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HARVARD UNIVERSITY
RE
LAS
LIBRARY
OF THE
MUSEUM OF COMPARATIVE ZOOLOGY
GIFT OF
The Nuttall Ornithological
Club
MUS. COMP. ZOOL,
LIBRAR
SEP 8 1972
HARVARD
UNIVERSITY,
7
BVIFAUNA OF THE EASTERN HIGHLANDS
OF NEW GUINEA
PUBLICATIONS OF THE NUTTALL ORNITHOLOGICAL CLUB, NO. 12
Editor, Raymond A. Paynter, Jr.
eo PALIN: Ol ICEL:
EASTERN HIGHLANDS OF
NEW GUINEA
JARED M. DIAMOND
PHYSIOLOGY DEPARTMENT
UNIVERSITY OF CALIFORNIA MEDICAL CENTER
Los ANGELES, CALIFORNIA 90024
Dedicated
in admiration and friendship
to
ROBERT MACARTHUR
CAMBRIDGE, MASSACHUSETTS
Published by the Club
1972
PUBLICATIONS OF THE NUTTALL ORNITHOLOGICAL CLUB
The Comparative Biology of the Meadowlarks (Sturnella) in Wisconsin, Wesley
E. Lanyon, 1957. 67 pp., 31 pls.
2. Comparative Breeding Behavior of Four Species of North American Herons.
Andrew J. Meyerriecks, 1960. 158 pp., 15 pls.
3. Structural Adaptations of the Head and Neck in the Black Skimmer, Rynchops
nigra Linnaeus. Richard L. Zusi, 1962. 101 pp., 44 figs.
4. The Rough-winged Swallow Stelgidopteryx ruficollis (Vieillot), A Study Based
on Its Breeding Biology in Michigan. William A. Lunk, 1962. 155 pp., 19 figs.,
3 pls.
5. Ecology and Bioenergetics of the Long-billed Marsh Wren, Telmatodytes palus-
tris griseus (Brewster), in Georgia Salt Marshes. Herbert W. Kale I, 1965.
142 pp., 61 tables, 22 figs.
6. Communications and Relationships in the Genus Tyrannus. W. John Smith,
1966. 250 pp., 3 tables, 51 figs.
7. Life Histories of Central American Highland Birds. Alexander F. Skutch, 1967.
213 pp., 1 table, 6 figs.
8. Breeding Biology of the Blue-faced Booby, Sula dactylatra personata, on Green
Island, Kure Atoll, Cameron B. Kepler, 1969. 97 pp., 42 tables, 32 figs.
9. Species Taxa of North American Birds. A Contribution to Comparative System-
atics. Ernst Mayr and Lester L. Short, 1970. 127 pp., 8 tables.
10. Studies of Tropical American Birds. Alexander F. Skutch, 1972. 228 pp., 2 tables,
15 figs.
11. Communication and Other Social Behavior in Parus carolinensis. Susan 'T.
Smith. 1972. 125 pp., 6 tables, 22 fies.
12. Avifauna of the Eastern Highlands of New Guinea. Jared M. Diamond. 1972.
438 pp., 19 tables, 42 figs., 4 maps.
Printed in the United States of America
© Copyright 1972
By Nuttall Ornithological Club
MUS. COMP. ZOOL,
LIBRARY
SEP 8 1972
HARVAKD
UNIVERSITY
Publications of the Nuttall Ornithological Club may be obtained from the Nuttall
Ornithological Club, c/o Museum of Comparative Zoology, Harvard University,
Cambridge, Massachusetts 02138
CONTENTS
MEN Paresh Tey LENG, URIS DINGS: fh cited Pee ey 2 oss 8 xin goatee «gegen gibt aed oN vine
I. ORNITHOLOGICAL STUDIES IN THE EASTERN
LEG SUL IN | Os aaa ae 2 a ae a ee gee ar or
SOUMREMG Om HINMORWEATION. 3) 5 cco dn eo) aA ware tahbon o ptoceid © KucGGip ager. o
AIP RUA ROTTS Qos Rae ete 0 ee eat ep Me ag BR SO lay ct «hae Nerve, Scars
EAPELO Gs eo eee eee eee ee We phen aot ee eo. ee
IDPSGRIPTION OF CORLEGTING LOGARITIBS, 4.505 ¢00 2 ce0e sess aslo
eMC Pall COTM TCG: s.< 75 ga anh Sees ee ie we es Me
Sybre GING MOOI ES, Sart Hie aus. vy PAS mene ivan inieus cele hoses
ACE POS COB bel 59 CAN ON if ro ee ane lee eo PRM Bl os, PEA Pe Ae
i SOME GENERAL FEATURES OF THE EASTERN
MC Teale AUN IDS PAWN © als ecole otip she aloe ol cate wp ete ais
EP UCEIENESS: OFF IDISTRIBU TION pe Gc 6 & obs %. deeiko.0-s ceded avg eles arses
“Drop-omis ay the Eastern ioinlands:...5.44 staves «cte< ea oes
Local “Drop-outs ,.and Rare‘or localized Species*...... «455.
BEGOLOGICATS SORDING IMIEGCHANISMS: .0.00 50022 verges cee s bese e
Pe patil SeOnC SAMO fae) ie amy. «wees oe oe eae
UNC (i016 (eR RS eee eee eer tc Sree i ar ee ee arr
eID MCciile> ava setes acs, ug Hare eons cet aue unin pe oe el oth, cota ee
Wertiicall cchistre Dion 20s ides ay hg edo ges ee chee 8
MUROD PULL Dts dre 6 fn ade Yee pes ea Gl ap ycdeah a Oetin Gs au ba! qesere
GheckensOara all OPAUly ims citn.c cose a his Ve Sash ee ee
2. Nonspatial Sorting Mechanisms (Food and Foraging Differ-
NIC.) SEE een <a ere rt ee Coe Seren ei ee eee Oar
pe, Memnperdl CONC OMMOM, fait ccd eet a orep tater os cae em
Some General Features of Ecological Sorting Mechanisms ....
ZOOGEOGRAPHICAL AFFINITIES OF THE EASTERN HIGHLANDS
SUT ERAS IO, OB ReaD TEE agit, ie A Ae oe eer eee OD ee Reha ee
Pee DING INON=BOREST AVINAUNA 4.5 5.050456 00s a wee aawea ye
Cres OMUMOSIMON, 2.7 Stix se ascelgihe ¥ haw) Wynn Se ewsh~ oN eves s
Origin of the Non-Forest Avifauna of the Eastern Highlands . .
Piomectine. Dy, INOM-FOLESG SPECIES! sicko d sin ia.s ew A ed has
Vv
46
a0
or
-_~ —_— -_~
SS Go =P)
QO 2) f
my Sy Soy) aa]
—
CONTENTS
MIGRANTS
RS He" 2 Sse ES, 4818 Fe gee ela seinios lei 6) 14, \@. ca! | sy i0) (ai! (o) 0) (9) ce) ui) ce bay Mn ca teal ee era lrah eli ava(el te. cib) eee
BirD FEEDING ASSEMBLAGES IN FRUITING AND FLOWERING TREES ..
RAINFALL
So et NS Bee SRY 1S) 0) le hee aA esi lee: | (0. /en'6) 16! eel <6) \6) wer ka: en eel ke Mega Mallen elisa eel ec ciate Menene
oe be ie 9! e, .s) @) Ven (me) eo! en Siig 8
278) ) 2 a8. 8) ce lee Eater Te) 6 ar Ney Alin t lol ie “Bs\'e/ 6) fe. ve” elim ts) Teltisivies (s)he: Leis) ta) late tei aeteuin) al fo elias ante
DY. SPECIES ACCOUNTS
Casuarlidae: cassowaries
IFoo ian of VSG oor Cais oY ae ny ee ere Cn er |
Phalaerocoracidie COnmOraMts .... i. 0.4. -sc8s o1 + ee ee
UMMMTONC ASS MINING AS. ses.a. s ests aa 2 adeoiae y's aa
PMuderdde: nerons aie bitters +....2em... sad eee ee
Threskiornithidae: ibises and spoonbills”.:.<aaaks os ta eee
/MreTHIG ACs CHUCK AIG: CCESEn i... en. Aes. he eee ee eee
ecipiecdde: hanks andieaeles: 2h sree .2< hie eek eee
RallGonne vetoes. Ses. ack oes coe Se ee eae
Mesapodidae: mesapodes or brushwtunkeys. ..< 047.2. 5ns 6.
Pbasianidae: quail! and: pheasants, 2 «.cikes 2 2she40 ast Se
cGommmerdac: duistand: Quail” yc 4 oi, soe «se oe eee Reena
ARCA lies" een SP exe. 4s, Sas. des akan eta gk ee A en ee
Caneel aes: OIOVENS. Ao. Adds cd esl oa ieee ope £ak nein ee
ScOlopacidae: saldpipers, cunlews ald: smipe = ..47...a.9 som
Glarcolidae: pratencoles: 72. ake. wi i etme Fe amon ea
Raidaes cull sci Wenn avis, < We aoe she areere et Aes bie een
COMMIT AS: PUG COMS wn tera a ee he eee mts Gatto yee ace tee
Fe SHC AGUCLACS PAM OUS: Sig ahr aie ayn aerety saa: <r arr ee ee ow Sere Noe
Crnremmlmelre se CME WOSe, Siccanexa RE as « Us SER eos Bekins eet
aieyro mid ae: arm OWS si snccnhs Bara sn 5 oe lh ie Ria en eee
SUMO TOT OWIS: fare et AUR acy He Nel + <2 urd a) EAion Seta eet acar secgetio
RoOcamordae: GCOP MOMS war yece keene Rave airtel «ve eR ree ang
ecouiclidae: Owletmionupans: ass. sea eta. ses oe ee
Capmmmmlgidac: moadtsuckers: 26 2-245 sos: 12s ee oe ee
ApOGIdae> “Swiis oy oo Rt wae ee cet es ee be Wem ee wale a
Hemiprocnidae: crested RVVILLLS Meecee ey AACR panies <, cd Meee ene ares Gee
Aliceciimmidae moms ene ki hate 4.0 hw Aes OR Se en ee
Mietopiciicn Wee Catcie™ (a a.ie stm. adhe ye=e-( aidee ne eee
COmICHOAC: DOUGIS: < oue ves o ks oe arias. Goel ioe Nek ere ee
Bircerotidaes MOM Olles 1. Sak bcd. acne Abe Does wae ae a ees
Pittidaes, PIS, ois Gace ee Es Ae as Vind eee
MO scliGlaes eTGhey pahuaes Sees ak ese eR eee Oa Re 2 ne
Plu WiC es SW AULO WES. «sock > somal 5 A.W wee cook Geet ae Gime, pe eee ee
Campephagidae: cuckoo-shrikes ......-....++ +000 ever ee.
Motacillidae: pipits and wagtails ......................04.
Vi
CONTENTS
Muscicapidae: old wore masect Caters ....,.40:c0rs err ae aes
CeCe chet. LULL INGE ate rea ace ov av ka ee we oe ee Pow ee ee os
SmenoMmchiNge: LOMMUMMEIS opines sat oe snes eco eae eee
IME MITER VON: WaT NGS co sucncus oak dco viacee ce dle aia nip cope ounce
Ses Wa OTS meting ay ies Ciba aly h'e aioummrpenel tue ares
Muscle@prpe: fivcatChens aise dsc ta wr. boa age ands mre es
eachivcermieMae: WINTSDLERG gaa sass Gene» coe pang # aR Rian we
Herpes, NITIROS ph cesenseimn ccs yews. <,', % DIESE osc oc PNe Ra wad Bie
PVE CMMIELAG= TWOOG. SUWIONYS: \. ks. tases + sieeve 4+ 0c siedngbuecaneues Alene
Pye CMEMGrneees SCAT MELE Nace Stirs hatentas ois (sac mists FO saad gs SAG ete OREN
(TES ta TSEC CoN is ee cn SP
Grabindae: mudinest. builders: .5..c20 4000s eee ns oo one bs see wee
Cracticidae: butcherbirds and bellazmappies .......2......05
RGR Mere (NOMOOS. 2hilaG-4, ane aakae <.bicwaach SPs sat poe aumaie nial 4 vere
(UGTA OE CS 61 1,0) a aa ee ee eee ee
Paradisaerdae> birds Of patadise a... et. cal wissen on ble wm ee
Generic revision of the Paradisaerdae .. 0.050. .0 0% 00 ee nee
Pirlomoraynchiciae: DOWErDIROS 4)... sy ssmess o> Sloe Aa ieee
INCOSTEIC de? Atistkaliam 1nthatChes: 2c... son Sis-ca0e.0n4 Mele « vale
PE CUROINING Gs GUNNS). ee Sera chan x coves aon SEN LRG eee
Michypriaciel ace” OME VGatenS: xg oh vis dn ous & aunts wt ephaclen wet
IGACICIAC? HOWErPCCKCNS: sh oc ar We = as 4 olka Ree eee
PG HET CO PUA WITCH EVES aye cac i icine es oss any oA Sranyid oe ea ee wi
Bscmudidae: waxbulls, amd matnikims. ......<. 0.1.6 oases ev.
eee Bags CRB GIG EDY 3435... paedinwwe aioe Aah en Gee eS
¥.
[NOLL Se ee ee A ee ee eae 8 a ee ee gee ee
Vil
INTRODUCTION
This monograph is a study of the avifauna of that part of New
Guinea known as the Eastern Highlands, i.e. the Central Range be-
tween longitudes 143°E and 146°E and above an altitude of 1,350 ft.
The discussion is based initially on my collections and studies made
during four expeditions to New Guinea between 1964 and 1969, but
the records of other workers in the Eastern Highlands are also sum-
marized. The first part of the text (I, pp. 3-15) gives a brief synopsis
of the ornithological exploration of the Eastern Highlands and de-
scribes my collecting itineraries, methods, and localities. ‘The second
part (II, pp. 17-94) analyses some features of general ecological and
evolutionary interest in the avifauna of the Eastern Highlands:
patchiness of distribution, ecological sorting mechanisms (niche differ-
ences between congeners), a speciation mechanism in montane birds,
zoogeographical affinities, altitudinal distribution, the breeding non-
forest avifauna, migration, bird assemblages in fruiting and flowering
trees, breeding patterns, and bird classification by New Guinea natives.
A summary of this second part is given on pp. 92-94. The third part
(II, pp. 95-412) consists of individual accounts of the 354 species
recorded to date from the Eastern Highlands. ‘This is followed by a
list of relevant literature (IV, pp. 413-419) and an index (V, pp. 421-
438).
‘The manuscript was accepted by the Nuttall Ornithological Club
on 20 September 1971.
I
ORNITHOLOGICAL STUDIES IN THE
EASTERN HIGHLANDS
SOURCES OF INFORMATION
The backbone of the island of New Guinea (Maps | and 2) is
formed by a central mountain range which rises to nearly 17,000 ft
and which runs uninterrupted for about 1,000 miles from near the
southeastern tip of New Guinea west to the head of Geelvink Bay
(longitude 135°E). There are no known north-south passes under
5,000 ft in the Central Range. In two areas, namely, in the Eastern
Highlands and in the Baliem and Haga Valleys of the Snow Mountains
of western New Guinea, the Central Range broadens into a system
of valleys with dense human populations. New Guinea’s remaining
mountains are the mountains of the Vogelkop Peninsula in the
far west, and a series of isolated ranges or “mountain islands” along
the north coast (Van Rees Mountains, Cyclops Mountains, North
Coastal Range, Adelbert Mountains, and mountains of the Huon
Peninsula) which are separated from the Central Range by lowland
basins. In this monograph the term “Eastern Highlands” is arbitrarily
taken to mean that portion of the Central Range between longitudes
143°E and 146°E. These limits are chosen partly for practical reasons
arising from availability of information, but they also correspond ap-
proximately to a natural zoogeographic unit which is weakly distinct
from the portions of the Central Range farther to the west and to the
east, as discussed on pp. 46-48. Coverage in this monograph is further
restricted to elevations above 1,350 ft, my lowest collecting station.
The area of New Guinea covered (Map 3) thus extends approxi-
mately from ‘Tari in the west to Kainantu 200 miles farther east, and
from the Schrader Range in the north to Lake Kutubu and Mt.
Karimui 120 miles farther south. ‘The highest peak of this area is Mt.
Wilhelm (ca. 15,000 ft), with numerous other peaks exceeding 12,000 ft
(e.g., Mt. Hagen, Mt. Giluwe, and Mt. Michael). The drainage is by
tributaries of the Sepik and Ramu Rivers in the north, and by
tributaries of the Purari and Kikori Rivers in the south. The principal
towns include Goroka, Mt. Hagen, Kainantu, ee Mendi, Minj,
and Wabag. The zoogeographical area which I call “Eastern High-
lands” includes, but is larger than, the political subdivision of the
‘Territory of Papua and New Guinea called the “Eastern Highlands
District.”
The following sources of information concerning the avifauna of
the Eastern Highlands were available to me:
The first systematic collections of birds in the Eastern Highlands
3
STUDIES IN THE EASTERN HIGHLANDS
‘pal eoIpur OsTe oie (SUTRJUNOW
yejty ‘surejunop neiuey) doyasoA oy) jo pue (a8uey uoweyM ‘sulejUNOy, SoztoFY “vaUIND, MON Ua}seaqyINOS
‘spur[Ysip{ Uloiseg ‘sureyuNOy Moug ‘sureJUNOJ puRyADAA) VSULY [VOUaD IY) JO suoTsa1 poureU ATLIeIQIGIe aUIOg “eTNS
-uludg UONFZ pur ‘suTeJUNO;L IWaqjapy ‘asury y[eiseoy YWAoN ‘sureyunop sdopAyD ‘surequnoy sooy uURA “eyNsut
-Udg UowuepueA, ‘doyfaso0A ay) :,,spuryst urejUNoU,, 10 sa8uRI parefOSI [RIAs a1v UONRAI[a IIMOT JO svaie Aq 1
wo pojeirdas pue asury [emusaDy ay} Jo You sy) OF, “sam ay) Ul Avg YUTATIeD Jo pray IY) 0} dn urd}seayINoOs
ay} wom paidniiajurun spuajyxa oSuey [enue oY], ‘veuINS MON Jo sasuei ureyunow [edpug ‘| av
DEUIND Man
jSDayNoS
‘SII BOZIaH
VIASNINSd NONH-~s
as _ “SLW SdOT0A0>"
x ‘SLW S3guy NVA
; VYIASNINAd NANWWVONVM 7
Q 4oyonbe
STUDIES IN THE EASTERN HIGHLANDS
‘}XO} OY] UL PoUONUSUT soanI[RI0O] SuNoaT[oo pue ‘spurysr ‘suMmo} ‘staat Tediourid ay} Jo awIOs Jo suoNwI0T °*Z dvjX
pinsuluad
yao, add9
. 2
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nd
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Apg jpjoquinyy<e Avg Spunist
: dae ea v) pk uodaniow y
Y3ANIY SYNENSAGI
H3AIN ONVESSNVN see
0 4oyonba nce s
> NY
400E!
STUDIES IN THE EASTERN HIGHLANDS
were carried out independently by three workers in the early 1950's in
the Wahgi Valley and surrounding mountains (see Map 3 for these
and other collecting stations in the Eastern Highlands). E. ‘T. Gilliard
of the American Museum of Natural History led expeditions in 1950
and 1952, the results of which were reported by Mayr and Gilliard
(1954). Gyldenstolpe (1955) described the collections he made for the
Stockholm Museum in 1951. Sims (1956) analysed birds collected by
Shaw-Mayer for the British Museum (Natural History) in 1950 and
1951. Almost all the material of these three collectors was obtained at
elevations above 5,000 ft. ‘Their surveys of the forest at higher eleva-
tions were sufficiently exhaustive that subsequent workers have been
able to add only two species to the forest avifauna breeding in the
Eastern Highlands above 6,000 ft.
In the Schrader Range on the northern rim of the Eastern High-
lands a small but important early collection on the 6,700-foot Schrader-
berg was made in 1913 by Biirgers as part of the Sepik River Expedi-
tion of the German colonial administration. It was studied by Strese-
mann (1923). ‘The remaining collections of this large expedition were
obtained at low elevations along the Sepik River and on mountains
(Hunsteinspitze, Lordberg, Etappenberg, Maanderberg) farther to the
west, which fall outside the area covered by this monograph. Another
collection in the Schrader Range was obtained by E. T. Gilliard in
1964 at 5,000-8,600 ft and was discussed by Gilliard and LeCroy (1968).
Collections for the Commonwealth Scientific and Industrial Re-
search Organization of Australia (C.S.I.R.O.) were made by Schodde
in 1961 at Lake Kutubu (2,450 ft) and Mt. Giluwe (ca. 5,900-11,500 ft)
and by Hitchcock in 1963 in the Kubor Range (ca. 6,400-12,000 ft).
The Lake Kutubu collection was described by Schodde and Hitchcock
(1968), while the Kubor collection was briefly discussed by Hitchcock
(1964), who kindly provided me with further unpublished records
based on it.
Unpublished information from other observers was also available
during the preparation of this report. Dr. R. N. H. Bulmer generously
provided detailed summaries of his extensive unpublished observations
between ca. 3,000 and 12,000 ft, principally from two areas: the
Kaironk Valley of the Schrader Range, and the Kyaka area from Baiyer
River to the northern slopes of Mt. Hagen. Major H. L. Bell furnished
records from several localities. ‘The New Guinea Bird Society News-
letter, published monthly since 1965, was the source of some additional
records by other naturalists resident in New Guinea.
My studies were carried out in the course of four expeditions (1964,
1965, 1966, 1969) to the southeastern part of the Eastern Highlands
at elevations between 1,350 and 12,300 ft. Most of the new distribu-
tional records and material of taxonomic interest were obtained below
5,000 ft, because the nearest localities at which the hill forest (2,000-
STUDIES IN THE EASTERN HIGHLANDS
5,000 ft) avifauna of the southern watershed of the Central Range had
been exhaustively surveyed were 400 miles to the west (Snow Moun-
tains) and 150 miles to the east (Wharton Range). Also, previous
taxonomic publications on Eastern Highlands material discussed little
material under 5,000 ft, except for the report of Schodde and Hitch-
cock (1968). A summary of my itineraries, methods, and collecting
stations follows.
