 to about 1,500 m in Xinjiang but up to 2,800 m in the Qaidam

Basin, Qinghai. Lop Nur, Bayingol, is the lowest known site for

this taxon but with the drying up of the lake and associated habitat

changes there in recent years, it is quite possible that it is no longer

present. It occurs alongside Eurasian Tree Sparrow Passer montanus.

Saxaul Sparrow P ammodendri , Desert Whitethroat Sylvia minula,

with which it shares a very similar breeding distribution (Olsson et

al. 2013), Isabelline Shrike Lanius isabellinus and even Biddulph’s

Ground Jay Podoces biddidphi.

DISCUSSION

Characters selected for comparison based on Tobias et al. (2010)

were assessed (Table 4). Among biometric characters, only wing

length was assessed because of the lack of clearly independent such

characters (see Tobias et al. 2010). In terms of vocalisations, peak

frequency and pace were selected; behavioural or ecological

differences were represented by innate habitat. Geographical

relationship (Tobias et al. 2010) is not applicable as the two taxa

are allopatric, although Vaurie (1955) maps locations of both

indicating that the two occur within approximately 300 km of each

other. Overall, a score of 13 easily surpasses the threshold score of

7 for species status set by Tobias et al. (2010).

Of the features listed above, the differences in iris colour is

considered major and therefore ranks highly. Iris colour varies with

PAULJ. LEADER PAUL J. LEADER

Forktail 29(2013)

Species limits within Rhopophilus pekinensis

35

Table 4. Characters selected for comparison of pekinensis and

albosuperciliaris based on Tobias et al. (2010), with score (see text) in

brackets.

age in many passerine species, typically being duller in juveniles, so

whilst it is possible that young pekinensis may show dull irides similar

in colour to those of albosuperciliaris , the difference between

pekinensis and albosuperciliaris appears to be consistent when

breeding season adults are compared. The dark iris of albosuperciliaris

was noted in the historical literature (Richmond 1896) but has been

overlooked in recent times (Collar & Robson 2007).

A comparable situation in two taxa closely related to

Rhopophilus exists in V inous-throated Parrotbill and Ashy-throated

Parrotbill P. alphonsianus , which have a dark brown and whitish

iris respectively (Robson 2007). Whilst usually treated as separate

species (Penhallurick & Robson 2009, Gill & Donsker 2012),

recent genetic studies (e.g. Crottini et al. 2010) found these two

taxa to be very closely related and suggested that alphonsianus may

be a clinal morph of P webbianus. In addition, in Silver-eared Mesia

Leiothrix argentauris, the subspecies laurinae from Sumatra is

unlike other subspecies in that it has pale irides, and the subspecies

orientalis (from south V ietnam and east Cambodia) of Blue-winged

Minla Minla cyanouroptera can also be distinguished from other

subspecies by its pale irides. However, species limits within

Silver-eared Mesia and the taxonomic status of orientalis require

further evaluation (Collar & Robson 2007). Other examples in

which iris colour varies between subspecies include Masked

Booby Sula dactylatra (O’Brien & Davies 1990), whilst Kemp &

Delport (2002) described a new subspecies of Red-billed Hornbill

Tockus erythrorhynchus largely on the basis of iris colour and

their proposal that the Red-billed Hornbill complex is better

treated as five separate species (based on consistent differences in

the colour of signal areas between geographically discrete

populations) has been adopted elsewhere (Gill & Donsker 2012).

In this study, it is noteworthy that even without the score for iris

colour a score of 10 would still readily achieve the threshold for

species status.

Based upon these results the following taxonomic treatment of

two monotypic species is proposed:

Beijing Babbler Rhopophilus pekinensis (Swinhoe, 1868)

Tarim Babbler Rhopophilus albosuperciliaris (Hume, 1873)

The English names reflect the geographical origin of the type

specimens and the use of ‘Babbler’ reflects recent taxonomic studies

which place Rhopophilus within the Timaliidae. ‘Bush-dweller’

(Collar & Robson 2007) is not adopted as we feel that ‘Babbler’ is

more accurate and that ‘Bush-dweller’ gives little or no insight into

the taxonomic relationships of the two species. We acknowledge

that some authorities treat the Sylviidae as a separate family rather

Figure 5. Map showing the approximate ranges of the two species Rhopophilus albosuperciliaris and R. pekinensis including the type localities.

Irkutsk

JTANA

ULAAN BAATAR

Harbin

Changchun

Urumqi

BISHKEK

KYRGYZSTAN

,Baotou

Hamhung

'ONGYANG

Dalian

SEOUL

Taejon

.Qingdao

Chonju

Gwangju

.Xuzhou

ISLAMABAD

Legend

Huainan.

Hefei

nwala

Xinyang

# Nanjing

Wuxi* .Sh

Lahore

Type localities

( T) pekinensis

@ albosuperciliaris

@ leptorhyrchus

(4) "bieki"

(5) "major"

.Ludhiana

Chengdu

Wuhan

Hangzf

.Chongqin;

Nanchang

.Lichuan

NEW DELHI

KATHMANDU

Jaipur

Lucknow

Guiyang

Fuzhou

’Kanpur

Allahabad •

Patna

Kunming

Varanasi

Rajshahi

.Liuzhou

- J

DHAKA

Geographic Range

V//A albosuperciliaris

V77A pekinensis

Bhopal Jabalpur

Guangzhou

Hong Kong

.Ahmadabad

Khulna

Kolkata • i

Nanning

Kaohsi

Chittagong

Indore

Mandalay

adodara

•Surat

.Nagpur

HANOI-

HainhnnP

36

PAUL J. LEADER, GEOFF J. CAREY & PAUL I. HOLT

Forktail 29 (2013)

than a subfamily within the Timaliidae (Gelang et al. 2009, Moyle

etal. 2012), but refer to the use of the English name Sylviid Babblers

for the Sylviidae (Gill & Donsker 2012) and note that the family

includes a number of species which have ‘Babbler’ in their English

name (e.g. African Hill Babbler Pseudoalcippe abyssinica).

The BeijingBabbler occurs from North Korea, north to southern

Jilin and then west across north China to Gansu and eastern Qinghai.

According to BirdLife International (2013a), the range continues

south through northern Sichuan, western Henan and north-eastern

Hubei as far as south-western Anhui; however, we are unaware of

any records from Sichuan, Hubei or Anhui and these provinces are

omitted by Zheng (201 1 ), although it has been recorded from Henan

since the 1930s (Fu 1937). The Tarim Babbler occurs in southern

Xinjiang from the western part of the Tarim Basin (restricted to the

rivers and oases around the margins of the Tarim Basin and avoiding

the Taklamakan Desert proper) east to the Qaidam Basin, Qinghai.

The ranges of the two species are shown in Figure 5.

Beijing Babbler is a fairly common and widespread species found

in shrubland and although its range has contracted and it is no longer

recorded in South Korea (Moores et al. 2009) and has declined in

North Korea (Duckworth 2006), it is probably not globally

threatened. Tarim Babbler, whilst sometimes locally common, is

probably facing similar threats to Biddulph’s Ground Jay and may

be declining due to fragmentation and degradation of desert habitats

caused by intensive grazing of livestock, extraction of fuelwood and

conversion of suitable habitat to irrigated land (BirdLife

International 2013b) and may qualify as Near Threatened.

ACKNOWLEDGEMENTS

We thank Lei Wei Dong, Gou Jun, Ma Ming and David Stanton for their help

in the field in Xinjiang and Wang Qingyu for her help in Xinjiang and Beijing.

Kadoorie Farm and Botanic Garden kindly funded the visit by PJL to the ZMB.

Richard Lewthwaite assisted with distributional information and both Michael

Leven and Per Alstrom provided useful comment on the English names of the

two species. Hincent Ng prepared Figure 5 and Urban Olsson assisted with

references. Mark Adams (NHMUK) and Sylke Frahnert (ZMB) kindly

permitted access to specimens in their collections. Will Duckworth and Nial

Moores commented on the status of pekinensis in Korea. Finally, Michael Leven

and Nigel Collar provided very helpful comments on a draft of this paper.

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classification of the avian superfamily Sylvioidea. Mol. Phylogenet. Evol.

38: 381-397.

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Collar, N. J. & Robson, C. (2007) Family Timaliidae (babblers). Pp.70-291 in

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world, 12. Barcelona: Lynx Edicions.

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Barbuto, M. & Casiraghi, M. (2010) Towards a resolution of a taxonomic

enigma: first genetic analyses of Paradoxornis webbianus and

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in DPR Korea. Bull. Brit. Orn. Club 1 26: 252-290.

Duckworth, J. W. & Moores, N. (2008). A re-evaluation of the pre-1 948 Korean

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24: 25-47

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Phylogeny of babblers (Aves, Passeriformes): major lineages, family limits

and classification. Zool. Scr. 35: 225-236.

Gill, F. & Donsker, D„ eds. (201 2) IOC World Bird Names (v 3.1 ). Available at

http://www.worldbirdnames.org [Accessed on 29/01/2013].

Kemp, A. C. & Delport, W. (2002) Comments on the status of subspecies in

the red-billed hornbill ( Tockus erythrorhynchus ) complex (Aves:

Bucerotidae), with the description of a new taxon endemic to Tanzania.

Ann. Transvaal Museum 39: 2-8.

Meise, W. (1937) in Stresemann, E., Meise, W. & Schonwetter, M. (1937) Aves

Beickianae.L Ornithol. 85: 375-576.

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2009.shtml [Accessed 28/04/2013],

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dactylatra from Lord Howe, Norfolk and Kermadec Islands. Mar. Ornithol.

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New insights into the intricate taxonomy and phylogeny of the Sylvia

curruca complex. Mol. Phylogenet. Evol. 67: 72-85.

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Ornithology.

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the world, 12. Barcelona: Lynx Edicions.

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Collar, N. J. (2010) Criteria for species delimitation based on phenotype.

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PaulJ. LEADER, AEC Ltd., 127 Commercial Centre, Palm Springs,

New Territories, Hong Kong. Email: pjl@aechk.hk

Geoff J. CAREY, AEC Ltd., 127 Commercial Centre, Palm Springs,

New Territories, Hong Kong. Email: gjc@aechk.hk

Paul I. HOLT, Bracken Dean, Pendleton, Clitheroe, Lancashire,

UK. Email: piholt@hotmail.com

FORKTAIL 29 (201 3): 37-42

Nesting of the Large-billed Reed Warbler

Acrocephalus orinus: a preliminary report

PAVEL KVARTALNOV, ABDULNAZAR ABDULNAZAROV, VERONIKA SAMOTSKAYA, JULIA POZNYAKOVA, IRINA ILYINA,

ANNA BANNIKOVA & EUGENIA SOLOVYEVA

Large-billed Reed Warbler Acrocephalus orinus has a limited breeding distribution. It is known to inhabit valleys of the Panj river and its

tributaries in Gorny Barakhshan Autonomous Republic (Tajikistan) and Badakhshan province (Afghanistan). Here we give descriptions of

nests and eggs of this species based on 18 fresh nests found in Panj and Ghund valleys (Tajikistan) in 201 1. Unlike the closely related species

A. dumetorum and A. scirpaceus, Large-billed Reed Warbler has nests built with a layer of wool and seed tufts. Nests are placed on twigs of

sea-buckthorn, willow and other bushes, herbs and reed stems over dry soil. Large-billed Reed Warbler clutch size is relatively small (on

average, 3.77 ± 0.83 eggs (n = 1 3)). The ground colour of eggs is usually white, not bluish, greenish or rosy as in the related species.

INTRODUCTION

Until recently Large-billed Reed Warbler Acrocephalus orinus

remained one of the least studied bird species of the Palaearctic

fauna. A. O. Hume discovered the first specimen on 1 1 November

1867 in the Sutlej valley, Himachal Pradesh, India (Hume 1869,

1870). He described the bird as Phyllopneuste macrorhynchus, and

later referred to it as A. macrorynchus. Oberholser (1905) changed

the name to A. orinus.

The taxonomic status of this form remained uncertain until

the beginning of the twenty-first century when Bensch & Pearson

(2002) studied the type specimen in detail, including sequencing

of mitochondrial and nuclear DNA. This study confirmed the

specific status of Large-billed Reed Warbler, although some doubts

remained (McCarthy 2006) until the moment when P. D. Round

caught a live bird near Bangkok, Thailand (Round et al. 2007).

Further studies of museum collections and searches for living

individuals helped to elucidate possible breeding, moulting and

wintering areas (Svensson et al. 2008, 2010, Timmins et al. 2009,

Koblik et al. 2010, 2011).

In 2009, a bird feeding fledglings was caught in south-east

Tajikistan, not far from the border with Afghanistan, in the

Shakhdara river valley (Aye et al. 2010). Museum specimens in the

Zoological Museum of Moscow University (Moscow, Russia) and

the Institute of Zoology and Parasitology (Dushanbe, Tajikistan)

reveal that previous records of Blyth’s Reed Warbler Acrocephalus

dumetorum breeding in the Vanj and Ghund river valleys in fact

refer to Large-billed Reed Warbler (Kvartalnov et al. 201 la, b,

Kvartalnov & Garibmamadov 2012). Although fledglings were

recorded in 1961 by A. V. Popov in the Vanj valley, near the village

ofGhijovast (Abdusalyamov 1973, Kvartalnov et al. 2011b) and in

2009 by R. Aye and colleagues (Aye etal. 2010), and birds collecting

nest material were observed in 1976 by V. V. Kashinin in the lower

Ghund valley, near the village of Barsem (unpublished manuscript

- see Kvartalnov et al. 2011a), no nests of this species have ever

been described.

The mystery of the Large-billed Reed Warbler could have been

solved in the mid-twentieth century. When in 1937 A. B.

Kistyakovsky took part in a Pamir expedition, he found that reed

warblers in the south-western Tajik Pamir mountains (in the

environs of the town of Khorog) were not typical Blyth’s Reed

Warblers. He therefore prepared a description of a new A.

dumetorum subspecies, but his manuscript and the intended type

series were destroyed in a fire together with all zoological collections

in Kiev University during the German occupation in the Second

World War (Nowak 200 1 ). Kistyakovsky ( 1 950) wrote his opinion

of the systematic position of this form, which he thought to be

endemic to Gorny Badakhshan, Tajikistan. Other naturalists who

had visited the Pamir mountains in the twentieth century did not

distinguish those birds from typical Blyth’s Reed Warblers.

In 2010-2011 we studied spring migration of Blyth’s Reed

Warbler and breeding biology of Large-billed Reed Warbler in

Tajikistan. Blyth’s Reed Warbler was found to be common during

spring passage in the vicinity of Dushanbe and in the south-west

part of the republic, but we failed to prove its breeding in Taj ikistan,

although this was suspected by Abdusalyamov (1973), Portenko

& Stubs (1976) and other ornithologists. There is no doubt that

Blyth’s Reed Warbler is a transient in all regions to the south of

Almaty in south-east Kazakhstan, and that all nests found there

that had been attributed to that species belong to others, including

Paddyfield Warbler Acrocephalus agricola and Sykes’s Warbler

Hippolais rama (Ivanitskii et al. 2012).

Data about phenology, breeding biology, social behaviour,

acoustics and morphometry of the Large-billed Reed Warbler

collected in 2011 are presented in this article with additional

data from 2012. This information is to help other

ornithologists to search for and distinguish nests of Large-billed

Reed Warbler.

MATERIALS AND METHODS

The main field observations were conducted in the Panj valley near

the village of Zumudg, Ishkashim region, Gorny Badakhshan

Autonomous Region, Tajikistan (36.9 17°N 72.183°E) between 10

June and 11 July 2011. Additional data were collected in the

Apharv forest area in the Panj valley (36. 800°N 71. 550°E) and near

the village of Langar in the lower Pamir river valley (37.033°N

72.667°E). AA inspected riverside forests in the Ghund valley near

the villages of Charthem (37.717°N 72.167°E), Vuzh (37.717°N

71.933°E) and Dehmiyona (37.700°N 71.917°E) (Figure 1). From

23 May to 23 July 2012 PK and colleagues studied breeding biology

and social behaviour of Large-billed Reed Warbler near the village

of Dehmiyona; the resulting data are not included here.

Nine adult Large-billed Reed Warblers were caught in mist-

nets and traps at nests near Zumudg (Plate 1). The birds were

identified by measurements of bills, wings, tails and legs according

to Svensson et al. (2010), Koblik et al. (2010) and from our

experience of working with series of Large-billed and Blyth’s Reed

Warblers in collections of the Zoological Museum of Moscow

University (Ivanitskii et al. 2012). Adults were marked with metal

and colour rings and by grease paint colouring on breast and head

for individual identification (a harmless method that we used

previously with other warbler species). Blood samples were taken

38

PAVEL KVARTALNOV etol.

Forktail 29 (2013)

Figure 1. A map of localities of

some historical and recent

observations of Large-billed Reed

Warbler A. orinus in Tajikistan.

A. orinus JQ651 381.1 Zumudg

A. orinus GU247955 Burma

A. orinus GU247957 India

A. orinus HM352785 Khorog. Tajikistan

A. orinus GU247958 India

A. orinus GU247952 Pakistan

A. orinus GU247953 Afghanistan

A. orinus GU247950 Afghanistan

A. orinus GU247954 Kazakhstan (?)

43

71

70

90

84

99

98

98

97

98

A.

A. orinus HM352786 Khorog, Tajikistan

~~ A. orinus GU247949 Afghanistan

A. orinus DQ681065 Thailand

A. orinus GU247956 Burma

A. orinus JQ651 384.1 Zumudg

A. orinus GU247951 Afghanistan

A. orinus JQ651380.1 Zumudg

A. orinus JQ651 382.1 Zumudg

A. orinus JQ651 383.1 Zumudg

“A. orinus HM352789 Shakhdara r.,

T ajik istan

scirpaceus

97

97

97

99

98

I A. concinens

^ A. dumetorum

\A palustris

97

97

j

^ A. agricola

' S. borin AY329474

0.01

Figure 2. The relationships

between Large-billed Reed

Warbler and closely related

species of the genus Acrocephalus

based on NJ and MP analyses.

Genbank accession numbers

for Eurasian Reed Warbler

A. scirpaceus, Marsh Warbler

A. palustris etc. are given in the

text.

from all caught adult birds and eight nestlings (also marked with

metal rings). Specific identification was supported by analysis of

mt DNA (Figure 2).

Total DNA was extracted from dried blood samples using the

standard protocol of proteinase K digestion, phenol-chloroform

deproteinisation and isopropanol precipitation (Sambrook et al.

1989). Mitochondrial DNA sequences were obtained from five

Large-billed Reed Warblers caught in 201 1 near Zumudg. The

partial cytochrome b gene (207 bp) was amplified in one polymerase

chain reaction (PCR) with the forward/reverse primer

combination L14841/H 1 5 149 (Kocher et al. 1989). Typical

conditions for cyt b amplification included initial denaturation at

PAVEL KVARTALNOV

Forktail 29 (2013)

Nesting of the Large-billed Reed Warbler Acrocephalus orinus: a preliminary report

39

Plate 1. Panj river valley near

Zumudg village, 25 June 201 1 .

94°C for 3 min, 35 cycles of 94°C for 30 s, annealing at 51°C for

1 min, and extension at 72°C for 1 min, followed by a final extension

at 72°C for 10 min and an indefinite hold at 4°C. PCR products

were visualised on 1 % agarose gel and then purified using DEAE

(Whatman) or NHJitOH. Approximately 10-50 ng of the

purified PCR product were used for sequencing with each primer

by the autosequencing system ABI 3 1 00-Avant in conjunction with

ABI PRISM’BigDyeTM Terminator, version 3.1.

Cytb sequences were aligned by eye using BioEdit 7.0. The final

alignment of the mitochondrial region included 207 bp, of which

54 sites were variable and 40 sites were parsimony-informative. For

the analysis we also used GenBank data (A. dumetorum,A. orinus ,

A. agricola , Eurasian Reed Warbler U. scirpaceus. Marsh Warbler

A. palustris and Blunt-winged Warbler H. concinens). Phylogenetic

neighbour-joining (NJ) and maximum parsimony (MP) analyses

were performed using MEGA 4.0.0.4083. The NJ tree was

reconstructed using the uncorrected p-distance. Unweighted MP

analysis was performed using heuristic search starting with stepwise

addition trees (random addition sequence, 10,000 replicates). To

assess clade stability in the MP and NJ trees, 1,000 bootstrap

pseudoreplicates were analysed.

GenBank accession numbers of obtained sequences are

JQ65 1 380-JQ65 1384. Genbank accession numbers of other

sequences used in this work comprise: A. agricola AJ004245-

AJ004248, AJ0043.30, AJ004331, AJ004775, AJ004776, FJ883021,

Y15694; A. concinens AJ004260-AJ004262, FJ883027; A.

dumetorum : AJ004263, AJ004264, AJ004336-AJ004340,

AJ004773, FJ883028; A. orinus -. DQ681065, GU247949-

GU247958, HM352785, HM352786, HM352789; A. palustris-

AJ004293, AJ004294, AJ004344, AJ004345, AJ004774, EU86 1 03 1 ,

FJ883036; A. scirpaceus- AJ004301-AJ004304, AJ004771,

AJ004772, AM889139, FJ883039, NC 010227, Z73483.

The identification of uncaught birds was based on characteristic

songs (Timmins et al. 2010, Ivanitskii et al. 2012). Recordings of

songs of five marked males proved that the song described by

Timmins et al. (2010) belongs to the Large-billed Reed Warbler

(Ivanitskii et al. 2012). Nests that AA found in the Ghund valley

were identified by comparison with known Large-billed Reed

Warbler nests from the Panj valley. For comparison we also used

unpublished data from 51 Blyth’s Reed Warbler nests found and

described by PK in 2007-2009 in the Kostroma region, Russia.

RESULTS AND DISCUSSION

The Large-billed Reed Warbler is a common species in suitable

habitat in the Panj, Ghund and lower Pamir valleys. We found nests

near the villages of Zumudg, Charthem, Vuzh and Dehmiyona,

observed actively singing males in the Apharv forest area, and saw

singing males and territorial pairs at the village of Langar (Figure

1). The birds inhabited thickets of sea-buckthorn Hippopbae

ramnoides and willow Salix turanica, S. shugnanica and S.

wilhemsiana intertwined with clematis Clematis hilariae , with

sparse ground cover of liquorice Glycyrrhiza uralensis , reed

Pbragmites australis and other species. Other bird species observed

in the same habitat of Panj valley include Hume’s Lesser

Whitethroat Sylvia althaea. Mountain Chiffchaff Pbylloscopus

sindianus. Common Rosefinch Carpodacus erythrinus, Cetti’s Bush

Warbler Cettia cetti , Common Nightingale Luscinia megarhyncbos ,

Bluethroat Luscinia svecica. Black-billed Magpie Pica pica, Isabelline

Shrike Lanius isabellinus phoenicuroides and Citrine Wagtail

Motacilla citreola calcar ata. The only other Acrocephalus warbler

recorded around Zumudg during our observations was Clamorous

Reed Warbler A. stentoreus (a single transient bird). According to

observations of PK in 20 1 2, in the Ghund valley Large-billed Reed

Warblers also breed in S. turanica, wild rose Rosa beggerana and

honeysuckle Lonicera stenantha thickets with Astragalus

longistipitatus, Potamogeton cariatum and other herbs along canals

among crop fields on alluvial fans.

Most Large-billed Reed Warblers were found near river banks,

canals or other wet localities. The birds breed in monogamous pairs,

although attempted extra-pair copulations by at least three paired

and two unmated territorial males were observed. We described 1 5

nests and one abandoned construction built in June andjuly 2011

in the Panj valley (Plates 2, 3), plus seven remains of nests built in

2009-2010 in the same area, and three recent nests in the Ghund

valley in 20 1 1 . Thirteen nests had complete clutches. Nests are built

by females (based on observations of building of nine nests in 2011,

including two nests observed from the first day of construction).

Most were in sea-buckthorn thickets, but one was in a willow bush.

Nearly all nests were placed over dry soil, except one that was built

on a branch over a canal temporarily filled by water.

Large-billed Reed Warblers attached nests to sea-buckthorn

twigs at forks (six nests), stems of Artemisia (three nests), liquorice

PAVEL KVARTALNOV PAVEL KVARTALNOV

40

PAVEL KVARTALNOV etal.

Forktail 29 (2013)

stems (three nests), thin willow stems (two nests), willow twigs at

forks (two nests), twigs of clematis at forks (two nests), reed stems

(one nest), reed stems and willow twigs (one nest), reed and

liquorice stems (one nest), liquorice stems and willow twigs (one

nest), sea-buckthorn and willow twigs (one nest), reed, liquorice

stems and a sea-buckthorn twig (one nest).

Females began nest construction with a platform of dry plant

debris, but from the first day they braided vertical supports (stems,

etc.) with plant fibres. Nests were fastened to stems more firmly

than the nests of Blyth’s Reed Warbler. The principal material

consists of bast and bark fibres of clematis, willow, liquorice and

Artemisia , fibres of reed sheath, dry leaves, stems and ears of grasses,

goat wool and clematis seed tufts. Bast strips also form an outer

covering that disguises the nest in thickets. An inner part of the

structural layer is made with wool and seed tufts, and rarely includes

bird feathers. Nests of Blyth’s Reed Warbler usually lack such soft

materials (Plate 5). The upper edges of Large-billed Reed Warbler

nest cups are made usually with ears of grasses. Nests are lined with

clematis bast fibres or (rarely) with thin dry grass stems, with the

addition of mammal hairs.

Nests found near Zumudg (n = 15) had the following

measurements (average and standard deviation): outer diameter

81.7 ± 8.0 mm; height 68.6 ± 9.4 mm; inner diameter 54.8 ±

2.0 mm; depth 45.1 ± 3.0 mm. The height of nests above ground or

water was 30-168 cm (average 82.1 cm). Nests inspected in the

Ghund valley in 201 1 were placed at 110-210 cm above ground.

Most full clutches in the Panj valley were of four eggs (n = 7),

rarely two (n = 1), three (n = 3) or five eggs (n = 2); on average,

3.77 ± 0.83 (n = 13), including a replacement clutch that

contained three eggs (the first clutch consisted of four eggs). We

also found a nest with three nestlings. Nests found in Ghund

valley had three (two nests) and four (one nest) eggs. The usual

clutch size of the Blyth’s Reed Warbler in Kostroma region was

5-6 eggs, rarely four eggs, on average (n = 37) 5.53 ± 0.56 eggs.