ITINERARIES
The 1964 expedition was conducted jointly with Dr. John W. Ter-
borgh. From 9 June to 28 June we worked in the vicinity of Okapa,
principally at the villages of Miarosa and Awande (Map 3). Between
29 June and 19 July we surveyed the avifauna of Mt. Michael, in-
cluding the collecting stations of Lufa and Mengino. On 20 July we
set out overland from Mengino through the territory of the Gimi
people for the Karimui Basin, which ‘Verborgh reached on 27 July.
I stopped en route to observe at a campsite on the Sena River between
25 and 28 July, and arrived at Karimui on 29 July. From then until
17 August we remained at Karimui Patrol Post, surveying the avi-
fauna of the Karimui Basin. We returned to Okapa for the period
18-27 August, spending part of this period at the nearby Okasa forest.
In 1965 I began observing in the Okapa area on 13 June and worked
there until 27 June, dividing the time between the village of Awande
and the Okasa forest. he remainder of the expedition, which lasted
until 15 September, was spent in the Karimui area. Studies were
carried out at Karimui Patrol Post itself 1-5 July, 10-18 July, 2-7
August, and 11-15 September. Bomai was reached by Cessna airplane
and observations made there between 6 and 9 July. My lowest-altitude
collecting station was Soliabeda, which I reached overland in two and
one-half days and where I worked from 19 July to 1 August. ‘The period
from 8 August to 9 September was devoted to altitudinal censuses on
the west peak of Mt. Karimui.
The 1966 expedition was mainly concerned with exploring the
North Coastal Range (Diamond, 1967b, 1968, 1969) and falls outside
the scope of this monograph. No collections were obtained in the
Eastern Highlands, but some observations were made at Karimui,
Okapa, and Goroka on 20-24 June and 14-15 September. During the
1969 expedition, studies were made at Astrolabe Bay, Karkar and
Bagabag Islands, New Britain, and Mt. Albert-Edward in southeastern
New Guinea, all lying outside the Eastern Highlands. However, the
avifauna of Mt. Albert-Edward is very similar at the species level to
that of the Eastern Highlands, and analyses of stomach contents and
some behavioral observations from Mt. Albert-Edward are cited in
this monograph.
STUDIES IN THE EASTERN HIGHLANDS
‘poulftopun are sornipesoy Apnys Aut jo sowieu oy, 7X9} ay} UT pauOoTjUaUT satjITRIOT SuTIa][OI pur
‘SUTRJUNOUL “staATt ‘sosey[IA Jediourid oy Jo suonvoo, ay] YIM “vauINS MON JO spur[YysipP, Ussisey oy, “| AVI
3.9 40S! XS aD Vy¥sesl IV 1 Jovi 5401
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STUDIES IN THE EASTERN HIGHLANDS
MrTHODS
In order to leave much of my time free for field observations, vir-
tually all specimens were collected and prepared by Eastern Highlands
natives: Paran, Koreko, and Esa of the Fore people, Gumini and Uro
of the Gimi people, and Kariniga and Omwai of the Karimui people.
Collecting was carried out with shotguns and with mistnets. I ex-
amined many additional specimens that had been obtained by local
people with bow and arrow, snares, sticks, and other less conventional
means. In the afternoon or evening I supervised the preparation of
specimens by Highlands assistants, sexing each specimen by dissection
and noting the condition of the gonads. In 1965, but not in 1964,
specimens were weighed prior to preparation. Most specimens were
prepared as skins, but some were made into skeletons or preserved in
formalin. Collections of frogs, lizards, snakes, and mammals (mostly
rodents and mistnetted bats, but also some marsupials) were brought
back in formalin. The bird skins are divided between the American
Museum of Natural History, the Harvard Museum of Comparative
Zoology, and the Los Angeles County Museum of Natural History;
all mammals are at the first-named institution, all birds in formalin
at Harvard; the frogs, lizards, and snakes are divided between the
American Museum of Natural History and Harvard; and the bird
skeletons are divided between these two institutions and the United
States National Museum. ‘Taxonomic evaluation of all the collected
bird material was undertaken in the Bird Department of the Ameri-
can Museum of Natural History, where the most extensive collection
of New Guinea birds is located. Some comparisons were also under-
taken at the Museum of Comparative Zoology.
Most mornings and parts of the afternoons were used for field ob-
servations. My observations were mainly devoted to determining, as
accurately as possible, the altitudinal ranges in undisturbed forest, to
identifying songs and calls, and to estimating relative abundances of
species by crude censusing techniques. ‘The most exact altitudinal
records were obtained on the west ridge of Mt. Karimui, which was
completely covered by undisturbed forest. ‘There were no man-made
clearings within two miles of this ridge. Visible evidence and reports
of local informants indicated that the summit of the ridge had not
been climbed previously and that the lower parts of the ridge were
visited only infrequently by native hunters. We made a footpath along
the crest of the ridge from its base at 4,000 ft to the summit at 8,165
ft; made horizontal paths to divide the mountain into eight altitudinal
zones of about 500 vertical feet each; selected convenient reference
marks to divide each zone into subzones ranging in vertical extension
from 60 to 200 ft; and placed camps at the top of Zone 1 (4,200 ft),
at the junction of Zones 2 and 3 (4,750 ft), at the junction of Zones
9
STUDIES IN THE EASTERN HIGHLANDS
3 and 4 (5,390 ft), and at the junction of Zones 5 and 6 (6,500 ft). The
altitude of each reference point was read to £5 ft on an aviation
altimeter on which the standard barometric pressure was set to 29.92
inches. “Phe average of numerous readings on different days under
varying atmospheric conditions was calculated for each point, since
daily standard barometer r sadings on Mt. Karimui were, of course,
unavailable. Upon retrieving a bird the collectors numbered the wrap-
ping paper of each specimen to indicate the subzone in which it had
been taken. By frequently noting the contents of the nets before my
assistants later came to empty them, I was able to determine that their
subzone assignments were completely reliable. I conducted daily cen-
suses of each subzone in turn, and recorded exact elevations whenever
a bird was encountered near a limit of its species range. An average
of four days was devoted to collecting and censusing in each of the
eight zones, and an average of a week was spent living at each of the
four camps. Thus, the relative abundance at vertical intervals of 60
to 200 ft was estimated for all species present on Mt. Karimui, and
the limits of some species (cf., Crateroscelis murina and C. robusta,
p. 27) were determined exactly.
Crude censuses of relative abundance (not of absolute abundance per
acre) were conducted by counting each bird heard or seen within a
horizontal distance of about 50 yards during several hours of observa-
tions each day. If a song was heard several times from the same locus,
suggesting that it came from the same individual, it was counted only
once. Individuals apparently more than about 50 yards away were
not counted, in order to reduce the extent to which the abundance
of conspicuous or noisy species might be overestimated. ‘The number
of individuals of a given species censused was divided by the total
number of individuals of all species censused in order to obtain one
estimate of the species’ relative abundance, expressed as a percentage
of the whole avifauna. A second estimate was derived from collected
specimens, by dividing the number of individuals of a given species
collected by the total number of all birds collected. ‘These two esti-
mates were then averaged, for each species and for each subzone or
collecting locality, to yield the relative abundances which will be cited
in the individual species accounts. For instance, the statement that a
given species “accounts for 2% of the local avifauna” means that two
of every 100 birds locally present are estimated to be of that species.
The relative abundances of the commonest species at a locality were
rarely above 10%, and were generally in the range 4-10%. ‘There were
about 21 species at Karimui Patrol Post (3,650 ft), and about 15 in
Zone 3 of Mt. Karimui (4,750-5,390 ft), with relative abundances
greater than 1%.
These crude estimates of local relative abundance are obviously
subject to systematic errors because of the methods used to derive
them. Species which rarely sing, or whose songs I failed to determine,
10
STUDIES IN THE EASTERN HIGHLANDS
or which tend to stay out of sight, were underestimated in field cen-
suses. Middlestory and upperstory species which are infrequently or
never caught in mistnets were underestimated in the collections. Some
protected species which my assistants were instructed not to collect
beyond a certain number (birds of paradise, bowerbirds, and the large
cockatoos) were underestimated in shotgun collections, but many of
these, fortunately, are noisy birds which were inevitably overestimated
in my field censuses. However, the combined estimate from censuses
and collecting should provide at least tentative first estimates of rela-
tive abundance. Comparisons of the relative abundance of the same
species at two different localities or altitudes should be freer from
distortion, since the same systematic errors would affect the species
at both localities.
DESCRIPTIONS OF COLLECTING LOCALITIES
General Remarks
Botanical descriptions of parts of the Eastern Highlands have been
given by Brass (1959), Hitchcock (1964), and Schodde and Hitchcock
(1968), and of montane vegetation in other parts of New Guinea by
Brass (1941), Archbold, Rand, and Brass (1942), and Gilliard and
LeCroy (1961). Briefly, the original vegetation over almost the whole
area of the Eastern Highlands was forest up to timberline at about
11,500 ft, above which alpine grassland takes over. ‘The total area oc-
cupied by natural non-forest habitats between 1,000 ft and timberline
is extremely small. It consists only of strips along the few lakes (Lake
Kutubu, Lake ‘Tebera, Lake Kandep) and numerous small rivers and
streams in this altitudinal range, plus occasional landslide, earthquake,
and tree-fall areas on mountain slopes. The clearing of the forest for
agricultural purposes has greatly expanded these open habitats. ‘Voday,
because of dense native population, the Eastern Highlands contain
the most extensive areas of (secondary) midmontane grassland in New
Guinea.
The forest itself gradually changes in composition with altitude, the
general succession being lowland rainforest at the lowest elevations,
midmontane forest dominated by oaks (Castanopsis, Quercus), forests
dominated by southern beeches (Nothofagus) at higher elevations, and
subalpine forest and shrubbery with numerous conifers (Podocarpus,
Libocedrus). Perhaps the most striking change in forest physiognomy
occurs at what may conveniently be termed the moss level. ‘The upper
elevations of New Guinea mountains are shrouded much of the time
in standing banks of clouds. Within this moist and poorly lit zone the
forest is stunted, and thick layers of moss cover much of the ground,
trunks, and branches. The lower limit of this heavily mossed zone,
which is said to coincide with the lower limit of the cloud banks, is
sometimes sufficiently distinct that it can be meaningfully defined to
11
STUDIES IN THE EASTERN HIGHLANDS
within 50 vertical feet. Its elevation varies considerably from mountain
to mountain, depending upon local cloud conditions. This moss level
occurs at about 6,500 ft on Mt. Karimui’s west mdse, abr 4,500 tt on
Mt. Karimui’s east ridge, at 8,700 ft on Mt. Michael, and at some
undetermined altitude about 7,500 ft in the Okapa area. Several birds,
such as Ptiloprora guisei, Oreocharis arfaki, Peneothello sigillatus, and
Lophorina (Pteridophora) alberti, have lower altitudinal limits which
correlate approximately with the moss level, while some other species
(p. 67) have ceilings at the moss level.
Specific Localities
Okapa area.—Collecting localities were Okapa Patrol Post (6,600 ft);
Awande (6,000 ft), a native village two miles east of Okapa; Miarosa
(5,800 ft), a native village two miles east of Okapa and one mile south
of Awande; and Okasa, a forestry camp near a village of the same
name, nine miles south of Okapa. The highest collecting elevation in
this area was at 7,500 ft on the ridge separating Miarosa from Awande,
and the lowest was at 3,550 ft on a small river below Okasa. This area
is inhabited by the Fore people, who have achieved medical promi-
nence through kuru or “laughing sickness” (Gajdusek and Zigas, 1957),
an incurable and invariably fatal neurological disease which is largely
restricted to the Fore and accounts for a high percentage of their
deaths. Much of the area around Okapa, Awande, and Miarosa has
been cleared for agriculture and exists as gardens, grassland, or second-
erowth, but the hill slopes behind Miarosa and Awande remain with
their original cover of midmontane forest. At 7,500 ft there is still little
moss in the forest. Correlated with this, Oreocharis arfaki and Peneo-
thello sigillatus are absent, and Ptiloprora guisei is very rare in the
Okapa area. Most collecting at Okasa was done between 3,550 and
4,250 ft in a forest with magnificent tall Araucaria pines. ‘This forest
is a reservoir of cerebral malaria and is presently uninhabited, but
large expanses of grassland around it testify to a formerly dense native
population.
Mt. Michael area.—Collecting localities were Mt. Michael itself, an
isolated 12,300-foot peak forested above 7,000 ft, below which altitude
the forest has been largely cleared; Lufa Patrol Post at 6,300 ft on the
north slope of Mt. Michael, situated in an area which has been largely
cleared; and Mengino, a village at 6,150 ft on the west slope of Mt.
Michael. Our principal camp was located in tall forest at 8,000 ft on
the northwest side of Mt. Michael, with a subsidiary camp in stunted
subalpine forest at 10,200 ft. Heavily mossed conditions are first en-
countered at 8,700 ft, and alpine grassland at 11,200 ft, but patches
and tongues of shrubbery in the alpine grassland are still found near
the summit at 12,300 ft. Terborgh and I reached alpine grassland and
the summit only on one day (10 July 1964), so that my experience of
alpine grassland in the Eastern Highlands is limited. Collecting at
12
STUDIES IN THE EASTERN HIGHLANDS
Mengino extended from the village at 6,150 ft down to a river at 4,600
ft, in partly cut forest interspersed with second-growth and gardens.
Gimi territory —The area between Mengino and the east rim of the
Karimui Basin is hilly and lies between 4,200 and 6,500 ft. Second-
erowth and gardens are found around several small villages of the
Gimi people, but most of the area is still forested. Little time was
available for collecting in this area. However, in passing through a
small patch of mossy forest at about 6,000 ft we met Pliloprora guiset
again, which was absent at the same and even higher elevations else-
where in areas with little mossing.
Karimui Basin —Most of the Eastern Highlands consists of rugged
and steep ridges and mountains. ‘The Karimui Basin, a volcanic plain
on the north side of Mt. Karimui, is of interest as one of the few large
flat expanses within the Eastern Highlands. ‘The basin floor is at about
3,500 ft, has no hills, and is flat except for a few narrow stream gorges.
As shown by several aerial reconnaissance flights, which I undertook in
1965, the basin is effectively sealed off from the rest of the Eastern
Highlands by a ring of mountains which rises to about 8,400 ft in
the central peak of Mt. Karimui and which stands at least 1,000 ft (in
most places much higher) above the basin floor on all sides except
where pierced by narrow river gorges. Thus, the Karimui Basin is an
island of tropical habitat isolated within the Eastern Highlands, a
fact which is reflected both in its endemic bird forms and in the
unusual composition of its avifauna (pp. 53-56). At the time of my
studies most of the basin floor was still covered by rainforest, which
unfortunately is rapidly being cleared in the vicinity of villages. In its
vegetational structure this basin forest is typical of the rainforest of
the flat lowlands near sea level, e.g., in the presence of emergent trees
up to 200 ft tall (“A story” in the sense of Richards (1952)), and differs
from the hill forest encountered at the same elevation at most other
localities in New Guinea. Collecting localities on the basin floor were
Karimui Patrol Post at 3,650 ft, henceforth simply called Karimui;
and Bomai, a small airstrip and native village at 3,250 ft about 12
miles west of Karimui. ‘The locality “Sena River” refers to a camp I
made in undisturbed forest beside a small stream at 4,500 ft. This
camp lay 15 miles east of Karimui and within the basin but part way
up its eastern wall. Soliabeda is a native village at 2,000 ft, nine miles
southeast of Karimui and beyond Mt. Karimui, i.e., outside of the
basin. ‘The village of Soliabeda was only a few years old, so that the
area of land which had been cleared for gardens was still small. The
terrain around Soliabeda is hilly, in contrast to the nearby Karimui
Basin. Below the village of Soliabeda the Wi River, flowing at 1,350
ft, represented the lowest altitude I reached in the Eastern Highlands.
Finally, collections were made on the west ridge of Mt. Karimui itself,
which was reached from the village of Iogoramalu, about six miles
west of Karimui Patrol Post. As described on p. 9, the ridge was
13
STUDIES IN THE EASTERN HIGHLANDS
divided into eight altitudinal zones with the following limits: Zone 1,
4,000-4,200 ft; Zone 2, 4,400-4,750 ft; Zone 3, 4,750-5,390 ft; Zone 4,
5,390-5,960 ft; Zone 5, 5,960-6,500 ft; Zone 6, 6,500-7,080 ft; Zone 7,
7,080-7,620 ft; Zone 8, 7,620 ft to the summit of the mdee at 365 it
The highest peak of Mt. Karimui is a few hundred feet higher than
the summit of this west ridge but is separated from it by a narrow
vertical-walled chasm several thousand feet deep and hence was in-
accessible to me. Zone | was still on the gently sloping edge of the basin
floor, while Zones 2 through 8 were on the ridge which rises con-
tinuously and steeply from the top of Zone | to the summit. A well-
defined moss level occurrs at 6,500 ft, above which the stunted summit
forest is a three-dimensional maze of fallen branches, low trees, and
bushes, all overgrown with cold, wet moss.
Goroka.—A few observations were made at this town, which is one
of the principal commercial centers in the Eastern Highlands, but no
specimens were collected. The elevation is 5,140 ft. All land in the
vicinity has been cleared, and no primary forest remains.
ACKNOWLEDGMENTS
These studies were made possible and were facilitated by generous
cooperation from many individuals and groups at all stages in the
work. It is a pleasure to record my debt to:
Paran, Koreko, and Esa of the Fore people, Gumini and Uro of the
Gimi people, and Kariniga and Omwai of the Karimui people, my
field associates, whose loyalty and efforts under sometimes difficult
circumstances were essential to the field work.
John Terborgh, who jointly led the 1964 expedition and permitted
discussion of his observations in this report.
G. Schmutterer and R. Hornabrook at Okapa, C. Campbell and J.
Burns at Lufa, A. Schultz at Agotu, and K. Mesplay, A. Wright, and
W. Metze at Karimui, whose hospitality and advice made much of the
field work both possible and enjoyable.
Members of the Department of Native Affairs and the Department
of Agriculture, Stock, and Fisheries, Territory of Papua and New
Guinea, for permitting and expediting these studies.
Dean Amadon, who made available the facilities of the Department
of Ornithology at the American Museum of Natural History.
R. A. Paynter, Jr., who made available the facilities of the Bird
Department at the Harvard Museum of Comparative Zoology, and
whose stimulation and advice were essential to the preparation of this
monograph.
Ernst Mayr and Mary LeCroy, who shared their knowledge of New
Guinea birds, and Carleton Gajdusek, Hobart Van Deusen, and Alan
Mann, who shared their experience of New Guinea.
R. Bulmer, W. B. Hitchcock, J. Kikkawa, and H. L. Bell, who fur-
14
STUDIES IN THE EASTERN HIGHLANDS
nished unpublished information concerning their bird studies in the
Eastern Highlands.
R. Mackay of the Territory Museum, who forwarded specimens.
Barbara Burgeson, who prepared the index and read proofs.
The National Geographic Society, Frank M. Chapman Fund of the
American Museum of Natural History, Explorers Club, American
Philosophical Society, Wenner-Gren Foundation, and Society of the
Sigma Xi, which generously provided financial support.
To these and many other individuals and groups, I am grateful.
II
SOME GENERAL FEATURES OF THE EASTERN
HIGHLANDS AVIFAUNA
PATCHINESS OF DISTRIBUTION
“Drop-outs”’ in the Eastern Highlands
The breeding avifauna of montane forest and alpine grassland of
the Eastern Highlands must now be considered well-explored ornitho-
logically. Since the surveys of Gilliard, Gyldenstolpe, and Shaw-Mayer
in the years 1950-1952, only two forest species appear to have been
added to the list of birds breeding or presumed breeding at elevations
above 6,000 ft, viz., the honeyeater Piiloprora meekiana and the man-
nikin Erythrura papuana. Most species breeding above 6,000 ft in the
Eastern Highlands have been recorded or collected there on at least
two separate instances, the sole exceptions being Pachycephala tene-
brosa and Ptiloprora meekiana, which are known in the Eastern High-
lands from a single specimen each. ‘There are undoubtedly further
breeding species to be recorded from the hill forests below 6,000 ft and
from grassland and disturbed areas above this elevation. More non-
breeding migrants and vagrants from the lowlands also remain to be
discovered. However, the addition of more than one or two species
to the breeding avifauna of undisturbed forest above 6,000 ft is un-
likely.
For these reasons it is significant that there are nine species that
have been recorded from the Central Range to the east and west of the
Eastern Highlands but not in the Eastern Highlands, even though the
Eastern Highlands provide suitable habitats. A few of these apparent
“drop-outs” may still turn up, but most of them will probably prove
to be absent. ‘hese absences, and several related phenomena involving
distributional patchiness in New Guinea, require detailed discussion
because of their evolutionary and zoogeographical significance. ‘The
nine apparent “drop-outs” are:
Mieraaetus morphnoides, Little Eagle (Accipitridae); a rare species
in New Guinea, recorded to date from the Central Range in western
New Guinea (Utakwa River, Baliem Valley, Idenburg River), the
Central Range in southeastern New Guinea (Aroa River, Hydro-
grapher Range, Sogeri District, Wau), two of the “north mountain
islands” (Adelbert Mountains, Huon Peninsula), and the lowlands
near Lae. Most of these records are from between 2,000 and 6,000 ft.
Coracina lineata, Barred Graybird (Campephagidae); a patchily dis-
tributed but locally common inhabitant of hill forest between 2,000
and 4,000 ft in the mountains of the Vogelkop, southeastern New
17
FEATURES OF THE EASTERN HIGHLANDS
Guinea, and three of the “north mountain islands’’ (North Coastal
Range, Adelbert Mountains, Huon Peninsula).
Amalocichla sclateriana, Greater New Guinea Thrush (Purdinae);
forests of the Central Range between 8,000 and 12,000 ft in western
and southeastern New Guinea.
Orthonyx temminckit, Logrunner (Orthonychinae); forest at 4,000-
10,000 ft on the Central Range in western and southeastern New
Guinea and on the Vogelkop.
Macgregoria pulchra, MacGregor’s Bird of Paradise (Paradisaeidae);
conspicuous and fairly common in high altitude forest (9,000-13,000 ft)
of the Central Range in west New Guinea, probably at ‘Velefolmin,
and in southeastern New Guinea. It strains one’s credulity to suppose
that Macgregoria pulchra could have escaped detection if it occurred
in the Eastern Highlands. Not only is it conspicuous, but most New
Guinea ornithologists have paid particular attention to birds of para-
dise. Gilliard, Bulmer, I, and others made a habit of exhaustively
quizzing local natives and would surely have learned of Macgregoria
in this fashion, as Gilliard did at ‘Velefolmin (Gilliard and LeCroy,
HYG, p. LO and p.-64).
Climacteris lewcophaea,! White-throated ‘Tree Creeper (Climacteri-
dae or (?) Certhiidae); the sole species of creeper in New Guinea;
forest from 5,000 to 10,000 ft on the Central Range (western and
southeastern New Guinea, Telefolmin) and the Vogelkop.
Ptiloprora plumbea, Leaden Honeyeater (Meliphagidae); a rare in-
habitant of the Central Range in forest at 4,000-6,000 ft (Utakwa River
of western New Guinea, ‘Velefolmin, and southeastern New Guinea).
Melanocharis arfakiana, Obscure Berrypecker (Dicaeidae); the sec-
ond rarest New Guinea bird, known from two specimens, one taken
on the Vogelkop at the western tip of New Guinea in 1867, and the
other taken 1,000 miles to the east in 1933 at 3,000 ft 1n southeastern
New Guinea.
Lonchura monticola and L. montana, Alpine Mannikin (Estrildidae),
a superspecies; alpine grassland at 9,000-14,000 ft in western and south-
eastern New Guinea, where it is common and occurs in flocks of up to
50 birds. ‘The only other ecologically related estrildid in New Guinea,
Oreostruthus fuliginosus, is uncommon and restricted to the edge be-
tween forest and alpine grassland. ‘The alpine grasslands of Mts.
Hagen, Wilhelm, and Giluwe have been explored by several collectors,
and this conspicuous species could hardly have been missed if present.
Of these nine “drop-outs,” the evidence is strongest for the absence
of Macgregoria pulchra and Lonchura monticola; nearly as strong for
Amalocichla sclateriana, Orthonyx temmincku, and Climacteris leuco-
phaea; less strong for Hieraaetus morphnoides, Coracina lineata, and
Ptiloprora plumbea; and inadequate concerning the mysterious Me-
1 Listed as Climacteris placens in Rand and Gilliard (1967).
18
FEATURES OF THE EASTERN HIGHLANDS
Map 4. Distribution of Climacteris leucophaea, the White-throated Tree Creeper.
This species is present on the Vogelkop and on the western and southeastern por-
tions of the Central Range but is absent from the Eastern Highlands, so that its
distribution is discontinuous despite the continuous expanse of montane habitat
on the Central Range.
lanocharis arfakiana. Since some of these drop-outs are uncommon or
rare, the obvious first question to ask is whether there has been enough
work in the Eastern Highlands to justify attaching significance to the
apparent absences. The following comparison shows that the ornitho-
logical exploration of the Eastern Highlands would have been more
than adequate to detect their presence in most cases. In southeastern
New Guinea the First Archbold Expedition collected 3,200 birds in
1933-1934; the collection included eight of the nine species that drop
out in the Eastern Highlands. In western New Guinea the Third
Archbold Expedition collected 4,846 birds in 1938-1939; the collection
included six of the nine Eastern Highlands drop-outs. ‘The total num-
ber of bird specimens collected to date in the Eastern Highlands is
considerably larger, viz., about 7,500, of which a higher proportion
came from medium and high altitudes (where most of the drop-outs
would be found) than in the case of the Archbold Expeditions. In
addition, workers in the Eastern Highlands have been able to supple-
ment their own observations with far more information gathered from
local natives than did the Archbold Expeditions. For instance, my
Fore and Daribi assistants gave me detailed descriptions of all the
birds that they or their relatives had at some time seen or heard of but
which I failed to collect (pp. 90-91). ‘These descriptions included even
inconspicuous, undistinctive, and rare warblers, and species of which
one individual had been caught 10 years ago but which had never been
19
FEATURES OF THE EASTERN HIGHLANDS
seen since. Most of these descriptions are identifiable, and none re-
sembles any of the drop-outs. The Fore and Daribi are unlikely to
have overlooked birds with habits as distinctive as those of Orthonyx
temmincku and Climacteris leucophaea while distinguishing sibling
species in the warbler genera Sericornis and Crateroscelis. Thus, I con-
sider it highly probable that most of the apparent drop-outs are
actually missing in the Eastern Highlands.
One must still ask whether the absences could be related to replace-
ment by closely similar competing species or to the absence of suitable
habitat. ‘The former explanation applies to a tenth species, Melidectes
ochromelas, which is also absent in the Eastern Highlands while pres-
ent to the east and west. As discussed (pp. 388-389), there is an unusual
competitive situation involving three congeneric species in genus Melli-
dectes, and the absence of M. ochromelas is correlated with the pres-
ence of two of its congeners. Coracina lineata, Ptiloprora plumbea,
and Melanocharis arfakiana are members of large genera, and com-
petition may also play a role in their absence, though not in any
simple fashion—i.e., their distributions are not complementary to the
distribution of congeners, as in the Melidectes case. No such factors
suggest themselves in the remaining cases. Elsewhere in New Guinea
Lonchura monticola is the common and sole alpine grassland man-
nikin; Macgregoria, a very distinct monotypic genus, is the common
and sole bird of paradise above 11,000 ft; the creeper Climacteris
leucophaea is the only representative of its family in New Guinea;
Orthonyx temmincki has distinctive habits and no congeners in New
Guinea; and the thrush Amalocichla sclateriana lives at altitudes above
its only congener and overlaps altitudinally only one other thrush
(Turdus poliocephalus), which has a quite different diet (mainly fruit
instead of insects). In the Eastern Highlands the niches of these drop-
outs remain unfilled in their absence.
Thus, as illustrated in Map 4 (p. 19) for Climacteris leucophaea, the
distribution of these drop-outs on the Central Range shows a gap of
about 250 miles in the Eastern Highlands. For the alpine species
Lonchura monticola and Macgregoria pulchra the available habitat
is discontinuous, but for the remaining drop-outs the Central Range
presents a 1,000-mile stretch of habitat with no major barriers. One is
accustomed to encountering drop-outs in the avifaunas of isolated
mountain ranges, but the finding of drop-outs in sections of a con-
tinuous habitat may come initially as a surprise. However, the other
parts of the Central Range also have drop-outs (pp. 46-47). South-
eastern New Guinea “inexplicably” lacks five or six montane species
that are present on the Central Range both in western New Guinea
and in the Eastern Highlands. ‘These are Porzana tabuensis, Pachy-
cephala tenebrosa, the Paradigalla superspecies, Lophorina (Pterido-
phora) alberti, and Archboldia papuensis; the Melidectes nouhuysi-M.
princeps superspecies is also missing but may be excluded by com-
20
FEATURES OF THE EASTERN HIGHLANDS
peting congeners, as discussed on p. 391, Ihe western part of the
Central Range also “inexplicably” lacks four montane species present
both to the east on the Central Range and to the west on the Vogelkop,
viz., Myzomela adolphinae, Erythrura papuana, Coracina lineata, and
Melanoc haris arfakiana. These lists exclude species which are absent
because their distribution complements that of a congener, or which
appear to have split off recently from a close relative and to be in the
process of expanding their range (cf., pp. 33-34). To set these numbers
in perspective, New Guinea has about 170 widespread montane species
or superspecies, so that the proportion of drop-outs in any given part
of the Central Range is only 3-5%. The proportion of widespread
species that drop out is much higher on the isolated mountain ranges
of the Vogelkop and “north mountain islands”: Vogelkop, ca. 8%;
Huon Peninsula, 8%; North Coastal Range, 58%; Cyclops Moun-
tains, 77%; Adelbert Mountains, 82%.
The drop-out phenomenon within New Guinea may be interpreted
along the lines developed by MacArthur and Wilson (1967) for island
faunas. To explain the fact that the number of species on an island
is observed to increase with increasing island size and decreasing
distance from the mainland, these authors postulated that the number
of species in an island fauna represents an equilibrium between ex-
tinction rates and immigration rates. Extinction rates are highest on
the smallest islands, where populations consist of the fewest individuals
and run the highest risk of being wiped out by random fluctuation in
numbers due to transiently poor conditions, such as disease, accidents,
failure in food supply, etc. Immigration rates increase with island size
and proximity to the mainland. ‘he same principles find immediate
application to the avifaunas of New Guinea’s isolated mountain ranges,
which for montane birds represent distributional islands of suitable
habitat in a “sea” of lowland. ‘These principles apply equally well to
any portion of the continuous Central Range, and the only differ-
ences from the situation on the isolated ranges are quantitative. A
species is more likely to be eliminated by chance, and extinction rates
are therefore higher, on a small mountain range with a small total
population than on a large mountain mass such as the Eastern High-
lands. When a species does disappear, immigration rates are higher on
the Central Range, where recolonization occurs from immediately
adjacent portions of the range, than on the isolated mountain ranges.
Thus, the percentage of drop-outs is lowest on the various parts of
the Central Range, higher on the large but isolated ranges of the
Vogelkop and Huon Peninsula, and highest on the smallest isolated
ranges (Cyclops Mountains and Adelbert Mountains). A similar phe-
nomenon is encountered in the distribution of New Guinea lowland
birds, because mountain ranges divide up the New Guinea lowlands
into habitat islands of various sizes. For instance, the flat lowland
strip hemmed in by mountains along the north coast of southeastern
21
FEATURES OF THE EASTERN HIGHLANDS
New Guinea lacks several species, e.g., Pitohut kirhocephalus, Mino
anais, Goura victoria, which are otherwise ubiquitous and common in
the New Guinea lowlands.
The drop-out phenomenon on the Central Range may provide one
of the keys to how the New Guinea montane avifauna evolved.
Speciation involves the breakup of what was originally one species
into two geographically isolated daughter populations, which may
diverge in isolation to the point where they are reproductively iso-
lated and ecologically distinct when they come into contact again. One
may therefore ask, “What provided the necessary isolation for specia-
tions in the mountains of New Guinea?” A glance at a map of New
Guinea (Map 1), with its continuous Central Range and isolated
smaller ranges, might suggest that divergences took place between
populations on the Central Range and populations on the isolated
ranges, with the latter then reinvading the former to generate new
species on the Central Range. If this were the case, one would expect
to find distributional patterns reflecting early stages in the develop-
ment of sympatry after reinvasion, such that one montane species
occurrs on the Central Range and a different but closely related species
occurrs on an isolated range plus the nearest portion of the Central
Range. However, there is not a single such pattern involving any of
the “north mountain islands,” and even the Vogelkop provides only
a few possible examples, viz., Peneothello cryptoleucus and P. cyanus;
Pachycephalopsis hattamensis and P. poliosoma; Ptiloprora erythro-
pleura and P. perstriata. ‘This suggests that the isolates of the small
detached ranges have rarely succeeded in reinvading the large Central
Range. In contrast, there are numerous examples of pairs of forms that
clearly represent every possible intermediate stage in speciation involv-
ing eastern and western isolates on the Central Range. ‘These examples
include: pairs of allopatric semispecies with one member in the
western part and the other in the eastern part of the Central Range
(e.g; Lophorina (Parotia) carolae and L. (Parotia) lawesi, Melidectes
princeps and M. nouhuysi, Lophorina (Astrapia) splendissima and
L. (Astrapia) mayeri and L. (Astrapia) stephaniae); pairs of closely
related species that are sympatric over much of the middle of the
Central Range but only one of which reaches the western tip and the
other the eastern tip (e.g., Rallicula rubra and R. forbesi, Psittacella
modesta and P. madaraszi, Epimachus fastosus and E. meyeri, Toxo-
rhamphus novaeguineae and T. poliopterus); and pairs of closely re-
lated species, one of which is confined to the eastern part of the Central
Range and the other of which is distributed over the whole length of
the Central Range (e.g., Pliloprora guise: and P. perstriata, Paradisaea
rudolphi and the P. raggiana superspecies, Amblyornis subalaris and
A. macgregoriae, and probably Cnemophilus macgregoriae and C.
(Loria) loriae).
These cases suggest that montane speciation in New Guinea can
aa
FEATURES OF THE EASTERN HIGHLANDS
take place, and in most cases has taken place, on the Central Range
alone, despite the absence of major east-west geographical barriers on
this range. The apparent paradox of speciation on a continuous
mountain range may be resolved by the drop-out phenomenon, which
provides the necessary isolation. Evidently the first stage in speciation
is the development of east-west clines and subspeciation on the Central
Range, as illustrated by the subspecies of most midmontane birds
(pp. 00-00). In the second stage an intermediate population dies out,
as illustrated by the Eastern Highlands drop-outs. ‘These local ex-
tinctions may have higher probabilities of occurring at times when the
area of suitable habitat has been reduced by climatic or vegetational
fluctuations, e.g., when a general cooling compresses forest altitudinal
bands or when dry conditions restrict the forest. In the third stage
the now-isolated eastern and western populations diverge to the point
where they become well-marked subspecies or distinct members of a
superspecies. ‘his stage is represented by the Alpine Mannikin, Lon-
chura monticola, which occurs in southeastern New Guinea, has no
population in the Eastern Highlands and is represented in western
New Guinea by the semispecies, or very distinct subspecies L. montana
(= L. monticola montana). ‘Vhe fourth stage is reached when these
isolates come into contact again (e.g., the Parotia, Melidectes, and
Astrapia superspecies cited above). In some cases the rejoined western
and eastern populations will not have acquired reproductive isolation
and will hybridize, leading to very distinct but interbreeding western
and eastern subspecies (e.g., Crateroscelis robusta robusta and C. r.
sanfordi, Rallicula forbesi forbesi and R. f. steint). In other cases,
however, such as those cited in the previous paragraph, reproductive
isolation will be perfected, and as the final stage in speciation the two
forms will gradually reinvade each other’s ranges, or else one will re-
invade the entire range of the other. ‘The ecological correlates of these
east-west reinvasions follow a consistent pattern and are discussed on
pp. 00-00. ‘These considerations may be relevant to speciation problems
in continuous habitats in other parts of the world, such as the problems
posed by the rich montane avifauna of the Andean chain of South
America.
Local “Drop-outs,” and Rare or Localized Species
If in New Guinea one surveys the avifaunas of different mountains
a few miles apart in the same range, one finds that a given mountain
does not support the entire montane avifauna of the region but rather
that some species are missing. Stein (1936) was struck by this point in
comparing Mt. Kunupi and Mt. Sumuri, adjacent peaks in the Wey-
land Mountains, and many examples turned up during my 1966 survey
of four peaks in the North Coastal Range (Diamond, 1968). ‘The pat-
tern of this effect is that the smaller the mountain, the more species
are found to be missing at a given altitude. That is, a mountain 5,000
23
FEATURES OF THE EASTERN HIGHLANDS
ft high obviously lacks all species restricted to elevations above 5,000
ft, but it also lacks some of the species found at 3,000-5,000 ft on larger
mountains. ‘he explanation is presumably the relation between species
diversity and area: the smaller the mountain, the less the area at a
given altitude, hence the fewer individuals in the local populations
and the greater the likelihood of temporary local extinctions. In the
Eastern Highlands this tendency is illustrated by Mt. Karimui, a peak
8,400 ft high rising above the Karimui Basin (3,500 ft) and about 10
miles from the nearest peak over 8,000 ft. We collected and netted
many specimens on Mt. Karimui, and my field observations there for
a month were supplemented by those of seven Fore assistants, three
Gimi assistants, and five Daribi assistants who were intimately familiar
with the New Guinea midmontane avifauna. Thus, our survey was
probably complete, and most of the expected midmontane forest
species including rare ones turned up. However, four species that are
widespread elsewhere in the Eastern Highlands were not found and
may be assumed to be absent on Mt. Karimui, viz., Ducula chalconota,
Melampitta lugubris, Heteromyias albispecularis, and Melanocharis
longicauda.
A related phenomenon involves birds whose distribution is patchy
to an extreme degree, such that well-established populations have
been found at a few widely scattered localities but the species is
absent in similar habitats over most of New Guinea. For instance, the
Yellow-breasted Bird of Paradise, Loboparadisea sericea, was found
to be fairly common at 3,000-6,000 ft at three far-flung localities (Mt.
Kunupi at the western end of the Central Range, the Utakwa River
in southwestern New Guinea, and Dawong in eastern New Guinea),
and was otherwise known only from a few specimens at five other
scattered localities until it turned up in numbers on the west ridge of
Mt. Karimui. Archboldia papuensis sanfordi, a striking race of Arch-
bold’s Bowerbird endemic to the Eastern Highlands, is known only
from two very local populations, one on the southwestern side of Mt.
Hagen and the other on Mt. Giluwe, despite extensive searches and
inquiries among hundreds of natives by several ornithologists. ‘The
Yellow Thicket Flycatcher, Poecilodryas placens, is present in six
scattered areas and common in most of these (Batanta Island off the
western end of New Guinea, the Menoo Valley of the Weyland Moun-
tains, Astrolabe Bay in southeastern New Guinea, Lake Kutubu, and
Karimui) but has been reported nowhere else. All three of these
species have been missed in exhaustive collections involving thousands
of specimens in numerous other areas.
These and other rare or patchily distributed birds in New Guinea
appear to fall into two categories. Some (c.g., Megatriorchis doriae,
Rhamphomantis megarhynchus, Uroglaux dimorpha, Gampochaera
sloetii, Androphobus viridis, Macgregoria pulchra, Loboparadisea
sericea, Archboldia papuensis, and Daphoenositta miranda) belong to
2A
FEATURES OF THE EASTERN HIGHLANDS
distinct monotypic genera without close relatives. Others (e.g., Ptilino-
pus nanus, Charmosyna wilhelminae, Charmosyna multistriata, Tany-
siptera nympha, Coracina lineata, Crateroscelis nigrorufa, Sericornis
nigroviridis, Gerygone chloronota, Poecilodryas placens, Pachycephala
aurea, Pachycephala tenebrosa, Meliphaga flavirictus, Ptiloprora plum-
bea, Ptiloprora meekiana, Pycnopygius stictocephalus, and Melano-
charis arfakiana) belong to large genera with many ecologically similar
species. In many cases the habits of these local or rare species are
sufficiently well known to rule out the suggestion that they have narrow
ecological requirements uniquely met by only a few far-flung localities
in New Guinea. Their fragmented, relict distributions suggest that
more likely they are slowly becoming extinct, either because they are
the last survivors of unsuccessful evolutionary lines (monotypic genera)
or because they are being out-competed by ecologically similar close
relatives in the same genus.
Thus, a tendency to distributional patchiness expresses itself at
several levels in the New Guinea avifauna: in differences between the
avifaunas of nearby mountains; in large geographical gaps in the dis-
tribution of otherwise widespread species; and in the highly frag-
mented, relict-like distributions of other species. Examples of any of
these phenomena may also be found in the temperate zones, but they
are more prominent in New Guinea and perhaps in the tropics in
general. The tendency to drop out or to be patchily distributed is as
much a characteristic of a given species as its abundance, and is dis-
tinct from abundance. For instance, the cuckoo Chrysococcyx meyerit,
the thrush Drymodes superciliaris, and the honeyeater Meliphaga
polygramma are always uncommon. Nevertheless, they turn up in low
numbers at almost any hill forest collecting station on the Central
Range and on most of the “north mountain islands” as well, on most
of which some much commoner Central Range species (Pachycephala
soror, the Melidectes honeyeaters) are absent. If one views local species
diversity as an equilibrium between local extinction rates and immigra-
tion rates, the greater patchiness of bird distribution in New Guinea
as compared to the temperate zones is most readily explained by
postulating lower dispersal rates in New Guinea. This postulate is
almost certainly valid, since there is little migration in New Guinea
forests and hence recolonization after local extinctions should be slow,
whereas each year in the temperate zones the annual migrations and
seasonal dispersals generate a flood of potential new colonists. Species
vary in their dispersive tendencies and abilities, and these differences
may partly underlie the tendency toward patchy distribution that is
characteristic of certain species. An additional contributing factor may
be the closer niche packing and increased interspecific competition
resulting from higher species diversity in the tropics, such that the
local survival of each species becomes more precarious (MacArthur,
i972, ‘chapter 2).
FEATURES OF THE EASTERN HIGHLANDS
In short, the distributional patchiness of many New Guinea birds
comes initially as a surprise to some temperate zone ornithologists,
whose first reaction may be to dismiss the phenomenon with a trivial
explanation such as that the patchiness is due to inadequate explora-
tion or highly specialized habits. Enough is known about distribution
in most cases, and about habits in many cases, to dismiss these explana-
tions. In fact, distributional patchiness is seen on reflection to be an
expected consequence of low dispersal rates and high species diversity
in the New Guinea forest and of the continuing occurrence of specia-
tion. In a saturated fauna an automatic corollary of the fact that new
species are constantly arising is that other species must constantly be
becoming extinct. The New Guinea avifauna provides clear examples
of intermediate stages both in speciation and in extinction.
ECOLOGICAL SORTING MECHANISMS
If two formerly conspecific forms have succeeded in perfecting
reproductive isolating mechanisms by the time contact is reestablished
after a period of geographic isolation and evolutionary divergence,
they then fulfill the definition of species. For the forms to succeed in
reinvading each other’s ranges and to become sympatric, they must in
addition develop ecological differences sufficient to permit coexistence.
That the development of reproductive isolating mechanisms and of
these ecological sorting mechanisms may proceed at different rates is
shown by the existence of superspecies, comprised of forms which are
perfectly, or nearly perfectly, isolated reproductively but which are
strictly allopatric in distribution, presumably because ecological dif-
ferences are still insufficient to permit reinvasion. Examples are: Chal-
copsitta; Psittaculirostris; Micropsitta keiensis and M. pusio; Lalage;
Parotia; Melipotes; the Melidectes ruforissalis group; and Melidectes
nouhuysi and M. princeps. In addition to being a prerequisite for
completion of the speciation process, ecological sorting mechanisms
also underlie the problem of species diversity, viz., the question of
what makes it possible for many species to coexist at a locality (Lack,
a):
Lack (1944) has analysed the avifauna of Great Britain, and Moreau
(1948) the avifauna of Tanganyika, by determining the ecological sort-
ing mechanisms between the members of all pairs of species in a given
family. In the present analysis of ecological sorting mechanisms in the
Eastern Highlands of New Guinea I shall adopt a narrower choice of
material for analysis and shall largely confine discussion to pairs of
congeneric rather than merely confamilial species. Confamilial species
of different genera presumably have been distinct longer, and might
be expected to have diverged ecologically from each other in more
ways, than have congeneric species. One, in fact, finds that most
congeneric pairs of species in New Guinea differ ecologically mainly
26
FEATURES OF THE EASTERN HIGHLANDS
in one or two simple respects. These ecological sorting mechanisms
may be classified under three headings: spatial differences (altitudinal
segregation, habitat preference, vertical segregation, allopatry, and
checkerboard allopatry) which lead to pairs of congeners having
mutually exclusive territories; differences in feeding or foraging be-
havior; and temporal differences (occupation of the same habitat at
different times). ‘These categories will be considered in turn.