Plate 2. Nest 5-1 1 of the Large-billed Reed Warbler A. orinus with full

clutch (two unhatched eggs from this nest are now held in the Natural

History Museum, Tring, UK), 21 June 201 1.

Plate 3. Nest 11-11 of the Large-billed Reed Warbler with full

clutch (now held in the Zoological Museum of Moscow University), 9

July 2011.

Plate 4. A clutch from nest 13-11 of the Large-billed Reed Warbler

showing markings on eggs, 1 1 July 201 1 .

Plate 5. A nest of Blyth's Reed Warbler A. dumetorum with full clutch,

Kostroma region, Russia, 14 June 2007.

PAVEL KVARTALNOV PAVEL KVARTALNOV

ABDULNAZAR ABDULNAZAROV PAVEL KVARTALNOV

Forktail 29 (2013)

Nesting of the Large-billed Reed Warbler Acrocephalus orinus: a preliminary report

41

Clutches of Blyth’s Reed Warbler from South Siberia had on

average 4. 8-5. 2 eggs (Totunov 1981) or 5.1-5.76 eggs (Kuranov

2008) in different years.

Eggs of Large-billed Reed Warbler have a dirty-white (rarely

creamy white or pure white) shell covered with small olive-brown

superficial spots that usually (but not always) form a sparse cap or

a ring at the larger end, and with more sparse dark-brown superficial

specks which rarely (as opposed to eggs of Blyth’s Reed Warbler)

lay over larger spots (Plates 4 & 5). Sparse and small deep bluish

spots also form a cap or a ring at the larger end. Some eggs are

covered not only with sparse spots but also with dense small olive

brown specks that can hide the basic shell colour. Fresh eggs are

dull or with a weak gloss. Eggs of Blyth’s Reed Warbler are greenish

or rosy, usually with no white background (Chernyshov 1998; our

data). Measurements of eggs (in mm; n = 58): 17.72 ± 0.55 (16.8-

19.1) x 13.14 ± 0.31 (12.3-13.8). Egg weight (n = 48): 1.51 ±

0.15 (1.13-1.76) g. One Large-billed Reed Warbler nest with a

Plate 6. A newly-hatched nestling of Large-billed Reed Warbler from

nest 5-1 1,3 July 2011.

I

Plate 7. Female Large-billed Reed Warbler feeding nestlings, nest 1-

11, 10 July 2011.

complete clutch (three eggs) is now deposited in the Zoological

Museum ofMoscow University (ZMMU Q-8036); two unhatched

eggs were sent to the Natural Elistory Museum, Tring, UK

(NHMUK E/2012.5.1).

Eurasian Reed Warbler has been found in Gorny Badakhshan

and nearby Afghanistan, although it is not known to breed there

(Abdusalyamov 1973, Timmins et al. 2010). It has nests without

the layer built with wool and seed tufts (Kvartalnov et al. 2006),

and eggs with bluish or olive-greenish basic shell colour, not white

(Nikiforov et al. 1989).

Only female Large-billed Reed Warblers have brood-patches,

but both partners incubate eggs and provide food to nestlings and

fledglings (Plates 6, 7, 8).

Large-billed Reed Warblers in the Panj valley have a relatively

wide range in dates of arrival on the breeding grounds (compared

with Mountain Chiffchaff and other passerines inhabiting the Pamir

Mountains: our observations). The nest found on 5 July with young

birds ready to fledge must have been built during the first days of

June. Most other nests near Zumudgwere built after 10 June. Several

new birds reached the breeding grounds at the end of June and the

beginning of July. The latest of the first clutches found near Zumudg

was finished on 7 July. On the last day of investigations (11 July)

we observed a female in the territory of a male which was singing

from 9 July. Of two nests built for replacement clutches, one was

still empty on 1 1 July. In the lower Pamir river valley on 2 July we

found several actively singing bachelor males, newly formed pairs

and apparently non-territorial birds. AA found nests with clutches

on 23 June, 16 July (nestlings hatched 17 July) and 17 July in the

Ghund valley. V. V. Kashinin in the lower Ghund valley (37.550°N

71.733°E) observed the peak of nest building to be in mid-June

(Kvartalnov et al. 201 lb). A. V. Popov saw fledglings on 22 June

1961 in the Vanj valley (38.550°N 71.733°E) (Abdusalyamov 1973,

Kvartalnov et al. 2011b); fledglings possibly of this species were

recorded by A. V. Popov in the Shakhdara valley on 26 July (birds

were not collected, and the year is unknown) (Abdusalyamov 1973).

Aye et al. (2010) saw fledglings near the Shakhdara River on 19

July 2009. According to museum collections (Koblik et al. 2010,

2011) and our observations in 2012, Large-billed Reed Warblers

reach breeding grounds in Gorny Badakhshan in the final third of

May. AA saw the last birds that he thought to be this species in the

first ten days of September in the Panj valley, but this needs to be

confirmed by mist-netted birds.

Plate 8. A young Large-billed Reed Warbler ready to fledge, nest 1 2-

11,5 July 2011.

PAVEL KVARTALNOV

42

PAVEL KVARTALNOV etal.

Forktail 29 (2013)

ACKNOWLEDGEMENTS

The authors are grateful to the Director of the Zoology and Parasitology

Institute (Dushanbe) Abdusattor Saidov for help in organising our

expeditions, to Shamsher Mirzobekov (Zumudg) and his family for their

hospitality, to Tamara Krutenko (Lomonosov MSU) for plant identification

and to Nigel Collar, Brian Sykes and two anonymous referees for their careful

work with the text of our paper. This work was supported by the Rufford

Small Grants Foundation and by Russian Fund for Basic Research (##10-04-

00483, 13-04-01771). The publication is dedicated to the memory of

Alexander Kistyakovsky (1904-1983).

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the enigmatic Large-billed Reed Warbler Acrocephalus orinus. J. Avian

Biol. 41: 452-459.

Bensch, S. & Pearson, D. (2002) The Large-billed Reed Warbler revisited. Ibis

144:259-267.

Chernyshov, V. M. (1998) [Character of inheritance of egg shell colour in

Blyth's Reed Warbler (Acrocephalus dumetorum)]. Pp. 18-20 in Actual' nye

problemyoologii. Lipetsk: Lipetsk State Pedagogical University Press. (In

Russian.)

Hume, A. O. (1869) To the Editor of The Ibis'. Ibis (2) 5: 355-357.

Hume, A. O. (1871) Stray notes on ornithology in India. No. VI. On certain

new or unrecorded birds. Ibis (3) 1 : 23-38.

Ivanitskii, V. V., Kvartalnov, P. V., Marova, I. M., Samotskaya, V. V. & Bannikova,

A. A. (2012) Modes of stay, vocalization patterns and morphological

variability of Blyth's Reed Warblers (Acrocephalus dumetorum) in Central

Asia. Zoologicheskii Zhurnal 91: 1459-1509. (In Russian with English

summary.)

Kistyakovsky, A. B. (1950) [Materials on the zoogeography of Pamir (based

on the distribution of land vertebrates)]. Proceedings of the Zoological

Museum of Shevchenko Kiev State University 2: 5-58. (In Russian.)

Koblik, E. A., Arkhipov, V. Yu. & Redkin, Ya. A. (2010) New data on distribution

of the Large-billed Reed Warbler Acrocephalus orinus Oberholser, 1905.

Russian Journal of Ornithology 19 (Express-issue 596): 1619-1633. (In

Russian.)

Koblik, E. A., Redkin, Ya. A., Meer, M. 5., Derelle, R., Golenkina, S. A.,

Kondrashov, F. A. & Arkhipov, V. Yu. (201 1 ). Acrocephalus orinus: A case

of mistaken identity. PLoS ONE 6: No. 4. - e17716. doi:10.1 371/

journal. pone.001 7716.

Kocher, T. D„ Thomas, W. K„ Meyer, A., Edwards, S. V., Paabo, S„ Villablanca,

F. X. & Wilson, A. C. (1 989) Dynamics of mitochondrial DNA evolution in

animals: amplification and sequencing with conserved primers. Proc.

Natn. Acad. Sci. USA 86: 6196-6200.

Kuranov, B. D. (2008) Peculiarities of nesting biology in the Blyth Reed Warbler

(. Acrocephalus dumetorum, Passeriformes, Sylviidae) in urban habitats.

Zoologicheskii Zhurnal 87: 466-475. (In Russian with English summary.)

Kvartalnov, P. V. & Garibmamadov, G. D. (201 2) Ornithological and oological

collections of E.N. Pavlovsky Zoology and Parasitology Institute,

Tajikistan: their history and present significance. Pp.1 62-1 65 in Nazemnye

Pozvonochnye Zhivotnye Aridnykh Ekosistem. Tashkent: Chinor Enk. (In

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behaviour of the Reed Warbler ( Acrocephalus scirpaceus) at the eastern

Azov sea coast. Ornitologia 33: 100-108. (In Russian with English

summary.)

Kvartalnov, P. V., Ivanitskii, V. V., Marova, I. M. & Samotskaya, V. V. (2011a) A

bewitched bird: history of Large-billed Reed Warbler. Priroda 6: 35-40.

(In Russian with English summary.)

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museum collections to live birds. Priroda 12: 54-56. (In Russian.)

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Nikiforov, M. E., Yaminskiy, B.V.& Shklyarov, L. P. (1 989) Ptitsy Belorussii [Birds

of Belorussia: a guide for nests and eggs], Minsk: Visheyshaya shkola. (In

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region, East Africa. Proc. US Natn. Mus. 28: 823-936.

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Verbreitung Palaearkischer Vogel 5: 1 -5.

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Lost and found: the enigmatic Large-billed Reed Warbler Acrocephalus

orinus rediscovered after 1 39 years. J. Avian Biol. 38: 1 33-1 38.

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manual. New York: Cold Spring Harbor Laboratory Press.

Svensson, L„ Prys-Jones, R., Rasmussen, P.C. & Olsson, U. (2008) Discovery of

ten new specimens of Large-billed Reed Warbler Acrocephalus orinus,

and new insights into its distributional range. J. Avian Biol. 39: 605-610.

Svensson, L„ Prys-Jones, R., Rasmussen, P. C. & Olsson, U. (2010) The

identification and distribution of the enigmatic Large-billed Reed Warbler

Acrocephalus orinus. Ibis 1 52: 323-334.

Timmins, R. J., Mostafawi, N., Rajabi, A. M., Noori, H., Ostrowski, S., Olsson, U.,

Svensson, L. & Poole, C. M. (2009) The discovery of Large-billed Reed

Warblers Acrocephalus orinus in north-eastern Afghanistan. BirdingASIA

12:42-45.

Timmins, R . J., Ostrowski, S., Mostafawi, N., Rajabi, A. M„ Svensson, L. & Olsson,

U. (201 0) New information on the Large-billed Reed Warbler Acrocephalus

orinus, including its song and breeding habitat in north-eastern

Afghanistan. Forktail 26: 9-23.

Totunov, V. M. (1981) [Breeding of the Blyth's Reed Warbler ( Acrocephalus

dumetorum Blyth.) in vicinities of Chany lake.] Pp. 160-165 in Ekologiyai

biotsenoticheskiye svyazi perelyotnykh ptits Zapadnoy Sibiri. Novosibirsk:

'Nauka' Press, Siberian Branch. (In Russian.)

Pavel KVARTALNOV, Vertebrate Zoology Department, Faculty

of Biology, Lomonosov Moscow State University, Leninskiye

Gory, Moscow 1 1 999 1, Russian Federation. Email : cettia@mail.ru

Abduinazar ABDULNAZAROV, Pamir Biological Institute,

Khorog 736002, Tajikistan. Email: abdu_70@mail.ru

Veronika SAMOTSKAYA, Vertebrate Zoology Department,

Faculty of Biology, Lomonosov Moscow State University,

Leninskiye Gory, Moscow 1 19991, Russian Federation

Julia POZNYAKOVA, Vertebrate Zoology Department, Faculty

of Biology, Lomonosov Moscow State University, Leninskiye

Gory, Moscow 1 19991, Russian Federation

Irina ILYINA, Vertebrate Zoology Department, Faculty of

Biology, Lomonosov Moscow State University, Leninskiye Gory,

Moscow 1 19991, Russian Federation

Anna BANNIKOVA, Vertebrate Zoology Department, Faculty

of Biology, Lomonosov Moscow State University, Leninskiye

Gory, Moscow 1 19991, Russian Federation

Eugenia SOLOVYEVA, Vertebrate Zoology Department, Faculty

of Biology, Lomonosov Moscow State University, Leninskiye

Gory, Moscow 1 1 999 1, Russian Federation

F. GERMI © NATURAL HISTORY MUSEUM, LONDON

FORKTAIL 29 (2013): 43-47

First records of Chinese Sparrowhawk Accipiter soloensis

wintering in Papua (Indonesian New Guinea)

FRANCESCO GERMI, AGUS SALIM & ANDREA MINGANTI

Chinese Sparrowhawks Accipiter soloensis were found for the first time wintering in mainland Papua (Indonesian New Guinea) during a field

survey carried out between December 2010 and March 201 1. A combination of 39 road, boat and foot transects were completed in the

provinces of Papua and West Papua, covering a total of 2,303 km, of which 1,948 km were on roads or footpaths and 355 km on rivers.

Transects were supplemented by frequent spot counts and stops to broadcast recordings of Chinese Sparrowhawk vocalisations. Routes

covered eight sample areas in the most representative habitats of the region. A total of 1 0 Chinese Sparrowhawks were recorded at four

locations, all close to the coast. The new records are up to 1,200 km east of the easternmost extent of the previously known wintering range,

thus proving that this species does winter in Indonesian New Guinea, although most likely at low density. Seventeen other raptor species

were recorded on the transects. In addition, 12 days were spent between 6 and 17 March 2011 at a suitable coastal watch site at the

westernmost point of West Papua, but no visible migration of Chinese Sparrowhawk was observed.

INTRODUCTION

The Chinese Sparrowhawk Accipiter soloensis is a small-sized

accipiter, whose breeding grounds, although imprecisely known,

are in China, Korea and Ussuriland, and whose wintering grounds

are thought to be mainly in the Philippines and eastern Indonesia.

Despite this being the most numerous migratory raptor in East

Asia, with an estimated global population running well into six

figures (Ferguson-Lees & Christie 2001), very few records exist

from the presumed wintering range, suggesting that the winter

distribution remains largely unknown (Wattel 1973, White 1976,

Mees 1982, White & Bruce 1986, Beehler et al. 1986).

Recent migration research (Germi 2005, Germi & Waluyo

200 6, Nijman et al. 2006, Germi et al. 2009) showed that at least

350,000 individuals of this species are streaming into eastern

Indonesia each autumn, through both the Sangihe-Talaud

Plate 1 . The five Chinese Sparrowhawks collected in New Guinea in the nineteenth century at unspecified localities, held at the Natural History

Museum, Tring, UK. From left to right: NHMUK 1955.6.N.20.2635; 1873.5.12.1638; 1955.6.N.20.2636; 1 955.6.N.20.2633 and 1955.6.N.20.2632.

44

FRANCESCO GERMI, AGUS SALIM & ANDREA MINGANTI

Forktail 29 (2013)

Archipelago in the north and Bali in the west. Nevertheless, other

than scattered records from Sulawesi’s northern peninsula, winter

observations of Chinese Sparrowhawk elsewhere on this island are

negligible, despite some areas having been relatively well surveyed

(Hartert 1896, Meyer & Wiglesworth 1898, Rozendaal & Dekker

1989). Likewise, there are very few winter records of this species

from the Moluccas and the Lesser Sundas. A handful of specimens

and records provide solid, albeit scarce, evidence that this species

occurs further to the east in Wallacea and at least in the extreme

western tip of New Guinea (Coates 1985, Beehler et al. 1986,

Coates & Bishop 1997, Gjershaug& Rov 2000). Only four records

from the islands off western Papua are known in the literature:

Meyer & Wiglesworth (1898) quoted Salvadori (1880-1882) with

two specimens collected at Gagi (by Bernstein) and Waigeo (by

Platen) islands and one from an unspecified locality in ‘ Nova

Guinea (by Wallace). Ripley (1964) reported one female collected

at Efman (Jefman) Island near Sorong. Additionally, we located

five specimens held in the ornithological collections at the Natural

History Museum (formerly British Museum of Natural History),

Tring, collected in New Guinea in the nineteenth century at

unspecified localities (Plate 1). These specimens appear to have

been overlooked in the relevant literature. As the islands of eastern

Indonesia are regularly visited by birding tours, it is remarkable

that such a large number of Chinese Sparrowhawks could have been

so consistently unrecorded during the wintering months.

Moreover, our observations on Sangihe Island (Germi et al. 2009)

showed that when present the Chinese Sparrowhawk is easy to

locate, both because of its strong diurnal vocal activity and when

perched or hunting from prominent tree branches. The possible

explanation for the fact that the species has been so widely

‘overlooked’ in Wallacea, may be: (1) it has been genuinely

unrecorded due to an absence of observers present at the right time

of year; or (2) the lack of suitable wintering areas in parts of

Wallacea, so that a large proportion of the migrants disperse into

remote and poorly surveyed areas as far as Papua (Indonesian New

Guinea), or both. In order to address the paucity of winter records

in Wallacea, we undertook this survey in Papua to establish if

Chinese Sparrowhawk overwinter on the island, simultaneously

carrying out the first raptor road counts in New Guinea.

METHODS

Study area

Papua, the Indonesian half of the island of New Guinea, covers an

area of 416,129 km2. It is still largely covered in relatively

undisturbed primary forests, the largest tropical forest wilderness

remaining in the Asia-Pacific region. For the past 50 years it was

essentially inaccessible to all but a few field researchers, and thus a

terra incognita (Marshall & Beehler 2007). Dominated by the

massive Central Cordillera (more than 3,000 m high) that generates

abundant rainfall, rivers drain into vast forested interior basins and

alluvial floodplains. In the far south-east, near the Papua New

Guinea border, is a swath of savannah known as the Trans-Fly,

ecologically resembling northern Australia rather than New

Guinea. To the west, the heavily logged Vogelkop and Bomberai

peninsulas are dominated by small mountain ranges and island

groups. Papua’s equatorial climate is dominated by the North-west

Figure 1. Map of Papua, Indonesia. Black lines show all transects carried out during December 2010 - March 201 1 in the numbered sample areas: (1)

Merauke; (2) Kimaam; (3) Asmat; (4) Timika; (5) Wamena; (6) Manokwari; (7) Sorong; (8) Biak. Black stars: new Chinese Sparrowhawk records January-

March 201 1. White stars: old Chinese Sparrowhawk records 1880-1964. Black asterisk: migration watch site at Tanjung Kasuari, March 201 1.

130°E 132°E 134° E 136'E 138“E 140°E 142°E

130°E 132*E 134°E 136°E 138*E 140°E 142*E

SoZ Sotr So9 So8

Forktail 29 (2013)

First records of Chinese Sparrowhawk wintering in Papua (Indonesian New Guinea)

45

Table 1 . Sample areas, transects and Chinese Sparrowhawk records, Papua, December 201 0-March 2011.

Habitat types: 1 Trans-Fly savannah, 2 Grassland, 3 Inundated grassland , 4 Swampland, 5 Monsoon forest , 6 Mangrove forest , 7 Lowland rainforest, 8 Montane forest, 9 Coastal forest, 10 Secondary forest , 1 1

Degraded woodland, 12 Semi-arid scrub, 13 Agricultural landscapes, 14 Villages, 15 Transmigration settlements.

Monsoon and the South-east T rade Winds. The main rainy season

occurs from November to March; however in the wetter areas the

seasons are reversed, and most rain falls in the April-October

period. The highest rainfall is recorded in the southern scarp of

the Central Cordillera, with more than 5,000 mm/year.

Temperature varies little, with elevation being the key variable. At

sea level, where most of our survey areas were located, the average

temperature is 25-27°C (Marshall & Beehler 2007).

Survey techniques

Our general methodology followed a combination of classic survey

techniques described in the literature (Fuller & Mosher 1987,

Bibby et al. 1998, Bibby etal. 2000, Bird & Bildstein 2007, Malan

2009). We also reviewed specific literature on previous raptor road

surveys from other parts of the world in order to adapt our own

protocols to local conditions (Millsap & LeFranc 1988, Ellis et al.

1990, Hanowsky & Niemi 1995, Vergara 2010).

Thirty-nine road, boat and foot transects were carried out in

eight sample areas in the Indonesian administrative provinces of

Papua and West Papua (Figure 1), chosen by habitat type and for

their accessibility. The following main habitats were surveyed:

Trans-Fly savannah and grassland, swamp forest, mangrove forest,

lowland rainforest, secondary forest and agricultural landscapes

(Table 1). Logistical difficulties such as lack of roads and suitable

transport, high cost of transport, access restrictions due to oil and

mining exploration and floods due to the rainy season hindered

the fieldwork. The resulting poor accessibility and the insecurity

of some areas restricted the number of study sites and the time

spent in them. All 39 transects were carried out in the middle of

the wet season (December-late February) and recorded as GPS

tracks, and all raptor sightings as GPS waypoints. The length of

routes was measured by using a GPS, the vehicle odometer and

topographic maps. As habitat structure, detectability of raptors and

driving conditions were too variable, we did not follow a strict

methodology protocol (i.e. strip, line or point transects), but a

combination of the three. To facilitate relocation of the routes for

future surveys, we downloaded all recorded tracks in a GPS

navigation software (Touratech QV). Sample segments were

arranged by habitat type. To minimise differences in detectability

among transects, we standardised the time of day and weather

conditions. Driving speed was 40-60 km/hr on paved road and

20-40 km/hr or less on dirt roads, although road conditions and

habitat type were too variable to permit a uniform driving speed.

Roadside counts were conducted by 1-2 observers assisted by one

driver. Observers looked carefully for accipiters while driving slowly

(<50 km/hr), and during frequent stops to identify distant birds

or scan the landscape, from after sunrise to before sunset (07h00-

18h00), using 10x binoculars and a 20-60x telescope. All

accipiters seen perched or flying were counted and identified if

possible. As habitat structure in the proximity of the roads affected

detectability of birds, we stopped at points spaced about 1-3 km

in suitable habitats to look and listen for accipiters during a 5-10

minute period, or to conduct foot transects when necessary. During

these stops we broadcast three different recordings of Chinese

Sparrowhawk vocalisations in an effort to elicit vocal responses or

approaches, using a standardised protocol (Parker 1991).

Twelve days from 6 to 17 March 2011 were spent at a

peninsular migration watch site, Tanjung Kasuari (0.823°S

131.231°E), near the westernmost point of West Papua, in the

proximity of the town of Sorong, to assess whether migratory

movements were visible at this site. Methods were similar to those

used in previous migration studies in Indonesia (Germi etal. 2009).

RESULTS

Transects covered a total length of 2,303 km, of which 1,948 km

were on roads or footpaths and 355 km on rivers. T ransects in forest

habitats resulted in very few raptor sightings, probably due to poor

detectability in dense forest, thus suggesting that this habitat type

requires a different methodological approach to carry out raptor

counts (Thiollay 1989, Whitacre 1991, Whitacre & Turley 1991).

Eighteen raptor species were recorded during transects, all at low

density (Table 2), and including three species (Black-shouldered

Kite Elanus caeruleus, Bat FFawk Macheiramphus alcinus and

Brown Goshawk Accipiter fasciatus ) from areas outside their known

distribution, thus extending their range within New Guinea.

Chinese Sparrowhawk was observed five times at four sites

(Kimaam, Timika, Manokwari and Biak), 10 individuals in all

(Table 1), up to 1,200 km east of the previously known limit of

the wintering range, thus proving that this species does winter in

Papua. These are the first records for the species in mainland New

Guinea. All five records were at sea level, near coasts, in degraded

habitats and swampland. Individuals were observed at sufficiently

close range to permit positive identification using diagnostic field

characters. The authors are very familiar with the identification of

this species from previous field studies in the region (Germi 2005,

Germi & Waluyo 2006, Germi et al. 2009).

New records

Kimaam, Dolok island: 1 0 January 2011, one adult male observed

and photographed at short distance, roosting on a light pole at

Kimaam airstrip (7.980°S 138.853°E) in Trans-Fly savannah.

Manokwari: 12 February 2011, one adult male roosting on a

tree branch in secondary coastal forest in the proximity of the town

of Manokwari (0.808°S 134.053°E).

46

FRANCESCO GERMI, AGUS SALIM & ANDREA MINGANTI

Forktail 29 (2013)

Tabie 2. Raptor species recorded during transects, Papua, December 201 0-March 2011.

Biak island: 14 and 15 February 2011, two single females

(possibly the same bird) observed at the same site, roosting on a

tree branch in degraded woodland (1.094°S 136.330°E).

Timika: 3 March 2011, a flock of six individuals soaring and

gliding at low altitude, apparently in westward migration, observed

and photographed over swampland (4.69 1°S 136.887°E) along the

mining road between the town of Timika and the port of

Amamapare.

From previous observation in Sangihe (Germi et al. 2009), we

found that Chinese Sparrowhawk can be conspicuously vocal when

perched on exposed tree branches, especially in the early morning.

Although we broadcast Chinese Sparrowhawk vocalisations on

numerous occasions in seven sample areas (excluding Biak) and

towards the individual observed in Manokwari, we never obtained

any response or approach.

During 1 2 days spent at a suitable watch site at the westernmost

point of Papua during the migration season, no movements of

Chinese Sparrowhawk were observed, although dates were chosen

to precede by a few days the known passage period in North Sulawesi.

DISCUSSION

W e plotted the known records of migrating Chinese Sparrowhawks

in Wallacea and the handful of historical winter records from the

islands off the westernmost tip of Papua. This hinted at a probable

winter distribution in poorly surveyed but ecologically suitable areas

in Papua, particularly in the open habitats of the south-east. As

this territory is subject to heavy flooding during the rainy season,

its wintering avifauna is poorly known, partly because of the

inaccessibility of much of the region. Moreover, very few formal

surveys have been published from Papua in the past 20 years and

only a handful of informal accounts from short visits by

ornithologists and birdwatchers have appeared (Marshall & Beehler

2007).

Records of wintering Chinese Sparrowhawks in other parts of

Indonesia show that this accipiter is not a closed-canopy forest

species; instead it occupies a variety of open habitats ranging

from forest edge, secondary forest and scrub to agricultural

landscapes. Previous observations in central Flores (Germi et al.