1. Spatial Segregation
Altitude——TVhe most important ecological sorting mechanism in the
New Guinea avifauna as a whole, and in the Eastern Highlands in
particular, is segregation by altitude. That is, more pairs of congeners
segregate by living at different altitudes than by any other means. ‘The
Eastern Highlands provide about 37 instances of pairs of similar and
congeneric species which live in the same gross habitat type but have
altitudinal ranges which are largely or completely mutually exclusive.
Some examples (citing the low-altitude form first in each case) are the
pigeons Ducula rufigaster and D. chalconota, the kingfishers Halcyon
torotoro and H. megarhyncha, the flycatchers Peltops blaimvillii and
P. montanus, the flycatchers Machaerirhynchus flaviventer and M.
nigripectus, the bowerbirds Ailuroedus buccoides and A. crassirostris,
and the honeyeaters Ptiloprora guisei and P. perstriata. In about 12
additional cases there are three similar congeners which replace each
other with altitude. Examples include the logrunners Eupetes caerules-
cens, E. castanonotus, and E. leucostictus and the flowerpeckers Melan-
ocharis nigra, M. longicauda, and M. verstert. Four congeneric species
replace each other successively in two cases (the warbler genus Seri-
cornis and the bird of paradise Lophorina), and in the whistler genus
Pachycephala there is a sequence of five species. Many similar cases
have been discovered by ‘Yerborgh (1971) in the avifauna of the Peru-
vian Andes.
In a considerable portion of these cases the altitudinal transition is
sharp, mutual exclusion of each species from the other’s range is com-
plete, there are no significant ecological differences between the species
except for those associated with altitudinal range, and the species are
quite similar to each other morphologically and apparently are the
product of a recent speciation. One case (p. 230) is provided by the
warblers Crateroscelis murina and C. robusta, which are the com-
monest insectivorous gleaners of the understory, resemble each other
in color, pattern, size, and song, and forage in virtually identical man-
ners between ground level and five feet above the ground. As we
ascended the west ridge of Mt. Karimui from its base, C. murina pro-
gressively increased in abundance with increasing altitude until it was
the second most abundant bird in Zone 3b (5,080 to 5,390 ft). It
abruptly disappeared at 5,390 ft. At 5,400 ft C. robusta abruptly ap-
Zi
FEATURES OF THE EASTERN HIGHLANDS
peared at its maximum abundance and progressively decreased in
abundance with increasing altitude towards the summit at 8,165 ft
(Fig. 1). No individual of either species was ever found within the
other's altitudinal zone. There was no vegetational discontinuity in
the forest at the transition altitude, and the ecotone at the lower level
of the moss forest lay at 6,500 ft, which is 1,100 ft higher. As shown
in Figure 2, which depicts the altitudinal ranges of the Crateroscelis
warblers at various localities at different longitudes on the Central
Range, the transition altitude varies locally between 4,500 and 5,500 ft,
but there is no consistent difference between eastern and western New
Guinea. Additional instances of similarly sharp transitions are discussed
DISTRIBUTION OF GENUS CRATEROSCELIS ON MT. KARIMUI
individual records relative abundance
summit
8000 -
7000 F
6000 F
5000 F
feet
above
sea 2 .
level’ 40007 eaeceacocsesescag
pEoRRRREQ BERR EO BARK
3000 fF
| 1 =I |
0 46 8 101
% of total bird individuals
Fic. 1. Distribution of the warblers Crateroscelis robusta (@) and C. murina
(QO) on Mt. Karimui as a function of altitude. On the left side each mark repre-
sents one individual heard, seen, or collected at the given altitude (the relative
paucity of records between 2,000 and 3,000 ft is due to my having spent only a
brief time at this altitude). The right side gives the relative abundance in the
whole avifauna, i.e., the percentage of bird individuals of all species estimated as
being C. robusta or C. murina (see p. 10). Note that the two species replace each
other sharply at 5,400 ft, and that each species reaches its maximum population
density just above or just below this altitude.
28
FEATURES OF THE EASTERN HIGHLANDS
in the text (cf., Melanocharis nigra and M. verstert, Coracina montana
and C. schisticeps, Rhipidura hy perythra and R. albolimbata, and the
genera Eupetes, Sericornis, Peltops, and Ailuroedus).
To varying degrees in other instances the altitudinal transition is
not sharp, ecological sorting mechanisms in addition to altitudinal
distribution are apparent, and the two or more species, while con-
generic, show sufficient morphological differences to suggest that speci-
ation occurred less recently. For instance, the pigeons Ptilinopus
ornatus and P. perlatus have similar color patterns suggesting recent
divergence, and in general represent each other altitudinally, P. perla-
tus being found at lower elevations. However, P. perlatus is less com-
mon, and the altitudinal ranges of the two species overlapped at
Karimui, an overlap made possible by P. perlatus’s larger size and its
preference for the forest edge over the forest interior. On the southern
slopes of the Eastern Highlands live three congeneric parrots, Char-
mosyna placentis occurring at low elevations, C. pulchella at medium
elevations, and C. papou at higher elevations. ‘he altitudinal ranges
are largely mutually exclusive, but each overlaps the next by up to
several hundred feet. C. placentis is more closely related to other
species in the genus than to C. pulchella or to C. papou; the latter two
share the same distinctive color pattern with each other but differ by
a factor of 2.6 in weight. The five whistlers Pachycephala griseiceps,
P. hyperythra, P. soror, P. schlegelii, and P. modesta are encountered
from sea level to timberline in that sequence, but their altitudinal
ALTITUDINAL RANGES IN GENUS CRATEROSCELIS
14000 5
I2000 4
feet
above [0,000 5
oS axe = C. robusta
level
6,000 54 O (C.nigrorufa)~
4000 +
2000 75 C. murina
O i
7 T ae iS 5 = ies al aed | ‘mpeel |
é
I32 134 I36 138 140 I42 144 146 148 I50
Longitude, °E
Fic. 2. Altitudinal ranges of Crateroscelis warblers as a function of longitude,
based on the records of collections at ten localities. Note that the altitudinal ranges
of C. murina and C. robusta are mutually exclusive, and that the transition altitude
varies locally between 4,500 and 5,500 ft but shows no systematic variation with
longitude. Where the rare C. nigrorufa is present, its range lies between those of
the other two species. None of the 10 expeditions on whose records the figure is
based collected continuously from sea level to timberline, so that the apparent gaps
in altitudinal distribution at some localities are artifacts caused by discontinuous
collecting transects.
29
FEATURES OF THE EASTERN HIGHLANDS
ranges overlap broadly: P. soror straddles the ranges of P. hyperythra
and P. schlegelii, which nearly meet; the range of V. schlegeli: uciudes
much or most of that of P. modesta; and that of P. griseiceps includes
most or all of that of P. hyperythra. No two of these five species are
as immediately derived from each other as are Crateroscelis murina
and C. 10busta, or Ptiiinopus ornatus and P. periatus. The additional
ecological sorung mechanism making altitudinal overlap possible is
a tendency towards vertical segregation: on the average P. griseiceps
forages higher in the vegetational column than does P. hyperythra,
P. hyperythra higher than P. soror, P. soror perhaps higher than P.
schlegelu, and P. schlegelii lower than P. modesta. ‘Vhe correlation
between degree of morphological similarity and strictness of altitudi-
nal segregation in bird altitudinal sequences in New Guinea suggests
that strict altitudinal segregation was the first sorting mechanism to
develop in these cases, and that some degree of altitudinal overlap
became possible only as further evolution and morphological di-
vergence led to other ecological differences. ‘his interpretation is
supported by the evidence presented on pp. 33-34.
Certain high-altitude species appear to take over the combined
niches of several low-altitude congeners. For instance, in the cuckoo
genus Cacomantis the single species found above 6,000 ft is C. pyr-
rhophanus, which inhabits the forest interior as well as second-growth
and gardens. Below this elevation it is replaced by two species, C.
castaneiventris in the forest interior and C. variolosus in second-
erowth and gardens. Up to six species of Piilinopus pigeons may be
found locally sympatric at low elevations, differing in size and prefer-
ence for the forest interior versus the forest edge. ‘These yield to one
medium-sized species, P. rivoli, at higher elevations.
I found that individuals collected outside a species’ usual alti-
tudinal range and in the range of a congener generally proved to be
immatures. A typical example arose on Mt. Karimui in the warbler
genus Sericornis, four species of which divided up the mountain into
mutually exclusive altitudinal bands; S. spilodera up to 4,240 ft, S.
arfakianus from 4,400 to 5,000 ft, S. perspicillatus from 5,100 to 6,350
ft, and S. papuensis from 6,450 ft to the summit at 8,165 ft. The only
violation of these limits detected came when two immature individuals
of S. spilodera were collected at 4,950 it, mean the wpper limit or thie
next higher species.
This tendency of immatures to be found at the peripheries of the
altitudinal range characterizes most species of the New Guinea mon-
tane avifauna, and not just those which belong to altitudinal se-
quences. The change in population structure with increasing altitude
is typically that immatures are found at the bottom of the altitudinal
range; somewhat higher one meets immatures plus birds in adult plum-
mage but in nonbreeding condition, with females usually appearing
at lower altitudes than males; next comes the optimal part of the
30
FEATURES OF THE EASTERN HIGHLANDS
species’ range, with singing males and adults of both sexes in breeding
condition; and, finally, another band of immatures but of few adults
appears at the upper altitudinal limit in some species. Examples of
this population structure discussed in detail in the species accounts
include Pachycephala leucostigma, Pachycephala schlegeli, Myzomela
rosenbergii, Melipotes fumigatus, and Ptiloprora guiset. ‘Vhis tendency
is manifest to an extreme degree in some species of birds of paradise,
in which displaying adult males may be compressed into the top 600
ft of the altitudinal range, with adult females and especially imma-
tures continuing to be found several thousand feet below the level
at which the lowest adult male is encountered (cf. Lophorina superba;
Lophorina (Diphyllodes) magnifica; Lophorina (Cicinnurus) regia;
and Loboparadisea sericea). Stein (1936) explicitly remarked on the
same population structure for the birds of paradise Cnemophilus
(Loria) loriae, Lophorina (Pteridophora) alberti, Epimachus fastosus,
and Epimachus meyeri in the Weyland Mountains.
Two explanations suggest themselves for the sharp transitions in
sequences of altitudinally segregating congeners. On the one hand, the
actual altitudinal limits of a species might be directly determined by
its adaptations, either to altitude itself, or else, probably more im-
portantly, to variables correlated with altitude, notably vegetation
and temperature. On this hypothesis each member of an altitudinal
sequence would be adapted only to a certain range of altitudes (or
forest types or temperatures) and would be unable to maintain stable
populations beyond its actual limits, even if the other species in the
sequence were absent. By this interpretation the actual altitudinal
limits and the physiologically possible altitudinal limits of a species
would be identical, and would be sharp only if there were a sharp
change in an adaptively critical environmental variable. ‘The alterna-
tive hypothesis is that the actual altitudinal limits of a species depend
upon competition with other species. Each species would be in-
trinsically capable of establishing itself over a broader altitudinal
range than that which it actually inhabits. However, in the presence
of ecologically very similar species each form would be limited to
that altitudinal range (or temperature range or vegetational type)
over which it was competitively superior, and would be excluded from
adjacent altitudinal bands in which the related species were competi-
tively superior.
While the first hypothesis (intrinsic survival ability) may prove valid
in some instances, competition is clearly the explanation in many
cases. To begin with, competition appears the only reasonable inter-
pretation when the population density of one species changes from
maximal to zero and the abundance of a closely related congener
changes from zero to maximal within the same 10 vertical feet in a
gradually changing forested habitat. Had sharp species transitions in
altitudinal sequences been due to a sharp vegetational discontinuity
31
FEATURES OF THE EASTERN HIGHLANDS
rather than to competition, then the bird transitions should have co-
incided with vegetational ecotones, and many bird transitions should
have clustered around the same altitude. ‘These expectations are not
fulfilled, as illustrated by Figure 3, which plots those transition alti-
tudes between congeneric birds on Mt. Karimui’s west ridge that I
was able to determine most accurately. ‘The sharpest ecotone was that
at 6,500 ft at the lower limit of the heavily mossed zone. No bird
transition coincided with this ecotone, and the nearest transition was
the one about 100 vertical feet below between Sericornis papuensis
SPECIES TRANSITIONS ON MT. KARIMUI
summit of mountain
8000
—~-Cnemophilus (Loria) loriae / C. macgregorii
7000 __ Penesthelle cyanus / P. sigillatus
—1--Sericornis perspicillatus / S. papuensis
6000 _Crateroscelis murina / C. robusta
+ Pitohui dichrous / P. nigrescens
— |-Pachycephalopsis poliosoma / Peneothello cyanus
—---Sericornis arfakianus / S. perspicillatus
==
SOOO _-Rhipidura hyperythra / R. albolimbata
feet =o /Melanocharis nigra / M. versteri
“ Sericornis spilodera / S. arfakianus
above = >-------Coracina schisticeps / C. montana
sea 4000 afer
«Menerche guttula / M. axillaris
level “I
“Meliphaga analoga / M. orientalis
3000
fe blainvillii / P. montanus
2000
lIOOO
O
Fic. 3. Some altitudes on Mt. Karimui’s west ridge at which pairs of congeneric
or closely related species replaced each other abruptly. In each case the first-named
species is the low-altitude form, the last-named species the high-altitude form. A
sharp line means that the transition altitude was identified exactly, while a bracket
means that the transition occurred at some altitude within the indicated band bux
was not determined more exactly.
32
FEATURES OF THE EASTERN HIGHLANDS
and S. perspicillatus. Figure 3 also shows that different bird transitions
occur at different altitudes, and that at least one transition is en-
countered every 600 feet in the well-studied region between 3,650 and
8,165 ft. Few transitions were identified between 3,250 and 2,000 it,
partly for the trivial reason that no collecting was carried out in this
band, partly because in the Karimui area this altitudinal band consti-
tutes part of a relatively well-defined ia ‘zone’ with few species
limits (pp. 67-70).
The facts that few or no vegetational ecotones in the New Guinea
forest are as sharp as many bird transitions in altitudinal sequences,
and that these sharp transitions do not coincide with ecotones, imply
that sharp transitions involve competition. To test this hypothesis,
one would like to find situations in which one member of a sequence
has been locally eliminated, and to examine whether its congener,
freed of the competition, expands into the missing species’ altitudinal
range. This test is made possible by two different circumstances: the
drop-out phenomenon and the reinvasion situation.
As discussed on pp. 21-24, small mountains and mountain ranges
have fewer bird species than large ones because localized populations
are most likely to become extinct. Under these circumstances one often
finds that congeners expand their altitudinal range at the expense of
the missing species (Diamond, 1970a, 1970b). For instance, in the
Okapa area the altitudinal ranges of the low-altitude flowerpecker
Melanocharis nigra and its high-altitude congener M. versteri were
separated by several thousand feet, and the intervening altitudes were
monopolized by the middle-altitude congener M. longicauda. On iso-
lated and small Mt. Karimui, M. longicauda was one of the species
that dropped out, and M. nigra expanded upwards and M. versteri
downwards to meet somewhere between 4,300 and 4,500 ft. In the
North Coastal Range the cuckoo-shrike Coracina schisticeps occurred
at most localities up to about 2,900 ft, above which altitude it was
replaced by C. montana. However, on Mt. ‘Turu, an isolated peak only
3,650 ft high, C. montana was absent, presumably because it could not
maintain a stable population within a band of only 750 vertical feet;
and C. schisticeps, freed of the competition, occurred up to Mt. ‘Turu’s
summit. Similar instances arise among altitudinally representative
species in Rallicula, Charmosyna, Psittacella, Crateroscelis, Pachyceph-
ala, and Melidectes.
The second kind of test for competition occurs during speciation,
when one form begins to reinvade the geographical range of another
to which it was formerly allopatric. As discussed on pp. 22-23, speci-
ation in the New Guinea mountains usually begins with the formation
of eastern and western isolates. There are 13 cases in which allied
eastern and western species are sympatric only over part of the geo-
graphical range, indicating that reinvasion is at an early stage. In 1]
of these cases the two species have mutually exclusive altitudinal
33
FEATURES OF THE EASTERN HIGHLANDS
ranges in the area of sympatry, and each has a broader altitudinal
range outside the area of sympatry. For instance, the honeyeater
Pile nnons guises occurs on the Central Range in eastern New Guinea
and on one “north mountain island,’ the Huon Peninsula. The sim-
ilar P. perstriata occurs on the Central Range in both western and
eastern New Guinea. In western New Guinea, where P. perstriata oc-
curs alone, it lives from about 5,000 to 12,000 ft. In the area of sym-
patry, eastern New Guinea, P. perstriata lives from about 9,000 to
12,000 ft, P. guisei from 5,000 to 9,000 ft, and the altitudinal ranges
are mutually exclusive. On the Huon Peninsula, where P. guise occurs
alone, it lives from 5,000 to apparently at least 10,500 ft. Thus, in the
area of sympatry the altitudinal ranges of both species are apparently
compressed by competition as compared to the range each can inhabit
when by itself. A clear example of the development of an ecological
sorting mechanism in the earliest stage of speciation is presented by
the birds of paradise Lophorina (Astrapia) mayert and L. (Astrapia)
stephaniae, between which reproductive isolation is not yet complete
and whose ranges are still largely allopatric. Sympatry has been re-
ported only from two mountains, Mts. Giluwe and Hagen, where the
two forms segregate altitudinally, L. (Astrapia) mayert being found at
higher elevations. Similar instances in the Eastern Highlands arise in
the bird of paradise genus Epimachus, in the honeyeater genus Tox-
orhamphus, and apparently in the megapode genus Talegalla.
Thus, the principal (but not the only) speciation mechanism in
the mountains of New Guinea appears to be the formation of western
and eastern isolates, followed by mutual reinvasion with altitudinal
displacement, until the two species occur sympatrically over all New
Guinea with mutually exclusive altitudinal ranges. In most cases alti-
tudinal segregation is the first ecological sorting mechanism to develop,
and other differences develop only later, after broad sympatry has
been achieved (cf. pp. 29-30).
In some genera this process has occurred two or three successive
times to give rise to three-species and four-species altitudinal se-
quences. However, two-species sequences are much commoner in New
Guinea. The larger sequences tend to be unstable and to lead to
elimination of the middle birds, whose narrow altitudinal ranges due
to competition from both above and below make them particularly
prone to drop out locally. The narrowest altitudinal ranges in the
New Guinea mountains prove to be those of middle birds in three-
and four-species sequences. For instance, on Mt. Karimui the warbler
Sericornis arfakianus, the second bird in a four-species sequence, was
compressed into a band of only 600 feet, while Melanocharis longi-
cauda, middle bird in a three-species sequence, was missing entirely,
having been squeezed out between its congeners. In the Cyclops Moun-
tains and North Coastal Range the whistler Pachycephala soror, third
34
FEATURES OF THE EASTERN HIGHLANDS
in a five-species sequence, has similarly been squeezed out between
P. hyperythra and P. schlegeli. ‘The warbler Crateroscelis nigrorufa,
middle bird in a three-species sequence, is very rare and local, presum-
ably because it has been eliminated at most localities. ‘The instability
of three-species sequences also appears responsible for the unique and
complex distributional pattern of compound checkerboard allopatry
in the Melidectes rufocrissalis-M. belfordi-M. ochromelas complex (pp.
388-389; 393-396).
To recapitulate, sharp altitudinal transitions between congeners are
due to competition, as shown by the facts that most vegetational eco-
tones are much less sharp; that the transitions do not coincide with
ecotones or with each other; that the component species expand their
ranges when one member drops out locally; and that they compress
their ranges in the zone of sympatry during reinvasion. In the Andes
of Peru ‘Terborgh (1971) has described essentially identical phenomena
exemplified by numerous altitudinal sequences of two to five species.
It is worth restating explicitly that the bearing of adaptations to alt-
tude, vegetation, and temperature on the sequences is not denied. ‘The
fact that Crateroscelis murina lives below C. robusta, and not vice
versa, can only mean that the former is better adapted to low altitude,
rainforest, and warm temperatures and that the latter is better adapted
to high altitude, Nothofagus forest, and cool temperatures. It is the
sharpness of the transitions, and the mutually exclusive altitudinal
ranges, in the absence of equally sharp changes in temperature and
forest composition, which must be explained in terms of competition.
Habitat—Congeners that live at the same altitude may differ in
respect to the gross habitat type preferred (forest, second-growth and
forest edge, grassland, savanna, swamp, or water). Since the fractional
area occupied by gross habitat types other than forest is small in un-
disturbed areas in most parts of New Guinea (especially in the moun-
tains) except in a low-rainfall band on the south coast, the proportion
the nonforest avifauna bears to the total avifauna in New Guinea is
low. ‘The nonforest avifauna of the Eastern Highlands is discussed
on pp. 70-79.
In the following 18 pairs or groups of species which occur in the
Eastern Highlands, the first member lives mainly at the forest edge, in
second-growth, or in gardens, while the second member or members
live mainly in the forest, but the birds are otherwise fairly similar
ecologically. Details are given under individual species accounts:
Accipiter fasciatus; A. melanochlamys and A. poliocephalus
Falco berigora; F. severus
Macropygia amboinensis; M. nigrirvostris (difference valid only above
5,000 ft)
Cacomantis variolosus; C. castaneiventris
35
FEATURES OF THE EASTERN HIGHLANDS
Centropus phasianinus (grassland); C. menbeki (forest)
Lyto capensis (grassland); T. alba (open wooded country); 7. tene-
bricosa (forest)
Halcyon sancta, H. macleayii (Australian wintering visitors); #7.
torotoro, H. megarhyncha (residents)
Coracina papuensis, C. tenuirostris; C. schisticeps, C. melaena, and
C. boyeri
Gerygone ruficollis; G. cinerea
Gerygone chloronota, G. magnirostris; G. chrysogaster
Rhipidura leucothorax; R. threnothorax
Rhipidura leucophrys (open country); R. rufiventris (forest edge);
R. hyperythra (forest interior)
Pachycephala ruftventris; several other species of Pachycephala
Colluricincla harmonica; C. megarhyncha
Cracticus cassicus; C. quoyi
Myzomela adolphinae; M. rosenbergii
Melidectes torquatus; M. rufocrissalis-M. belfordi
Meliphaga auga; M. mimikae
The rail Rallus pectoralis occurs in dry grassland, while the similar
Rallus philippensis is found in wet and swampy grassland.
In the hills the swiftlet Collocalia esculenta is confined to the air
space above small streams and forest glades, while C. hirundinacea
forages over extensively cleared areas.
Vertical distribution.—Many New Guinea forest species show strict
preferences for the height in the vegetational column at which they
forage. For example, the warblers Crateroscelis robusta and C. murina
are almost always seen within five feet of the ground, the thicket fly-
catcher Pachycephalopsis poliosoma within three feet of the ground,
and the logrunners Eupetes castanonotus and E. lewcostictus remain
on the ground or on fallen branches. Conversely, the warbler Gerygone
palpebrosa and the flycatcher Monarcha frater often forage down to
within about 12 ft of the ground but never within five feet of the
ground. These vertical preferences are well illustrated by results of
mistnetting, since our mistnets generally extended from near the
ground to six feet above the ground. Despite intensive use of mistnets
(about 30 in operation simultaneously) many common forest species
were never netted, and some species (e.g., the thicket flycatcher
Pencothello cyanus and the honeyeater Toxorhamphus poliopterus)
were netted much more often in proportion to their censused abundance
than others. I shall frequently cite netting results in the individual
species accounts as an indication of vertical foraging preference. While
temperate zone species, of course, also have vertical preferences, these
are less strict and absolute than in species-rich areas of the tropics.
For instance, if one operates mistnets for a long time, one eventually
catches every species of bird present in New Zealand forest or in east-
36
FEATURES OF THE EASTERN HIGHLANDS
ern North American forest, but litthe more than half of the species
present in New Guinea (in my experience) or in Peru (Terborgh, pers.
comm. ).
The following 19 forest species in the Eastern Highlands apparently
forage strictly or almost strictly on the ground: the cassowaries Casu-
arius bennettii and C. casuarius, the megapodes Megapodius freycinet,
Talegalla sp., and Aepypodius arfakianus, the rail Rallicula forbesi,
the woodcock Scolopax saturata, the ground doves Gallicolumba bec-
carii, G. rufigula, G. jobiensis, and Chalcophaps stephani, the pittas
Pitta erythrogaster and P. sordida, the thrushes Zoothera dauma, Am-
alocichla incerta, and Drymodes superciliaris, and the logrunners Eu-
petes castanonotus, E. leucostictus, and Melampitta lugubris.
Three additional forest species apparently forage on the ground
and in undergrowth within a few feet of the ground: the thicket fly-
catchers Heteromyias albispecularis and Pachycephalopsis poliosoma,
and the whistler Pitohwi cristatus.
Eight species forage in the understory within five feet, or in a few
cases ten feet, of the ground but usually not on the ground itself: the
warblers Crateroscelis murina, C. nigrorufa, and C. robusta, the fly-
catcher Rhipidura threnothorax, and the thicket flycatchers Pene-
othello sigillatus, P. cyanus, P. bimaculatus, and Poecilodryas placens.
At the other extreme are the numerous forest species which keep to
the middle and upper stories and rarely descend to the lower story. ‘To
quantitate this phenomenon by mistnet results, let us exclude from
consideration birds weighing over 250 g (hence possibly strong enough
to break out of the mistnets I was using, although a few larger birds
were in fact netted), and consider only species of which I saw, heard,
or collected at least 10 individuals. ‘Vhis leaves 137 forest species. Of
these 137 species, 77 (569%) were netted at least once, while 60 (44%)
were never netted. ‘The list of 60 species not netted includes most
pigeons except for the four small ground doves of the genera Galli-
columba and Chalcophaps; most parrots except Psittacella; the cuckoo
Eudynamis scolopacea; all campephagids without exception; the war-
blers Phylloscopus trivirgatus and Gerygone ruficollis; the flycatchers
Peltops montanus and P. blainvillit, and Monarcha frater and M.
chrysomela; the whistler Pachycephala hyperythra; the oriole Oriolus
szalayt; the birds of paradise Manucodia (including “Phonygammus’’),
Epimachus (including “Drepanornis’), Paradisaea, and Loboparadisea,
and Lophorina (Parotia) lawesi (despite the male’s terrestrial display
court) and Lophorina (Ptiloris) magnifica; the honeyeaters Myzomela
eques, M. nigrita, M. cruentata, Oedistoma pygmaeum, Meliphaga
subfrenata and M. analoga, and Pycnopygius ixoides; and the dicaeids
Dicaeum geelvinkianum and Oreocharis arfaki. In addition, I netted
females but never males of the honeyeater Meliphaga obscura, al-
though males were collected by other means. There are a couple of
these species that I netted elsewhere in New Guinea or observed in the
37
FEATURES OF THE EASTERN HIGHLANDS
understory despite my failure to net them. Most of them, however, are
species that I saw dozens or hundreds of times and which simply do
not descend to the understory, unless under the unusual conditions
noted below.
Of the 77 species netted, seven rarely use the understory and were
netted only under one of three unusual conditions. First, birds flying
from one side of a ridge to another often prefer to fly through a notch
or dip in the ridge, and if one sets up nets in such a notch, anything
may turn up. For instance, one such net on Mt. Menawa in the North
Coastal Range caught a flock of 25 Charmosyna parrots, which feed in
the crowns of flowering trees. Secondly, at high elevations where the
fovest becomes stunted, species that remain in the middle- and upper-
stories of tall forest turn up in nets, simply because the middle- and
upperstories disappear at high elevations. For instance, the parrot
Oreopsittacus arfaki and the honeyeater Myzomela rosenbergii were
netted in the stunted summit forests of Mt. Michael and Mt. Karimui
but not in the taller forest at lower elevations. Finally, some species
present both in the forest interior and at the edge are netted much less
often in the interior than at the edge (e.g., the honeyeaters Melipotes
fumigatus, Melidectes rufocrissalis, and Meliphaga orientalis).
‘Thus, almost half of the forest species in the Eastern Highlands have
sufficiently marked middle- and upperstory preferences that they are
rarely or never netted in the forest.
‘There are about eight pairs or groups of congeneric forest species in
the Eastern Highlands for which differences in vertical preference pro-
vide an important or the most important ecological sorting mechanism,
one species foraging largely in the lower story and the other largely in
the middle- and upperstories. All eight cases involve small passerines.
In four of these cases the upperstory species was never taken in mist-
nets, but even in these four instances the vertical segregation was not
absolute because the lowerstory species was occasionally or often seen
in the middlestory. The eight cases, listing the lowerstory bird or birds
first in each case, are:
Sericornis nouhuyst; S. arfakianus, S. perspicillatus, S. papuensis
Rhipidura threnothorax; R. hyperythra
Rhipidura atra, R. brachyrhyncha; R. albolimbata
Monarcha axillaris; M. frater
Monarcha guttula; M. chrysomela
Pachycephala soror; P. hy perythra
Pachycephala schlegeli; P. modesta
Meliphaga mimikae, M. aruensis; M. orientalis, M. analoga
Allopatry—New Guinea provides numerous examples of super-
species, i.e., pairs or groups of forms that already are, or may be
assumed to be, reproductively isolated but whose distributions are still
largely or strictly allopatric. Superspecies consist of isolates (‘“semi-
38
FEATURES OF THE EASTERN HIGHLANDS
species”) between which geographical contact may have been reestab-
lished but which are presumably still too similar ecologically to
reinvade each other’s ranges. The Eastern Highlands contain about
eight pairs of allopatric semispecies, which fall into two categories.
The first category consists of lowland or hill forest species to which
the Central Range, running east-west, presents a major barrier, so
that one form has been isolated on the northern slopes of the Central
Range and the other on the southern slopes. Four examples, listing
the northern form first and the southern form second in each case, are
the crowned pigeons Goura victoria and G. scheepmakeri; the fig par-
rots Psittaculirostris edwardsti and P. desmarestti; the trillers Lalage
atrovirens and L. leucomela; and the red birds of paradise Paradisaea
minor and P. raggiana. In one additional case, the cassowary Casu-
arius casuarius, one member of a superspecies has been found on the
southern slopes, while the corresponding form of the northern slopes
(Casuarius unappendiculatus) has not yet been reported from the
Eastern Highlands but will undoubtedly turn up when the northern
slopes are explored at low altitudes.
The other category consists of midmontane species for which the
Central Range does not provide a distributional barrier but which
have formed eastern and western semispecies by the mechanism dis-
cussed on pp. 22-23. In this category fall two pairs of birds of paradise,
Lophorina (Astrapia) mayerit and L. (Astrapia) stephaniae, and Lo-
phorina (Parotia) carolae and L. (Parotia) lawesi (the western form
is listed first in each case). ‘he astrapias provide a fascinating example
of the development of ecological sorting mechanisms in the early stages
of reinvasion, and are discussed further on p. 34 and pp. 338-339.
Finally, the megapodes Talegalla jobiensis and T. fuscirostris pro-
vide at first glance another pair of northern and southern semispecies
which are still allopatric, but the evidence discussed on p. 000 suggests
that the development of sympatry and ecological sorting may actually
be far advanced in this instance.
Checkerboard allopatry—In the previously cited examples of al-
lopatry, each member of the superspecies has a coherent and exclusive
range, and a continuous geographical line running north-south or east-
west divides it from its relative. here are in addition five puzzling
cases of forms whose distributions appear to be largely or wholly
allopatric but which replace each other in checkerboard fashion. ‘Thus,
the range of each species is broken into discrete geographical areas
separated from each other by areas inhabited by the other species.
‘Three of these cases involve pairs of forms: the fruit doves Ptilinopus
pulchellus and P. coronulatus, the owlet-nightjars Aegotheles albertisii
and A. archboldi, and the mannikins Erythrura trichroa and E. papu-
ana. A fourth case (pp. 388-389) consists of three honeyeaters (Meli-
dectes rufocrissalis, M. belfordi, and M. ochromelas) and is more
complex: most New Guinea mountains support two of these three
39
FEATURES OF THE EASTERN HIGHLANDS
species, and the identity of the locally missing species varies geo-
graphically in checkerboard fashion. The last case involves eight man-
nikins of the genus Lonchura, which have colonized the midmontane
erasslands in an irregular checkerboard, each area usually supporting
only one species but the same species recurring in widely separated
areas. Further details of all these cases are given under the individual
species accounts. At least three different phenomena (habitat differ-
ences, colonization, and recent range expansion) appear to be involved,
as discussed in the following paragraphs.
The geographical ranges of the fruit doves Ptilinopus pulchellus
and P. coronulatus (pp. 124-125) both include essentially the whole of
New Guinea. In some cases they replace each other over large areas.
For instance, P. coronulatus is apparently found over the whole lower
and middle Fly Rivers, while P. pulchellus is found on the upper Fly
River; and P. coronulatus is on the northern slopes of the Eastern
Highlands, P. pulchellus on the southern slopes. However, there are
also areas, as on the Vogelkop, where these species replace each other
over much shorter distances and inhabit smaller patches. One suspects
that these two species recognize some consistent difference in the en-
vironment or habitat and that a competitive equilibrium has been
reached, each species being able indefinitely to exclude the other from
the type of habitat in which it is competitively superior. However, the
basis of this postulated habitat difference is unknown, and I cannot
detect any feature which consistently characterizes the areas chosen by
P. pulchellus and distinguishes them from the areas chosen by P.
coronulatus.
‘The checkerboard colonization of the midmontane grasslands by the
eight Lonchura mannikins appears to have been settled on a “‘first
come first serve” basis, and the diversity of habitats colonized by each
species makes it unlikely that superior adaptations to subtle habitat
differences determined the local winner. For instance, L. spectabilis
is the sole mannikin for a distance of at least 150 miles in the mid-
montane grassland of the Eastern Highlands over a considerable range
of altitudes, climatic conditions, and grass types and heights; L. cani-
ceps is equally well entrenched in midmontane grassland of south-
eastern New Guinea presenting a similar but equally varied range of
conditions; and L. tristissima fills this niche on several of the “north
mountain islands” (see pp. 409-410 for details). The midmontane grass-
lands in their present form are recent by-products of native agricul-
ture, and much of their grassland avifauna consists of colonists from
the lowlands. The Lonchura mannikins appear to colonize only with
difficulty, so that for them the colonization of these grassland islands
is a slow and random process. Once a given species arrives, it becomes
established over the whole area and excludes potential future colonists.
The identity of the locally successful colonist is usually as unpredict-
40
FEATURES OF THE EASTERN HIGHLANDS
able as trying to guess which of several related species will become
established on a particular oceanic island or on a particular mountain.
The remaining three cases appear to involve recent range expan-
sions due to invasions or speciations. In the genus Erythrura the
species E. papuana is endemic to New Guinea, where it is known
only from a handful of scattered localities. The very similar E. trichroa
occurs from Celebes to Australia, the Solomon Islands, and Micronesia,
and has been found at most midmontane localities in IwWew Guinea
except those where E. papuana occurs (p. 000). Presumably £. papwana
is the older species in New Guinea and has been eliminated at all but
a few localities by E. trichroa, a recent invader from the outside. The
other two cases (Aegotheles and Melidectes) suggest recent speciations
within New Guinea itself. Aegotheles archboldi and A. albertisi re-
place each other in checkerboard fashion in western New Guinea,
while A. albertisii has the Vogelkop and eastern New Guinea to itself
(p. 177). Sympatry has been demonstrated at one locality and may de-
pend on different altitudinal preferences. Sympatry between those two
of the three Melidectes forms present at a given locality depends also
upon altitudinal exclusion. Apparently two successive speciations pro-
duced three honeyeaters, and since altitudinal sequences of three spe-
cies tend to be unstable in New Guinea (p. 34), one of the three species
(but not always the same one) disappeared at each locality.
Thus, the Erythrura case suggests a speciation and on-going rein-
vasion during which an ecological sorting mechanism adequate to
permit sympatry has not developed and the older form has been re-
duced to a fragmented and possibly shrinking range. In the Aegotheles
and Melidectes instances a sorting mechanism, viz., altitudinal segre-
gation, has developed in at least a few areas, but either has failed to
develop in most areas, or else was inadequate to guarantee stable sym-
patric populations of both forms and resulted in local elimination
of one.
2. Nonspatial Sorting Mechanisms (Food and Foraging
Differences)
‘The sorting mechanisms discussed so far (altitude, habitat, vertical
distribution, allopatry, and checkerboard allopatry) have as their usual
result that the spatial overlap between territories of congeners is
minimal or nonexistent. We shall now consider those factors, mainly
related to food consumed and the means of obtaining it, that permit
congeneric species to occupy spatially overlapping territories. It should
at once be mentioned that it requires careful observation to decide in
some cases whether segregation is mainly spatial or nonspatial. For
instance, even when two congeners occur at the same locality, at the
same altitude, and in the same gross habitat type and forage at the
same height, they may still be interspecifically territorial due to their
4]
FEATURES OF THE EASTERN HIGHLANDS
recognizing finer habitat categories, such as denser and less dense
forest. ‘This is likely to prove true, for example, of the two ground
doves Gallicolumba jobiensis and G. rufigula. Conversely, in the East-
ern Highlands those congeners which have markedly different vertical
foraging preferences appear in no case to have foraging spheres which
completely exclude each other vertically, and other factors also con-
tribute to niche differences. For instance, the honeyeater Meliphaga
aruensis spends more time in the lowerstory than does M. analoga but
also has a stouter bill; and the warbler Sericornis nouwhuysi spends
more time in the lowerstory than do its congeners but is also a larger
bird.
The commonest type of nonspatial segregation among congeners in
the Eastern Highlands depends upon a difference in body size, imply-
ing an average difference in size of food taken or in places food is
sought (a large bird can take larger food, but a light bird can forage
on smaller twigs and perches). ‘There are 15 genera containing two or
more species that overlap broadly in habitat and altitudinal prefer-
ence but differ in size. The prize instance of segregation by size in
New Guinea is provided by the pigeon genus Ptilinopus, which is rep-
resented on the New Guinea mainland by 11 species, all of them fruit
eaters, arboreal, and colored largely green. ‘The five that occur sym-
patrically at Karimui form a graded size series, each about 1.5 times
heavier than the next: P. nanus (average weight 49 ¢), P. pulchellus
(76 g), P. superbus (123 g), P. ornatus (163 g), and P. perlatus (245 g).
A sequence of three arises in the hawk genus Accipiter at Karimui;
A. poliocephalus < A. novahollandiae < A. buergersi. Vhe other cases
are listed in ‘Table 1.
It will be seen from Table 1 that the weight ratio averages 1.9 and
always falls between 1.33 and 2.73, usually between 1.5 and 2.5. ‘he
fact that the values show only this limited spread about the optimal
ratio is due to a balance of selective pressures. Birds with a lower
weight ratio would be too similar to coexist and would have to segre-
gate by some other means (altitude, habitat, allopatry, etc). Birds with
a higher weight ratio would leave an intermediate niche empty: 1.e., a
medium-sized bird could evolve or invade and not be too close in size
to either the heavier or the lighter bird. For instance, if there were
two otherwise similar birds with relative weights of 3.0 and 1.0, both
could also coexist with a medium-sized bird whose relative weight
was 30S ive
Several pairs of congeners differ in other simple and obvious re-
spects related to obtaining food. ‘The whistler Pachycephala leuco-
stigma eats mainly fruit, while its congeners are largely insectivorous.
The various species of honeyeaters of the Meliphaga analoga complex
differ in the extent to which they use flowering trees.
There remain several groups of congeners that share largely or in
part the same gross habitat and are of similar sizes but between which
42
FEATURES OF THE EASTERN HIGHLANDS
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FEATURES OF THE EASTERN HIGHLANDS
the ecological sorting mechanisms cannot be adequately defined at
present. Presumably foraging differences remain to be identified i
most of these cases, while interspecific territoriality based on fine
habitat distinctions may be operative in a few: the hawks Accipiter
novachollandiae and A. fasciatus, the pigeons Ducula zoeae and D.
rufigaster, the ground doves Gallicolumba jobiensis and G. rufigula,
the cuckoo-shrikes Coracina boyeri, C. melaena, C. schisticeps, and C.
morio, the whistlers Pitohui kirhocephalus, P. dichrous, and: i. yer-
rugineus, the honeyeaters Melidectes fuscus and M. princeps, and the
flowerpeckers Melanocharis striativentris and M. longicauda or M.
verstert.
3. ‘Temporal Segregation
In principle, two species could forage in similar manners in the
same habitat if they occupied the habitat at different times of the year
(or, conceivably, of the day). In practice, there are few cases suggestive
of temporal segregation among New Guinea birds, since seasonal
variation is so limited. The pigeon Ptilinopus superbus was breeding
in the Eastern Highlands at a time when its congeners were not breed-
ing. In the south coastal savanna at Merauke, where seasonal changes
may be more marked than in rainforest, the harrier Circus spilonotus
is present in the wet season, Circus approximans in the dry season
(Hoogerwerf, 1964). The postulated local migrations of some montane
lories (p. 80), if they really do take place, may result in temporal re-
placement. The kingfisher Melidora macrorhina is partly or mainly
nocturnal, whereas other kingfishers are diurnal. It would be worth
noting whether those few New Guinea forest species (e.g., Halcyon
megarhyncha, Eurystomus orientalis waigiouensis) which are conge-
neric or conspecific with an Australian form wintering in open habi-
tats in New Guinea spend more time in open habitats after the
migrants leave.
Some General Features of Ecological Sorting Mechanisms
Two general remarks are appropriate at this point.
‘There are systematic changes with altitude in the fineness of niche
differences and in the relative importance of differing sorting mecha-
nisms. Species cliversity decreases with increasing altitude, and niche
differences become correspondingly coarser. At altitudes above 8,500
ft, even in tall Nothofagus forest, most species in the impoverished
avifauna belong to different genera or even families; no genus is rep-
resented by three species living at the same altitude; and only seven
genera (Psittacella, Aegotheles, Collocalia, Sericornis, Rhipidura,
Pachycephala, Melidectes) are represented by two species at the same
altitude. In contrast, below 3,000 ft there are nine genera in which
one may find between four and eight congeners occurring sympatri-
cally at the same altitude. While altitude provides the most important
45
FEATURES OF THE EASTERN HIGHLANDS
sorting mechanism for the New Guinea avifauna taken as a whole, it
is Obviously of no importance in the avifauna of the flat lowlands,
where there are no differences in altitude. A different picture of the
relative importance of various sorting mechanisms might also be ob-
tained if one compared all species pairs in the same family, as Lack
(1944) did for England and as Moreau (1948) did for Tanganyika,
rather than just pairs in the same genus, as done here for the Eastern
Highlands.
The discussion of strict altitudinal segregation applies only to large
areas of extensive virgin forest undisturbed by man, where ecological
interrelationships are characterized by a beauty and simplicity lost in
disturbed areas. Near human settlement one will look in vain for con-
geners replacing each other within 10 vertical feet. For instance, the
whistlers Pitohui nigrescens and P. dichrous had mutually exclusive
altitudinal ranges on Mt. Karimui but were netted at the same altitude
at the forest edge behind Awande. Vertical stratification also breaks
down in ecologically disturbed areas. For instance, the honeyeater
Meliphaga orientalis remains in the middle- and upperstories in un-
disturbed forest and was never caught in mistnets on Mt. Karimui
but was netted twice at Awande. Finally, even reproductive isolating
mechanisms may break down in disturbed areas, as has occurred on
a large scale in the case of the honeyeaters Melidectes belfordi and
M. rufocrissalis.
ZOOGEOGRAPHICAL AFFINITIES OF THE EASTERN
HIGHLANDS AVIFAUNA
The affinities of the Eastern Highlands nongrassland avifauna will
be discussed, first at the species level, then at the subspecies level. ‘Uhis
section extends a previous analysis by Mayr and Gilliard (1954, pp.
328-329), which was based on Gilliard’s collections in the Wahgi
region.