2009) showed that wintering Chinese Sparrowhawks feed

primarily on cicadas, and the majority ot birds leave the island in

late November once cicada emergences cease, and only small

numbers remain through the winter. Interestingly, large emergences

of cicadas are known to occur at the beginning ol the rains in

October-November in Papua and in October-February in the

Moluccas, suggesting that Chinese Sparrowhawks move east when

insect abundance shifts with the advancing rainy season (Germi et

al. 2009). The south-east ol Papua, part of the Greater Trans-Fly,

is characterised by an extensive mosaic of monsoon forest,

secondary forest, swampland and swathes of savannah, thus

potentially ideal wintering habitat. Although this region was

initially the focus of our study, lack of accessibility and the

subsequent paucity of records (only one individual at Kimaam,

Dolok island) prompted us to extend the survey into different parts

of Papua. Most areas surveyed presented several logistical

constraints and difficult access. Suitable habitat covered large

inaccessible areas, and the scarcity of records during this study

suggests that the species disperses at low density within the vast

region. However our records, although small in number, clearly

indicate that an unknown proportion of this accipiter population

overwinters in Papua, apparently in coastal areas.

The absence of migrating Chinese Sparrowhawks in March

(migration season) at the watch site of Tanjung Kasuari (Figure 1)

might reflect the use of a different route further to the south. We

assumed that migrants would travel from the Sorong area to

Waigeo and Halmahera islands, but the lack of migrants at Tanjung

Kasuari could indicates that migrants leave Papua via Salawati

Island, where shorter water crossings are available, and continue

east through Kofiau, Obi and the Sula islands before reaching

Forktail 29(2013)

First records of Chinese Sparrowhawk wintering in Papua (Indonesian New Guinea)

47

Sulawesi. Access restrictions on the Papua coast in the proximity

of Salawati prevented us from testing this hypothesis.

ACKNOWLEDGEMENTS

This survey was funded by The Peregrine Fund (USA) and Hawk Mountain

Sanctuary (USA). We wish to thank Cheryl Tipp (British Library of Wildlife

Sounds) and Yasunori Nitani (Asian Raptor Research and Conservation

Network) for providing Chinese Sparrowhawk recordings; Mark Adams at

the Natural History Museum, Tring, for access to specimens; Daniel Natush

and Jessica Lyons for their help in Timika and for sharing raptor records;

Benja Mambai and Marco Wattimena (WWF Papua) for logistical assistance;

Rick Watson and Keith Bildstein for reviewing an earlier draft of the

manuscript and two anonymous referees for helpful comments. We are also

grateful to the following people for their kind cooperation at various stages

of the study: Stephen Bird, Giuseppe Carpaneto (Department of

Environmental Biology, Rome University), Tom Edwards, James Ferguson-

Lees, Lloyd Kiff (Global Raptor Information Network/The Peregrine Fund),

Marco Panella (Italian Ministry of Forests) and Michael Schneider.

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Francesco GERMI, 46 (2F2) Elm Row, Edinburgh EH7 4AH, UK.

Email: fgermi@yahoo.co.uk

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16113, West Java, Indonesia. Email: salim@asalim.org

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Italy. Email: andming@alice.it

FORKTAIL 29 (201 3): 48-51

Spatial distribution of the Tawny Fish Owl Ketupa flavipes

shaped by natural and man-made factors in Taiwan

SHIAO-YU HONG, YUAN-H5UN SUN, HSIN-JU WU & CHAO-CHIEH CHEN

This study investigated the distribution of the Tawny Fish Owl Ketupa flavipes, a rare top predator in Taiwan, and examined natural and

man-made factors that affect it. Records of Tawny Fish Owls from 1 993 to 2006 were compiled from field studies, literature surveys, museum

notes and specimens, and interviews with researchers, birdwatchers and indigenous hunters. In total, 91 Tawny Fish Owl territories were

identified, widely distributed along mountain streams in the Central Mountain Range between 48 and 2,407 m — more than half of them

were below 700 m. The upper altitudinal range of the owls is probably limited by food availability and stream size. Territories were on

average 431 m higher on the west side of the Central Mountain Range than on the east. Habitat selection analysis further indicated that, in

proportion to the land area available, the Tawny Fish Owl was absent from areas below 500 m.This is apparently due to extensive deforestation

of lowlands for agriculture and urbanisation on the west side of the island. It is recommended that a protected area be established in the

north-east part of Taiwan, to preserve the remaining lowland streams and riparian forests still inhabited by the species.

INTRODUCTION

Fish owls are top predators in freshwater food chains and are

important indicator species for healthy stream ecosystems (Duncan

2003, Wu et dl. 2006). The Tawny Fish Owl Ketupaflavipes is widely

distributed in the Himalayas, eastern Indochina, south China and

Taiwan (Voous 1988, Marks et al. 1999). It reaches 58 cm in body

length (Sun 1996), making it one of the largest raptors within its

range. However, it is so rarely observed in the wild that it is

considered to be rare over most of its range (Marks et al. 1999). It

is currently listed in CITES Appendix II (UNEP-WCMC 2009).

The species was first reported from Taiwan by Kuroda (1916).

Since then its natural history has remained poorly known (Voous

1988). In 1989, when the Wildlife Conservation Law was first

implemented in Taiwan, the Tawny Fish Owl was listed as an

Endangered Species (Class I) due to poaching (Sun 1996) and

destruction of riparian forests (Severinghaus 1987). Even though

its population was still considered very small (Fang 2005), Tawny

Fish Owl was down-listed to Rare and Precious Species (Class II)

in 2008, due to better knowledge of its population size (Sun 1996,

Sun etal. 2000,2004).

As the distribution and population size of the Tawny Fish Owl

in Taiwan remained unclear, further investigation was needed to

correctly assess its protection status. The aims of this study were to

determine its spatial distribution and to evaluate factors affecting

its distribution by identifying locations throughout Taiwan where

Tawny Fish Owl had been recorded recently.

METHODS

Study area

Taiwan is a mountainous island lying between 21 and 25°N and 120

and 122°E in subtropical East Asia, about 150 km off the coast of

south-east China. It is 394 km long and 144 km wide with a total

area of 36,000 km2. Hills and mountains with elevations higher than

100 m cover about 70% of the island (Taiwan Forestry Bureau 1995).

The Central Mountain Range, with more than 200 peaks over

3,000 m in height, runs from north to south. The rivers that originate

from the mountain range are short and steep with rapid currents,

particularly on the east side. A lower Coastal Mountain Range is

present between the east coast and the Central Mountain Range.

Forests cover 59% of the island, of which 73% is natural forest

(44% of the island) (Taiwan Forestry Bureau 1995). Broad-leaved

forests predominate below 1,500 m, coniferous forests above 2,500

m, with mixed forests in between. The coastal plains are much wider

on the west side than on the east, and most of them have been

converted to urban or agricultural areas. Only small remnants of

the original riparian forests remain on the peripheral hilly areas on

the west side of the mountain range, whereas most forests on the

east side remain little affected because of access difficulties (Taiwan

Forestry Bureau 1995). Annual precipitation is between 1,000 mm

and 4,800 mm, and decreases gradually from north-east to south¬

west because of monsoon rainfalls and incidence of typhoons (Chiu

2006). Annual average temperature is about 25°C at sea level and

4°C at about 3,800 m (Chiu 2006).

Data collection and analysis

Tawny Fish Owl records between 1993 and 2006 were collected

using the following approaches to identify locations where it

occurred. First, faunal reports and the bird-sighting databases of

wild bird societies were searched for records, and the collection data

for Tawny Fish Owl specimens preserved in the Academia Sinica

(AS), the National Taiwan Museum (NTM), National Museum of

Natural Science (NMNS), and Taiwan Endemic Species Research

Institute (TESRI) were examined. A total of 42 records were

obtained: nine records from faunal reports, 15 from wild bird

societies, two from AS, five from NTM, six from NMNS, and five

from TESRI.

Second, 19 birdwatchers and 1 19 hunters from 27 indigenous

villages located near rivers where there were no or very few records

of Tawny Fish Owls were interviewed. All information indicating

the presence of owls was collected, including hunting captures,

sightings, calling birds, pellets, feathers, droppings and food

remains. Tawny Fish Owl calls are easily distinguished from those

of other owls in Taiwan (Sun 1996). Interviewees had to describe

the sound of the owl clearly before each record was considered valid.

As a fish-eater, the droppings and food remains of Tawny Fish Owls

were easily identifiable (Wu et al. 2006). We obtained 52 records

from the interviewees.

Third, field surveys were conducted to identify Tawny Fish Owl

sites, using the same clues as above to detect the presence of owls.

The sites surveyed included 15 coldwater fish farms (Rainbow

Trout Oncorhynchus mykiss and smelt [Osmeridae spp. ] ) near

mountain streams, mostly in northern Taiwan. The owls preyed

on fish in the farms and often left remains, such as fish scales,

bloodstains and internal fish organs, along with their own feathers,

near the predation sites (Sun et al. 2004).

The Tawny Fish Owl has a strong territorial habit (Fogden

1973, Sun 1996, Sun et al. 2000) . A territory of a pair of owls was

Forktail 29(2013)

Spatial distribution of the Tawny Fish Owl Ketupa flavipes in Taiwan

49

estimated to be about 6.2 km along a river and less than 550 m

from the bank, derived from two territories measured by radio¬

tracking (Sun et al. 2000). In this study any owl records obtained

within a 6.2 km length of river were considered to refer to a single

pair, and the midpoint of the two outermost records was used to

represent the territorial site.

To construct the Tawny Fish Owl’s altitudinal distribution chart,

the elevation of each of the owl territories was obtained by using

coordinates under Identify (ArcGIS) on a Digital Terrain Model

(DTM) of Taiwan (precision 40 m x 40 m). The perpendicular

distance of each owl territory from the crest line of the Central

Mountain Range was calculated, to illustrate how altitudinal

distribution varied across the island. A 50 m interval contour map

was created from the DTM by 3D Analyst, and then transformed

to a 3D topographic chart in ArcScene. The 3D topographic chart

was rotated to calculate the altitudinal profile of the island (between

25 and 375 km south of the northern tip of the island), as viewed by

an observer standing on the south end of the island. Elevations of

territories to the east and west sides of the Central Mountain Range

were compared with an independent r-test.

The altitudinal range of the island from sea level to 3,950 m

was divided into four bands: <500 m, 500-1,000 m, 1,000-

1,500 m and >1,500 m. The proportion of owl territories in each

altitude band was compared with the proportion of the land area

in each altitude band. A use and availability analysis (Litvaitis etal.

1994) was conducted using a chi-squared test and multiple

simultaneous comparisons between Bonferroni confidence

intervals of observed use and the proportion of land area in each

altitude band. All statistical analyses were conducted with SPSS

10.0.7C for Windows with an a-level set at 0.05.

RESULTS

A total of 153 owl records were obtained. Of these, 15 museum

specimens and 1 1 records from wild bird societies had no clear

location and were excluded from the study. The 127 valid records

consisted of 37 from birdwatchers, 28 from indigenous hunters,

35 from field surveys along rivers, 1 5 from fish farms, 9 from fauna

reports and 3 from museum collections. Of these, 39 records were

direct sightings of owls, 35 hunting captures or owls found dead,

28 sites with owl pellets, 17 calling birds and 8 records of feathers,

food remains, droppings or other signs (Table 1).

Taking owl records found within a 6.2 km section of a river to

represent single pairs, the 127 valid records indicate 91 owl

territories. Two of these territories had four records and 70

Table 1. Sources and categories of 127 Tawny Fish Owl records

obtained in 1993-2006 for this study. Captured birds included those

caught by hunters or found dead by birdwatchers and those preserved

as specimens in museums and research institutions.

Sources

Figure 1. Locations of 91 verified Tawny Fish Owl territories in Taiwan,

1993 to 2006.

120°0'0"E DIWE 122 0'0"E

25 0’0"N

24 0'0"N

23 0'0"N

22“0'0"N

Elevation (m)

0-500

501 - 1,500

1 ,50 1 - 2,500

01020 40 60 80

territories contained only one. The territories were distributed along

mountain streams in the Central Mountain Range (Figure 1 ). There

were no records from either of the coastal plains or the Coastal

Mountain Range in the east.

The altitudinal distribution of the owl territories ranged

between 48 m and 2,407 m with mean 687 m and the middle 50%

range between 300 m and 1,100 m (Figure 2). The distribution

was skewed toward low elevations (skewness = 0.97), implying that

most of the owl territories were located at lower altitudes.

The transverse altitudinal distribution of the owl territories in

relation to the Central Mountain Range is shown in Figure 3. There

was significant difference {t = 3.04, df = 55 ,P < 0.01) between the

Figure 2. Frequency distribution of the elevations of 91 verified Tawny

Fish Owl territories in Taiwan, 1 993-2006.

in

<L>

CD

>

o

c

CD

Z3

cr

CD

LL

14 ]

12

10

8

0

1,000 1,500 2,000 2,500

Elevation (m)

50

SHIAO-YU HONG etal.

Forktail 29(2013)

Figure 3. Altitudinal distribution of Tawny Fish Owl territories relative

to the crest line of the Central Mountain Range. The grey outline shows

the altitudinal profile of the island, viewed from the south end. (The

vertical scale is magnified 30-fold as the elevation of Mount Yushan,

the highest peak of Taiwan, is relatively small [4 km] compared with

the width [144 km] of the island.)

Crest line of Central

Mountain Range

0 1 — - - 1 - 1 - 1 - 1 — - - 1 - 1 — - - 1 — 1 - 1

0 20 40 60 80 100 120 140 160

Transverse distance (km)

east and west side of the mountain range. Most of the territories on

the west side were above 500 m, whereas more than half of the

territories on the east side were below 500 m. The former averaged

431 m higher than the latter. Territories were generally situated

in deep valleys, well below the surrounding mountain ridges

(Figure 3).

Tawny Fish Owls did not use the four altitude bands in

proportion to the land area of the island (%2 = 41.57, df = 3, P <

0.0001). Multiple comparisons showed that the owls appeared in

land below 500 m and above 1,500 m in a significantly lower

proportion compared with the land area of these altitude ranges

(Table 2). In contrast, the proportion of owl territories was twice

the proportion of the land area between 500 and 1,500 m,

indicating that the owls selected this altitude range.

DISCUSSION

The Tawny Fish Owl mainly takes prey from streams (Sun 1996,

Wu et al. 2006). Its upper altitudinal range may be constrained by

the distribution of stream fishes (Voous 1988, Marks et al. 1999).

In Taiwan, the highest reported elevation for stream fish is

2,400 m (Tzeng 1986, Wang 20 10), which coincides with the upper

limit of the owl’s altitudinal range (Figure 2). Fish abundance and

diversity increase at lower elevations, for example 10 fish species

occur at 1,500 m (Wang 2010) and only six owl territories were

recorded above 1,500 m suggesting that fish abundance and

diversity is important to the species. However, amphibians and

crabs were found to be the most important prey items for Tawny

Fish Owls at Sakatang Stream, eastern Taiwan (Wu et al. 2006),

and the abundance and diversity of these taxa also increase at lower

altitudes.

The occurrence of the Tawny Fish Owl was also related to the

distribution of mountain streams. Streams with stable flow above

1,500 m are only found in central regions where large mountains

occur. Tawny Fish Owls seldom forage in small creeks less than

5 m wide, perhaps due to food scarcity and the poor

manoeuvrability of a bird with a wingspan of 1.5 m (Sun et al. 2000).

As a result, the highest parts of the range mainly occurred around

the largest mountains. The latter phenomenon was possibly

enhanced by the Massenerhebung effect, which predicts that

locations at the same elevation are warmer on large mountains

compared to smaller ones, because the large mountains release heat

more slowly (Flenley 1994).

Conversely, local differences between the altitudinal range of

the owls and the surrounding mountain ridges were larger in small

mountains and mountains with steep slopes. For example, the owl’s

upper altitudinal limit on the east side of the Central Mountain

Range, where rivers are much smaller, steeper and often dry up,

was lower than that on the west side. This may partly explain why

owl territories on the east side of the island were lower than those

on the west side. In addition, no Tawny Fish Owls were recorded

in the Coastal Mountain Range where rivers are short and small.

Similarly, the owls were seldom found in streams on low mountains

in the south of Taiwan. In contrast, streams in the north of Taiwan

have regular water flow all year round, due to the north-east

monsoon rainfalls (4,000 mm/year) (Chiu 2006), so the owls

occurred further upstream, closer to the mountain ridge.

The distribution of Tawny Fish Owls in Taiwan was also

influenced greatly by habitat loss due to human activities.

Destruction of lowland habitat is widespread, especially on the west

side of the island, due to the conversion of natural forests to farmland

and urbanisation over the last 200 years. The habitat selection

analysis indicated that there were fewer owls in lowland areas, even

though lowlands should have more suitable streams than the hills —

wider, with more reliable flow, and greater prey abundance and

diversity. Most Tawny Fish Owl territories to the east of the Central

Mountain Range were below 500 m, but only two owl territories

were found below that elevation on the west side. The lower

altitudinal limit on the west side is likely to have retreated into higher,

mountainous areas, as is the case for many forest birds and mammals

on the island (Liu et al. 2003, Shiu 2003). It is likely that the loss of

natural forests below 500 m has eradicated Tawny Fish Owls from

the western lowlands, since riparian natural forests are the main

habitat type used by fish owls (Hayashi 1997, Sun et al. 2000).

Poaching is one of the most serious threats to the Tawny Fish

Owl population (Severinghaus 1987) and at least 61 records of

hunting have been reported since the species was legally protected

in Taiwan in 1989 (Wang et al. 1995). Of the 28 owl records

obtained from indigenous hunters, 54% were the result of poaching.

Only a small proportion of indigenous hunters were interviewed,

suggesting that poaching is likely to be widespread across the island.

More than 60% of the owl territories found in this study were

situated outside areas protected for wildlife (Hong 2007). The

lowlands are largely unprotected and those on the developed

western part of the island are already deforested, therefore we

recommend the establishment of a protected area for the Tawny

Fish Owl along lowland streams in north-east Taiwan, where the

original riparian forests still remain nearly intact (Taiwan Forestry

Table 2. The effects of altitude on territory selection by Tawny Fish Owls in Taiwan (%2= 41 .57, P < 0.0001). The Bonferroni confidence intervals

show that the % use of each altitude band is higher (selected) or lower (avoided) than the % availability.

Forktail 29(2013)

Spatial distribution of the Tawny Fish Owl Ketupa flavipes in Taiwan

51

Bureau 1995). This is possibly the only place in Taiwan where the

Tawny Fish Owl range still reaches the coastline. This

recommendation is in accordance with other studies that have

recommended low-elevation protected areas for threatened species

including Clouded Leopard Neofelis nebulosa , Leopard Cat

Prionailurus bengalensis , Small Indian Civet Viverricula indica,

Formosan Pangolin Manis pentadactyla and Fairy Pitta Pitta

nympha (Liu et al. 2003, Chiang & Pei 2004, Lee et al. 2006).

ACKNOWLEDGEMENTS

We thank C. L. Bridgman and L. C. Lo for providing useful suggestions, and

C.-F. Tsai for editorial help during the preparation of this paper. C. T. Yao

and Y. J. Chen helped locate specimen records. We are also indebted to

numerous birdwatchers and hunters who were willing to share with us their

information on the Tawny Fish Owl.

REFERENCES

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Clouded Leopard and other medium-to-large mammals at Tawu Nature

Reserve and vicinities (III). Taipei: Taiwan Forestry Bureau. (In Chinese

with English abstract.)

Chiu, C.-A. (2006) Applying the ecoclimatic indices to predict the potential

natural vegetation of Taiwan. PhD thesis, National Chung-Hsing

University, Taichung, Taiwan. (In Chinese with English abstract.)

Duncan, J. R. (2003) Owls of the world: their lives, behavior and survival. New

York: Firefly Books.

Fang, W.-H. (2005) A guide to threatened birds of Taiwan. Taipei: Owl

Publishing House. (In Chinese with English abstract.)

Flenley, J. R. (1994) Cloud forest, Massenerhebung effect and ultraviolet

insolation. Pp.1 50-55 in L. S. Hamilton, J. O. Juvik & F. N. Scatena, eds.

Tropical montane cloud forests. New York: Springer-Verlag.

Fogden, M. (1973) Fish-owls, eagle owls and the Snowy Owl. Pp. 53-85 in J.

A. Burton, ed. Owls of the world: their evolution, structure, and ecology.

New York: A. W. Visual Library.

Hayashi, Y. (1997) Home range, habitat use and natal dispersal of Blakiston's

Fish-Owls. J. Raptor Research 31: 283-285.

Hong, S.-Y. (2007) Distribution pattern of Tawny Fish Owls ( Ketupa flavipes)

in Taiwan. MS thesis, National Pingtung University of Science and

Technology, Pingtung, Taiwan. (In Chinese with English abstract.)

Kuroda, N. (1916) Rare species of avifauna of Formosa . Zoological Magazine

Tokyo 28: 263-264. (In Japanese.)

Lee, P.-F., Bai, M.-L. & Lin, R.-S. (2006) Habitat preference and distribution

prediction of vulnerable Fairy Pitta (Pitta nympha) in Taiwan by remote

sensing and GIS. Taipei: Council of Agriculture. (In Chinese with English

abstract.)

Litvaitis, J. A., Titus, K. & Anderson, E. M. (1994) Measuring vertebrate use of

terrestrial habitats and food. Pp. 254-274 inT. A. Bookhout, ed. Research

and management techniques for wildlife and habitats. Fifth edition.

Bethesda, MD: The Wildlife Society.

Liu, C.-N., Liu, C.-H. & Chang, C.-H. (2003) Current condition and

conservation of medium-to-large mammals at low altitude. Natural

Conservation Quarterly 43: 61-66. (In Chinese.)

Marks, J. S., Canning, R. J. & Mikkola, H. (1999) Family Strigidae (typical owls).

Pp. 76-242 in J. del Hoyo, A. Elliott & J. Sargatal, eds. Handbook of the

birds of the world, 5. Barcelona: Lynx Edicions.

Severinghaus, L. L. (1987) The Tawny Fish Owl. Pp.354-355 in A.W. Diamond,

L. L. Severinghaus & C. Chen, eds.Sove the birds. Frankfurt, Germany:

Pro Nature.

Shiu, H.-J. (2003) Spatial and seasonal variations in avian assemblages in

Taiwan. PhD thesis, National Taiwan University, Taipei, Taiwan. (In

Chinese with English Abstract.)

Sun, Y.-H. (1996) Ecology and conservation of Tawny Fish Owl in Taiwan.

PhD thesis, Texas A & M University, Texas, USA.

Sun, Y.-H., Wang, Y. & Lee, C.-F. (2000) Habitat selection by Tawny Fish-Owls

( Ketupa flavipes) in Taiwan. J. Raptor Research 34: 102-107.

Sun, Y.-H., Wu, H.-J. & Wang, Y. (2004) Tawny Fish-Owl predation at fish farms

in Taiwan. J. Raptor Research 38: 326-333.

Taiwan Forestry Bureau (1995) Thethird forest resource and land use inventory

in Taiwan. Taipei: Taiwan Forest Bureau. (In Chinese.)

Tzeng, C.-S. (1986) Distribution of the freshwater fishes of Taiwan. J. Taiwan

Museum 39: 127-146.

UNEP-WCMC (2009) UNEP-WCMC species database: CITES-listed species.

Available online: http://www.unep-wcmc.org. Accessed 22 November,

2010.

Voous, K. H. (1988) Owls of the Northern Hemisphere. London: Collins.

Wang, H.-W. (2010) Ecoregion classification by using drainage fish

community in Taiwan. MS thesis, National Kaohsiung Normal University,

Kaohsiung, Taiwan. (In Chinese with English abstract.)

Wang, Y., Sun, Y.-H. & Wu, H.-J. (1995) The distribution and use of the listed

endangered birds by indigenous peoples and the ecology of Tawny Fish-

Owl in Taiwan. Taipei: Taiwan Normal University. (In Chinese with English

abstract.)

Wu, H.-J., Sun, Y.-H., Wang, Y. &Tseng, Y.-S. (2006) Food habits ofTawny Fish-

Owls in Sakatang Stream, Taiwan. J. Raptor Research 40: 111-119.

Shiao-Yu HONG, Institute of Wildlife Conservation and

Graduate Institute of Bioresources, National Pingtung

University of Science and Technology, Pingtung 9(2, Taiwan

Yuan-Hsun SUN, Institute of Wildlife Conservation, National

Pingtung University of Science and Technology, Pingtung 912,

Taiwan

Hsin-Ju WU, Department of Life Science, National Taiwan

Normal University, Taipei 1 17, Taiwan

Chao-Chieh CHEN (Corresponding author) Department of

Biomedical Science and Environmental Biology, Kaohsiung

Medical University, Kaohsiung 807, Taiwan. Email:

chen51 23@kmu.edu.tw or chenkmu@gmail.com

FORKTAIL 29 (2013): 52-56

Population, breeding and threats to the White-rumped

Vulture Gyps bengalensis in Bangladesh

M.MONIRULH. KHAN

The population of the White-rumped Vulture Gyps bengalensis in Bangladesh has declined very rapidly in recent years, so a research-cum-

conservation project was launched in July 2008 that continued until June 201 2. Three species of vultures were found during the survey —

White-rumped Vulture, Himalayan Vulture Gyps himalayensis and Cinereous Vulture Aegypius monachus. Based on nesting sites and frequent

sightings of vultures, a total of six 'hotspots' were identified in the areas of Moulvibazar, Habiganj, Haor Basin, Mymensingh, Sundarbans

(northern end) and Barisal.The total population of the White-rumped Vulture in suitable habitats across the country shows that numbers

have drastically declined from 1,972 to 816 (nearly 60% drop) in four years. In two consecutive breeding seasons only 5 out of 32 and 8 out

of 31 nests were successful in producing fledglings (one from each nest). The overall breeding success was very low (1 5.6-25.8%). The

reason for such poor breeding success was sudden death or disappearance of parent birds, apparently due to poisoning by diclofenac, a

veterinary drug used to treat livestock ailments. The project identified poisoning as the principal cause of vulture decline. Although the

Government of Bangladesh banned use of veterinary diclofenac from 25 October 2010, 53% of the veterinary drug stores still sell it illegally.

Awareness campaigns have made people aware of vulture conservation and the adverse effects of diclofenac.

INTRODUCTION

Historically, seven species of vultures — White-rumped Gyps

bengalensis , Himalayan G. himalayensis , Griffon G.fulvus , Slender-

billed G. tenuirostris , Cinereous Aegypius monachus, Red-headed

Sarcogyps calvus and Egyptian Neophron percnopterus — have been

recorded in Bangladesh (Khan 2008, Siddiqui et al. 2008).

However, only the White-rumped Vulture, Himalayan Vulture and

Cinereous Vulture have been seen in the last four years and only

the White-rumped Vulture is now known to breed in Bangladesh.

Populations of the White-rumped Vulture and other resident

Gyps vulture species have declined very rapidly since the mid- 1 990s

across the Indian subcontinent (Prakash 1999, Gilbert et al. 2006,

Prakash et al. 2007, Chaudhary et al. 2012). Declines in numbers

of the White-rumped Vulture have exceeded 99.9% in India

(Prakash et al. 2007) and the species is classified as Critically

Endangered (BirdLife International 2001, BirdLife International

2012). If the rate of decline cannot be arrested, the species will

disappear from the Indian subcontinent in the next few years. This

species not only plays a key role as a scavenger but is also part of the

heritage of the Bengal region. The people of Bangladesh are not

hostile to vultures, but they have been almost totally unaware of the

threats and dire situation that vultures are facing. A decade ago the

White-rumped Vulture was a common and widely distributed bird

in Bangladesh (Harvey 1990, Thompson &Johnson 1996). Recent

studies in India, Nepal and Pakistan confirm that vultures are

poisoned when they feed on the carcasses of cattle treated with the

veterinary drug diclofenac shortly before their death (Green et al.

2004, Oaks et al. 2004, Shultz et al. 2004). This is also likely to be

the main cause of the decline in the vulture population in

Bangladesh, but there may be other factors contributing to the

decline.

Because the White-rumped Vulture is a globally and nationally

threatened species (BirdLife International 2001, BirdLife

International 2012) it was necessary to take measures to save it from

local extinction. Here the findings of a research-cum-conservation

project focusing on this species, started in 2008, are reported. The

aim was to understand the conservation status of the White-

rumped Vulture in Bangladesh and to implement ways of reducing

population decline. The specific objectives were to estimate the

relative abundance of populations in different parts of the country,

assess the population trend, identify important vulture ‘hotspots’

(where they nest and are frequently sighted), record breeding

success and assess the threats (focusing mainly on the availability

and use of veterinary diclofenac). Moreover, vulture conservation

awareness programmes were conducted to spread knowledge of the

dire situation faced by the vulture population and discourage the

use of diclofenac to treat cattle.

METHODS

Study area

The project was implemented in different parts of the country, but

focused on Greater Sylhet (north-east), Greater Khulna (south¬

west), Greater Mymensingh (north) and Greater Barisal (south)

Figure 1. Bangladesh showing hotspots (shaded black) for the White-

rumped Vulture Gyps bengalensis.

Forktail 29 (2013) Population, breeding and threats to the White-rumped Vulture Gyps bengalensis in Bangladesh

53

where the White-rumped Vulture was known to nest and roost

regularly (Figure 1 ). Geographically, Bangladesh is located between

20.567°-26.550°N and 88.017°-92.683°E. The total area of the

country is 147,570 km2, with a population of around 160 million

people. The climate is tropical monsoon, characterised by marked

seasonal variations. Abundant rainfall during the monsoon (July-

October) is followed by a cool winter period (November-February),

and then a hot and dry summer (March-June). Bangladesh can be

divided into three main physiographic divisions — Tertiary hills,

Pleistocene terraces and recent plains (Khan 2008).

Field methods

A small team of researchers carried out the surveys, but local people

were also involved in most areas. Five members of the research team

had been trained in different aspects of wildlife biology. The project

team worked closely with villagers and labourers on tea plantations,

since the vultures mainly occur and breed in and around villages

and tea estates. Journalists, veterinarians and other professional

people were also involved with various activities. The project team

liaised closely with the Bangladesh Forest Department. Work was

carried out from July 2008 to June 2012 (hence 2008-2009 is the

period between July 2008 and June 2009, so that each breeding

season falls in one slot), but not all the activities were carried out

each year of the project. Some data were collected during 2005-

2008 to confirm the relative abundance of vultures in different areas

and find the hotspots.

Surveys to assess relative abundance and identify vulture

hotspots

Between 2005 and 2008, vulture sightings by 50 local people were

recorded every year in each of the seven administrative divisions

(Dhaka, Chittagong, Sylhet, Rajshahi, Rangpur, Khulna and

Barisal) so that division-wise the relative abundance of vulture

populations could be estimated. In order to get the best output from

the limited resources available for awareness and conservation

activities, hotspots were identified on the basis of the occurrence

of nesting sites and frequent sightings (where interviewees have seen

vultures at least once in every two-month period).

Population surveys

To determine the population trend, the team selected potential

survey areas throughout the country and systematically visited the

known roosting sites in the morning and in the afternoon to count

White-rumped Vultures in roosting colonies. Sightings of other

vulture species were recorded during these surveys. The survey team

interviewed local people about their vulture sightings and, in areas

where interviewees claimed recent sightings of roosting vultures, the

survey team stayed and counted the vultures when they returned to

roost. In areas where the interviewees said that there were no recent

sightings at roosting colonies, the survey team quickly moved on to

new areas. The same roosts visited during 2008-2009 were visited

again in 2009-20 10,2010-2011 and 2011-2012. Since the count

was conducted in all the potential sites for vultures, although not in

each and every part of the country, the annual counts can be

considered as total counts for the country or a very close

representation of it. More importantly, however, the count has been

repeated in a standardised way so that the data are comparable year

on year.

Monitoring breeding success

The breeding success of vultures was documented by periodically

(at least once every two weeks) observing every known nest during

the breeding season (dry season: October to March). Telescopes

and binoculars were used for these observations. A commercially

made camouflaged hide was often used so that the nesting vultures

were not disturbed by the presence of observers. The main

information recorded was the fate of nests — if the nestling from a

particular nest flew (fledged), the nest was treated as successful.

Information on nesting trees (species, nest height from the ground)

was also recorded.

Surveys for diclofenac

The availability of veterinary diclofenac and other non-steroidal anti¬

inflammatory drugs (NSAIDs) was assessed in two ways. First,

undercover surveys (posing as buyers) of the veterinary drug stores

throughout the country were undertaken (70 drugstores every year),

from 2008-2009 to 20 1 1 -20 1 2. Second, the use of diclofenac was

recorded by interviewing local cattle-owners (86 individuals in total)

in different areas of the country. This was done openly (since the

cattle-owners were not as secretive as the veterinary drug salesmen)

by using. a standard questionnaire that also included a question on

what the cattle-owners do with dead cattle.

RESULTS

Based on sightings of the White-rumped Vulture during 2005-

2008 surveys by local people in suitable habitats of the seven

administrative divisions, the highest relative abundance was found

to be in Sylhet (5.1 sightings/interviewee/year) and the lowest in

Chittagong (0.7 sightings/interviewee/year) (Table 1). This gives

an average for the country of 2. 8 sightings/interviewee/year. Based

on nesting sites and frequency of sightings, a total of six hotspots

were identified: the areas of Moulvibazar, Habiganj, Haor Basin,

Mymensingh, Sundarbans (northern end) and Barisal (Figure 1).

Table 1. Relative abundance of the White-rumped Vulture in suitable

areas of seven divisions of Bangladesh.

Survey data from 2008-2009 to 201 1-2012 demonstrate that

the White-rumped Vulture declined by nearly 60% over the four-

year study period throughout the country, and that other species of

vultures are extremely rare. The total population of the White-

rumped Vulture in suitable habitats across the country (which can

be treated as the total Bangladesh population) shows that the

population drastically declined from an estimated 1,972 in 2008-

2009, 1456 in 2009-2010, 991 in 2010-2011 and 816 in 2011-

2012.

Other than the White-rumped Vulture there were 16 sightings

of Himalayan Vulture from Moulvibazar (north-east), Sunamganj

(north-east), Habiganj (north-east), Jamalpur (north), Sirajganj

(west) and Rangpur (north-west). All sightings were between

October and March indicating that they were winter visitors. Most

of the sightings were small flocks, but the largest roosting flock of

21 birds was sighted in a tall Silk Cotton Salmalia sp. tree at

Dawrachara Tea Estate, Moulvibazar, on 3 March 20 1 2. Himalayan

Vultures were seen roosting, soaring, or feeding with White-rumped

Vultures six times. There were only four sightings of Cinereous

Vulture from Rajshahi (north-west), Madhupur Tract (central),

Narshingdi (north-east) and Moulvibazar (north-east). All were

solitary birds, including two juveniles, sighted at different seasons,

indicating that they were probably vagrant individuals.

54

M. MONIRUL H. KHAN

Forktail 29 (2013)

Table 2. Breeding status of the White-rumped Vulture in seven divisions of Bangladesh

Division

Number of nests observed

Season-1 Season-2

Unsuccessful

(nesting birds died/ vanished)

Season-1 Season-2

Status of breeding

Successful (nestling left nest)

Season-1 Season-2

Breeding success (%)

Season-1 Season-2

NB. Season 1 (October 2009-March 2010) and Season 2 (October 2010-March 2011) were the two consecutive breeding seasons of vultures.

Breeding success of the White-rumped Vulture for all nests in

all the known breeding areas in Bangladesh was documented. In

Season 1 (October 2009 to March 2010), a total of 32 nests were

observed of which only five birds from five nests successfully fledged

(clutch size is one), giving an overall breeding success of 1 5.6% (Table

2). In Season 2 (October 2010 to March 2011), a total of 31 nests

were observed from which eight birds fledged (25.8% success rate).

At unsuccessful nests the parent birds were found dead, either on

the ground near the nest (n = 1 1) or on/beside the nest (n = 4), or

the parent birds just vanished suddenly (n = 35), indicating that

they had probably died elsewhere. Although no post-mortems were

undertaken, the dead vultures were apparently in good health,

indicating sudden death that can be caused by poisoning. It was

observed that, whether the nest had an egg or nestling, one parent

almost always attended the nest to guard against crows (House Crow

Corvus splendens , Large-billed Crow C. macrorbynchos ) and other

raptors (Pallas’s Fish Eagle Haliaeetus leucoryphus , Steppe Eagle

Aquila nipalensis). The parents take shifts so that both can feed and

bring food for the nestling.

Vulture nests were found in the following trees -.Albizia lebbeck ,

Albizia procera, Antbocephalus chinensis , Bombax ceiba, Borassus

flabellifer. Cocos nucifera , Ficus benghalensis , Ficus religiosa, Mangifera

indica, Sivietenia mabagoni. Vultures had no preference for nesting

tree species, but all nesting trees were large. Nests were constructed

at heights ranging from 7 to 17 m.

The veterinary drug stores were surveyed for four years, from

2008-2009 to 201 1-2012, and it was found that the availability of

diclofenac in stores has decreased from 100% to 53% and the

availability of three other NSAIDs (meloxicam, ketoprofen and

sodium salicylate) had increased (Figure 2). This is in response to

the ban on production of veterinary diclofenac from 25 October

2010 and the banning of the use of veterinary diclofenac from six

Figure 2. Availability of different NSAIDs in veterinary drug stores in

Bangladesh. N.B. Otherthan NSAIDs, paracetamol and dexamethasone

were available, which are used for the same purposes.

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months after the banning of production (25 April 2011). Diclofenac

is most commonly used in the form of injection vials, but it is also

available as a suppository and tablet. Each 30-ml vial costs Taka 80

(US$1). Today, meloxicam, the safe alternative to diclofenac

(introduced after the diclofenac ban), is available in the market, but

diclofenac is still illegally sold in veterinary drug stores (Figure 2,

20 1 1 -20 1 2). Other than NSAIDs, paracetamol and dexamethasone

are also available in veterinary drug stores as anti-inflammatory

drugs. Various steroids and ureas for cattle fattening are also available,

which are normally used before the cattle are sold, so the highest

use is before the Eid-ul-Azha when many cattle are ceremonially

slaughtered by Muslims.

During 2008-2009, i.e., before the diclofenac ban, cattle-owners

from different areas were interviewed and it was found that 57% of

them used diclofenac when the cattle suffered from fever or at the

early stage of any disease. Except in remote areas, and areas where

cattle are scarce (in Chittagong Hill Tracts and parts of Chittagong

and Cox’s Bazar), diclofenac was found to be the most commonly

used cattle medicine. In 59% of cases the cattle owners said they use

diclofenac on their own without following a veterinarian’s

prescription. Although other similar drugs are available, diclofenac

is more popular because people believe that it is a more effective

cure. This medicine was used throughout the year, but most

commonly in early monsoon (April-May) when cattle diseases are

common.

Alarmingly, a very high proportion (61%) of cattle-owners said

that when cattle died they skinned them, sold the hide and buried

the body in order to control the smell. In the past, the smell would

not reach human habitations because there were many distant open

areas, but today there are very few areas as such where dead cattle

can be disposed in the open. Some cattle owners (9%) also use the

dead cattle as food for prawn and catfish. Therefore, it is likely that

a reduction in vulture food supply has also taken place. In some of

the areas that were surveyed there were very few large trees suitable

for vultures to nest, but this scarcity did not appear critical in limiting

the breeding of vultures.

DISCUSSION

Previous reports on the status of the White-rumped Vulture mainly

quote the sizes of flocks that were regularly sighted at particular

roosting and nesting areas (Sarker 1983, 1987, Harvey 1990,

Thompson et al. 1993, Khan 2009, 2011), so it is difficult to

compare the present population or relative abundance with those

of the past. However, based on the scattered records mentioned

above it is certain that the present population, estimated to be around

816 birds, is considerably less than the population of 20 years ago.

Sarker (1987) arbitrarily estimated that the population density in

the Sundarbans was 0.03 to 0.91 individuals/km2, which is higher

than the inferred density today. It is not surprising that the lowest

Forktail 29 (2013) Population, breeding and threats to the White-rumped Vulture Gyps bengalensis in Bangladesh

55

relative abundance was estimated in the Chittagong division.

Mountfort & Poore (1968) had reported that vultures are common

in all areas except the Chittagong hills. This is probably because

there are fewer cattle in the hilly areas of Chittagong.

Areas like the Sundarbans interior and Aricha (Manikganj),

where vultures used to breed (Sarker 1983, 1987), were checked and

it was found that they are no longer resident there. Vultures were

reported to breed colonially (Sarker 1987, Ali & Ripley 1989,

Grimmett etal. 1 998), but during this survey it was found that they

nest either alone or there are just a few nests in an area, but not very

close together (except in one case). This is probably simply the result

of the severe decline in the population. Like the previous reports

(Sarker 1987, Ali & Ripley 1989, Grimmett etal. 1998), it was found

that the White-rumped Vulture traditionally uses trees for nesting

and the breeding peak is October to March. Even in the 1980s the

breeding success of vultures was reported to be low (Sarker 1983),

but this might be the result of a limited field survey.

The slowing of the rate of decline of the White-rumped Vulture

population appears to be the result of lower availability of diclofenac

in veterinary drugstores (Figure 2) following the ban. Similar results

were found in India (Cuthbert etal. 20 1 la,b). Following the decrease

in availability of diclofenac, however, the availability of other

NSAIDs, including ketoprofen that is also known to be poisonous

for Gyps vultures (Naidoo et al. 2009, Taggart et al. 2009), has

increased. Sodium salicylate is believed to be harmless, but it has

never been tested on vultures. It is now very well established that

diclofenac poisons and causes the death of Gyps vultures (Green et

al. 2004, Oakses al. 2004, Shultz etal. 2004, Cuthbert etal. 2011c),

so based on wide availability, even after the ban, it can be concluded

that veterinary diclofenac is the biggest threat to vultures in

Bangladesh. Although steroids and urea are widely used for cattle

fattening, these are not known to be poisonous to vultures (although

not tested thoroughly), and the fattened cattle are mainly consumed

by humans, so these are not major problems for vultures even if the

fattening doses have adverse effect on vultures.

The manufacturers of diclofenac for cattle have been against the

ban and have argued in different meetings that vultures might have

declined for other reasons such as food shortage. Our evidence shows

that illegal production (information gathered from some drugstores

suggested that small factories illegally produce diclofenac for

veterinary use), sale and use are still going on, and veterinary

diclofenac is still available despite the ban. The support of the media

is crucial to stop the use of veterinary diclofenac. It is important,

however, that the media employees are properly motivated and

informed so that they can convey the right message. Moreover,

proactive government support is needed to take legal action against

those who violate the ban on veterinary diclofenac.

During the field surveys few dead cattle were found and

interviewees have reported that dead cattle are often buried or used

as food in shrimp farms, suggesting that food shortage might also

be contributing to the decline of vultures. Food shortage as a cause

of vulture decline has also been pointed out by others (Round &

Chantrasmi 1985, Sarker 1987), and in Bangladesh wild mammal

carcasses are a very scarce food resource for vultures (Sarker 1987).

Even a decade ago, the human population density of the country

was much lower and there were many open areas and extensive

wetlands where the dead cattle could be disposed of in the open.

People should be encouraged to leave their dead cattle and dogs

(which have not been treated with diclofenac or ketoprofen) in the

open so that vultures can eat them.

Large trees suitable for nesting were rare in some areas and

disturbance of vultures by people was rarely reported, and probably

neither of these factors have made a significant contribution to the

rapid decline of vultures. In the Sundarbans the scarcity of nesting

trees was reported by Sarker (1987), but was probably exaggerated

since we found that the Sundarbans have many trees suitable for

vultures to nest. Other potential threats like poisoning by

insecticides (Sayer & Han 1983, Rahmani 1998, Ghatak 1999),

disease (Risebrough 2000, Prakash & Rahmani 2000) and nest

predation (Sarker 1987) might also contribute to the decline of

vultures in Bangladesh, but these could not be assessed and are

certainly less immediate threats than diclofenac. The existing nesting

trees should be preserved and, in areas where large trees are scarce,

some trees should be allowed to grow so that vultures and other

large birds can make nests.

Since vultures mainly occur outside designated protected areas,

i.e. areas not protected under the Bangladesh Wildlife Act, and often

outside natural forests and wetlands, thus, outside the jurisdiction

of government departments, and usually make their nest in large

trees on private properties, it is difficult to ensure their protection

and control human activities. The only way forward is through mass

motivation and awareness in the vulture hotspots, which we began

and succeeded in gaining some public support. We approached and

convinced a few key people in the locality and with their help we

reached the mass community. This approach was efficient and

successful, because these few key people in the locality are more

trusted by the vast majority living there than are outsiders such as

ourselves.

Both formal and informal awareness programmes were

conducted to educate people (especially children, cattle owners,

journalists and veterinarians) about the adverse effect of diclofenac,

and actions that can help vultures, such as not disturbing nesting

birds. Awareness programmes included popular lectures, interactive

discussions, quizzes, drawing contests for children and posters and

other material were distributed. An informative book with

photographs of vultures has also been published (Khan 2012).

In response to our (together with other bird-lovers and

concerned people) repeated appeals in the media (television and

newspapers) and meetings, the government banned veterinary

diclofenac formulations, although there is the risk that human

formulations might be used instead, as in India (Cuthbert et al.

2011b). More challenging, however, will be the full implementation

of this ban, because more than 50% of the veterinary drug stores

still illegally sell diclofenac, which probably has been produced

illegally in Bangladesh by small-scale manufacturers. The interviews

that were given on local television channels and in newspapers have

helped bring about the reduction in the use of diclofenac.

Monitoring of vulture numbers and breeding, at least in vulture

hotspots, should be continued so that the population trends are

known. The identified hotspots should be declared vulture

sanctuaries or more precisely, following the guidelines of Saving Asia’s

Vultures from Extinction (SAVE 2012), as Vulture Conservation

Areas, in order to attain Vulture Safe Zone status in the future.

Special measures should be taken for collaborative management

involving local communities. Moreover, regular monitoring should

take place on the use of different cattle medicines, including the

illegal use of diclofenac and use of the safe alternative meloxicam.

Proactive measures by the government authorities will be required

to ensure that diclofenac and any other similar drugs that are likely

to be dangerous to the vultures are removed from veterinary practice.

We also need to work on developing captive breeding capacity of

vultures at Dhaka Zoo, where a small captive population already

exists and one pair often breeds successfully without any special

assistance. Wider involvement of institutions and individuals is

urgently needed to take up actions to conserve vultures. A national

committee should be formed in order to draw up a Bangladesh

Vulture Action Plan, and coordinate and advise on activities related

to vulture conservation and management in Bangladesh. This

committee can function as the coordinating body, guiding the

relevant governmental and non-governmental activities. Moreover,

regional initiatives should be taken so that vulture populations can

be monitored and conserved across the region. Thankfully, the

56

M. MONIRUL H. KHAN

Forktail 29 (2013)

process has already been started and the governments of Bangladesh,

India, Nepal and Pakistan signed the Regional Declaration on the

Conservation of South Asia’s Critically Endangered Vulture Species

in May 2012.

ACKNOWLEDGEMENTS

The author is grateful to Conservation Leadership Programme (CLP) of

BirdLife International, Conservation International, Fauna & Flora International

and Wildlife Conservation Society; and the Forktail Zeiss Award of the Oriental

Bird Club for providing financial support to conduct the study of the White -

rumped Vulture. Chris Bowden (Royal Society for the Protection of Birds),

Nick Lindsey (Zoological Society of London), David Chivers (University of

Cambridge) and Pamela Rasmussen (Michigan State University) kindly

provided technical support to this work. Richard Cuthbert and Chris Bowden

of RSPB kindly reviewed the manuscript and made significant improvements.

Sincere thanks to all research students (Ashikur Rahman, Bayezid Khan, Shibli

Sadik and Mohammad Moniruzzaman) of the Department of Zoology,

Jahangirnagar University, who were involved in all the field activities. Special

thanks to Munir Khan and Tania Khan for supporting the fieldwork. Thanks

also to the local people and journalists of print and electronic media for help to

raise public awareness for vulture conservation.

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University, Savar, Dhaka 1342, Bangladesh. Email:

mmhkhan@hotmail.com

FORKTAIL 29 (2013): 57-63

The birds of the Longbao National Nature Reserve

and surrounding basin, Yushu county, Qinghai, China

JOHN D. FARRINGTON, ZHANG XIULEI & ZHANG MIN

The Longbao National Nature Reserve lies at an elevation of 4,200 m on the Tibetan Plateau of southern Qinghai province, China. The

reserve was established in 1986 as Qinghai's second national nature reserve and is the third most important known breeding ground of the

Black-necked Crane Grus nigricollis. From October 2010 to July 2012, the authors conducted 18 complete and several part circuits of the

main Longbao wetland at various times of year from early April to mid-November, the main residence period of migratory birds at the

wetland. During these circuits, counts of all cranes sighted on the main wetland were made, as were extensive counts of other waterbird

and non-waterbird species. In addition, avifauna on the Longbao wetland and in the surrounding basin was observed extensively on non¬

survey circuit days. In total, 67 bird species were recorded in the Longbao basin over the 21 -month survey period, including 39 species with

no known earlier records, extending the list of birds recorded at Longbao to 71. Direct threats to breeding waterbirds at Longbao include

loose and feral herding dogs and recently erected powerlines that, at present, lack markers to increase their visibility to large passing birds.

Indirect threats to waterfowl breeding on the Longbao wetland include increased livestock grazing pressure on the more resilient pastures

of the main wetland, as pasture conditions on the hill slopes surrounding the wetland rapidly deteriorate, and climate change, which is

believed to be causing the large-scale drying up of shallow wetlands in the region, primarily as a result of permafrost degradation and

increased evaporation. The sum total of peak counts of individual waterbird species at Longbao in 201 1 was 1 1,266. Notably, the Black¬

necked Crane population summering at the reserve roughly quadrupled between 1 984 and 201 1 . Evidence is presented that the Longbao

National Nature Reserve qualifies for Ramsar status under Ramsar Criteria 2 and 6.

INTRODUCTION

The Longbao National Nature Reserve (NNR) (33.183°N

96.583°E) is located on the Tibetan Plateau in southern Qinghai

province, about 50 km north-west of the town of Jiegu in Yushu

county, beside the highway connecting Yushu and Zhiduo counties.

The Longbao reserve was established as Qinghai province’s second

NNR in 1986 and is the third most important known breeding

ground of the Black-necked Crane Grus nigricollis after the

Ruoergai Wetlands in northern Sichuan and the remote Seling Co

Lake Region of the central Tibet autonomous region (Bishop

1996). The reserve covers roughly 100 km2 comprising the boggy

bottom of a long, fairly broad mountain basin, with a maximum

length of 30 km and maximum width of 5 km. The reserve’s primary

ecological importance is its main wetland, which forms the eastern

half of the reserve and is approximately 14 km long with a maximum

width of about 3.5 km and is located at about 4,200 m. The entire

nature reserve is flanked by steep ridges that typically rise up to

750 m above the valley floor. While lying outside the boundaries

of the Longbao NNR, these surrounding mountains are nominally

protected as part of the administratively distinct Sanjiangyuan

NNR that protects the vast Yellow, Yangtze and Mekong river

source regions of Qinghai province.

The main Longbao wetland is fed by groundwater, streams,

precipitation and snow melt, and is frozen over from about late

November to early March. The entire Longbao basin drains into

the Yi Chu River, located north-west of the main wetland, a short

tributary of the nearby Tongtian (upper Yangtze) River.

Ecosystems in the reserve are primarily a mix of wet and dry

Kobresial Carex sedge meadows, but also include aquatic ecosystems

such as lake, pond, marsh and riparian corridor-type ecosystems

(Li & Zhou 1985, Li & Li 2005). Some limited willow Salix

shrublands occur on hill slopes along the south-west of the main

wetland. Mammals frequently seen on the wetland during this

survey included the Tibetan Fox Vulpes ferrilata. Red Fox V. vulpes ,

Altai Weasel Mustela altaica, Himalayan Marmot Marmota

himalayana , Woolly Hare Lepus oiostolus , Black-lipped Pika

Ochotona curzoniae and a vole Microtus. Grey Wolf Canus lupus,

Tibetan Gazelle Procapra picticaudata , White-lipped Deer Cervus

albirostris. Blue Sheep Pseudois nayaur and Snow Leopard Uncia

uncia inhabit the mountains ringing the Longbao basin. Eurasian

Otter Lutra lutra and various rodents additional to the foregoing

have also been recorded at Longbao (Li & Zhou 1985).