As already mentioned, the Central Range provides an uninterrupted
expanse of montane habitat with no known major east-west barriers
between about longitudes 135°E and 150°E. Most montane species are
distributed over the whole extent of the Central Range. Within the
Central Range three lesser subdivisions may be recognized, most un-
equivocally on the basis of species ranges (presence of a few endemic
species, absence of a few widespread species) but also on the basis of
subspecies:
1. Southeastern New Guinea, extending from long. 150°E west to
the Herzog Mountains (long. 146°30’E). ‘This region is characterized
by six drop-outs, i.e., by the absence of six widespread species (Porzana
tabuensis, Pachycephala tenebrosa, the Paradigalla superspecies, Lo-
phorina (Pteridophora) alberti, Archboldia papuensis, and the Meli-
dectes nouhuysi-M. princeps superspecies); by the absence of
46
FEATURES OF THE EASTERN HIGHLANDS
Epimachus fastosus, a western product of a recent speciation that has
yet to reach the southeast; by the presence of an endemic species,
Amblyornis subalaris, the product of a recent speciation; and by the
presence of five other species or semispecies resulting from recent
speciations, shared with the Eastern Highlands and discussed 1m the
next paragraph.
2. Eastern Highlands, extending east at least to the Okapa area
(long. 145°30’E) and west at least to Tari (long. 143°E) and probably
at least to long. 142°30’E. This region is characterized by the absence
of the nine drop-outs listed on pp. 00-00, and by the absence
of Melidectes ochromelas, which is kept out by competitive exclusion
(pp. 388-389); by the presence of two endemic semispecies, Lophorina
(Astrapia) mayeri and Melidectes princeps; by the presence of two
endemic semispecies shared with southeastern New Guinea, Lophorina
(Parotia) lawesi and Lophorina (Astrapia) stephaniae; and by the pres-
ense of three endemic species shared with southeastern New Guinea,
all products of recent speciation (Cnemophilus macgregori, Para-
disaea rudolphi, and Ptiloprora guiset).
3. Western New Guinea, extending from long. 135°E east at least
to the Telefolmin region (long. 141°30’E). ‘This region contains the
highest mountains of New Guinea and has the most distinctive and
numerous endemic species. here are three endemic species in mono-
typic genera, two of them alpine birds: Anurophasis monorthonyx
(alpine), Androphobus viridis, and Oreornis chrysogenys (alpine).
‘There are four other endemic species, Aegotheles archboldi, Petroica
archboldi (alpine), Pachycephala lorentzi, and Lonchura teerinki, and
two endemic semispecies, Lophorina (Astrapia) splendissima and Meli-
dectes nouhwysi. Seven other species, five of them apparently products
of recent speciation, are shared with the Vogelkop but are absent from
the Eastern Highlands and southeastern New Guinea: Rallicula rubra,
Charmosyna josefinae, Melampitta gigantea, Peneothello cryptoleu-
cus, Pachycephalopsis hattamensis, Ptiloprora erythropleura, and Zos-
terops fuscicapilla, There are five drop-outs present on the Vogelkop
and in eastern New Guinea but not on the western part of the Central
Range (Coracina lineata, Myzomela adolphinae, Melanocharis arfaki-
ana, Erythrura papuana, and Zosterops novaeguineae; the absence of
the last-named species may be correlated with the presence of Zosterops
fuscicapilla). Accipiter buergersi should perhaps also be considered a
drop-out, since it occurs in southeastern New Guinea, the Eastern
Highlands, and the Huon Peninsula but not in western New Guinea,
has no close relatives, and must be a relatively old species.
No systematic collecting has been done between long. 141°30’E and
long. 143°E, or between long. 145°30’E and long 146°30’E. It remains
to be seen how sharp these transition zones are, i.e., whether a region’s
endemic species will disappear and its drop-outs reappear within a
47
FEATURES OF THE EASTERN HIGHLANDS
short distance in these uncollected gaps. ‘These two transition areas
are apparently less than 70-100 miles in extent, while the three regions
are 200-440 miles in extent.
The Eastern Highlands appear to be a fairly uniform region with
respect to species and subspecies distributions above 5,000 ft. The
exceptions to this statement are sufficiently few to be worth detailing.
The only well-established limit of a midmontane species within the
Eastern Highlands is that of the semispecies Lophorina (Astrapia)
mayert and Lophorina (Astrapia) stephaniae which replace each other
around long. 144°E (Mts. Hagen and Giluwe). Mts. Hagen and Giluwe
also have the easternmost known colonies of Archboldia papuensis,
which then apparently disappears again for 350 miles to the west until
Mt. Wilhelmina is reached. The eastern and western semispecies
Lophorina (Parotia) lawesi and Lophorina (Parotia) carolae also prob-
ably meet in the Eastern Highlands, but the details are still unknown.
The only midmontane species which appears to be represented in the
Eastern Highlands by well-marked eastern and western races is Pene-
othello sigillatus. ‘Vhree other midmontane species (Coracina caeruleo-
grisea, Sericornis nouhuysi, and Paradisaea rudolphi) are represented
in the Eastern Highlands by thinly differentiated eastern and western
races which intergrade clinally. In the cases of three species (Lophorina
(Astrapia) stephaniae, Melidectes rufocrissalis, and Melidectes bel-
fordi) the race on the outlying Schrader Range differs from that in
the remainder of the Eastern Highlands.
For analysis of subspecific affinities the Eastern Highlands avifauna
will be broken down into five somewhat arbitrary categories based on
altitude, since several different patterns emerge:
1. Alpine species: species confined to alpine gre assland or the adyja-
cent forest edge, rarely descending below 10,000 ft in the Eastern High-
lands. ‘These species live only on the highest mountains in New
Guinea and show the highest degree of endemism, as expected from
the broken and isolated nature of their habitat. Of the seven alpine
species in the Eastern Highlands, one (Melidectes princeps) is an en-
demic semispecies; three (Megalurus timoriensis montanus, Turdus
poliocephalus erebus, Oreostruthus fuliginosus hagenensis) are repre-
sented by endemic subspecies, the latter two nearest the southeastern
New Guinea forms; two (Petroica bivattata bivittata and Melidectes
fuscus fuscus) belong to the southeastern New Guinea race, while the
western New Guinea race differs; and one (Anthus gutturalis rhodo-
dendri) belongs to the Huon Peninsula race, which is nearer the south-
eastern New Guinea race than the western New Guinea race. The
reason for these southeastern affinities of the alpine avifauna of the
Eastern Highlands is probably that the alpine grasslands of south-
eastern New Guinea are less distant than those of the Snow Mountains
of western New Guinea.
2. Midmontane species: birds that do not descend below 5,000 ft,
48
FEATURES OF THE EASTERN HIGHLANDS
excepting alpine species, which have already been considered. Grass-
land and swamp species will also be considered separately (pp. 71-72).
Twenty-seven midmontane species provide no indication of Z00-
geographical affinities, either because all New Guinea populations
belong to one race, or else because all Central Range populations
belong to one race and differentiation occurs only on isolated moun-
tain “islands” (Vogelkop, Huon Peninsula, etc.). Of the remaining
species, ten (Psittacella picta excelsa, Psittacella madaraszi hallstromt,
Epimachus meyeri bloodi, Cnemophilus (Loria) loriae amethystina,
Cnemophilus macgregorii sanguineus, Archboldia papuensis sanfordi,
Daphnoenositta miranda kuboriensis, Melidectes rufocrissalis thomas,
Ptiloprora guisei uwmbrosa, Zosterops novaeguineae wahgiensis) are
represented by valid endemic subspecies, with two additional endemic
subspecies on the Schrader Range (Lophorina (Astrapia) stephaniae
feminina and Melidectes belfordi schraderensis). In the cases of 19
species of which there are two races, a western and an eastern race, on
the Central Range, racial determinations are available for the popula-
tions of the Snow Mountains (about 400 miles west of the Eastern
Highlands), ‘Velefolmin (about 200 miles west of the Eastern High-
lands), the Eastern Highlands, and the Wharton Range of southeastern
New Guinea (about 200 miles east of the Eastern Highlands). In eight
of these 19 cases the racial division falls between southeastern New
Guinea and the Eastern Highlands; in five cases, between the Eastern
Highlands and ‘Telefolmin; and in six cases, between ‘Telefolmin and
the Snow Mountains. In the four additional cases mentioned on p. 48,
western and eastern races divide within the Eastern Highlands. ‘The
Eastern Highlands population of Pachycephala modesta belongs to
the race of the Huon Peninsula mountain “island,” and the south-
eastern and ‘Telefolmin populations on the Central Range are both
racially distinct.
‘Thus, almost all geographic variation of midmontane species on the
Central Range is east-west variation, and affinity decreases with in-
creasing distance. ‘Vhere are somewhat closer affinities with the popula-
tions to the east in southeastern New Guinea than with the more
distant Snow Mountains of western New Guinea, as concluded pre-
viously by Mayr and Gilliard (1954, p. 329) and as also true of the
alpine avifauna. Except for the three isolates of the Schrader Range
(p. 48), there are no racial differences between the northern and
southern slopes. ‘his is as one would expect, since the Central Range
extends much further east-to-west than north-to-south and presents no
major barriers to a midmontane species. ‘The dissected terrain consists
of mountains separated by valleys which provide minor barriers and
whose floors along the watershed divide lie at 5,000 ft or higher. A
species will therefore find suitable habitat patchier the higher its
altitudinal range lies, and the group with the highest percentage of
differentiated forms is the alpine avifauna.
49
FEATURES OF THE EASTERN HIGHLANDS
3. Tropical species: birds ranging from sea level up to a ceiling
which is lower than 5,000 ft. Since the Central Range has no passes
under 5,000 ft, it acts as a barrier which completely stops direct contact
between populations in the northern New Guinea lowlands and
southern New Guinea lowlands. Two lengthy indirect paths are avail-
able for north-to-south gene flow: around the tip of southeastern
New Guinea in the extreme east, and across the Geelvink Gap or
around the Vogelkop in the extreme west. Thus, the populations of
tropical species form a continuous or broken ring in the lowlands
around the periphery of New Guinea. The tropical avifauna on the
southern slopes of the Eastern Highlands is well known taxonomically
as a result of my collections and those of Schodde and Hitchcock,
whereas little material from below 5,000 ft on the northern slopes
has been evaluated taxonomically. ‘The following comments therefore
apply only to the southern slopes unless stated otherwise, and mainly
to my material from the Karimui area and Okasa. ‘The conclusions
reached by Schodde and Hitchcock concerning Lake Kutubu will be
cited separately for comparison.
‘The races of 42 tropical species replace each other consecutively in
simple fashion as one proceeds around the lowlands periphery of New
Guinea. There are, of course, other species in which all New Guinea
lowlands populations belong to the same race. Regarding east-west
differences, there are 24 of these 42 cases in which the southeastern,
Eastern Highlands, and southwestern populations belong to the same
race; 1] cases in which one race occurs in southwestern New Guinea,
the Fly River, and Eastern Highlands and a different race in south-
eastern New Guinea; five cases in which one race occurs in south-
western New Guinea and on the Fly River and a different one in the
Eastern Highlands and southeastern New Guinea; and two cases in
which one race occurs in southeastern New Guinea, the Eastern High-
lands, and part of the Fly River, while a different race occurs on
another part of the Fly River and in western New Guinea. Thus, the
Karimui-Okasa avifauna is slightly closer to that of the Fly River to
the west than to that of southeastern New Guinea to the east, as
Schodde and Hitchcock (1968, p. 11) also showed for the Lake Kutubu
avifauna. Regarding north-south differences, there are four instances
(the Goura superspecies, the Chalcopsitta superspecies, the Psittacult-
rostris superspecies, and the Paradisaea raggiana-P. minor superspecies)
in which one semispecies occurs in southern New Guinea and the
Eastern Highlands, a different semispecies in northern New Guinea;
34 cases in which one subspecies occurs in southeastern New Guinea
and the Eastern Highlands, a different subspecies in northern New
Guinea; only five cases where the southern New Guinea, northern New
Guinea, and Eastern Highlands populations all belong to the same
subspecies; and no instance in which the Eastern Highlands population
belongs to a northern New Guinea race and differs from a southern
50
FEATURES OF THE EASTERN HIGHLANDS
New Guinea race. Thus, the affinities of the avifauna on the southern
slopes of the Eastern Highlands are exclusively southern, as expected.
That north-south differentiation is more frequent than east-west dif-
ferentiation is also as expected, since the distances within the southern
New Guinea lowlands or northern New Guinea lowlands are less (and
the opportunities for gene flow therefore greater) than between
northern and southern New Guinea around the eastern and western
tips of the Central Range.
Gilliard, Gyldenstolpe, and Shaw-Mayer obtained specimens of seven
tropical species from the northern slopes of the Eastern Highlands.
These all belong to northern semispecies (Goura victoria, Psittaculi-
rostris edwardsi, Paradisaea minor) or subspecies (Gallicolumba ruft-
gula septentrionalis, Ceyx azureus ochrogaster, Lophorina (Cicin-
nurus) regia similis, and Ailuroedus buccoides geislerorum).
Four tropical species are represented by endemic subspecies which
probably have narrow ranges on the southern slopes of the Eastern
Highlands (Domicella lory somu, Aegotheles bennetti terborghi, Rhi-
pidura leucothorax clamosa, Myzomela eques karimuitensis).
A different pattern of variation appears in four other tropical species:
one race is widespread over most of New Guinea, and a different race
is confined to the Fly River lowlands. ‘The Eastern Highlands popula-
tion belongs to the widespread race in three cases (Crateroscelis murina
murina, Microeca flavovirescens cuicui, Myzomela nigrita meyert) and
to the Fly River race in one case (Amaurornis olivaceus ruficrissum).
Finally, there are three cases in which the racial affinities of the
Eastern Highlands population are neither with southeastern New
‘Guinea directly to the east nor with the Fly River directly to the west,
but with the southern slopes of the Snow Mountains farther to the
west, giving a checkerboard distribution (Coracina melaena melaena,
Pitohui ferrugineus ferrugineus, Meliphaga flaviventer rubiensis).
4. Hill forest species: birds whose ceilings lie below 5,000 ft but
which do not descend as far as sea level. Just as for tropical species,
the Central Range is a complete barrier, except at its ends, to the
north-south movement of birds in this category. Consideration. is
again limited to forms on the southern slopes of the Eastern Highlands
unless stated otherwise.
Eleven hill forest species show no geographical variation on the
Central Range. In nine species variation fits a simple east-west and/or
north-south pattern. Regarding east-west differences, the southeastern,
Eastern Highlands, and southwestern (i.e., southern slopes of the
Snow Mountains) populations belong to the same race in three cases;
in five cases the southeastern and Eastern Highlands populations be-
long to one race, the southwestern population to another; and in one
case (Pachycephala hyperythra) the southeastern and Eastern High-
lands populations belong to the same race, but the species is absent
from southwest New Guinea. ‘The hill forests at the head of the Fly
51
FEATURES OF THE EASTERN HIGHLANDS
River cannot be included in this comparison because they are virtually
unexplored ornithologically. Regarding north-south differences, the
Eastern Highlands and southeastern (or southern) populations belong
to the same race in all nine cases, while the species is absent from
northern New Guinea in one case (Meliphaga auga) and is represented
by a different population in northern New Guinea in all eight other
cases. In three instances (Manucodia (Phonygammus) keraudrenit,
Lophorina (Diphyllodes) magnifica, Ailuroedus crassirostris) Gilliard
or Gyldenstolpe took specimens from the northern slopes of the Eastern
Highlands and found them to belong to the northern race. Thus, the
main pattern of geographic variation for hill forest species in the
Eastern Highlands is identical to that for tropical species.
More complex patterns are met in four hill forest species. Pachycare
flavogrisea shows clinal east to west variation, and it proves convenient
to recognize the boundary between two races as occurring in the
Eastern Highlands. Peneothello bimaculatus is represented by one race
in the northwestern, southwestern, Eastern Highlands, and southern
slopes of southeastern New Guinea, by another race on the northern
slopes of southeastern New Guinea and on the Huon and Adelbert
“north mountain islands.” ‘The situation is reversed for Zosterops
atrifrons, of which the populations of the Eastern Highlands (southern
slopes) and the northern slopes of southeastern New Guinea are con-
racial, while the populations of the southern slopes of southeastern
New Guinea and the southwest are racially distinct. Finally, Pachy-
cephalopsis poliosoma from the Eastern Highlands (southern slopes)
proves, surprisingly, to belong to the race albigularis from the northern
slopes at the western extremity of the Central Range, with distinct
races intervening in southeastern, southwestern, and northeastern New
Guinea. ‘There are no endemic hill forest subspecies in the Eastern
Highlands.
5. Birds ranging both above and below 5,000 ft. Only five species
in this category exhibit north-south racial differences in eastern New
Guinea. ‘Three populations (Cacomantis variolosus oreophilus, Meli-
phaga orientalis facialis, Melanocharis striativentris striativentris)
belong to the southern race, two (Opopsitta diophthalma festetichi,
Trichoglossus haematodus intermedius) to the northern race. ‘The ceil-
ings of these five species vary locally between about 5,500 and 6,800 ft,
and the number of north-south passes at these altitudes is limited, so
that the Central Range, although not a complete barrier to north-south
gene flow, is still somewhat of a barrier.
Simple east-to-west variation involving two races on the Central
Range is encountered in six cases. ‘The division falls between south-
eastern New Guinea and the Eastern Highlands in three cases (Ralli-
cula forbesi, Alisterus chloropterus, Pachycephala soror), between the
Eastern Highlands and ‘Telefolmin in one case (Dicaeum geelvin-
kianum), and between Telefolmin and the Snow Mountains in two
52
FEATURES OF THE EASTERN HIGHLANDS
cases (Cacomantis castaneiventris, Amblyornis macgregoriac). In two
instances (Coracina caeruleogrisea, Paradisaea rudolpht) eastern and
western races meet within the Eastern Highlands.
The Eastern Highlands populations of three species belong to
endemic subspecies (Caswarius bennettii shawmayeri, Tregellasta leu-
cops wahgiensis, Paradisaea rudolphi margaritae, plus Lophorina su-
perba pseudoparotia on the Schrader Range). .
In two instances (Lophorina superba latipennis, Melanocharis longt-
cauda captata) Eastern Highlands populations belong to the subspecies
of the Huon Peninsula mountain “island” and differ racially from the
Central Range populations to the east and west. These patterns almost
surely arose from populations differentiating in the Eastern Highlands
and then invading the Huon Peninsula, not yice versa (cl., also,
Pachycephala modesta, p. 49).
The Eastern Highlands population of Meliphaga orientalis is derived
from the southern slopes of western New Guinea (facials) rather than
the southern slopes of southeastern New Guinea (orientalis) and differs
from the northwestern (citreola) and northeastern (beck) populations.
Colluricincla megarhyncha is represented in the Wahgi Valley by
the northeastern New Guinea race tappenbecki and on the southern
slopes of the Eastern Highlands by the Herzog Mountains race nea.
The populations from southeastern New Guinea and the Fly River
differ.
Summary of zoogeographic affinities. ‘Lhe hindrances to gene flow
implicit in geographic variation result in New Guinea either from
geographical barriers, hence a discontinuity in the distribution of a
species, or else simply from distance. ‘he Central Range runs east-west
and lacks passes under 5,000 ft. For tropical and hill forest species,
which are absent above 5,000 ft, the Central Range functions as a
complete or nearly complete barrier, so that there are marked differ-
ences between northern and southern populations, as well as east-west
variation resulting from distance. For midmontane species, which do
not descend below 5,000 ft, north-south population differences are
virtually nonexistent, and variation is east-to-west. ‘The higher a species’
altitudinal lower limit, the patchier the environment becomes to the
species, so that the highest proportion of endemic forms is encountered
among alpine birds. Eastern and western distributional limits of species
and semispecies can be used to divide the Central Range into three
sections, of which the Eastern Highlands as defined in this monograph
is one.
AVIFAUNA OF THE KARIMUI BASIN
The unusual bird distribution patterns in the Karimui Basin re-
quire separate discussion. As already mentioned, Karimui is a flat basin
with a floor lying at about 3,500 ft, and is effectively sealed off from
53
FEATURES OF THE EASTERN HIGHLANDS
the outside by a ring of mountains. The Karimui avifauna is marked
by the presence of many lowland species whose altitudinal ceiling else-
where lies considerably below 3,500 ft; by the absence or rarity of some
hill forest species normally encountered at this altitude; and by the
presence of three strikingly distinct endemic forms. The avifauna we
found at Karimui Patrol Post (3,650 ft) contrasted with that of the
Okasa forest (3,550-4,250 ft), which is at the same altitude but in hilly
terrain typical for this altitude in most of New Guinea.
The following 26 or 27 lowland forest species were present at Kari-
mui but rarely reach this altitude elsewhere in New Guinea. Some of
these species, in fact, normally disappear as soon as one leaves the flat
lowlands at sea level. None was present at the same elevation at Okasa,
and Schodde and Hitchcock met only 11 at Lake Kutubu despite the
elevation there (2,450 ft) being 1,200 ft lower than at Karimui Patrol
Post: Aviceda subcristata, Rallina tricolor, Ptilinopus nanus, Megalo-
prepia magnifica, Chalcophaps stephani, Charmosyna placentis, Psit-
taculirostris desmarestt1, Opopsitta gulielmiterti, Geoffroyus geoffroyt,
Eudynamis scolopacea, Centropus menbeki, Aegotheles bennettii, Hal-
cyon torotoro, Campochaera sloeti, Coracina boyeri, Sericornis spilo-
dera, Rhipidura threnothorax, Monarcha chrysomela, Microeca flavo-
virescens, Poecilodryas placens, Pitohui kirhocephalus, Pitohwui
ferrugineus, Lophorina (Cicinnurus) regia, Ailuroedus buccoides, My-
zomela eques, Pycnopygius ixoides, and probably Casuarius casuarius.
The following nine lowland species of the forest edge, second-
growth, and open country were present at Karimui somewhat above
their normal altitudinal ceiling. All were absent at Okasa, but Schodde
and Hitchcock found all except Circus approximans at Lake Kutubu
(2,450 ft): Circus approximans, Amaurornis olivaceus, Ptilinopus per-
latus, Dacelo gaudichaud, Rhipidura leucothorax, Mino dumontii,
Cracticus cassicus, Dicrurus hottentottus, and Nectarinia sericea.
The following ten hill forest species which would normally be
encountered at the elevation of Karimui were absent: Caswarius ben-
nettii, Charmosyna pulchella, Halcyon megarhyncha, Sericornis arfaki-
anus, Phylloscopus trivirgatus, Rhipidura atra, Microeca griseoceps,
Lophorina (Parotia) lawesi, Lophorina superba, and Ailuroedus crassi-
rostris. In the case of three additional hill forest species expected at
3,500 ft, one or more immatures were taken at Karimui but no adults:
Monarcha axillaris, Pachycephala soror, and Manucodia (Phonygam-
mus) keraudrenin. Adults of five additional hill forest species were
taken at Karimui but were much rarer than expected at 3,500 ft:
Tregellasia leucops, Pachycephalopsis poliosoma, Pachycephala leuco-
stigma, Melidectes torquatus, and Meliphaga orientalis. Fifteen of
these 18 hill forest species were present at Okasa and include all of
the commonest Okasa birds.
In seven of these cases a species unexpectedly present and a species
unexpectedly absent at Karimui are successive members of an altitudinal
54
FEATURES OF THE EASTERN HIGHLANDS
sequence (p. 27), so that the “wrong” member of the sequence was
present: Charmosyna placentis vs. C. pulchella, Halcyon torotoro vs.
H. megarhyncha, Sericornis spilodera vs. S. arfakianus, Rhipidura
threnothorax vs. R. atra, Microeca flavovirescens vs. M. griseoceps,
Ailuroedus buccoides vs. A. crassirostris, and probably Caswarius casu-
arius vs. C. bennettii.
‘Two kinds of evidence indicate that the distributional anomalies at
Karimui are ultimately due to the flatness of the basin floor. ‘The first
line of evidence is based on the changes in the avifauna on the lower
slopes of Mt. Karimui, which rises out of the basin. Near the base of
Mt. Karimui’s west ridge the basin floor begins to slope gently upward
as one goes towards the mountain, until at an elevation of 4,200 ft the
foot of the west ridge is encountered and the mountain abruptly rises
at a steeper angle. Of the eight altitudinal zones into which I arbi-
trarily divided Mt. Karimui, the lowest, Zone 1, was on the sloping
basin edge just at the foot of the ridge and spanned an altitude range
from 4,000 to 4,200 ft. Although Zone | began at an altitude only 350
ft above Karimui Patrol Post (3,650 ft), my main collecting station on
the flat basin floor, all but six of the 26 or 27 lowland forest species
present at Karimui above their normal ceiling were absent from Zone
1, and five of these six disappeared as soon as the west ridge was
reached, i.e., in Zone 2. Conversely, of the 18 hill forest species absent
or rare on the basin floor, 10 were present and generally common
already in Zone 1, and seven of the remaining eight appeared in Zone
2. ‘Thus, the distributional anomalies of the flat basin floor were re-
versed as soon as sloping terrain was encountered even though there
had been only a small change in altitude itself.
‘The second and more striking line of evidence that the anomalous
lowlands avifauna of the Karimui Basin is due to its flatness was
provided by the discontinuous altitudinal ranges of at least three hill
forest species. ‘he flycatcher Rhipidura atra was present on the slopes
of Mt. Karimui from 4,090 ft upward and was absent on the basin
floor. When I crossed the mountain wall ringing the basin and de-
scended into hilly country en route to Soliabeda, Rhipidura atra
reappeared at 3,230 and 2,770 ft. On the slopes of Mt. Karimui the
parrot Charmosyna pulchella was present from 4,400 ft upward, and
the bird of paradise Loboparadisea sericea from 4,000 ft upwards. Both
were absent on the basin floor but reappeared at Soliabeda, which is
at 2,000 ft in hilly country. The absence of these three species in the
basin thus could not have been due to altitude, since they were present
at both lower and higher altitudes in hilly terrain, and must have been
due to the flatness. Had more time been spent en route between
Karimui and Soliabeda, and had the difficulty and circumstances of
the route not been so unconducive to bird-watching, I suspect that
similar instances of discontinuous altitudinal distributions would
have been unmasked in some other hill forest species.
55
FEATURES OF THE EASTERN HIGHLANDS
‘The explanation these facts suggest for the anomalous avifauna of
Karimui is that the flatness of the basin floor has caused it to fill up
with the rainforest, and hence many of the birds, characteristic of the
flat lowlands rather than of the hill slopes. This interpretation seems
reasonable, since the slope governs the effectiveness of drainage and
hence the plant communities, and since quite a few New Guinea bird
species (¢.g., Otidiphaps nobilis, Gymnophaps albertisii, Lophorina
(Diphyllodes) magnifica), are absent from the flat lowlands but present
in hilly terrain even near sea level. What makes Karimui unique is
the general ruggedness of New Guinea’s hills and mountains, so that
one rarely, or nowhere else, finds a flat area at 3,500 ft as extensive
as at Karimui.
Three of these isolated populations trapped within the Karimui
Basin have differentiated to yield strikingly distinct endemic forms, all
tending towards melanism (Diamond, 1967a). ‘The unique type of the
owlet-nightjar, Aegotheles bennetti terborghi, not only represents an
altitudinal record for this lowland rainforest species but also is 25%
larger in linear dimensions than the other races (hence presumably
twice as heavy) and much darker. The goshawk Accipiter novaehol-
landiae exists elsewhere in two color phases, one pure white, the other
gray and rufous, while the Karimui population is melanistic, being
uniform dark gray-brown. The mannikin Lonchura_ spectabilis
gajduseki has buff underparts as opposed to white in other races. ‘The
distinctness of these endemic forms suggests that strong selective
pressures are operative at Karimui and that the mountain ring is an
effective barrier to gene flow for some species. For such species the
Karimui Basin represents an isolated island of tropical habitat,
analogous to the evolutionary significance of isolated high mountain
summits for alpine species.
ALTITUDINAL DISTRIBUTION
In this section the results of my altitudinal censuses in the Karimui
area (Soliabeda, Bomat, Karimui, Mt. Karimui) will be analysed. Pre-
vious analyses of the altitudinal distribution of New Guinea birds
include those by Stresemann (1923, pp. 7-15), Archbold and Rand
(1935, pp. 535-543), and Archbold, Rand, and Brass (1942, p. 285).
I identified 242 bird species in the Karimui area. Of these, 36 were
nonforest species, strictly confined to grassland, second-growth, or
water. Such habitats do not exist on Mt. Karimui, and these 36 species
are therefore excluded from consideration in the present analysis. Of
the 206 forest species, there were 40 of which I obtained too few records
to work out the altitudinal range satisfactorily.
Figure 4 presents the altitudinal ranges of the 166 forest species for
which my data are adequate. All altitudinal ranges were assumed to be
continuous; if a bird was recorded above and below but not at a cer-
56
FEATURES OF THE EASTERN HIGHLANDS
tain elevation, it was assumed to be present at that elevation but to
have been overlooked. This assumption is probably valid except in
the cases of a few hill forest species present above and below but not
in the Karimui Basin (pp. 55-56). Birds present at Soliabeda, my lowest
station (1,350-2,000 ft), are arbitrarily depicted in Figure 4 either as
continuing down to sea level or else as having their lower limits at
Soliabeda, the decision being made on the basis of my ex perience
elsewhere in New Guinea or on the basis of published information.
On Mt. Karimui, Zone 1 (4,000-4,200 ft), Zone 2 (4,400-4,750 ft), Zone
3a (4,750-5,080 ft), Zone 3b (5,080-5,390 ft), Zone 4a (5,390-5,780 ft),
Zone 4b (5,780-5,960 ft), Zone 5a (5,960-6,250 ft), Zone 5b (6,250-6,500
ft), Zone 6a (6,500-6,770 ft), Zone 6b (6,770-7,080 ft), Zone 7a (7,080-
7,280 ft), Zone 7b and 8a (7,280-7,620 ft), and Zone 8b (7,620-8,165 ft,
the summit) were each considered as units in constructing Figure 4:
i.e., if a species was recorded anywhere within the unit, it is depicted
as being present throughout the unit.
The following conclusions may be extracted from Figure 4:
l. Species diversity (Fig. 5). “The number of forest species recorded
at a given altitude is relatively constant at about 100 species from
3,650 ft (Karimui) and 3,250 ft (Bomai) down to at least 1,350 ft (the
lower limit of collecting at Soliabeda). On Mt. Karimui the number
of species decreases regularly with increasing altitude, from 77 at
4,000-4,200 ft (Zone 1) to 16 in the summit zone (7,620-8,165 ft). Pos-
sibly some of the difference between the species totals for the lowest
elevation on Mt. Karamui (77 species) and the three village locations
at lower elevations (ca. 100 species) is because larger and more diverse
areas of forest were surveyed at a given elevation at the village locations
than on Mt. Karimui. The apparent slight maximum in Figure 5 at
3,650 ft (Karimui) is probably an artifact of more time spent collecting
at Karimui than at other localities. It should be mentioned again
that the actual number of forest species is about 25% higher than
depicted in Figure 5, since ranges for only 166 of the 206 forest species
in the area were used to construct Figure 5.
Three factors contribute to the decrease in species diversity with
altitude depicted in Figure 5. First, with increasing elevation the forest
becomes more stunted, and MacArthur, Recher, and Cody (1966) have
shown that bird species diversity among different habitats in the same
area correlates well with the foliage profile diversity. Second, the total
area in New Guinea at a given altitude decreases with increasing
altitude. Over long times, the total number of species that can evolve
to utilize a given type of habitat should be larger, the greater the
available area of the habitat, and one would therefore expect fewer
high-altitude than low-altitude species in New Guinea even if the
foliage did not change. For example, there is a much greater area of
alpine grassland, and consequently much greater variety of alpine
grassland bird species, in the South American Andes than in New
57
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FEATURES OF THE EASTERN HIGHLANDS
SPECIES DIVERSITY, MT. KARIMUI
1205
e)
1004
80
number
of 60
species |
40-4
20-4
= Lee ia TIF T Ie le
O |IOOO 2000 3000 4000 5000 6000 7000 8000
altitude (feet) summit
Fic. 5. Species diversity on Mt. Karimui. Each point gives the number of forest
species recorded at the given elevation. Species diversity decreases with increasing
altitude.
Guinea. Third, as discussed previously (p. 24), Mt. Karimui is a
small, isolated mountain and does not have all the species present on
larger New Guinea mountains due to drop-outs. Mt. Karimui supports
the smallest fraction of the available species pool at the highest eleva-
tions, where the effects of isolation and small land mass are most
pronounced.
2. Species with broad altitudinal ranges. Figure 4 shows that most
species have ranges of less than 5,000 but more than 1,000 feet. Only
19 species have ranges greater than 5,000 ft, and only six of these
exceed 6,500 ft. All 19 of these wide-ranging species descend to sea
level. No montane species (i.e., a species that fails to descend to sea
level) has a range greater than 4,200 ft.
Of the 19 wide-ranging species, 12 have no congeners in the Eastern
Highlands, either because they belong to monotypic genera or else have
congeners restricted to other islands or other parts of New Guinea.
This correlation is another expression of the fact that altitudinal
segregation is the most important ecological sorting mechanism in
New Guinea that permits sympatry between congeners. Hence species
without congeners are the ones most likely to have a broad altitudinal
range.
3. Species with narrow altitudinal ranges. ‘Vhere are two classes of
species with ranges narrower than 1,000 feet. One of these consists of
66
FEATURES OF THE EASTERN HIGHLANDS
high altitude species for which Mt. Karimui is barely high enough
and which can exist only near the summit. For instance, the bird of
paradise Cnemophilus macgregorii was confined to the top 545 {t,
and the flowerpecker Paramythia montium was not seen more than
40 ft below the summit. The other class consists of species whose
ranges fall in the middle of the mountain and in some cases are com-
pressed by congeners living at higher or lower altitudes. For instance,
the total range of the warbler Sericornis arfakianus 1s 600 feet, the
whistler Pitohui nigrescens is 900 feet, and the bird of paradise
Manucodia (Phonygammus) keraudrenti is 750 feet. ‘These narrow
altitudinal bands imply excessively small populations which might
easily be eliminated by an unusual environmental fluctuation (a poor
breeding season, disease, expansion of another species, etc.). Such
species are most like to drop out on Mt. Karimui or to have difficulty
becoming established in the first place.
4. Altitudinal “zones.” If the altitudinal limits of many species
tended to coincide, this fact could be expressed by recognizing altitudi-
nal zones of faunal distribution. If limits of many species did not
coincide, zones would be arbitrary and not worth recognizing. ‘This
question is examined in Figures 6-9.
Figure 6 plots, as a function of altitude, the absolute number of
species per 100 feet reaching their upper altitudinal limit in a given
zone. Figure 7 reexpresses the same numbers as the percentage of the
species in a given zone which reach their upper limit in each 100 feet of
the zone. It is apparent that few species (only seven) reach their ceilings
below Karimui (3,650 ft). ‘here would, of course, be some further
species present at sea level whose ceilings lie below Soliabeda and
which were therefore not observed in these studies. Species ceilings
are well distributed from 3,650 ft to the summit, with some concentra-
tion at three maxima: viz., between the flat basin floor at Karimui
(3,650 ft) and the lowest slopes of Mt. Karimui (4,000 ft), where 35
species drop out; at the moss level (6,500 ft), where nine species (20%,
of the avifauna at 6,350-6,500 ft) drop out; and above 7,280 ft, where
13 species drop out before the summit is reached and 16 species con-
tinue up to the summit and necessarily drop out there. Seventy-two
species have ceilings between 4,000 ft and the moss level on Mt.
Karimui, but there are no well-defined maxima in this range.
Figures 8 and 9 are the corresponding figures for the absolute and
relative numbers of species reaching their lower limits per 100 feet.
Below Bomai (3,250 ft) species drop out at a low rate (24 species, or
somewhat less than one species or 19% of all species present dropping
out per 100 vertical feet). Above Bomai the frequency of species lower
limits, whether expressed absolutely or relatively, first increases and
then decreases with increasing elevation and has a very broad maxi-
mum around 5,000 ft. “Phe apparent increase again at the highest
altitude (farthest right-hand point of Figures 8 and 9) reflects the
67
FEATURES OF THE EASTERN HIGHLANDS
SPECIES UPPER LIMITS (ABSOLUTE), MT. KARIMUI
number
per ©
lO@: 54
feet
E hae anes ! eae eee =a
© 1000 2000 3000 4000 5000 6000 7000 8000
altitude (feet) summit
Fic. 6. Absolute frequencies of the upper altitudinal limits of forest species on
Mt. Karimui, as a function of altitude. The ordinate gives the number of species
whose upper altitudinal limits lay in a vertical band of 100 feet centered about
the altitude given on the abscissa. Species ceilings are well distributed over the
whole mountain above about 3,500 ft, with an accumulation at 4,000 ft (transition
between flat basin floor and hill slopes) and a lesser accumulation at 6,500 ft (the
moss level).
appearance of only two new species in the depauperate avifauna of
Zone 8.
One could therefore speak of a relatively uniform tropical zone ex-
tending from the Karimui Basin (3,650 ft) down to Soliabeda (1,350-
2,000 ft) and (by extrapolation) to sea level, where the total number
of species remains quite constant and few species have their ceilings
or lower limits. Subdivision of Mt. Karimui (>4,000 ft), which lies
above this tropical zone but still contains a majority of tropical species
for some distance above 4,000 ft, into several zones is not possible.
Instead, new species gradually come in with increasing altitude, with
a very broad maximum around 5,000 ft, and species gradually reach
their ceilings, with a slight maximum at the moss level ang. an ac
celeration towards the summit.
Previous authors have discussed a “sharp avifaunal break which
occurs at about 4,500-5,000 ft in the altitudinal zonation of New
Guinea birds” (Gilliard and LeCroy, 1961, p. 22; cf., also, Stresemann,
1923, who places the break at 1,500 m = 4,920 ft, and Archbold and
Rand, 1935, who place it at 1,700 m = 5,580 ft). In the field I had the
strong impression that such a sharp break did exist. ‘This impression
was due partly to the rapid species turnover on Mt. Karimui (e.g., only
FZ species are found at both 3,650 and 6,000 ft); and partly because
68
FEATURES OF THE EASTERN HIGHLANDS
SPECIES UPPER LIMITS (RELATIVE), MT. KARIMUI
IS
%o
per
100
feet
O |O0O0 2000 3000 4000 5000 6000 7000 8000
altitude (feet) summit
Fic. 7. Relative frequencies of the upper altitudinal limits of forest species on
Mt. Karimui, as a function of altitude. The ordinate gives the percentage of species
present at a given elevation whose upper altitudinal limits lay in a vertical band
of 100 feet centered about the altitude given on the abscissa.
some common and conspicuous species reached their upper limit
(Crateroscelis murina, Pachycephalopsis poliosoma, Meliphaga flavi-
venter) or lower limit (Crateroscelis robusta, Sericornis perspicillatus,
Rhipidura albolimbata, Peneothello cyanus) near 5,000 ft. However,
the detailed results presented in Figures 6-9 show that one cannot speak
of a sharp break at 5,000 ft on Mt. Karimui: there is no increased
frequency of ceilings and only a gentle maximum in frequency of lower
limits. If one had to designate any break on Mt. Karimui, it would be
at 3,650-4,000 ft, and even that one would involve only a minority of
the species present.
‘These findings concerning Mt. Karimui cannot be assumed valid
for other New Guinea mountains without further testing, since dis-
tributional patterns on Mt. Karimui are unusual in three respects.
First, the intrusion of the flat basin has caused a conspicuous tropical
element to persist in undisturbed forest up to 3,650 ft to a degree for
which I know of no parallel elsewhere in New Guinea. Elsewhere I
69
FEATURES OF THE EASTERN HIGHLANDS
SPECIES LOWER LIMITS (ABSOLUTE), MT. KARIMUI
2.55
20-
number 1.574
per
lOO
feet lo4
Olas
: i |
O 1000 2000 3000 4000 5000 6000 7000 8000
altitude (feet) summit
Fic. 8. Absolute frequencies of the lower altitudinal limits of forest species on
Mt. Karimui, as a function of altitude. The ordinate gives the number of species
whose lower altitudinal limits lay in a vertical band of 100 feet centered about
the altitude given on the abscissa. The frequency of lower limits reaches a very
broad maximum around 5,000 ft.
would not expect so little change in the avifauna between 1,350 and
3,650 ft and so marked an accumulation of species ceilings at 3,650-
4,000 ft, i.e., so marked a tropical “zone.” Second, moss forest descends
lower on Mt. Karimui than on the highest central peaks of New
Guinea, though it descends still lower on isolated peaks nearer the
sea. The avifauna at 7,500 ft on Mt. Karimui is similar to that at
10,000 ft on Mt. Michael, and birds compressed between 4,000 ft and
6,500 ft on Mt. Karimui would range from 3,000 to 8,500 ft on larger
mountains. Finally, Mt. Karimui is too low to support the depauperate
New Guinea alpine grassland and timberline avifauna.
BREEDING NON-FoREST AVIFAUNA
Species Composition
In areas undisturbed by man the original vegetation is forest over
most of New Guinea. The only extensive area in which the climax
vegetation is not forest is the low-rainfall savanna strip on the south
coast, where most birds are Australian rather than typical New Guinea
species. Elsewhere original non-forest vegetation consists of alpine
orassland, naturally disturbed areas (due to landslides, earthquakes, or
fallen trees), swamps and marshes, and the edges of streams and rivers.
70
FEATURES OF THE EASTERN HIGHLANDS
SPECIES LOWER LIMITS (RELATIVE), MT. KARIMUI
Sa7
205
Zo]
05>
4 Se f T
O 1IOOO 2000 3000 4000 5000 6000 7000 8000
altitude (feet) summit
Fic. 9. Relative frequencies of the lower altitudinal limits of forest species on
Mt. Karimui, as a function of altitude. The ordinate gives the percentage of species
present at a given elevation whose lower altitudinal limits lay in a vertical band
of 100 feet centered about the altitude given on the abscissa.
The total unforested area in undisturbed regions at middle elevations
is minute. As a result the breeding non-forest avifauna of the Eastern
Highlands is limited in variety (about 80 species in all types of non-
forest habitats at all elevations above 1,350 ft). In areas settled by man
non-forest habitat is greatly increased. It is of interest to examine
the list of breeding bird species characteristic of each of six types of
non-forest habitat in the Eastern Highlands. It should be remembered
(see pp. 79-80) that migrants from outside New Guinea make a con-
spicuous but nonbreeding contribution to the non-forest avifauna,
especially to category 6 (open country and forest edge).
1. Alpine grassland.—Alpine grassland occurs naturally above about
11,500 ft and is separated by a forest belt from the man-made mid-
montane grassland, whose upper limit varies locally up to about
8,000 ft. Only four species occur in the alpine grassland, all repre-
sented by endemic subspecies: the pipit Anthus gutturalis, which is
strictly confined to the grassland; the thrush Turdus poliocephalus
and the grass finch Oreostruthus fuliginosus, which live in the grass-
71
FEATURES OF THE EASTERN HIGHLANDS
land and adjacent forest edge (Oreostruthus barely gets into the grass-
land and is mainly in the forest edge); and the grass warbler Megalurus
tumorensis, which has isolated and racially distinct populations in low-
land, midmontane, and alpine grassland.
2. Water.—Bodies of water consist of a few lakes (Kutubu, Tebera,
Kandep, and the small alpine lakes on Mt. Wilhelm), a few small
rivers, and numerous brooks and small streams. The breeding water
birds are the grebe Podiceps novachollandiae, the ducks Anas super-
ciliosa and Salvadorina waigiuensis, and the kingfisher Ceyx azureus.
All occur in the much more extensive waterways of the lowlands
except for Salvadorina. There are two strictly riparian passerines, both
absent from the lowlands: the flycatcher Monachella muelleriana and
the ‘Yorrent Lark, Grallina bruijni. The duck Nettapus coromande-
lianus has been reported from Lake Kandep, the duck Anas gibberi-
frons from the Wahgi River and Mt. Giluwe, and several cormorants
and herons at Lake Kutubu, but I do not know if these species breed.
The cormorant Phalacrocorax melanoleucos has been found breeding
on the Baiyer River. Salvadorina waigiuensis is apparently the sole
aquatic bird in the alpine lakes of Mt. Wilhelm.
3. Swamps.—Small swamps border watercourses and harbor up to
five rails: Rallus philippensis, Porzana tabuensis, Porzana pusilla (one
locality), Porphyrio porphyrio, and Gallinula tenebrosa (one record).