Human presence in the Longbao NNR includes about 1 50 yak-

herding households residing on or within 1 km of the perimeter of

the main wetland, who all rely heavily on the wetland’s pasture for

much of the year. Most of the wetland has been divided into

individual household pasture allotments, and since 1997 these

single-family allotments have been fenced off, although some tracts

of unfenced wetland pasture used collectively by groups of families

remain. These households own an average of about 40 yaks each

with a few households owning small numbers of sheep, goats and

horses. In addition, a large village lies at the eastern end of the

reserve, just north of the highway to Yushu, while Longbao, the

local administrative town, is located on the highway at the western

end of the reserve.

Because of its easy access, narrow geography and large intact

wetland, the Longbao NNR is the premier site in Yushu prefecture

for observing waterfowl, with the first geese and ducks arriving in

early March and the last departing in mid to late November.

H owever, until the present study, no in depth research on

Longbao’s avifauna had been conducted throughout the eight-

month annual residence of migratory birds. Earlier knowledge of

bird life at Longbao is based largely on Black-necked Crane breeding

studies conducted during the 1980s by Li & Zhou (1985) and Li

& Ma (1989), with the former having compiled a list of 31 bird

species in spring and summer 1 984; a single additional species was

recorded by Li & Ma in summer 1988 (Table 1). Lin (2003) stated

that there are 30 species of birds in the reserve, but gave no species

list.

The avifauna survey described here took place over parts of the

annual occurrence of migratory waterfowl in 2010, 201 1 and 20 12.

The impetus behind this survey was to improve protection of cranes

in the reserve and to determine if it qualified for designation as a

Ramsar Wetland of International Importance based on its

waterbirds. All bird species seen during the surveys were recorded,

with detailed counts of all species observed on the main wetland in

2011.

58

JOHN D. FARRINGTON, ZHANG XIULEI & ZHANG MIN

Forktail 29 (2013)

METHODS

A preliminary survey on 28 October 2010 consisted of a cursory

‘binoculars only’ practice count of Black-necked Cranes on the main

Longbao wetland. The wetland was circled clockwise by car; 118

Black-necked Cranes were counted, with eight other bird species

recorded. This practice count established the basic survey

methodology for the 2011 survey, which again consisted of driving

clockwise around the entire perimeter (about 35 km) of the main

wetland. This circuit started from the reserve headquarters and

generally involved counting all birds seen at each of 22 set survey

points established at good viewing locations giving full coverage of

the wetland.

Between 6 April and 1 6 November 20 1 1 , a rotating team of three

counters made 15 full and two part circuits, using spotting scopes

and binoculars (Appendices 1 & 2). No count was made during

August. In general, the entire circuit was counted in one day although

one circuit was divided between two days on 12 and 13 May

(Appendices 1 & 2). In addition to the formal survey circuits,

supplementary observations were made during short walks along

the wetland and in the surrounding mountains. Birds were identified

using MacKinnon & Phillipps (2000) and Grimmett et al. (1999).

RESULTS

In total, 67 bird species were recorded in the Longbao basin over

the nearly 21 -month survey period, including 39 species with no

known earlier records there. This expanded the list of birds

recorded there from 32 species to 71 (Table 1); other unpublished

records may exist. The counts during 2011 indicate how waterbird

populations change on the main wetland from spring to autumn

(Appendix 1). These were separated from counts made of non-

waterbirds in order to determine if Longbao NNR qualifies for

Ramsar designation based on Ramsar Criterion 5 (Appendix 2).

Several bird species seen in 2011 do not appear in the bird count

tables because they were not seen during counts. In 2012, full

circuits of the wetland were made on 20 April and 30 June. Full

counts were only made of Black-necked Cranes, although other

bird species sighted were noted; counts were as follows: on 20 April

- adult Black-necked Crane 178, Common Crane 7, on 30 June -

adult Black-necked Crane 128, Black-necked Crane chicks 21,

Common Crane 6, Cattle Egret 130.

Notes on species of interest at Longbao with respect to

conservation and/or changing status follow.

Table 1 . List of bird species recorded at the Longbao National Nature Reserve, 1 984-201 2.

Seasonality of occurrence: S V = Summer Visitor, PM = Passage Migrant, R = Resident (including altitudinal migrants).

Abundance categories: A = Abundant: regular in very large numbers, C = Common: regular in large numbers, F = Fairly common: regular in

moderate numbers, U - Uncommon: somewhat regular in small numbers, R = Rare: occasional in very small numbers.

'First record by Li & Zhou (1985), 1984. "Only record by Li & Zhou (1985), 1984. ’First record by Li & Ma (1989), 1988.

Forktail 29 (2013) Birds of Longbao National Nature Reserve and surrounding basin, Yushu county, Qinghai, China

59

Bar-headed Goose Anser indicus

The common goose of the Tibetan Plateau, this is by far the most

numerous species at Longbao, with a high count of 8,282 at the

height of the spring migration on 5 May 2011. Numbers quickly

declined: much lower numbers, both breeding and non-breeding,

summered (Appendix 1). On 16 November 2011, only three

remained on the main wetland; these presumably left during the

third week of November. In 2012, the first returning Bar-headed

Goose was recorded by the reserve staff on 4 March.

Ruddy Shelduck Tadorna ferruginea

The common duck of the Tibetan Plateau, this is the second most

numerous bird species at Longbao. A high count of 1,560 on 5

May 20 1 1 declined precipitously the following week. Much lower

numbers remained to breed (Appendix l).On 16 November 2011,

1 10 remained, when the main wetland was about 90% frozen over;

these presumably had departed by the end of November. In 2012,

the first returning Ruddy Shelduck was recorded by the reserve

staff on 3 March.

Ferruginous Pochard Aythya nyroca

Near Threatened (BirdLife International 2013a). Nevertheless, it

is locally common at Longbao, with 257 recorded during spring

migration on 5 May 2011, 392 at the height of autumn migration

on 6 November 2011, and smaller numbers believed to breed

(Appendix 1).

Black-necked Crane Grus nigricollis

Vulnerable (BirdLife International 2013a) because of its single

small population that may be in decline through loss and

degradation of wetlands and changing agricultural practices in its

breeding and wintering grounds (Bishop et al. 2000, Yan & Wu

2005, Wang et al. 2006, Farrington 2009, Ma et al. 2009). In July

1984, the peak count of adult and returning young Black-necked

Cranes on the Longbao wetland was 24 (Li & Zhou 1985).

Following 25 years of concerted protection at Longbao and

throughout its range, a historical high count of 216 was made at

Longbao on 25 April 2011 (Appendix 1; also Farrington & Zhang

2013). These numbers more than halved by the beginning of

summer. On the main Longbao wetland in 20 1 1 , 29 pairs produced

43 offspring, as counted on 28 July (Appendix 1).

Black-tailed Godwit Limosa limosa

Near Threatened (BirdLife International 2013a). This species was

recorded just once at Longbao: on 5 May 201 1 a group of 80 was

seen on a stopover.

Cinereous Vulture Aegypius monachus

Near Threatened (BirdLife International 2013a). Although

apparently resident in very small numbers in eastern Yushu

prefecture, the only record on this survey was of one individual at

Longbao on 17 November 201 1, feeding on carrion next to the

highway at the eastern end of the reserve with a small group of

Himalayan Griffons Gyps himalayensis.

Pallas's Fish Eagle Haliaeetus leucoryphus

Vulnerable (BirdLife International 2013a). The only known record

at Longbao is a sighting by Li & Zhou (1985) in 1984. JDF’s only

sighting on the Tibetan Plateau was on 1 8 May 20 1 2: one individual

near the mouth of the Yike W ulan Chu River on the northern shore

of Qinghai Lake. Given this sighting north of Longbao, the

occasional Pallas’s Fish Eagle may well stop over at the Longbao

wetland while on migration (BirdLife International 2013b).

Saker Falcon Falco cherrug

Endangered (BirdLife International 2013a). The most common

falcon of the Longbao basin and on the grasslands of Yushu

prefecture during this survey, it was regularly sighted in small

numbers in the Longbao NNR and in the surrounding mountains

where it is known to breed (Appendix 2).

Little Egret Egretta garzetta

Although very conspicuous, Little Egret was not noted by Li &

Zhou (1985) in 1984. More commonly associated with low-lying

areas of southern China, South Asia, South-East and East Asia and

elsewhere, the numbers of Little Egrets summering on the Tibetan

Plateau may be increasing. In 201 1, it was sighted on the Longbao

wetland four times between 12 May and 4 July with the highest

count being two on 27 May (Appendix 1). A group of 11 were

sighted next to the highway beside the outflow marsh at the western

end of the wetland on 30 June 2012, and four were seen nearby on

11 July 2012.

JDF’s first sighting of Little Egret on the Tibetan Plateau was

on 11 July 2009: eight in Lhasa’s Lhalu wetland. These birds

apparently summered there; JDF’s last sighting (of five) on the

Lhalu that year was on 17 October. An exhaustive search of

English and Chinese literature revealed no earlier records for the

Lhasa area, and no mention of Little Egret on the Tibetan Plateau

is made in the comprehensive works of Vaurie (1972) and Zheng

etal. (1983).

Cattle Egret Bubulcus ibis

Cattle Egret is another very conspicuous species not noted by Li &

Zhou (1985) in 1984. In 2011, it was regularly seen at Longbao

from 5 May until 6 November, with the highest count being 106

on 4 July 201 1 (Appendix 1). On 30 June 2012, 130 were counted

on the Longbao wetland, corroborating the opinion of reserve staff

that the species has increased significantly in recent years.

Black Stork Ciconia nigra

The first Black Stork record at the Longbao wetland was made by

Li & Ma (1989). About 30 stopped over in the relatively drier

eastern end of the reserve for about a month in spring 1988. Eight

were seen on 4 May 1988 being driven away from a Black-necked

Crane nesting site by the male crane (Li & Ma 1989). Since that

time, Black Stork numbers at Longbao have declined dramatically,

and none was seen in 20 1 1. The species was in decline in China as

early as 2000 (MacKinnon & Phillipps 2000). Prior to 2012, the

last known sighting at Longbao was in June 2008: one individual

seen by reserve staff just east of the reserve headquarters. On 15

June 2012, four individuals were seen just below the highway on

the north side of the main wetland about 3 km north-west of the

Longbao Monastery at the eastern end of the reserve. Five were

sighted there on 27 June 20 12. JDF’s only other sighting in Qinghai

province was on 19 November 2011: a pair in southbound flight

over a forested ridge at about 2,900 m near the town of Dongxia in

Datong county, about 45 km north of the provincial capital,

Xining.

Black Drongo Dicrurus macrocercus

In eastern Qinghai, Black Drongo is presumably a rare spring

passage migrant and possibly a summer resident. In the Longbao

basin, it was sighted once: on 3 June 2011 next to the reserve

headquarters. Elsewhere in eastern Qinghai, JDF has seen a single

bird on two other occasions in June 2011 and July 2012.

Hodgson's Bushchat Saxicola insignis

V ulnerable (BirdLife International 20 1 3a) . A lone male was sighted

on the south side of the Longbao NNR on 25 April 2011,

presumably while stopping over on migration from the plains of

the northern Indian subcontinent to its main breeding grounds in

western Mongolia.

60

JOHN D. FARRINGTON, ZHANG XIULEI & ZHANG MIN

Forktail 29 (2013)

DISCUSSION

During the survey, 67 bird species were recorded in the Longbao

wetland basin. With the four species recorded by Li & Zhou (1985)

not sighted on the current survey (Pallas’s Fish Eagle, Hodgson’s

Redstart Phoenicurus hodosoni , Brandt’s Mountain Finch Leucosticte

o

brandti and Great Rosefinch Carpodacus rubicilla),7\ bird species

have been formally recorded at Longbao. These include three Near

Threatened species — Ferruginous Pochard, Black-tailed Godwit

and Cinereous Vulture; three Vulnerable species — Black-necked

Crane, Pallas’s Fish Eagle and Hodgson’s Bushchat — and one

Endangered species — Saker Falcon (Birdlife International 2013a).

Categorising the seasonal occurrence of birds of the Longbao basin

is obviously fraught with uncertainty in the absence of winter

observations; tentative classifications based on current observations

in the basin itself, not the broader eastern Tibetan Plateau region,

are provided in Table 1. This suggests that 24 species are resident

(including altitudinal migrants), 21 are summer visitors, and 26 are

passage migrants; many individuals of several species that are

summer visitors simply use the basin as a migration stopover site.

Formerly the largest direct threat to local waterfowl was the

widespread collection of eggs, particularly from Bar-headed Goose,

Ruddy Shelduck and Black-necked Crane, by both locals and outsiders

for personal consumption. Such collection was largely stopped

following the establishment of the Longbao NNR in 1986 and the

subsequent enforcement of regulations banning the practice. Loose

and feral Tibetan mastiff herding dogs were regularly seen on the

wetland during the survey, with six counted on 23 October 2011.

These are a threat primarily to eggs, unfledged chicks and injured birds.

A new threat to large waterfowl at Longbao is the erection (in late

2011) of high-voltage powerlines along the entire north side of the

wetland. Powerline strikes by Black-necked Cranes were documented

by Li et al. (2011) although this hazard can be mitigated easily, by

installing coloured markers on the cables (Li 2002, Li et al. 2011).

The primary threat to habitat on the Longbao wetland is

degradation through grazing damage and climate change. In recent

decades, upland summer pastures on hill slopes surrounding the main

wetland have been severely degraded. This probably results from

overgrazing, consequent to overstocking in the basin during the

collective period from the 1960s to the 1980s. This degradation is

believed to have been exacerbated by recent climate change, such as

permafrost degradation, which can result in reduced seep areas and

lower soil moisture, and the recent intensification of precipitation in

the region, which accelerates erosion of degraded slopes (Wang et al.

2006, Farrington 2009, Qiu 2012). Consequently, household pastures

on the more resilient Longbao wetland are now grazed intensively

for up to 10-12 months per year by thousands of domestic yaks and

smaller numbers of sheep, goats and horses. Notably, livestock

numbers on the wetland peak in May when the first shoots of green

grass in the basin appear at precisely the same time as waterfowl begin

nesting. During this survey, a high count of 5,357 yaks was made on

the main Longbao wetland on 1 1 May 2011.

Pasture conditions on the wetland remain fairly good, yet all

local herders interviewed indicated that grass height and pasture

productivity have declined in recent years. Presumably, disturbance

to nesting birds has increased with increased grazing pressure on

the wetland (Li & Ma 1989). Rising temperatures and permafrost

degradation on the Tibetan Plateau has led to widespread drying

up of shallow, permafrost-controlled wetlands in the Longbao

region and elsewhere on the plateau. As temperatures rise, the upper

permafrost melts, allowing surface water to percolate into the

ground (Wang et al. 2006, Farrington 2009, Ma et al. 2009, Qiu

2012, Farrington & Zhang 2013). Ultimately such drying of

shallow wetlands must negatively affect plateau waterbirds.

A main objective of this survey was to determine if the Longbao

NNR qualified for Ramsar Wetland of International Importance

designation based on its waterbird populations. Under Ramsar

Criterion 5, a wetland is considered internationally important if it

regularly supports at least 20,000 waterbirds (Ramsar 2009). In

order to see if Longbao met this criterion, the highest counts of

each waterbird species in 201 1 were totalled, regardless of when

the peaks occurred. The 2011 total of 32 waterbird species came

to 1 1,266, far short of meeting Criterion 5.

However, under Ramsar Criterion 6, a wetland is considered

internationally important if it regularly supports 1% of the

individuals in a population of one species of waterbird (Ramsar

2009). The most recent estimate of the total Black-necked Crane

population is 1 1,000 (Bishop & Drolma 2007). The average total

of 1 32 Black-necked Cranes counted over 20 survey circuits during

2010-2012 therefore qualifies the Longbao wetland for Ramsar

designation under Criterion 6 (Farrington & Zhang 20 13). Black¬

necked Crane is listed as Vulnerable on the IL1CN Red List, so

Longbao wetland also qualifies for Ramsar designation under

Criterion 2, whereby a wetland is considered internationally

important if it supports Vulnerable, Endangered or Critically

Endangered species (Ramsar 2009).

In addition to qualifying for Ramsar designation based on its

Black-necked Crane population, according to 2012 estimates of

waterbird populations compiled by Wetlands International (20 1 3),

the Longbao wetland also qualifies for Ramsar designation under

Criterion 6 based on its Bar-headed Goose, Ruddy Shelduck and

Black Stork populations (Table 2).

Notably absent during this survey were two bird species seen

by JDF along rivers elsewhere in Yushu county but not in the

Longbao basin itself, Ibisbill Ibidorhyncha struthersii and Red-

rurnped Swallow Hirundo daurica , as well as a number of waterbird

species commonly sighted throughout much of the Tibetan

Plateau, including Indian Spot-billed Duck Anas poecilorhyncha.

Common Teal Anas crecca and Red-crested Pochard Netta rufina.

Inevitably other bird species will be recorded at Longbao by future

observers. Finally, this survey should provide a baseline for gauging

future changes in the avifauna of the Longbao wetland, especially

given that this remarkable, yet fragile, high-altitude wetland will

continue to undergo profound ecological shifts resulting from

climate change in years to come.

Table 2. Summary of 1% thresholds for Ramsar Criterion 6 qualification

for four species of waterbirds at the Longbao wetland and the high

counts of these species made in 201 1-2012.

ACKNOWLEDGEMENTS

This study was funded by the WWF Network and was only made possible

through the generous support of the Qinghai Forestry Department, the

leadership of which has shown remarkable foresight in their commendable

efforts to protect Qinghai Province’s many unique and ecologically and socio¬

economically important high altitude wetland areas.

REFERENCES

BirdLife International (2013a) IUCN Red List for birds. Downloaded from

http://www.birdlife.org on 07/05/2013.

Forktail 29 (2013) Birds of Longbao National Nature Reserve and surrounding basin, Yushu county, Qinghai, China

61

Bi rd Life International (2013b) Species factsheet Pallas's Fish Eagle:

Haliaeetus leucoryphus. Downloaded from http://www.birdlife.org on

07/05/2013.

Bishop, M. A. (1996) Black-necked Crane ( Grus nigricollis). In C. D. Meine

and G. W. Archibald, eds. The cranes: status survey and conservation

action plan. Gland & Cambridge UK: IUCN/SSC Crane Specialist Group.

Online version: Jamestown: USGS Northern Prairie Wildlife Research

Center, March 2, 1998. http://www.npwrc.usgs.gov/resource/birds/

cranes/index. htm; downloaded on 21 August 2012.

Bishop, M. A. & Drolma, T. (2007) Tibet Autonomous Region January 2007

survey for Black-necked Crane, Common Crane, and Bar-headed Goose.

China Crane News 1 1 : 23-25.

Bishop, M. A., Harris, J. & Canjue, Z. (2000) Agricultural management zones

for Bar-headed Geese and Black-necked Cranes in Tibet. Pp.55-60 in

N.Wu, D. Miller, Z. Lu & J. Springer, eds. Tibet's biodiversity conservation

and management. Beijing: China Forestry Publishing House.

Farrington, J. D., ed. (2009) Impacts of climate change on the Yangtze source

region and adjacent areas. Beijing: China Meteorological Press.

Farrington, J. D. & Zhang X. L. (2013) The Black-necked Cranes of the

Longbao National Nature Reserve, Qinghai, China: current status and

conservation issues. Mountain Research and Development 33(3): 305-

313.

Grimmett, R., Inskipp, C. & Inskipp, T. (1999) Pocket guide to the birds of the

Indian subcontinent. New Delhi: Oxford University Press.

Li, D. H. & Zhou, Z. J. (1985) Behaviour of the Black-necked Crane at

Longbaotan during the breeding period. Chinese Wildlife 6: 4-9. (In

Chinese.)

Li, F. S. (2002) Two Black-necked Cranes die from power line strike at Caohai

Nature Reserve. China Crane News 6: 24-25.

Li, F. S., Bishop, M. A. & Drolma, T. (201 1 ) Power line strikes by Black-necked

Cranes and Bar-headed Geese in Tibet Autonomous Region. Chinese

Birds 2(4): 167-173.

Li, F. S. & Ma, J. Z. (1989) A study on the Black-necked Crane's behavior in

incubation period at Longboatan, China. Baraboo: International Crane

Foundation.

Li, Z. M. & Li, F. S. (2005) Research on the Black-necked Crane. Shanghai:

Shanghai Science, Technology, and Education Press. (In Chinese with

English abstract.)

Lin, J. C. (2003) Biological diversity conservation in Yushu Longbao National

Nature Reserve of Qinghai Province. Jilin Forestry Science and Technology

32(14): 37-40. (In Chinese with English abstract.)

Ma, Z. T., Li, F. X., Li, F. & Xiao J. S. (2009) The dynamic change research of

ecological environment in Longbao region of Yushu state in Qinghai.

Pratacultural Science 26(7): 6-1 1 . (In Chinese with English abstract.)

MacKinnon, J. & Phillipps, K. (2000) A field guide to the birds of China. Oxford:

Oxford University Press.

Qiu, J. (201 2) Thawing permafrost reduces river runoff: China's Yangtze River

is receiving less water as climate warms. Nature News (6 January 2012)

doi:10. 1038/nature. 2012.9749 Downloaded on 7 March 2012.

Ramsar (2009) Information Sheet on Ramsar Wetlands (RIS) 2009-2012

version. Gland: Ramsar Secretariat.

Vaurie, C. (1972) Tibet and its birds. London: H.F. & G. Witherby.

Wang, G. X., Li, Y. S., Wu, Q. B. & Wang, Y. B. (2006) Impacts of permafrost

changes on alpine ecosystem in Qinghai-Tibet Plateau. Science in China,

Series D: Earth Sciences 49: 1 1 56-1 1 69.

Wetlands International (201 3) Waterbird population estimates.

Wageningen: Wetlands International, http://wpe.wetlands.org.

Downloaded on 7 May 2013.

Yan, Z. & Wu, N. (2005) Rangeland privatisation and its impacts on the Zoige

Wetlands on the eastern Tibetan Plateau. Journal of Mountain Science

2: 105-115.

Zheng, Z. X., Li, D. H„ Wang, Z. X., Wang, Z. Y„ Jiang, Z. H. & Lu, T. C. (1983)

The avifauna of Xizang. Beijing: Science Press. (In Chinese.)

John D. FARRINGTON, World Wildlife Fund, P.O. Box 210,

Kawojangso, Thimpu, Bhutan. Email: doeage@gmail.com

ZHANG Xiulei, Community Development Consultant to WWF

& ZHANG Min, Wildlife Photography Consultant to WWF, 51

Bayi Lu, Apt. 401, Xining, Qinghai 810007, China.

62

JOHN D. FARRINGTON, ZHANG XIULEI & ZHANG MIN

Forktail 29(2013)

Appendix 1. 201 1 counts of waterbird species at the Longbao wetland.

Highest count for each species is shown in bold. NC= No count made.

1 Due to time restrictions, only counts of Black-necked Cranes and Great Crested Grebes were made.

2 Count made from 21 of 22 survey points only.

3 Count made from the 1 1 survey points along the highway on the north side of the wetland only due to deep snow covering the jeep track on the south side.

Forktail 29 (2013) Birds of Longbao National Nature Reserve and surrounding basin, Yushu county, Qinghai, China

63

Appendix 2. 2011 counts of non-waterbird species at the Longbao Wetland.

Highest count for each species is shown in bold. NC= No count made.

' Due to time restrictions, only Black-necked Cranes and Great Crested Grebes were counted.

2 Count made from 21 of 22 survey points only.

5 Count made from the 11 survey points on the north side of the wetland only due to deep snow on the south side.

FORKTAIL 29 (2013): 64-70

Habitat specialisation in the Reed Parrotbill Paradoxornis

heudei — evidence from its distribution and habitat use

LI-HU XIONG &JIAN-JIAN LU

The Reed Parrotbill Paradoxornis heudei is found in habitats dominated by Common Reed Phragmites australis in East Asia. This project was

designed to test whether the Reed Parrotbill is a specialist of reed-dominated habitats, using data collected through literature review and

field observations. About 87% of academic publications describing Reed Parrotbill habitat report an association with reeds, and the species

was recorded in reeds at 92% of sites where it occurred. On Chongming Island, birds were only recorded in transects covered with reeds or

transects with scattered reeds close to large reedbeds. At the Chongxi Wetland Research Centre, monthly monitoring over three years also

showed that the species was not recorded in habitats without reeds. The density of Reed Parrotbills was higher in reedbeds than mixed

vegetation (reeds with planted trees) and small patches of reeds. The species rarely appeared in mixed habitat after reeds disappeared.

These results confirm that the species is a reed specialist and highlights that conservation of reed-dominated habitat is a precondition to

conserve the Reed Parrotbill.

INTRODUCTION

Habitat specialisation results in some species having a close

relationship with only a few habitat types (Futuyma & Moreno

1988), and habitat specialists have some specific life-history

characteristics, for example, they often have weak dispersal abilities

(Krauss et al. 2003) making them sensitive to disturbance or

fragmentation (Sol et al. 2002, Krauss etal. 2003, Cofre etal. 2007).

Therefore, knowing if a species is a habitat specialist is important

for predicting population distribution, understanding the

relationship between life-history characteristics and habitat, and

providing a basis for a species protection strategy (Futuyma &

Moreno 1988, Julliard etal. 2006).

Reed Parrotbill Paradoxornis heudei is an insectivorous,

resident species restricted to east China, east Mongolia and south¬

east Russia (MacKinnon & Phillipps 2000, Lei & Lu 2006, BirdLife

International 2013). Two subspecies are recognised, P. h. heudei

and P. h.polivanovi ; subspecies heudei is mainly found in the middle

and lower reaches of Changjiang and polivanovi is mainly found in

north-east China (Lei & Lu 2006). It has been assumed that Reed

Parrotbill’s reedbed specialisation is responsible for its narrow

distributional area and small population (BirdLife International

2013). This is partly because reedbeds in East Asia are threatened

by commercial harvesting, wetland reclamation and sewage

discharges, which have resulted in declines in the quality and area

of beds, potentially threatening the Reed Parrotbill, and the species

is classified as Near Threatened (BirdLife International 2013).

Therefore it is important to understand the relationship between

the Reed Parrotbill and the Common Reed Phragmites australis

to understand species’s life history characteristics and advance its

conservation.

Most papers covering the species detail new distributional

records and descriptions of breeding biology (Wang& Tian 1988,

Bai & Bai 1993, Wang et al. 2011) and a few describe its ecology

(Ma 1988, Xiong et al. 2007). Studies in a tidal reedbed in the

Changjiang estuary found that Reed Parrotbills fed on insects on

reeds all year round (Xiong etal. 2007) and that reed shoots made

up more than 89% of the nest mass (Xiong & Lu 2013). In some

papers, the Reed Parrotbill has been declared to be a reed-

dominated habitat specialist (Xiong et al. 2007, Gan et al. 2009).

However, habitat specialisation of the species has not been critically

tested. In this paper, published and unpublished information on

the distribution and habitat use of the Reed Parrotbill in China is

reviewed, and two new field studies which test whether it is a reed-

dominated habitat specialist are reported and discussed.

METHODS

Three sets of information on Reed Parrotbill distribution and

habitat use were used: (1) distribution and habitat use data in the

Chinese part of its range, collated from academic publications, web

news, communication with birdwatchers and personal

observations, (2) the distribution of the species obtained through

transect observations on Chongming Island (Figure 1), one of its

main strongholds, (3) three years monitoring Reed Parrotbill

distribution in reed-dominated habitats (reedbeds, smaller reed

patches, reeds with dense trees and reeds with sparse trees) and

neighbouring habitats without reeds at Chongxi Wetland Research

Centre (31.700°N 121.200°E), located on the west end of

Chongming Island (Figure 1).

Figure 1. Location of Chongxi Wetland Research Centre, transects with

and without Reed Parrotbills in Chongming island and the location of

Chongming island in China.

Distribution and habitat use of the Reed Parrotbill in China

Online databases were searched — China Academic Journal

(CNKI), ISI Web of Science, Biological Abstracts and BIOSIS

Previews. The references in papers already retrieved and previous

review articles about the Reed Parrotbill were checked. The internet

was searched, birdwatchers contacted and personal records were

also included for new distributional records of the species (Table

1). The final online search was conducted in August 2012. The

following information was collated: locations where Reed

Forktail 29 (2013)

Habitat specialisation in the Reed Parrotbill Porodoxornis heudei

65

Parrotbills have been recorded (province and sites or region), dates

of records and descriptions of habitat use, paying special attention

to whether this included reed vegetation.

All sites in China where Reed Parrotbills have been recorded

were mapped, distinguishing sites where they were only recorded

before 1980 from sites where they only occurred after 1980 and

sites where they were present both before and after 1980. Records

of habitat use by the species were analysed, using reports in academic

publications and sites where Reed Parrotbills were recorded. As

there are few records before 1980, habitat use analysis was focused

on those after 1980. The proportion of publications with

descriptions of vegetation type and the proportion of publications

indicating that Reed Parrotbill used reed vegetation were

determined, as was the percentage of recording sites for which

descriptions of vegetation were available and the percentage of

recording sites where reed vegetation was present. Published habitat

use reports and site records repeated in edited books, review articles

and bird lists (Cheng 1987, Zheng 2005, Lei & Lu 2006) were not

duplicated in these analyses.

Distribution of Reed Parrotbills at Chongming Island

Chongming Island, about 1,200 km2 in area, the largest alluvial

island in the world, is located in the Changjiang estuary (Figure 1).

The intertidal wetland and the adjacent newly reclaimed area on

the island includes about 4,590 ha of reed vegetation (Huang etal.

2005). The island was divided into 10 km2 grid-squares with one

transect located in each square and a total of 118 transects (excluding

some which were inaccessible) were visited once in August 2005

(Figure 1). Each transect covered 10 ha, and they varied in length

from 1 to 2.5 km and from 40 m to 1 00 m in width. The number of

Reed Parrotbills and the vegetation they were using were recorded.

Habitat selection of Reed Parrotbills at Chongxi Wetland

Research Centre

The Chongxi Wetland Research Centre supports a 3 km2 tidal

marsh dominated by monospecific stands of Common Reed

(reedbeds) along the estuary. There were also two types of forested

wetlands by the dyke along the estuary: (1) reeds with dense trees

mostly Salix matsudana, planted in winter 2003; during 2004 and

2005, the trees and Common Reeds grew together, but by 2006

the trees were much taller than the reeds, which died back during

2007, and (2) reeds with sparse trees, planted in winter 2005. These

trees were located among reedbeds as patches, enabling trees and

reeds to coexist (Liu et al. 2009). There were also patches of

Common Reed near the main reedbeds and close to open water.

From 2006 to 2008, these patches of reeds expanded and merged

forming reedbeds. Other habitats present included protective

forest-belts, aquaculture ponds, and farmland inside the estuary

dyke; there were no reeds in these habitats.

From 2006 to 2008, all the above areas were surveyed for Reed

Parrotbills monthly, using six fixed transects, controlled to cover

about 1 ha each, with point counts located along the transects in

each area of habitat. The species has a far-carrying call, but it proved

difficult to estimate the distance of birds that could not be seen, so

in order to reduce the resulting bias in detection between seasons

and areas, individuals that were only heard were excluded. Non-

parametric tests were used to test for differences in Reed Parrotbill

densities in different habitats.

RESULTS

Distribution and habitat use of the Reed Parrotbill in China

Before 1980, the species was reported at only a few sites (Table 1,

Figure 2), mainly by foreign scholars. Since 1980, it has been

reported more widely (Table 1, Figure 2). The known distribution

has expanded from sites in Jiangsu, Jiangxi and Zhejiangprovinces

to over 60 sites in 10 provinces (Figure 2), but there have been no

records since 1980 from Jiujiang city near Poyang Lake in Jiangxi

province, Hangzhou city in Zhejiang province or Jiangyin city in

Jiangsu province, where they had previously been recorded (Gould

1874, Gee etal. 1929, Shaw 1934, Rank 1989). In the Chinese part

of its range, the species mainly inhabits coastal, lakeside and

riverside wetlands where the Common Reed is found. Some early

publications indicated that the species was recorded in reeds (David

1872, Lynes 1914, Gee & Moffett 1917, Table 1).

Of the 37 academic publications published after 1980 examined

in the study, 3 1 mentioned the vegetation used by Reed Parrotbills,

27 indicating that the species used reeds or reed-dominated

vegetation. The other four indicated that the species used habitats

that included ‘farmland and protective forest-belts near residential

area’ (Su et al. 1987), ‘dense bushes near stream and marsh’ (Hou

Figure 2. Distribution of Reed Parrotbill in China. See Table 1 for details of marked sites.

80°E 90°E 100°E 110'E 120”E 130°E

66

LI-HU XIONG &JIAN-JIAN LU

Forktail 29 (201 3)

Forktail 29 (2013) Habitat specialisation in the Reed Parrotbill Paradoxornis heudei 67

68

LI-HU XIONG & JIAN-JIAN LU

Forktail 29 (2013)

et al. 1997a), ‘bushes along the sea dike’ (Zhao et al. 2004) and

‘dense bushes near streams and marshes’ (Han et al. 2007).

The species has been recorded in 57 localities since 1980 in

academic publications (26 sites), our observations (16 sites),

personal communications (10 sites) and internet news (5 sites)

(Table 1). In 39 of the 57 sites or areas, the habitat types were

described and in 36 of them the habitat used by Reed Parrotbills

was reeds or reed-dominated vegetation.

Distribution of Reed Parrotbills at Chongmang Island

A total of 625 Reed Parrotbills was recorded in 22 transects (Figure

1). Nineteen of these transects were in intertidal mudflats and the

other three in newly reclaimed areas close to intertidal mudflats.

All the birds were recorded in reeds. Transects in intertidal

mudflats were covered with reedbeds and those in newly reclaimed

areas had reeds scattered around aquaculture ponds or in planted

woodland. No Reed Parrotbills were recorded in the remaining

transects, which included farmland, riversides, residential areas,

woodlands and aquaculture ponds away from the intertidal

mudflats. These transects either had no reedbeds or only had small

patches of reeds isolated from larger reedbeds.

Habitat selection of Reed Parrotbills at Chongxi Wetland

Research Centre

During three years observation, no Reed Parrotbills were recorded

in areas without reed vegetation (farmland, aquaculture ponds and

protective forest belts) . The species was only recorded in areas with

reed vegetation: including reedbeds, reeds with dense trees (RDT),

reeds with sparse trees (RST) and patches of Common Reeds

(PCR). In these areas, Reed Parrotbills was recorded almost year

round (Figure 3). There were significant differences in Reed

Parrotbill density between the four types of reed habitat (Friedman

test, Chi-square = 60.729, df = 3, P<0.001). Wilcoxon Signed

Ranks Test indicated that Reed Parrotbill density in reedbeds was

significantly higher than in PCR, RDT and RST (allP< 0.01), but

there were no significant differences among PCR, RDT and RST

(all P> 0.05). In RDT, the Reed Parrotbill was not recorded after

May 2007 when the reeds died back, but it was recorded in almost

all months in RST (Figure 3), indicating that presence of the

Common Reed is a necessary precondition for survival of the Reed

Parrotbill.

DISCUSSION

The use of reed vegetation by Reed Parrotbills is reported frequently

in published literature. These reports, combined with the

observations from Chongming Island, indicate that the species uses

reedbeds or reed-dominated vegetation as habitat throughout its

Chinese distribution. Detailed local observations at Chongming

Island and Chongxi Wetland Research Centre also indicated that

the species was almost exclusively associated with reeds and that

birds were not found in nearby areas without reeds. It was

concluded that the Reed Parrotbill is a reed-dominated habitat

specialist.

The species’s distribution map for China (Figure 2) includes

many new records. In the south, the range extends to the southern

shore of Hangzhou Bay but the northern extent cannot be

determined yet, as there are records of Reed Parrotbills in south¬

east Russia. This study revealed that the species has a larger range

than previously thought. Further changes to the known range are

anticipated, as research and hireling activities increase.

At Chongxi Wetland Research Centre, when reed shoots

disappeared from mixed vegetation, Reed Parrotbills were rarely

recorded in the reed-free vegetation. They were unable to persist in

these areas by utilising more distant patches oi reeds and disappeared

along with the reeds. This implies that the species is dependent on

reed vegetation and this dependence on reeds might constrain their

ability to use other vegetation in the absence oi reeds. There are

published accounts of Reed Parrotbills using non-reed vegetation

close to large areas of reeds (Su et al. 1987, Hou et al. 1997a, Zhao

et al. 2004, Han et al. 2007). The species has been observed using

bushes and woodland close to reeds when the reed-vegetation was

disturbed by, for example, reed harvesting (La Touche 1906, Wang

& Tian 1988). In the absence oi disturbance, birds occasionally visit

nearby non-reed vegetation, for example Spartina, but the number

oi individuals and their density were much lower than in reeds

(Dong etal. 2010, Gan et al. 2010). Birds evidently disperse readily

into non-reed vegetation near reeds, but it is not known whether

they use resources within the non-reed vegetation or make only

transitory visits. Birds visiting Spartina close to reeds only spent

very short periods of time there (Dong et al. 2010).

Records of the species in non-reed vegetation indicate that the

degree of habitat specialisation is not fully understood. It is

Figure 3. Monthly variation of Reed Parrotbill density (±SE) in reedbeds, reed patches, reeds with dense trees and reeds with sparse trees during

2006 to 2008, at Chongxi Wetland Research Centre.

Forktail 29 (2013)

Habitat specialisation in the Reed Parrotbill Porodoxornis heudei

69

important to understand the relationships between the species and

the reed vegetation. Studies have shown that Reed Parrotbills feed

on insects, insect eggs and larvae on reed shoots year round, in a

tidal marsh in Changjiang Estuary (Xiong etal. 2007, 2010). The

species may have special morphological adaptations in its bill, which

facilitate breaking reed stems to retrieve insects within (Xiong et

al. 2010) but which compromise its ability to use food resources in

other vegetation. Similar extreme specialisation is seen amongst the

bamboo-specialist insectivores, which feed on insects in, on and

around living bamboo (Cockle et al. 2009). Although the Reed

Parrotbill is limited to reed vegetation and it feeds on insects in

and on reeds (Xiong et al. 2007), it is not yet certain that their

prey is also restricted to reed vegetation.

It is not known whether the two recognised subspecies differ

in habitat use or other life history characteristics. Subspecies beudei

occurs on Chongming Island, where the fieldwork was carried out.

At sites where polivanovi is found, such as Zhalong National Nature

Reserve (site 3 in Figure 2), Longfeng Wetland Nature Reserve (4)

and Xingkaihu National Nature Reserve (6), the Reed Parrotbill

habitat was described as reed vegetation or reed marsh. Thus, it seems

likely that the two subspecies have the same habitat requirements.

CONCLUSIONS

This review of Reed Parrotbill distribution in China revealed that

its range and the number of locations where it occurs are larger

than previously thought. This does not indicate an improvement

in the conservation status of the Reed Parrotbill, as the study also

confirmed the species’s strict habitat specialisation. Given that

existence of Common Reeds is a precondition for Reed Parrotbills

to survive, more attention must be paid to the conservation of reed-

dominated habitat, such as coastal wetlands, lakeside wetland and

marshes, and corridors in reed-dominated habitats should be

designed and maintained to reduce the effects of habitat

fragmentation.

Corridors might also be useful to link areas of reedbeds without

Reed Parrotbill populations to nearby locations that are already

populated and this could be used in the selection and development

of protected areas.

It would be useful to learn more about how the Reed Parrotbill

uses and has adapted to reed vegetation, to help understand the

evolutionary history of the species and likely threat mechanisms.

The distribution of Reed Parrotbills may be predicted based on its

strong relationship to the Common Reed. Large areas of reed

vegetation close to or linked by corridors to reed vegetation with

Reed Parrotbill populations could be potential habitat.

ACKNOWLEDGEMENTS

We thank Jia Liu, Lingdong Li, Guolang Zhang for sharing their birding

records. We thank the staff at Chongxi Wetland Research Centre for

assistance in the field. We especially thank two anonymous reviewers and the

editors for their valuable comments.

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FORKTAIL 29 (2013): 71-77

Increasing variation in population size and species

composition ratio in mixed-species heron colonies in Japan

MIYUKI MASHIKO& YUKIHIKOTOQUENAGA

Long-term population dynamics of colonial herons and egrets are well documented in Europe and the USA, but not in East and South-East

Asia. Here the population dynamics of mixed-species colonies from 2002 to 201 1 in Ibaraki prefecture, east Kanto, Japan, are reported.

From censuses based on a combination of aerial and ground surveys, the number of breeding colonies was found to vary from 1 5 to 20. The

population dynamics of Great Egret Casmerodius albus and Black-crowned Night Heron Nycticorax nycticorax remained relatively constant,

while Grey Heron Ardea cinerea and Intermediate Egret Mesophoyx intermedia increased, but Little Egret Egretta garzetta and Cattle Egret

Bubulcus ibis decreased. When data for the six species were combined, the sum of their populations was almost constant, but variation

increased in colony size, species composition ratio and the number of years that individual colonies existed. The population of colonies

typically ranged from 200 to 2,000 individuals up to 2004, but smaller (under 200 individuals) and larger (over 2,000 individuals) colonies

appeared after 2006. Increased variation in the number of consecutive years colonies existed was closely related to increased variation in

colony size. Increased variation in species composition ratios in colonies was not a by-product of the increased variation in colony size, and

the occurrence of colonies dominated by Grey Heron, Intermediate Egret or Black-crowned Night Heron after 2006 played an important

role in the structural changes of mixed-species colonies.

INTRODUCTION

H erons and egrets (Ardeidae) are commonly found in aquatic

habitats worldwide (Kushlan & Hafner 2000). In Europe, long¬

term population trends of such species have been well investigated,

and some factors that explain how and why population sizes

fluctuate at regional level have been revealed: cold winters (Stafford

1971, Reynolds 1979, Hafner & Fasola 1997, Fasola et at. 2010),

rainfall (McKilligan 2001), water level (Griill & Ranner 1998),

habitat conditions (Tourenq etal. 2000, 2004), aquaculture (Fleury

& Sherry 1995) and human disturbance (Fasola et al. 2010). In

East and South-East Asia, long-term records of breeding

populations of colonial nesting herons and egrets only exist in Hong

Kong and Vietnam (Kushlan & Hafner 2000, Wong & Young

2006). Fack of local information makes it difficult to assess the

current status of these birds.

In Japan Grey Heron Ardea cinerea. Great Egret Casmerodius

albus, Intermediate Egret Mesophoyx intermedia. Little Egret

Egretta garzetta , Cattle Egret Bubulcus ibis and Black-crowned

N ight Heron Nycticorax nycticorax breed in mixed-species colonies.

Nationwide research was carried out in 1980 and 1992 (Research

Division of the Wild Bird Society of Japan 1981, Environmental

Agency ofjapan 1994), and it was reported that single- and mixed-

species colonies were distributed throughout Japan’s lowlands.

Although there are many observations of colonies in various areas,

long-term local population trends have only been reported by

Narusue (1992) and Matsunaga et al. (2000).

Narusue (1992) argued that both the populations and the

average colony size of these species declined from the 1 940s to 1992

in Saitama prefecture, west Kanto Plain, due to loss of foraging

areas and use of agricultural chemicals. Changes in the irrigation

of rice fields from shallow earth ditches to deep concrete -walled

channels and the decline in aquatic prey caused the decline of

Intermediate Egret (Narusue & Uchida 1993, Lane & Fujioka

1998), the commonest egret until the 1960s, but now categorised

as ‘near threatened’ (Ministry of the Environment 2002). In

contrast, a long-term study of Grey Herons in Hokkaido by

Matsunga et al (2000) reported an increase in population and the

number of colonies. There are currently no other reliable data to

assess population trends of herons and egrets in Japan.

The Environmental Agency ofjapan (1994) showed that in

Ibaraki prefecture, east Kanto, Japan, both the average colony size

and the population of Intermediate Egret were large compared to

elsewhere in Japan, and this suggested that data from this area could

provide important information for future assessment of

populations of colonial breeding herons and egrets in Japan and

other parts of Asia. In this study, colony censuses were carried out

from 2002 to 2011 in Ibaraki prefecture to investigate trends in

these populations using a combination of aerial and ground surveys.

The changes are discussed here with reference to the trends in

population dynamics of each species, changes in the nesting

vegetation and the number of consecutive years that colonies

existed.

METHODS

Study area

The study focused on Ibaraki prefecture and parts of Tochigi and

Chiba prefectures in Honshu, central Japan (35.783°-36.767°N

139.767°-140.683°E) (Figure 1). The area is in the east Kanto

Plain, near Lake Kasumigaura, and includes six major rivers: Kuji,

Naka, Sakura, Kokai, Kinu and Tone. The north is mountainous,

but the predominant land use in other areas is farming, with large

areas along the rivers being used for rice production. There are also

lotus fields near Lake Kasumigaura, areas of lowland forest and

human habitations. Japan started a national project to consolidate

rice production in 1 963; this included extending irrigation ditches,

improving service roads, and enlarging fields to facilitate

mechanised farming equipment. It was largely complete by 1980

(Himiyama & Kikuchi 2007), but continued in part ol the study

area into the last decade, being 78% complete by 2010 in Ibaraki

prefecture. The climate of the region is moderate with an annual

average air temperature of 14.0 ± 0.1°C and an annual precipitation

of 1,388.2 ± 54.3 mm. Despite a small annual decrease in rice

cultivation, neither climate nor land use showed obvious changes

during the study period (Figure 2).

The herons and egrets breed from March to August, but there

is considerable variation from species to species (Figure 3). The

Grey Heron arrives first in March, Great Egret, Little Egret, and

Black-crowned Night Heron arrive in April; these species are

residents and wanderers, and some individuals winter in this area.

Finally, the migrant species arrive. Intermediate Egret in late April

and Cattle Egret by early May (A. Abe in litt. 200 6). Usually Grey

72

MIYUKI MASHIKO& YUKIHIKO TOQUENAGA

Forktail 29 (2013)

Figure 1. Locations of colonies from 2002 to 201 1 . Grey regions show

an altitude greater than 1 00 m where the distribution of egrets is lower.

Dots enclosed by a circle are considered to be historically identical

colonies. Exceptionally, there are two cases in which the nearest-

neighbour distance is shorter than 6.47 km; ( 1 ) l20 — L30: because L was

newly established in 2010 and consisted of Grey Herons Ardea cinerea

and Great Cormorants Phalacrocorax carbo, we assumed L was different

from I. (2) Q40-R41 42: Q was newly established in 2006, whereas Koshida

(2007) reported that R has existed since 1984. It is difficult to accept

that Q and R are one colony.

N

Figure 2. Changes in a. mean temperature, b. annual rainfall, and

c. areas of six land-use types from 2002 to 201 1 in Ibaraki prefecture,

which was the main region of the study area (6,096 km2). 'Paddy field'

includes both rice paddies and lotus fields, and 'others' includes parks,

golf courses, and uncultivated fields. Data were downloaded 8 June

2013 from http: //www. data.jma.go.jp/obd/ stats/etrn/index.php for

climate and http: //www. pref.ibaraki.jp/bukyoku/kikaku/mizuto/

ibarakinotochi/25/ibarakinotochi.htm for land use.

Year

Heron and Black-crowned Night Heron are nocturnal, but during

the breeding season they are also active during the day.

Colonies were located in bamboo thickets, trees or a mixture

of both. Bamboo thickets were composed of Moso Bamboo

Phyllostachys pubescens, Simon Bamboo Pleioblastus simonii or

Dwarf Bamboo P. chino. Coniferous tree sites consisted mainly of

Japanese Red Pine Pinus densiflora, Japanese Cedar Cryptomeria

japonica and Japanese Cypress Cbamaecyparis obtusa-, broadleaftree

sites were mainly Japanese Zelkova Zelkova serrata, Japanese Oak

Qiiercus serrata and Yoshino Cherry Prunus x yedoensis.

Census of colonies

Colonies have been recorded in the area over the last 25 years

(Koshida 2007) and have high site fidelity (Custer et al. 1980,

Frederick et al. 1996); between March and early May, 93 ± 0.02%

of the colony sites were found by checking the places where colonies

had been located in previous years. When a colony was abandoned,

checks were made to determine whether other colonies had formed

nearby. Local literature and personal communication were used to

locate colonies that had not been found during the authors’ own

field work. Site vegetation was recorded by identifying bamboo or

tree species holding at least one nest.

In small colonies of fewer than about 50 nests, if all the nests

were visible from outside or within the site, the nests of each species

were counted directly from the ground, and the breeding

population estimated by doubling the number of nests counted.

In most cases, ground-based counts were impossible due to colony

size, the impenetrable nature of dense bamboo thickets, or other

vegetation, such as tall trees, that made nests invisible. Hence,

counts were made using a combination of aerial and ground surveys,

following the method of Fujioka et al. (2001).

For aerial surveys, a small ‘Sky Surfer’ radio-controlled

paraglider was used (Green Corporation, Japan, Plate la). This

equipment is quieter than fixed-wing aircraft or helicopters

commonly used for bird colony censuses (Kushlan 1979, Rodgers

et al. 2005), and very suitable in this case where more than 80% of

the colonies were close to residential areas. Aerial photographs of

each colony were taken at an altitude of 30- 50 m just before sunrise

(about 04h00) when most birds were in the colony. Photography

was started in mid-May after arrival of Cattle Egrets, and was

continued until early July when distinguishing between growing

nestlings and adults became difficult (Figure 3). Aerial photographs

were taken once during that period at each site. All individuals of

the four light-coloured species (Great Egret, Intermediate Egret,

Little Egret, and Cattle Egret) in the images were counted (Plate

lb). For large colonies, several photographs were used to obtain a

complete composite image of the colony.

Figure 3. Breeding period of each species in the study area showing

the timetable of aerial surveying and species composition counts.

White, grey, and black shading show arrival and nest building,

incubation, and chick-rearing periods, respectively. The parallelogram

shapes indicate the variation in individual breeding periods. After

breeding is over, some birds continue to roost in the colonies but all

disperse by October.

Mar. Apr. May Jun. Jul. Aug.

Grey Heron

Black-crowned Night Heron^-

Great Egret, Little Egret s'

Intermediate

Egr^^ZI

Cattle Egret ^^s'

Aerial photography

Species composition counting

GREEN CORPORATION

Forktail 29 (2013) Variation in population size and species composition ratio in mixed-species heron colonies in Japan

73

Plate 1.

a. Sky Surfer in flight.

b. An example of aerial

photographs that show one part

of a composite photograph.

Because it was not feasible to identify the light-coloured species

only from the photographs, and because the two dark-coloured

species (Grey Heron and Black-crowned Night Heron) were

difficult to count in the photographs, species composition ratios —

the proportion of each species within a colony — were estimated

using data from ground surveys. The number of individuals of all

species going in and out of each colony for a period of 30 minutes

in the daytime were counted using binoculars. It was very difficult

to identify to species level white egrets flying in and out at the same

time, so the viewing range of each observer was restricted by setting

a common range of observation, approximately a 30° field of vision.

In the case of large colonies, surveys were carried out from two or

three different directions. Ground surveys were made once or twice

in June, the peak chick-rearing period (Figure 3), when all species

engage in frequent foraging flights. Thirty minutes is much shorter

than the typical duration of one foraging flight and it was assumed

that each individual counted, whether arriving or departing, was

observed only once during the observation period and therefore

the observed proportion of each species reflects the species

composition ratio of the colony.

The total estimated number of individuals in the colony

(T = colony size) was calculated using t = -- where A is the actual

number of light-coloured individuals counted from aerial

photographs, and x is the sum of the proportion of Grey Herons

and Black-crowned Night Herons obtained from the ground

survey. The estimated population size of each species in the colony

was obtained by multiplying the colony size by the proportion of

each species.

Data analyses

To determine the number of colonies each year, the number of

observed colony sites was first counted. But the number of sites

itself was not taken to be the number of colonies because a few

colony sites were very close to each other despite foraging ranges

having radii of about 10 km, and sometimes over 20 km (Nabeya

2011). An earlier study showed that heron colonies are evenly

distributed to avoid overlap of foraging sites (Gibbs et al. 1987).

Consequently if colony sites were located close together, they were

grouped together and counted as a single colony because their

foraging areas overlapped substantially. To determine which sites

should be counted as a single colony, the half of the mean nearest-

neighbour distance (ND) of observed colony sites for each year

was used. If more than one colony site was located within the ND,

colony censuses were carried out at each site, the data were

combined and it was counted as a single colony.

Colonial birds have been found to have high site fidelity (Custer

et al. 1980, Frederick et al. 1996); every year some colonies

returned to the same locations as the previous year, some birds

were abandoned and new ones were established. To obtain the

number of consecutive years (NCY) each colony existed, the

number of years from first establishment at the location was

counted. The movement of a colony was also considered and

when abandonment and new establishment occurred in

neighbouring locations in successive years, the new site was assumed

to be a descendant of the abandoned one, e.g. abandonment was

sometimes caused by vegetation loss through natural causes or

felling and the colony was often re-established nearby. The ND

was used to determine a reasonable displacement distance of a

colony and it was assumed that each colony had a domain of

attraction of half the average distance between the next nearest sites.

Thus, colonies consecutively established at the same site or at a

different site within a radius of the ND were counted as a single

colony. Koshida (2007) was used as data source of the NCY of

colonies established before 2002; consequently the NCY ranged

from 1 to 36 years rather than being limited to the period of

this study.

The population of each species was calculated annually using

the mean population size per colony rather than summing the

population sizes for all colonies with census data. This approach

was used because aerial and ground surveys produced only partial

data due to practical difficulties — problems in taking aerial

photographs and/or delays in detecting colony sites. The simple

sum of colony population sizes would have been inappropriate

because it is an increasing function of the number of colonies with

census data. The percentage of colonies surveyed increased from

78% in 2002-2004 to 94% in 2006-201 1. (Data from 2005 were

excluded because aerial and ground surveys were limited to only 5

out of 1 8 colonies.) Thus, the overall total population of the target

M. MASHIKO

74

MIYUKI MASHIKO& YUKIHIKOTOQUENAGA

Forktail 29(2013)

species reflect the mean colony sizes rather than the total number

of individual birds in the study area.

To evaluate difference in colony size, species composition ratio,

and NCY among colonies, the coefficient of variance (CV) for each

year was calculated. For species composition ratio, the proportional

similarity index (Whittaker 1952) was calculated for each colony

every year as Lif,'0~f)-> where p, is the proportion of species i in one

colony and p~t is the mean proportion of that species in all colonies

surveyed in that year. The index ranges between zero and unity:

zero means completely different and unity means completely equal.

Then the CV of proportional similarity of the species composition

ratio was obtained for each year.

Ten years is too short for ordinal time series analyses, so

randomisation tests were done to assess temporal trends in the

number of colonies, population sizes of each species, sum of the

population sizes of the six species, colony sizes and CVs of three

variables (colony size, species composition ratio and NCY). In a

randomisation test, the linear regression coefficient ((3) of a target

variable based on the original data was obtained first. Next the data

were shuffled 30,000 times and compared beta with the linear

regression coefficients (|3’s) of the shuffled data to obtain one-sided

P-values to assess whether the target variable was increasing or

decreasing. Sensitivity analyses of the population of each species

against the three CVs (colony size, species composition ratio and

NCY) were performed. Generalised linear models specifying

population sizes of the species as explanatory variables and CVs as

dependent variables were constructed, using Gaussian distribution

with an identity link function for all model fitting. The most

suitable models based on Akaike’s information criterion values were

chosen and the coefficients of explanatory variables of the models

as sensitivity against dependent variables were considered.

If the CVs of colony size and species composition ratio show

parallel changes, there is a possibility that the variation in species

composition ratios increased as a by-product of the increase of

variation in colony sizes. To examine this possibility, a

randomisation test was performed to determine whether the

variation in species composition ratios was solely caused by a

sampling bias according to the variation in colony size. First a

hypothetical total number of herons that consisted of the six species

was prepared. The species composition ratio of the whole number

of herons was arbitrary. Next multiple colonies with equal colony

sizes from the total number of herons were sampled. Then

proportional similarities of species composition ratios of these

hypothetical colonies against the species composition ratio of the

whole number of herons were calculated. Proportional similarities

for hypothetical colonies of the same number but with different

colony sizes were also calculated. Finally, the variance of the

proportional similarities between equal and unequal size colonies

were compared, and the probability that proportional similarities

of unequal size colonies were larger than or equal to those of equal

size colonies with 10,000 iterations was obtained.

To evaluate changes in nesting vegetation, the Friedman test

was used to analyse whether the vegetation of colony sites changed

from year to year. Nesting vegetation consisted of one or a mixture

of the following three types: bamboo thickets, coniferous trees and

broadleaf trees; there were seven types in total.

Finally, a randomisation test was performed to determine

whether there was a positive correlation between NCY and colony

size among colonies by reshuffling the year record so as to randomise

the consecutive colony-size dynamics of each colony.

All statistical analyses were conducted using R ver. 2.13.0 (R

Development Core Team 2011). Data are presented as mean ± SE

throughout. The randomisation test on the relationship between

the variation of population sizes and that of species proportion

ratios was also conducted with R. All R scripts for the above

statistical analyses are available from the authors.

RESULTS

During the 10-year period, there was an average of 19 colony sites

in the study area every year (19.10 ± 0.72 colony sites, n = 10);

cumulatively 191 colony sites were used over the 10 years. Some

colonies were in the same locations for more than one year, and a

total ol 62 colony sites were used (1 to 62 in Figure 1). Colony

sites were separated by an average of 13 km (mean ND over 10

years 12.95 ± S.39 km, n — 191), so the ND was defined as within

6.47 km. Hence, these 62 colony sites were categorised into 27

colonies (A to a in Figure 1) because colony sites consecutively

established at different locations within a 6.47 km radius were

considered a single colony. Six of 27 colonies were made up of two

or three colony sites in at least one breeding season, and the median

distance between them was 1.44 km (range: 0.32-5.12 km, 10

combinations of colony sites in all). Finally, the annual number of

colonies increased gradually ((3 = 0.382, P = 0.006) from 15 to 20

(Figure 4a).

In the case of Intermediate Egret and Black-crowned Night

Heron, the average population per colony was relatively large (about

300 individuals) and these species remained dominant throughout

the 10-year period (Figure 4b). Conversely, it was small (about 50

individuals) for Great Egret and Grey Heron, and intermediate

(about 100 individuals) for Cattle Egret and Little Egret. The sum

of the population of the six species (mean colony size) ranged from

726 to 966 individuals and remained almost constant ((3 = -4.301,

P = 0.342).

The population trends of each species varied (Figure 4b). Grey

Heron and Intermediate Egret increased (Grey Heron: (3 = 9.575,

P < 0.001; Intermediate Egret: (3 = 9-519, P = 0.033), whilst Little

Egret and Cattle Egret decreased steadily (Little Egret: (3 = -2.069,

P = 0.002; Cattle Egret: (3 = -20.672, P < 0.001). The Black-

crowned Night Heron population fluctuated over the years but

remained almost constant ([3 = 9.3 1 1 , P = 0. 145). The Great Egret

population was small but almost constant ((3 = 0.036, P = 0.492).

Colonies were very variable in size, and the CV of colony size

continuously increased (Figure 4c) over the ten years ((3 = 7.510,

P < 0.001). Colonies ranged from 200 to 2,000 individuals until

2004, while smaller (under 200 with minimum 8 individuals) and

larger (over 2,000 with maximum 3,280 individuals) colonies

appeared after 2006. Between 2008 and 20 1 1, the smaller and larger

colonies increased from 33% to 41% of colonies surveyed.

In parallel with the increase in the CV of colony size, the CV

of proportional similarity of species composition ratios increased

(Figure 4c), especially after 2006 ((3 = 7.002, P < 0.001). Until

2004, most colonies consisted of five species (Great Egret,

Intermediate Egret, Little Egret, Cattle Egret and Black-crowned

Night Heron), and the composition ratio was similar among

surveyed colonies (mean proportional similarity = 0.86 ± 0.02,

n = 43). Grey Heron bred in only three, six and seven colonies in

2002, 2003 and 2004, respectively. Until 2004, the composition

ratios of the Intermediate Egret and the Cattle Egret were higher

than those of other species in half of the surveyed colonies in

accordance with their large population (Figure 4b), but no species

became dominant (over 50% of the composition ratio). After 2006,

37% of all surveyed colonies were dominated by the Grey Heron,

Intermediate Egret or Black-crowned Night Heron, and differences

in the species composition ratios among colonies increased.

The CV of the NCY also increased gradually (Figure 4c) ((3 =

2.453, P < 0.001). Eight colonies persisted between 2002 and 2011;

the remainder were abandoned or newly established. Every year

1-4 colonies were abandoned and 0-3 were established.

Considering the period prior to this study, 14 out of 27 colonies

had existed before 2002 and 4 had persisted for over 25 years.

Table 1 shows the results of the sensitivity analyses of

population of each target species against three CVs. Increase in the

Forktail 29 (2013) Variation in population size and species composition ratio in mixed-species heron colonies in Japan

75

Figure 4.

a. Changes in the number of colonies between 2002 and 2011.

b. Changes in population of each species per colony of and the sum of

the six species.

c. Changes in the coefficient of variation (CV) of colony sizes, number

of consecutive years (NCY), and species composition ratios. For the

changes in population size and CVs of colony sizes and species

composition ratios, the year 2005 is not shown because aerial and

ground surveys were limited to only 5 out of 18 colonies.

Year

CV of colony sizes was explained by the increasing Grey Heron

population, and the increase in the CV of species composition ratios

was also explained by the increasing Grey Heron population, and

marginally explained by the increasing Intermediate Egret

population. The increase in the CV of NCY was explained by the

increasing Grey Heron population and the fluctuating, though

statistically constant overall, population trend of the Black-

crowned Night Heron.

The randomisation test to determine whether the variation in

species composition ratios increased as a by-product of the increase

in variation of colony sizes did not reveal a significant result: the

probability that the proportional similarities between unequal size

colonies would be larger than or equal to those of equal size colonies

was almost even (0.538). The increase of variation in species

composition ratios could not solely be caused by the increase of

variation in colony sizes.

Changes in vegetation of the colonies were significant over the

years (%: = 25.2, df = 6, P < 0.001) (Figure 5). While the vegetation

in most colonies included bamboo until 2004, after 2008 more than

half the colonies were located in trees.

The slope obtained by a linear regression analysis of colony sizes

against NCY (43.05 ± 5.43) was significantly larger (P < 0.001)

than slopes obtained by the randomisation test where the year

record was shuffled for each colony so as to randomise consecutive

colony-size dynamics (Figure 6). This randomisation test indicates

that there was a positive correlation between colony sizes and the

NCY for the colonies.

Table 1. Sensitivity of population sizes against CVs. [5s are coefficients

of the best fit generalised linear model with Gaussian distribution and

identity link function. R2 = (null deviance - residual deviance)/(null

deviance). CS: colony size, SCR: species composition ratio, NCY: number

of consecutive years.

Figure 5. Changes in colony vegetation. See description of study area

for details of species.

Bamboo

tree

2002 2003 2004 2005 2006 2007 2008

Year

Figure 6. Relationship between colony size and longevity. Each dot

represents a colony censused in a particular year (n = 141). The

regression line was obtained by a linear regression analysis of colony

sizes against longevity assuming that each annual colony was

established independently.

76

MIYUKI MASHIKO& YUKIHIKOTOQUENAGA

Forktail 29 (201 3)

DISCUSSION

The survey in Ibaraki prefecture from 2002 to 20 1 1 indicated that

the number of breeding colonies (average 19) increased slightly and

mean colony size was almost constant. These results accord well

with the report by the Environmental Agency of Japan (1994):

there were 20 colonies in 1992 in Ibaraki prefecture, ranging in

size from 15 to 2,990 individuals (CV = 1 12.5), and the population

of these species has been relatively constant in the area for at least

two decades to 2011. However, variations in size, species

composition ratio and NCY among colonies increased significantly.

Colony vegetation changed from predominantly bamboo thickets

to tall trees. Trends in population dynamics differed: Grey Heron

and Intermediate Egret increased. Little Egret and Cattle Egret

decreased; and Great Egrets and Black-crowned Night Herons were

relatively constant — the population of both the latter species were

similar to previous reports (Research Division of the Wild Bird

Society of Japan 1981, Environmental Agency of Japan 1994) and

unchanged for three decades. Overall, there was no significant

change in population of these colonial species in the study area

during the decade, but variation in the structure of colonies and

population dynamics clearly increased.

In contrast to Great Egret and Black-crowned Night Heron,

the population of the other four species changed during the period

(Figure 4b). Grey Heron showed the greatest population growth,

which is in line with earlier reports that its population is growing

in other parts of Japan (Narusue 1992, Environmental Agency of

Japan 1994, Matsunaga et al. 2000, Sasaki 2001). In Hokkaido,

Matsunaga et al. (2000) suggested that recent climatic warming

and increase in aquaculture have provided the species with

additional food resources. It is not known whether the increase of

this species in other more temperate parts of Japan also depends

on these factors, but its ability to respond quickly to changes in

food availability (Adams & Mitchell 1995) would be expected to

boost populations. The other increasing species, Intermediate

Egret, was a predominant species in this area even though it has

been designated as a ‘near threatened’ species in Japan (Ministry

of the Environment 2002). Owing to the lack of current data from

other parts of Japan, it is not clear whether the population has been

recovering, but the abundant population in this area may be of

conservation significance in Japan; monitoring of this species

should continue.

Little Egret and Cattle Egret both showed a steady decline over

the period; the Environmental Agency ofjapan ( 1994) considered

them to be predominant and numerous throughout Japan,

including Ibaraki prefecture in 1992, and the population of both

has decreased during the last two decades. Although mild winter

weather contributed to their increase in France (Hafner & Fasola

1997) and rainfall drove the changes in Cattle Egret population in

Australia (McKilligan 2001) and Hong Kong (Wong & Young

2006), climatic variables are unrelated to the decrease of these

species in the study area because both temperature and rainfall have

been almost constant (Figure 2). It seems likely that changes in

food resources or foraging habitats may be contributory factors. In

northern Japan, Shimada et al. (2005) suggested that Little Egrets

might be strongly affected by the increase in population of the

introduced Black Bass Micropterus psalmoides, which has caused a

decrease of the smaller native fish species they prefer. In the absence

of historical and quantitative data in Japan, monitoring studies in

other regions are needed to make a complete assessment of

population dynamics of these declining species.

During the study period, variations in size, species composition

ratio and NCY increased (Figure 4c). Since these temporal trends

showed parallel changes, there is a possibility that the variation in

species composition ratios increased as a by-product of the increased

variation in colony sizes; but a randomisation test contradicted this

possibility, and it was concluded that the observed increased

variation in proportional similarities of species composition ratio

could not be solely caused by the increased variation in colony size.

Another change that coincided with the study period was the

change in nesting vegetation; the majority of colonies changed from

bamboo thickets to trees (Figure 5). More colonies were newly

established in tall trees even though bamboo thickets persisted in

the area. The decrease in the number of colonies in bamboo may

be due to the increase in Grey Herons because they prefer to nest

near the top of tall trees. However, those results contradict the

general knowledge that the target species use a wide range of nest

sites, including trees, bushes, reeds and on the ground. No other

species shows a particular preference for specific substrates

(Kushlan & Hancock 2005). Hence, there is no strong support for

the possibility that the vegetation of established colony sites

affected the size or species composition ratios of colonies.

Increasing variation in NCY may help explain the increased

variation in colony size and species composition ratios. These

results showed that the variation in the NCY among colonies grew

from year to year (Figure 4c), and there was a significant positive

correlation between colony size and the NCY that a colony existed

(Figure 6). Although food availability, measured as the area of

potential foraging habitat around the colony, has often been

thought to be the most important Factor affecting colony size

(Fasola & Barbieri 1978, Gibbs et al. 1987, Gibbs 1991, Baxter &

Fairweather 1998), previous studies in this locality showed that

variables related to foraging sites (areas around ponds, rivers, paddy

fields and lotus fields) did not have a major impact on colony size;

instead the NCY had a significant positive relationship with colony

sizes (Fujioka et al. 2001, Tohyama 2005). Increasing variation in

a colony’s size is therefore closely related to the colony’s longevity.

As for increasing variation in species composition ratio among

colonies, variations were due to the occurrence of colonies

dominated by Grey Heron, Intermediate Egret, or Black-crowned

Night Heron after 2006. In particular, the dominance of Grey

Heron was notablet in small, recently established colonies (Figure

6). It is well known that the Grey Heron often breeds in small

colonies of only 2-10 nests, while the other five species are more

gregarious and usually breed in large mixed-species colonies

(Kushlan & Hancock 2005). Thus, the Grey Heron population

growth after 2007 might contribute significantly to the increasing

variation in the species composition ratio and colony size despite

its relatively small overall population (Table 1).

Overall, the local population of herons and egrets in eastern

Japan seems to have remained constant for at least the last decade,

in parallel with the constant climate and land use variables.

Nonetheless, population dynamics of constituent species have been

changing, and variations in colony sizes and species composition

ratios have also increased. Such changes are revealed only by long¬

term and comprehensive colony census. Continuing studies are

required not only to reveal the factors affecting the population

dynamics of each species at a regional level, but also to establish a

better understanding of relationships between each species’s

population and the sizes or composition ratios of mixed-species

colonies.

ACKNOWLEDGEMENTS

We thank T. Tohyama, T. Yamaguchi, R. Yamagishi, C. Koshida, H. Takeda,

A. Abe, K. Nabeya and L. T. Carrasco for colony censuses, and S. Ikeno, M.

Seido and K. Takeda for information on the location of colonies. We especially

thank Y. Asano and M. Kobayashi for their mechanical advice and

maintenance of the Sky Surfer, and K. Ohashi and M. Fujioka for helpful

comments on an early draft. We also thank the Kashima Kyodo Shisetsu and

the Kubota Tsukuba Plant for access to their premises to make colony censuses.

Forktail 29 (2013) Variation in population size and species composition ratio in mixed-species heron colonies in Japan

77

We are grateful to our anonymous reviewers for useful comments. This study

was partly funded by Global Environment Research Fund from the Japanese

Ministry of the Environment and by a Grant-In-Aid for Scientific Survey

from the Japanese Ministry ofEducation, Science and Culture (No. 13740433

and 19570014).

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Miyuki MASHIKO and Yukihiko TOQUENAGA, Graduate

School of Life and Environmental Sciences, University

of Tsukuba, Ibaraki 305-8572, Japan. Email:

mashiko@pe.ies.Hfe.tsukuba.ac.jp

FORKTAIL 29 (2013): 78-87

Phenotypic evidence for the specific and generic validity

of Heteroglaux blewitti

P. C. RASMUSSEN & N. J. COLLAR

The genus Heteroglaux was established for the Forest Owlet H. blewitti when the species was first described, but owing to certain similarities

with Spotted Owlet Athene brama, the use of Heteroglaux fell into disuse in the twentieth century until the species was rediscovered in

1 997, and is still not universal; moreover, perceptions appear to linger that blewitti might even be conspecific with brama owing to a recent

claim of interbreeding. In reality blewitti is distinct from brama on external morphology (plumage described elsewhere; narial position

related to bill width; bill height; more heavily feathered toes; length of middle and hind claws; wing formula) and osteology, in which

blewitti is distinct from all three species of Athene (multiple cranial elements, especially the greatly widened and inflated frontal, and the

extremely stout tarsometatarsus). Lateral tail-flicking and direct, non-undulating flight further support generic separation.

INTRODUCTION

For well over a century the Forest Owlet Heteroglaux \Athene ]

blewitti remained as much a taxonomic as a conservation

enigma. It was discovered in central India 140 years ago, and at

once placed confidently in its own genus Heteroglaux by Flume

(1873);

At first sight it would certainly be classed as an Athene-, but

the head is much smaller [ sic. evidently a lapsus for ‘larger’]

than in any of the Athene's I possess, viz., brama, radiata,

malabarica, cuculoides, castaneonota. The nostrils are not

pierced from the front, backwards at the margin of a swollen

cere, but are well inside the margin, and are pierced straight

in. The upper surfaces of the toes, too, are not covered with

bristles, but thickly feathered.

In the decade that followed, only six further specimens were

taken (enumerated in Rasmussen & Collar 1998), and then the

species disappeared. Perhaps as a consequence of this, its generic

placement by Hume was never widely accepted. Although

Heteroglaux continued to be used by Hume himself and some

contemporaries (e.g. Hume 1879, Murray 1887, Sharpe 1891,

1899), from an early stage the Forest Owlet was also treated as

congeneric with the Little Owl Athene noctua and Spotted Owlet

A. brama, first in the genus Carine — even as early as Ball (1878),

writing in Hume’s own Stray Feathers — and more recently Athene.

Although Gurney (1894) retained it in Heteroglaux, he cited

Hume’s (1873) view that it looks much like A. brama and added

that Blanford considered it to belong to Athene-, and the following

year Blanford’s (1895) treatment as such appeared. Dubois (1904)

retained it in Heteroglaux, albeit without comment, but within a

few decades virtually all works treated blewitti as an Athene (e.g.

Baker 1934, Peters 1940, Biswas 1953, Ripley 1961, Abdulali 1972,

Marshall & King 1988). Without explanation, Wolters (1975)

assigned both blewitti and brama to the subgenus Heteroglaux, with

noctua and Burrowing Owl A. ( Speotyto ) cunicularia occupying

separate subgenera within Athene-, otherwise Heteroglaux has only

ever been used for blewitti. Voous (1989: 191) suggested that

blewitti ‘might provide a clue to understanding these relationships

[between Athene, Ninox and Glaucidium], though the Forest Owlet

may already be too close to the Spotted Owlet for that purpose’.

Recent molecular phylogenies of owls (e.g. Wink etal. 2004, 2009)

have not included blewitti.

Indeed, some considered the similarity of brama and blewitti

so great as to render them conspecific. Baker (1923) treated

blewitti as well as most subspecies of brama — all except, inexplicably

(perhaps as a lapsus), A. brama tarayensis of the north-western

areas of the subcontinent — as races of A. noctua. This view,

although not elsewhere accepted, may explain the listing in

the NHMUK specimen register of the Davidson specimen

(NHMUK 1925.12.23.958) that was stolen and remade by

Richard Meinertzhagen (Rasmussen & Collar 1999) as ‘ Carine

noctua blewetti' [sic], although the other blewitti in the same

accession (then NHMUK 1925.12.23.1, now MCZ 236630) was

listed as Athene blewitti. The fictitious locality of the stolen

specimen, which came to the (now) Natural History Museum,

Tring, UK, in the late 1960s in Meinertzhagen’s posthumous

bequest (Rasmussen & Collar 1999), may in turn explain why it

took until 1997 before the Forest Owlet was seen in the twentieth

century (King & Rasmussen 1998). Over much of the intervening

period, however, in the absence of clear diagnostic illustrations and

texts, the species was speculated or judged to be so close in

appearance to A. brama that it would be difficult and perhaps

impossible to distinguish it (Ripley 1976, Ali 1978, Ali & Ripley

1981), and consequently the few reports or claims of blewitti that

appeared in the interim were shown upon scrutiny to be brama

(Rasmussen & Collar 1998).

Following the rediscovery of the species there has been a degree

of conservation-oriented research focusing on its distribution and

ecology (Jathar & Rahmani 2002, 2004, Rahmani & Jathar 2004,

Ishtiaq & Rahmani 2005, Kasambe et al. 2005, Mehta et al. 2008,

Chavan & Rithe 2009, Yosef et al. 2010). However, one aspect of

its resurrection has remained unexplored: the issue of its generic

identity. Publications at the time of the rediscovery and in its

immediate aftermath mentioned both Athene and Heteroglaux in

their titles (King & Rasmussen 1998, Rasmussen & Collar 1998,

1999, Rasmussen & Ishtiaq 1999). The two major monographic

treatments of owls that appeared at this time (del Hoyo etal. 1999,

Konig et al. 1999), both of which cited the preceding references,

elected to retain the species in Athene, although the latter entered

a caveat that its tail-flicking habit ‘argues against a close relationship

with other Athene owls and suggests closer affinity with pygmy owls

[Glaucidium)’ , and consequently proposed ‘placing this species in

the subgenus Heteroglaux’ .

In the twenty-first century the trend has clearly been towards

accepting Heteroglaux as a valid monotypic genus. Collar et al.

(2001: 1775) remarked that despite the species’s ‘strong

superficial resemblance’ toAthene'its original placement in its own

genus appears well justified based on osteological evidence

(Rasmussen & Collar in prep.) and on recent behavioural

observations including flight pattern and song (Rasmussen &

Ishtiaq 1999)’. Thereafter, world lists (Dickinson 2003, Gill &

Wright 2006), Indian avifaunas (Rasmussen & Anderton 2005,

Forktail 29 (2013)

Phenotypic evidence for the specific and generic validity of Heteroglaux blewitti

79

Grimmett etal. 2011), one monograph (Mikkola 2012) and many

journal papers and reports (preceding paragraph) have used

Heteroglaux. Nevertheless, some sources have retained Athene (e.g.

Clements 2007, Konigetrt/. 2008, Yosef etal. 2010), one even with

the cryptic entry 'Remarks: Spurious use of the generic name

Heteroglaux' (Weick 2006). This is perhaps unsurprising given that

a clear case for the acceptance of this genus has never been made,

and the osteological evidence referred to above never published.

Here we seek to rectify these deficiencies.

This need is rendered all the more pressing following a recent

report (Pande et al. 2011), albeit rejected (Ishtiaq 201 1, Jathar &

Patil 2011), of hybrid Forest Owlets x Spotted Owlets. For this

reason, we also consider the extensive structural differences between

blewitti and brama beyond the plumage distinctions established

in Rasmussen & Collar (1998). However, the exercise further

requires the osteological analysis to extend beyond differences

between these two species to cover not only all members of Athene

but key representatives of other related owl genera (including

Surnia, Glaucidium , Xenoglaux , Micrathene , Athene, Aegolius and

Ninox: Ford 1967, del Hoyo etal. 1999).

METHODS

We considered two types of evidence: external morphology

(focusing on the differences between blewitti and brama) and

osteology (considering the differences between blewitti and Athene,

thence to other genera) . Plumage comparisons between blewitti and

brama have previously been presented in Rasmussen & Collar

( 1 998), and we therefore here restrict our comparisons of external

morphology to mensural characters. We also briefly review data

reported elsewhere for acoustics and behaviour.

External morphology of blewitti and brama

For the external morphological analysis we assembled for

examination at the Natural History Museum, UK (NHMUK) all

known specimens of blewitti (seven; four males, three females), and

used the opportunity to compare them with other owl species, most

importantly Athene brama, with which blewitti is ostensibly so

closely allied as to have been considered conspecific (as noted

above). We measured all specimens of blewitti (data in Rasmussen

& Collar 1998) and specimens of brama at NHMUK; American

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

Natural Sciences of Philadelphia (ANSP); Museum of

Comparative Zoology, Harvard University (MCZ); National

Museum of Natural History, Smithsonian Institution,

Washington, DC (USNM); University of Michigan Museum of

Zoology, Ann Arbor (UMMZ); and Zoological Survey of India,

Calcutta (ZSI). This sample includes numerous representatives of

each of the races of brama recognised by Peters (1940).

The specimens of blewitti were measured, x-rayed,

photographed and videotaped in detail. Comparative

measurements were also taken from 84 brama skins (37 males, 35

females, 12 unsexed) at AMNH (n = 19), ZSI (10), NHMUK (26)

and USNM (29). Of the brama skins measured, 27 originated near

known localities for blewitti, but specimens were included from

throughout the range of brama. Measurements (in mm) taken from

skin specimens were: culmen from base of skull; culmen from distal

edge of cere; minimum width between nares; height of upper

mandible at distal edge of cere; length of longest rictal bristle

(straightened); tarsus; wing (straightened and flattened); tail

(callipers inserted between central rectrices at insertion point);

middle claw (digit 3) and hindclaw (digit 1, both claw

measurements taken from the distal edge of scutes).

To compare wing formulae, shortfalls from the wing-tip of

each of the primaries (P1-P10, numbered from the outside)

were measured (in mm) for six blewitti (one blewitti, NHMUK

1886.2.1.544, was excluded as its wing-tip is heavily worn) and

23 brama from USNM. The distances from the notches in the

inner web to the tip of each of the outer four primaries (P1-P4)

were also measured (in mm), as was the distance from the distal

end (narrowest point) of the emargination on the outer webs

to the feather tip for P2-P4. The notch for P4 was often not

obvious in specimens of brama, and in these cases it was not

measured.

Osteology

Measurements of skeletal features were taken directly from x-rays

of blewitti and brama specimens, in which multiple views taken

at various angles allowed direct comparisons with skeletal

elements of brama and with the actual skin specimens x-rayed

to ensure that the bones were oriented along the correct axes to

avoid size distortion due to foreshortening. Only elements lying

close to the film surface were measured to minimise parallax.

Measurement options were limited by bone preservation and the

fact that they are articulated in skin specimens of blewitti.

Measurements taken were: greatest width of skull; lengths of

humerus and ulna; length of carpometacarpus from proximal end

to distal articular surface; length and minimum width of tibiotarsus,

and width oi its condylar end; and length and minimum width of

tarsometatarsus.

Univariate statistics and principal components analysis (PCA)

using correlation matrices were done separately on external, skeletal

and wing formula measurements using SYSTAT for Windows

(version 5). Variables used in PCA were chosen partially to

maximise the number of specimens of blewitti that could be

included without estimation of missing data. Because of the small

sample of blewitti, sexes were combined.

Intergeneric skeletal comparisons

To allow osteological comparisons, several skeletal elements (the

entire humerus, radius, ulna, tibiotarsus and tarsometatarsus; a

femur missing the head; and the skull missing part of the posterior

and caudal regions) were removed by J. P. Angle from the left side

of a blewitti skin specimen (NHMUK 1886.2.1.546) using the

techniques in Olson et al. (1987); casts were retained at USNM,

as USNM 261299. These elements were compared directly with

USNM skeletons of A. brama (n = 6); Little OwlH. noctua (10);

Burrowing Owl A. cunicularia (5); Jungle Owlet Glaucidium

cuculoides (7); White-browed Owl Ninox superciliaris (l); Brown

Hawk OwlIV. scutulata (3); Boreal Owl Aegolius fiunereus (1); and

indirectly with noctua (4) and brama (2) from UMMZ. The

UMMZ osteological specimens were examined the week following

the USNM comparisons, and were videotaped to allow further

study. In addition, important osteological features that were

observed in the extracted blewitti skeletal elements were then

examined (as possible) in the x-rays of all seven blewitti specimens

and the x-rayed brama. Osteological terminology follows Howard

(1929) and Baumel & Winner (1993). Measurements taken of the

above specimens, along with brama (9), noctua (4), N. scutulata

(1) and Philippine Hawk Owl N. philippensis centralis (1) were:

skull (including culmen) length; minimum widths of the frontal

both anterior and posterior to the supraorbital processes; maximum

skull width; height of lateral rim of frontal; width of distal half of

lacrimal (maximum medio-lateral width); length of lacrimal

(maximum antero-posterior length of caudal edge); maximum

width across both palatines in situ ; maximum length and minimum

width of ulna; for humerus, femur, tibiotarsus and tarsometatarsus,

maximum lengths, minimum widths and distal widths, and for the

last two elements maximum proximal widths as well. For

tibiotarsus, length was from the proximal articular surface, and

proximal width did not include the fibula.

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P. C. RASMUSSEN & N. J. COLLAR

Forktail 29(2013)

RESULTS

Externa! morphology of blewitti and brama

The characters that separate blewitti from brama in the held are

summarised in Rasmussen & Collar (1998). Even within a race,

brama presents great variability in plumage and in most (but not

all) ol the characters distinguishing the two species a few individuals

of brama closely approach the condition in blewitti, especially when

the latter species is in worn plumage.

The nares are situated more widely apart in blewitti than in

brama, owing largely to the broader culmen ridge of blewitti (Table

1). Moreover, the nares of blewitti are positioned more obliquely,

not facing directly anteriad as in brama. The cere of blewitti is less

inflated and the nares are situated well inside the cere, instead of

right at the edge of the more swollen cere, as in brama.

Compared to brama, blewitti has more heavily feathered toes

(Hume 1873, Rasmussen & Collar 1998), except in the extremely

worn specimen (NHMUK 1886.2.1.544). In most blewitti the

white tarsal feathering continues uninterrupted onto the toes, while

in brama the more mottled, dingier tarsal feathering stops more

abruptly at the top of the toes, with only sparser bristles on the

toes themselves. Although the extent of feathering on the tarsus

and toes is often highly variable within an owl species (as it is in

both Little and Burrowing Owls), the difference in this feature

between the Forest and Spotted Owlets seems quite constant,

allowing for the effects of wear. The toes and claws of blewitti

appear noticeably heavier (and the latter longer; see below) than

those of brama.

Although the four traditional external measurements (culmen,

wing, tarsus and tail lengths) overlap broadly between blewitti and

Table 1. Summary statistics for measurements3 (mm) of Heteroglaux blewitti and Athene brama skin specimens (sexes combined) and results of

Principal Components Analysis'3 on these variables.

a I = length, w = width, h = height

b Eigenvalues and percent variance explained for PCI-3 on external measures: 4.0, 44.3%; 1 .6, 17.9%; 1 .3, 14.3%, respectively; for PCI-2 on skeletal measures: 3.1, 61.6%; 1.1, 23.0%, respectively; for PCI-3 on wing

formula measures: 8.7, 54.4%; 2.5, 1 5.6%; 1 .6, 9.8%, respectively.

Forktail 29(2013)

81

Phenotypic evidence for the specific and generic validity of Heteroglaux blewitti

2 -

rsi 1 -

u_ 0 -

-1 -

-2

Factor 1

Figure 1. Graph of individual component scores on PC 1 and 2 for

principal components analysis on measurements of Heteroglaux

blewitti (diamonds) and Athene brama (circles). (A) external; (B) wing

formula; (C) skeletal from x-rays.

Figure 2. Wing formulae of Heteroglaux blewitti (solid lines, A,C,E) and

Athene brama (dotted lines, B,D,F). (A,B) shortfalls from wing point of

PI-10; distance from tips of individual feathers to (C,D) notches on

inner webs of PI -4 and (E,F) emarginations on outer webs of P2-5

(descriptive statistics presented in Table 1).

brama, the taxa differ strongly in several other external

mensural characters, even though sexes were combined owing

to the small sample of blewitti (Table 1). External measurements

that do not overlap between the two species are: width between

nares, bill height and lengths of middle and hindclaws (Table 1).

A PCA of external measurements (Table 1, Figure 1A) shows

that by far the greatest proportion of the variance is explained on

PC 1 by a contrast between middle claw length, width between

nares, and upper mandible height vs tail and wing lengths.

Complete separation between the species is attained on this axis

(FigurelA).

Several differences exist between the wing formulae of blewitti

and brama, although most measurements overlap at least minimally

between the species, and the sample of blewitti is small (Table 1,

Figure IB). In blewitti, P7-P10 each have a smaller shortfall, i.e.

the feather tips fall closer to the wing-point, making the inner wing

broader than in brama (Figure 2A,B). In blewitti, P7 is never as

short as PI, while in brama PI and P7 are approximately equal.

The emarginations on the outer webs of P2-P4 and the notches

on the inner webs of P 1 -P4 are all closer to the tips of the individual

feathers in blewitti (Figure 2C-F); measurements of emargination

position did not even overlap between the species (Table 1). Finally,

14 of 23 brama do not show a distinct notch on P4, whilst all six

blewitti examined have a definite notch on the inner web of this

primary.

A PCA of wing formulae showed that by far the greatest

percentage of the variance was explained by Factor 1 (Table 1),

which was mainly a size axis, on which shortfalls of P3 and P4 were

not strongly correlated, and those of PI and P2 were weakly

negatively correlated. All blewitti had negative Factor 1 scores, while

the scores of all brama fell above -1 on Factor 1, reflecting the

smaller shortfalls of the inner primaries and notch and

emargination distances of blewitti (Figure IB).

82

P. C. RASMUSSEN & N. J. COLLAR

Forktail 29 (2013)

Osteology

Despite the similarity in plumage of blewitti and brama , there are

several major osteological differences (Plate 1 ) between blewitti and

all three species normally recognised in Athene (including the highly

polytypic A. \Speotyto\ cunicularia).

Relative to Athene , the nasal process of the premaxillary of

blewitti (Plate 1 A,B) is expanded anteriorly; the culmen ridge (Os

nasale) of the premaxillary is more arched; the distal tip of the

premaxillary is longer and more caudally directed, so the rostrum

of blewitti is heavier and more strongly hooked; the narial openings

are larger and more ovoid; the mandibular symphysis is broader;

and the entire mandible is somewhat heavier.

The frontals of blewitti are much broader both anterior and

posterior to the supraorbital process than for any Athene (Plate 1

A,B), so that the skull of blewitti strikingly resembles that of

Glaucidium and Ninox superciliaris ; the latter Malagasy species has

been wrongly placed in Ninox , and is closer to Athene-. H. F. James

and S. L. Olson, pers. comm. 1997; Wink etal. 2004). The posterior

portion of the interorbital roof of blewitti is not wider than the

anterior portion, unlike Athene. The lateral rim of the frontal

anterior to the supraorbital process is greatly inflated in blewitti

compared to members of Athene, similar to but even more so than

in G. cuculoides, N. superciliaris and most other small owls. The great

inflation of this region is visible in x-rays of other blewitti specimens

as well. The lacrimals of blewitti are very large relative to those of

Athene, but like them (Ford 1967) are short and do not contact the

jugal bar; those of the extracted skull of blewitti are detached from

the skull, but their position relative to the jugal bar is confirmed by

x-rays of all blewitti specimens. The maxillopalatines of blewitti are

large, with straight medial edges that nearly contact each other for

most of their length, unlike in Athene , where the maxillopalatines

are more triangular in shape so that they only contact each other at

the apex. The palatines of blewitti are relatively short

anteroposteriorly as in Athene but are more expanded posteriorly,

in the latter respect being similar to G. cuculoides. The supraorbital

processes of blewitti are better developed than in most other Athene

specimens we examined. The temporal fossa is much deeper in

posterior view in blewitti than in brama.

The quadrate of blewitti has no intercapitular groove, the lack

of which is apparently otherwise autapomorphic for Athene

including Speotyto (Ford 1967); but the articular surface of the

external capitulum is longer than in Athene, being similar to that

of G. cuculoides. The otic process of the quadrate of blewitti is longer

than in other owls examined, and the mandibular articulation is

broad. The socket for the quadratojugal of blewitti is long and more

strongly twisted externally than in brama, similar to that of G.

cuculoides.

The humerus of blewitti is slightly longer and heavier than in

brama, while the ulna of blewitti , although not longer, is

substantially more robust. The leg proportions of blewitti (Table

2, Plate 1 C,D) are unusual in that the hindlimb is much more

gracile proximally than distally: the femur and the proximal end of

the tibiotarsus are heavier than those of brama, but not markedly

so; however, the distal end of the tibiotarsus and the entire

tarsometatarsus are greatly enlarged and especially broadened

relative to those of brama. In addition, the single measurable femur

of blewitti is longer than that of brama-, the tibiotarsus is

approximately the same length in both; and the tarsometatarsus of

blewitti is shorter than that of brama, while the combined length

of these three elements is roughly the same for the two species.

The shaft widths of each of the leg elements in brama are very

similar to one another, in strong contrast to the situation of blewitti,

in which the tarsometatarsus shaft is much wider than that of the

femur. In comparison to blewitti , G. cuculoides\\2.s all leg elements

more uniformly stout; the femur and proximal tibiotarsus heavier,

the tarsometatarsus similar in breadth but considerably longer. The

Factor 1

Factor 1

Figure 3. PCAs of skeletal measurements of selected surniine owls.

Athene noctua, noc; A. brama, bra; Heteroglaux blewitti, ble; A.

cunicularia, cun; Ninox philippensis, phi; N. solomonis, sol; N. scutulata,

scu -, Aegolius funereus, fun; and Glaucidium cuculoides, cue. (A) skull;

(B) forelimb; (C) hindlimb.

H. TAYLOR, ©NATURAL HISTORY MUSEUM, LONDON

Forktail 29(2013)

Phenotypic evidence for the specific and generic validity of Heteroglaux blewitti

83

Table 2. Limb proportions of Heteroglaux blewitti, Athene brama, Athene noctua, Glaucidium cuculoides, and Ninox superciliaris. Ratios are of

mean measurements.

‘Humerus I + ulna I + carpometacarpus I

Temur I + tibiotarsus I + tarsometatarsus

Plate 1. Comparisons between skeletal elements of Heteroglaux blewitti (NHMUK 1886.2.1.546)

and Athene brama (NHMUK S/1 989.25.4). (A,B) Skulls of (A) blewitti and (B) brama in cranial (top),

lateral (middle), and caudal (bottom) views; (C,D) left femora, tibiotarsi, and tarsometatarsi of

(C) blewitti and (D) brama in posterior view.

H. TAYLOR, ©NATURAL HISTORY MUSEUM, LONDON

84

P. C. RASMUSSEN & N. J. COLLAR

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Phenotypic evidence for the specific and generic validity of Heterogtaux blewitti

85

tibiotarsus and tarsometatarsus ofiV. superciliaris are much longer

and more gracile than for any of the above species. The only

osteological feature of the limbs listed by Ford (1967) as diagnostic

o i Athene (including Speotyto) is the pointed posterior edge of the

outer rim of the middle trochlea of the tarsometatarsus; however,

unlike Athene and like other owls examined, this is rounded in the

single blewitti specimen.

Skull widths, length of wing elements (humerus, ulna and

carpometacarpus) and widths of leg elements (tibiotarsus and

tarsometatarsus) are all considerably larger in blewitti than in brama

(Tables 1 and 3, Figure 1C). The tarsometatarsi of blewitti are no

longer (averaging shorter) than in brama, but are considerably more

robust (Tables 1 and 2). In a PCA of skeletal measurements (Table

1, Figure 1C), Factor 1 is astrongsize axis on which tarsometatarsus

length is negatively correlated; all blewitti scores fall above 0 on

Factor 1, while most brama scores fall below 0.

Intergeneric skeletal comparisons

In PCAs of skull and hindlimb measures (Table 3, Figure 3A-C),

scores for the single available blewitti skeleton fall well apart from

those of any Athene species. On skull measurements (Table 3),

blewitti is closest to Aegolius funereus and Glaucidium cuculoides ,

both of which have relatively large measurements on the variables

included in the analysis (particularly so for width of frontal posterior

to supraorbital process and height of lateral rim of frontal), and all

have Factor 1 scores well above 0. Athene species, conversely, have

small measurements on these variables, and all but a few large

cunicularia fall below 0 on Factor 1. Factor 2 principally contrasts

skull length with posterior frontal width and lateral rim height, and

on this axis blewitti and A. funereus differ strongly from large

cunicularia.

Wingproportions of Athene species and blewitti are very similar

(Table 2, Figure 3B), with scores of all but large cunicularia falling

below 0 on Factor 1, a strong size axis on which skull width is

uncorrelated, indicating that, compared to other genera sampled,

most Athene and blewitti have small wings relative to head size.

On a PCA of hindlimb measures (Table 3, Figure 3C), Factor

1 is a general size axis on which tarsometatarsus explains most

variance, and on this axis cunicularia is the most distinctive group

due to its extremely long legs, while blewitti is well separated from

Athene and close to G. cuculoides. Factor 2 is basically a contrast

between tibiotarsus and tarsometatarsus length with

tarsometatarsus width, and on this axis blewitti has the highest

score, again reflecting the stoutness of this element (see Table 2).

DISCUSSION

The specific validity of blewitti

There ought to be no question about the status of blewitti as a full

species, but in the light of a recent report of a pair composed of a

male brama and a female blewitti producing a supposedly fertile

offspring, and indeed of a population of hybrids which ‘may have a

much wider distribution that could equal or surpass the very limited

one of the Forest Owlet’ (Pande et al. 2011), all of which could be

taken to imply the conspecificity of the taxa, we briefly here

recapitulate and expand the evidence.

First, blewitti differs in plumage and external structure from

brama on multiple characters, including the narial and cere

characters given in the species description by Hume (1873), and

others enumerated in Rasmussen & Collar (1998); our elaboration

above of the external structural differences involves much greater

width between nares, bill height and claw length, plus a somewhat

different wing-shape. It is interesting to note how, albeit with tiny

sample sizes, the seemingly small size advantage of blewitti over

brama translates into a doubling of body mass (241.0 g,n= 1 [based

on ‘8.5 oz’ on label of type specimen and in Hume 1873] vs 110-

1 14 g, n = 2) (Dunning 1993). Second, blewitti possesses a wide

range of osteological distinctions from not only brama but all

members of Athene, including cunicularia. These involve many

cranial characters (some on fused elements) on the nares,

premaxillary, frontal, mandible, lacrimal, maxillopalatine and

quadrate. Of these, the much broader frontal (on cranial view) with

its greatly inflated lateral rim anteriorly is most striking. There are

also differences between blewitti and Athene in hindlimb

proportions, especially the short, very stout tarsometatarsus of

blewitti, and in the conformation of the middle trochlea of the

tarsometatarsus. Third, the song of blewitti is very dissimilar to

vocalisations of brama, and does not support its treatment as a close

relative (Rasmussen & Ishtiaq 1999, Jathar & Rahmani 2002,

Rasmussen & Anderton 2005). Fourth, behavioural differences

include the direct flight and lateral tail-flicking habits of blewitti

(King& Rasmussen 1998, Konig^r al. 1999, Rasmussen & Ishtiaq

1999, Pande et al. 2011, Mikkola 2012).

Finally, even if these many differences between the taxa are

somehow not considered sufficient justification for the specific

distinctness of blewitti from brama, the two forms are broadly

sympatric: brama occurs wherever blewitti occurs. While the

microhabitat where he collected them probably differed, J.

Davidson collected at least 1 1 brama (all now in NHMUK) in the

same region where he took his five blewitti (and other unattributed

Khandesh and ‘Candesh’ specimens in the Hume and Seebohm

collections may have also been collected by Davidson). Although

Davidson took numerous egg sets of brama (Davidson MS) he

never found a nest of blewitti (Barnes 1888), unsurprisingly since

even in Khandesh in the 1880s the Spotted Owlet was clearly much

the commoner species.

Sympatric occurrence is, of course, a prerequisite of

hybridisation, but the evidence presented in Pande et al. (2011) is

impossible to interpret owing to shortcomings in figure labelling,

description and photograph quality. In our experience no specimens

of blewitti or brama can be said to be intermediate in more than a

few characters, and we have seen no specimens of either for which

there is any doubt as to their specific identity. Based on the data

presented above on plumage, other external morphological,

mensural, mass and osteological differences, and the lack oi

intermediates, it is evident that blewitti is a well-marked, distinct

species. Its coexistence with brama in areas where both are resident

strongly reinforces this view. Thus, under any widely accepted

species concept, the Forest Owlet must be considered specifically

distinct from the Spotted Owlet. Until far better evidence is

produced we take the view that hybridisation is unproven and, given

the wide range of differences between the taxa, very unlikely. Even

if occasional hybridisation were proven, wild intrageneric hybrids

are known for several avian taxa (see McCarthy 2006), and these

are not generally taken as evidence of exceptionally close

relationship.

Baker’s (1923) notion of the conspecificity of blewitti with

noctua has never been taken seriously elsewhere, but has never been

explicitly dealt with; hence, we do so here. Although A. noctua is a

variable species, it is considerably more different in appearance from

blewitti than is brama. This is most evident in: the pattern oi the

underparts (streaked in noctua, barred in blewitti ); crown pattern

(streaked in noctua, nearly unmarked in blewitti)-, broader, spotted

frontal semi-collar; less white on face; less banded wings and tail;

smaller bill and claws; and tarsal length and shape (long and much

more gracile in noctua, short and stout in blewitti). We can find no

features in which blewitti resembles noctua more than it does

brama, except for the streaked underparts of juvenile blewitti

(Rasmussen & Anderton 2005). In osteology, brama, noctua and,

except ior tarsometatarsus length, cunicularia are very similar to

each other, far more so than any is to blewitti.

86

P. C. RASMUSSEN & N. J. COLLAR

Forktail 29(2013)

The generic validity of Heteroglaux

Clearly the plumage differences between blewitti and established

members of Athene are insufficient alone to justify the maintenance

o f Heteroglaux. Moreover, while the song of blewitti is very different

from brama in its high pitch, tone and modulation, it does resemble

the male song of noctua in overall quality; and even the song of

cunicularia is somewhat intermediate between blewitti and brama.

As a consequence we abandon the pursuit of generic limits in

blewitti through acoustic evidence: such a line of taxonomic inquiry

is untried elsewhere for fairly obvious reasons of interpretation

(especially as convergence may play a part), and would require the

discovery of very strong differences to be considered in any way

informative. However, we note that while some vocalisations have

been documented for blewitti (Rasmussen & Ishtiaq 1999, Ishtiaq

& Rahmani 2005), to our knowledge only one recording is available

online (AV 16764; http://avocet.zoology.msu.edu/ recordings/

16764), so detailed further analysis of vocalisations of blewitti

cannot in any case be made until a more complete sample of

recordings becomes available.

This then leaves morphological and behavioural differences to

consider. In terms of external morphology, blewitti would not

appear out of place within Athene. However, many osteological

differences involving multiple cranial elements (especially the

greatly widened and inflated frontal and the large, straight-edged

maxillopalatines) and the hindlimb (the extremely stout

tarsometatarsus) separate it from other Athene, and indicate that

plumage convergence or perhaps even mimicry may have resulted

in the relative similarity of external phenotype between blewitti

and brama. It is unusual to find many marked qualitative (as

opposed to mensural or quantitative) osteological differences

within an avian genus, and it is even occasionally difficult to

distinguish closely related genera osteologically.

Extinct island owls placed in Athene show great variation in

size and length of extremities: one {A. cretensis of Crete) was

relatively large, with very long tarsometatarsi (Weesie 1982), while

another {A. angelis of Corsica) had unusually long femora and

robust tarsometatarsi (Mourer-Chauvire et al. 1997), and a third

(A. vallgornerensis of Mallorca) was small, with short, robust

tarsometatarsi (Guerra etal. 2012). However, judging from figures

(Guerra et al. 2012), even A. vallgornerensis had a distinctly less

robust tarsometatarsus than does blewitti. Living and fossil island

owls show no trend in overall size, but do tend to have somewhat

larger feet and claws than closely allied continental species

(Louchart 2005), as does the mainland blewitti to a striking degree.

Behaviourally, blewitti differs from other members of Athene

in its direct, non-undulating flight, and in its lateral tail-flicking.

While these differences by themselves may not suggest distinctness

at the generic level, they provide significant corroborating evidence

to the osteological data.

In summary, the numerous (and in some cases major)

differences in skull and tarsometatarsus morphology between

blewitti and all other species of Athene (including Speotyto ) —

involving many cranial elements, especially the frontals, and the

extremely stout tarsometatarsus — indicate that (although a rapid

evolution cannot be excluded) blewitti seems likely to be distantly

related to the others. Because of this distinctness, coupled with its

unusual flight and tail-flicking behaviours, we consider the

resurrection of the monotypic genus Heteroglaux fully justified,

and far from ‘spurious’ (Weick 2006). Even those predisposed to

recognise very broad genera, and who may thus prefer to maintain

blewitti in Athene , should at least be aware that it is osteologically

much the most distinctive of the group and quite possibly evolved

from an ancient divergence event. Further study involving more

owl taxa, such as phylogenetic analyses based on morphology and /

or DNA, is likely to shed more light on the relationships of blewitti ,

but in the meantime we contend that the generic distinctiveness

of blewitti only increases the urgency with which the conservation

needs of this Critically Endangered species must be addressed.

ACKNOWLEDGEMENTS

Special thanks are due staff of the N atural H istory Museum, UK, for allowing

extraction of skeletal elements from one blewitti specimen, and toj. P. Angle

for removing the blewitti elements. Assistance of various kinds was given by

R. P. Prys-Jones, M. P. Walters, !. H. Cooper, H. van Grouwand O. Crimmen,

the Natural History Museum, UK; M. LeCroy, P. Sweet, G. Barrowclough

and M. N. Feinberg, American Museum of Natural History; the late R. A.

Paynter and A. Pirie, Museum of Comparative Zoology, Harvard University;

S. L. Olson, G. R. Graves, R. L. Zusi, H. F. James, J. P. Angle, F. Grady, C. M.

Milensky and J. T. Marshall, Jr., National Museum of Natural History,

Smithsonian Institution; the late R. W. Storer andj. Hinshaw, University of

Michigan Museum of Zoology; and M. Holmes, National Museum of Ireland.

Funding for travel to museums was provided by the Research Opportunities

Fund of the National Museum of Natural History, and by British Airways,

through M. Sitnick and T. Shille, Office of Biodiversity Programs,

Smithsonian Institution. We are most grateful to referees J. H. Cooper and