All five species have wide ranges outside New Guinea. P. pusilla is
confined to middle elevations in New Guinea, and Rallus phillippensis
and Porzana tabuensis are represented in the Eastern Highlands by
thinly differentiated endemic subspecies.
4. Air.—In the air, remote from forest, are 14 species, of which six
swifts and swallows take aerial food, while eight hawks merely hunt
from the air. Six of these 14 species—the hawk Henicopernis longi-
cauda, the swifts Collocalia esculenta, C. hirundinacea, and C. white-
head, the crested swift Hemiprocne mystacea, and the wood swallow
Artamus maximus—also occur over hill or montane forest and streams
in areas undisturbed by man; Henicopernis also hunts inside the forest.
I suspect that seven of the other species have begun to appear in the
Eastern Highlands from the lowlands only since man began extensive
clearing of the forest, viz., the hawks Milvus migrans, Haliastur indus,
Accipiter fasciatus, Circus spilonotus, Circus approximans (only Kari-
mui), and Falco berigora and the swallow Hirundo tahitica. The hawk
Elanus caeruleus, whose range includes much of Africa and Eurasia, is
confined in New Guinea to extensively deforested areas at midmontane
elevations. The New Guinea population is subspecifically distinct (E. c.
wahgiensis) but has never been found in areas of New Guinea un-
disturbed by man, so that its origins are obscure.
5. Midmontane grasslands—Midmontane grasslands in their pres-
ent extent are a product of native agriculture. If one defines grassland
species as those that live in the grass and do not require other elements
12,
FEATURES OF THE EASTERN HIGHLANDS
(trees, fences, bushes) for perches, then 16 grassland species occur 1M
the Eastern Highlands. Most of these 16 species are confined in the
Eastern Highlands to areas with extensive destruction of forest by man.
Seven of these also occur in the lowlands and do not have endemic
midmontane races: the cuckoo Centropus phasianinus, the nightjar
Caprimulgus macrurus, the lark Mirafra javanica, the grass warblers
Acrocephalus arundinaceus and Cisticola exilis, the bowerbird Chlamy-
dera lauterbachi, and the mannikin Lonchura grandis (only at Okasa).
While Chlamydera lauterbachi is widespread, the other six species are
each known only from a few localities or areas in the Eastern High-
lands, making one suspect that they have just begun to colonize the
midmontane grasslands. Six grassland species also occur in the low-
lands, but the midmontane population is subspecifically distinct: the
quail Synoicus ypsilophorus and Coturnix chinensis, the bustard-quail
Turnix maculosa, the wren-warbler Malurus alboscapulatus (two races),
the grass warbler Megalurus timoriensis, and the mannikin Lonchura
spectabilis (two races). The races Synoicus ypsilophorus lamonti,
Turnix maculosa gilwwensis, Megalurus timoriensis wahgiensis, Mal-
urus alboscapulatus kutubu, and Lonchura spectabilis gajduseki are
endemic to the Eastern Highlands, while Coturnix chinensis novae-
guineae is shared with the midmontane grasslands of western New
Guinea, Malurus alboscapulatus mafulu is shared with the midmon-
tane grasslands of southeastern New Guinea, and Lonchura spectabilis
wahgiensis is shared with the midmontane grasslands of the Huon
Peninsula. The remaining three species—the rail Rallus pectoralis,
the owl Tyto capensis, and the pipit Anthus novaeseelandiae—have
wide geographic ranges outside New Guinea, but the New Guinea
populations are subspecifically distinct and confined to the midmontane
grasslands (absent from the lowlands, except for one localized popula-
tion of the pipit).
6. Open country and forest edge ‘Open country and forest edge”’
may be used as the name for a category that excludes swamps and
pure grassland but includes native gardens, grassland with occasional
trees or fences to provide perches, open vegetation around villages and
towns, planted groves of trees remote from forest, regenerating vegeta-
tion in a cleared and fallow area, and the forest edge. Many forest
species may, of course, also be found at the forest edge, but if discussion
is restricted to birds regularly found remote from forest and to birds
which are much commoner at the forest edge than in the forest interior,
then one arrives at a somewhat arbitrary list of 33 species. About 17 of
these species are encountered above 5,000 ft, 16 only below 5,000 ft,
and 24 at sea level. ‘This list is probably not exhaustive, since I spent
little time in towns remote from forest.
Two of these 33 species (the cuckoo Cacomantis pyrrhophanus and
the warbler Gerygone ruficollis) occur only above 5,000 ft. Both may
be found in the interior of undisturbed forest but have become much
73
FEATURES OF THE EASTERN HIGHLANDS
commoner in trees of towns and gardens, perhaps because they have
so few competitors in these habitats.
Five species are absent from the lowlands and live mainly between
about 3,000 and 7,000 ft: the flycatcher Peltops montanus, the shrike
Lanius schach, and the honeyeaters Myzomela adolphinae, Melidectes
torquatus, and Meliphaga auga. Four of these five species (Peltops
montanus and the three honeyeaters) are endemic to New Guinea and
also live in forest or in natural forest edge habitats, so that their origin
poses no problems. Lanius schach is confined to man-made habitats
(gardens and grassland with perches) and is represented in New Guinea
by an endemic race of a species with a widespread extralimital range.
‘Ten species (the pigeon Macropygia amboinensis, the parrot Tricho-
glossus haematodus, the cuckoo Cacomantis variolosus, the Barn Owl,
Tyto alba, the chat Saxicola caprata, the flycatchers Rhipidura rufi-
ventris and Rhipidura leucophrys, the whistlers Pachycephala ruft-
ventris and Colluricincla harmonica, and the bird of paradise super-
species Paradisaea minor-P. raggiana) occur in the lowlands and are
found only below 5,000 ft in undisturbed areas but have ceilings,
depending on the species, between 5,500 and 7,000 ft in settled areas.
Four of these species (the pigeon, the parrot, the bird of paradise, and
Rhipidura rufiventris) are regularly found in forest up to about 4,500
ft, so that only a small upwards step was required to colonize the
settled areas above 5,000 ft. ‘The cuckoo is mainly, the other five
species are strictly, forest edge and open country birds, and all six are
absent at elevations above a few thousand feet in undisturbed areas.
The remaining 16 species range from the lowlands up to ceilings
varying between about 2,000 and 4,000 ft: the rail Amaurornis ollt-
vaceus, the kingfisher Dacelo guadichaud, the roller Ewrystomus
orientalis waigiouensis, the cuckoo-shrikes Coracina papuensis and C.
tenuirostris, the wren-warbler Todopsis cyanocephala, the warbler
Gerygone magnirostris, the flycatchers Peltops blainvilli, Rhipidura
leucothorax, and Myiagra alecto, the starling Mino dumontii, the
oriole Oriolus szalayi, the butcherbird Cracticus cassicus, the drongo
Dicrurus hottentottus, the sunbird Nectarinia sericea, and the honey-
eater Philemon novaeguineaec. Three if these species (Dacelo gaudi-
chaud, Oriolus szalayi, Philemon novaeguineae) also live in the forest
interior, while Amaurornis olivaceus lives in garden edges and grass
and the remaining species live at the forest edge or in partly cut forest.
For most of these species the ceilings in the disturbed areas of the
Eastern Highlands (2,000-4,000 ft) are somewhat above the ceilings in
undisturbed areas.
Origin of the Non-Forest Avifauna of the Eastern Highlands
There appear to have been five sources of the nonforest bird species
in the Eastern Highlands.
1. Evolution in pre-existing specialized habitats—Three types of
74
FEATURES OF THE EASTERN HIGHLANDS
specialized habitats, inhabited by specialized birds already listed, must
always have existed in the Eastern Highlands: alpine grassland, bodies
of water, and swamps. ‘The total areas occupied by these specialized
habitats are very small now, have been little affected by man, and
probably have not been significantly larger in recent geological history,
except for expansion of alpine grassland during cooler parts of the
Pleistocene. The endemic montane species and subspecies now living
in these habitats have had an indefinite amount of time to evolve in
them. The very small number of such endemic montane forms (four
in alpine grassland, three on waterways, two in swamps) 1s what one
would expect from the small area of such habitats in the New Guinea
mountains. Some of the water and swamp species also live in the low-
lands and are not represented by endemic montane races. ‘They prob-
ably evolved mainly in the lowlands, where the area occupied by
waterways and swamps is larger.
2. Evolution in pre-existing forest edge habitats —Even in montane
areas undisturbed by man, natural forest edge habitats adjacent to
open spaces exist, notably in permanent form along streams, but also
transiently on steep slopes with landslides, as a result of trees falling
in the forest, and as a result of earthquakes. ‘The total area of forest
edge habitat in undisturbed montane areas is very small and increases
vastly in regions disturbed by man. ‘The population densities of forest
edge species in settled regions are now high in many cases, but before
native agriculture these species must have evolved and eked out “a
precarious existence in the shifting bits of habitat suitable for them”
(Rand and Brass, 1940, p. 377). As illustrations I shall cite several
instances in which I obtained records in montane areas remote from
human disturbance for birds which are completely absent from the
interior of closed forest. At the Sena River the Crested Swift, Hemz-
procne mystacea, perched in riverside trees and sallied into the open
space over the stream. In the river gorges below Karimui, Bomai, and
Soliabeda and on the Sena River I met the swift Collocalia esculenta
hawking over the streams, and JI have seen both C. esculenta and C.
hirundinacea over ravines and the sides of forested ridges on Mts.
Karimui, Michael, Menawa, and Turu. On Mt. Menawa I saw the
wood swallow Arfamus maximus perched in trees on the crest of a
steep forested ridge. The butcherbird Cracticus cassicus was present
in the river gorges below Soliabeda and Bomai. ‘The flycatcher Peltops
montanus sallied over the Sena River from trees on the banks, and I
have regularly found it in trees on the crests of steep mountain ridges
(Mts. Karimui, Somoro, Nibo, Menawa). Once I saw the sunbird
Nectarinia sericea in the tree canopy on the steep slopes of the Nami
Gorge in the Karimui Basin. I found a nest of the honeyeater Meli-
phaga auga in a bamboo thicket near a small stream below Soliabeda.
3. Evolution in forest—Five of the seven montane species that
occur in open country and forest edge habitats and do not descend to
75
FEATURES OF THE EASTERN HIGHLANDS
sea level also occur in the interior of montane forest, where they
undoubtedly evolved. These are Cacomantis pyrrhophanus, Gerygone
ruficollis, Pitohui dichrous, Myzomela adolphinae, and Melidectes
torquatus. Gerygone ruficollis and Melidectes torquatus are very un-
common in the forest interior but are the two most abundant species
in trees in open country around 5,000 ft, and their numbers must have
undergone an enormous explosion as a result of native agriculture.
4. Recent expansion from the lowlands.—Because there are many
more rivers, marshes, and swamps in the lowlands than in the moun-
tains, more species of forest edge and grassland birds have evolved in
the lowlands than in the mountains. With the recent spread of open
country in the mountains, some lowlands species may have begun to
colonize the midmontane open country only recently. Though it is,
of course, impossible to be certain about the time of arrival, a recent
arrival must be suspected for species whose midmontane populations
are not subspecifically distinct and which are absent from the moun-
tains in areas undisturbed by man, particularly if their distribution in
midmontane open country is very local. In this category probably fall
the swallow and six hawks listed on p. 72, most of the seven grassland
birds listed on p. 73, and most of the 26 open country and forest edge
birds listed on pp. 73-74 which are shared with the lowlands and are
not subspecifically distinct.
5. Rapid recent evolution?—The evolution of non-forest lowland
species poses no problems because there is a modest amount of non-
forest habitat in the lowlands and these species can be seen in these
natural habitats. The evolution of the few montane forest edge species
also poses no problems because, although the amount of such natural
habitat is small, one can find these species in natural forest edge
habitats in undisturbed areas, as illustrated on p. 75. The main
problem arises in connection with the evolution of the endemic mid-
montane grassland populations, because the amount of natural mid-
montane grassland (along streams, in landslide and earthquake areas)
is minute, and because most of the endemic forms have never been
recorded in these natural habitats in which they might be postulated
to have evolved. Specifically, for the following 10 species with endemic
midmontane races every midmontane record that has come to my
attention is from a man-made habitat: Synoicus ypsilophorus, Coturnix
chinensis, Turnix maculosa, Tyto capensis, Anthus novaeseelandiae,
Saxicola caprata, Malurus alboscapulatus, Megalurus timoriensis,
Lanius schach, and Lonchura spectabilis. Most of the species to which
these endemic races belong range through the Indonesian Archipelago
or as far as Asia and are presumably of non-Papuan origin, the ex-
ceptions being Malurus alboscapulatus (confined to New Guinea) and
Lonchura spectabilis (New Guinea and New Britain). ‘The only New
Guinea midmontane grassland forms that are endemic to this habitat
76
FEATURES OF THE EASTERN HIGHLANDS
at the species level are Lonchura teerinki of western New Guinea and
Lonchura vana of the Vogelkop. ;
‘There are two possible explanations for these cases. First, some of
these species may have been present for a long time in the tiny amount
of natural midmontane grassland habitat, may have evolved there into
endemic races, and may simply not have been detected yet in such
habitats. In other words, they would have the same history as mid-
montane forest edge species, which had to eke out a precarious exis-
tence before the advent of agriculture. Second, they may have evolved
in the short time since native agriculture began to create significant
areas of secondary midmontane grassland. Present evidence suggests
that agriculture on a limited scale may have become possible in New
Guinea about five millenia ago (Bulmer, 1964). However, extensive
clearing of land began with the introduction of the sweet potato,
which appears to have reached the Eastern Highlands only a few
centuries ago. Thus, the time scale for evolution of the midmontane
grassland races would be of the order of several centuries up to a few
millenia. Until recently such a short time scale for subspeciation
would have appeared unlikely, but the demonstration by Johnston and
Selander (1964) of geographical differentiation in the House Sparrow
Passer domesticus since its introduction into the New World 137 years
ago shows that this is perfectly feasible in some cases. It is important
to remember that population changes will occur much more rapidly
when breeding stock is transplanted into a new type of environment
and isolated from the parent population than in the cases of estab-
lished populations; and, also, that severe selection pressures may
operate on new colonists of the New Guinea midmontane grasslands,
whether they arrive from the New Guinea lowlands or from outside
New Guinea.
Probably some of the 10 races listed on p. 76 have the first type of
history, some the second. Midmontane grassland forms that surely have
a long history include the sole two endemic species, the distinct man-
nikin species Lonchura teerinki of western New Guinea and the semi-
species Lonchura vana of the Vogelkop, which probably evolved in
the marshes of the Baliem River and the Anggi Lakes, respectively.
I would not be surprised to see the midmontane race of Malurus
alboscapulatus eventually turn up in a marsh beside some Eastern
Highlands stream remote from areas of human influence. At the
other extreme, the mannikin Lonchura spectabilis gajduseki requires
large expanses of grassland and is confined to the Karimui Basin,
whose topography and history of human settlement are sufficiently
well known to make it unlikely that this mannikin population could
have existed anywhere in the Basin before the present century.
Lonchura spectabilis gajduseki is, accordingly, a likely candidate to
have differentiated rapidly and recently.
FEATURES OF THE EASTERN HIGHLANDS
Pioneering by Non-Forest Species
When natives clear a new village and garden area in the middle of
the forest, an island of habitat is created for grassland and second-
growth birds, surrounded by a barrier of unsuitable forest habitat. The
colonization of these grassland islands poses the same problems as those
involved in the zoogeography of oceanic islands. One of my collecting
stations, Soliabeda, was just such a non-forest island that had been
cleared by natives a few years previously and lay several miles from the
next such island. Consideration of the non-forest avifauna at Soliabeda
is interesting because of its relevance to the question of what makes
a good colonist.
Although Soliabeda and Karimui shared most forest species in com-
mon, Soliabeda had only part of the non-forest avifauna of Karimui.
All four of the small aerial species of Karimui (Hemiprocne mystacea,
Collocalia esculenta, C. hirundinacea, Artamus maximus) were present
at Soliabeda. ‘hese species are strong fliers, and the first two would
have been present along forest streams even before the village site was
cleared. Of the four large aerial species of Karimui (the open country
hawks Haliastur indus, Accipiter fasciatus, Circus approximans, Falco
berigora), only the first was present, perhaps because the cleared area
at Soliabeda was marginally small to provide even a single feeding
territory. Of the 16 species in the category “open country and forest
edge” at Karimui which also utilized the forest edge or forest interior
(Macropygia amboinensis, Trichoglossus haematodus, Gacomantis vari-
olosus, Dacelo gaudichaud, Eurystomus orientalis waigiouensis, Rhipi-
dura leucothorax, Rhipidura rufiventris, Pachycephala rufwentris,
Pitohui dichrous, Oriolus szalayi, Mino dumonti, Cracticus cassicus,
Dicrurus hottentottus, Paradisaea raggiana, Nectarinia sericea, Meli-
phaga auga), all except two (Eurystomus orientalis waigiouensis,
Pachycephala rufiventris) were present at Soliabeda. Of the three
species of “open country and forest edge” at Karimui which could not
utilize the forest interior or forest edge (Amaurornis olivaceus, Saxi-
cola caprata, Rhipidura leucophrys), only the rail Amaurornis oliva-
ceus was present at Soliabeda. Of the four grassland species at Karimui
(Synoicus ypstlophorus, Malurus alboscapulatus, Megalurus timoriensis,
Lonchura spectabilis), only the wren-warbler Malurus alboscapulatus
was at Soliabeda.
Generalizing, one can say that open country species that can also
use the forest and forest edge make the best colonists, partly because
they can disperse most easily (through the forest or by using forest edge
habitats along rivers as stepping stones), partly because more habitat
is available to them for building up a stable population than to a bird
restricted to grassland and open country. At a disadvantage are species
with a large territory requirement, a category including not only
hawks but perhaps also the Lonchura mannikins, which normally as-
78
FEATURES OF THE EASTERN HIGHLANDS
sociate in large flocks. Of the species restricted to grassland and open
country, the rail and the wren-warbler are apparently good colonists.
The superior dispersal ability of the rail is in keeping with the ability
of rails to reach the most remote oceanic islands.
MIGRANTS
Eleven species that breed in the palearctic region have been recorded
from the Eastern Highlands as winter visitors, with most of the records
between September and April. Eight of these 11 species are shorebirds
(Charadriidae, Scolopacidae): Pluvialis dominica fulva, Numenius
minutus, Tringa hypoleucos, Tringa brevipes, Gallinago megala, Gal-
linago hardwicku, Calidris ruficollis, and Calidris acuminata. The
other three are the duck Anas querquedula, the cuckoo Cuculus
saturatus horsfieldi, and the wagtail Motacilla cinerea caspica.
Thirteen species that breed in Australia have been recorded from
the Eastern Highlands as winter visitors, with most of the records
between April and October: the spoonbill Platalea regia, the falcon
Falco cenchroides cenchroides, the pratincole Glareola isabella, the
cuckoos Cacomantis pyrrhophanus prionurus and Chrysococcyx lucidus
plagosus, the nightjar Ewrostopodus albogularis albogularis, the king-
fishers Halcyon macleayu incincta and Halcyon sancta sancta, the bee-
eater Merops ornatus, the dollarbird Eurystomus orientalis pacificus,
the swallow Hirundo nigricans nigricans, the cuckoo-shrike Coracina
novaehollandiae melanops, and the flycatcher Myiagra cyanoleuca.
The most striking fact about these 24 palearctic and Australian
migrants in New Guinea is that their distribution is strictly confined
to open habitats, second-growth, and the forest edge. I am unaware
of a single record of a palearctic or Australian migrant taken inside
undisturbed forest anywhere in the Eastern Highlands. ‘There is no
record above 9,000 ft, where there is little cleared land except natural
alpine grassland, which the migrants evidently do not utilize. ‘The
implication is that migrants have been coming to the Eastern High-
lands in significant numbers only since the advent of extensive native
agriculture, i.e., probably within the last millennium (p. 77). The
palearctic migrants wintering in Africa similarly avoid the interior
of the rainforest (Moreau, 1966). By contrast, wintering North Ameri-
can species comprise a significant percentage of bird individuals inside
the tropical rainforest of Central America.
Because of New Guinea’s equatorial position New Guinea breeding
species do not migrate from the island. However, there is evidence
for local seasonal migration within New Guinea in the cases of a few
species. Four categories may be distinguished. (1) Some species which
breed in the lowlands may undertake postbreeding migrations to the
Eastern Highlands. For instance, large postbreeding flocks of the
starling Aplonis cantoroides have been recorded at Nondugl in the
79
FEATURES OF THE EASTERN HIGHLANDS
Wahgi Valley. Records of the starling Aplonis metallica, of cormo-
rants, anhingas, and herons, and possibly of some hawks (Milvus
migrans, Circus approximans, Haliastur indus ?) may also represent
postbreeding stragglers from the lowlands. As in the case of palearctic
and Australian migrants, these lowland migrants are found mainly in
cleared and second-growth areas in the Eastern Highlands. (2) Some
species, par ticularly lories and myzomelid honeyeaters, which breed
in the Eastern Highlands and which depend for food on transient
sources of flowers and fruit, may undertake local seasonal migrations
in search of food. For instance, Fore native informants told me that
the parrots Pseudeos fuscata, Psitteuteles goldiei, and Geoffroyus
simplex breed and are conspicuous at Okapa in the wet season, but I
found them uncommon or absent in the dry season. The erratic dis-
tribution of the honeyeater Myzomela adolphinae, which depends
upon flowering trees for food, also suggests local migration. (3) The
open country shrike Lanius schach breeds at Okapa in the dry season,
but native informants said that it disappears during the wet season.
Similarly, at Lake Kutubu, Schodde and Hitchcock were told by native
informants that the open country honeyeater Philemon novaeguineae
is absent during the wet season. ‘The cuckoo Eudynamis scolopacea
was apparently absent from Karimui during the driest months of
1965. (4) Some stragglers may be driven into the Eastern Highlands
by storms, an example being the appearance of Sooty ‘Yerns, Sterna
fuscata, at Mt. Hagen township in June 1967.
Little or nothing is known about the details of these local migra-
tions. Further examples of palearctic, Australian, and New Guinea
lowland species which migrate or straggle to the Eastern Highlands
undoubtedly will be uncovered and probably will provide most of
the future additions to the list of species recorded from the Eastern
Highlands.
Birp FEEDING ASSEMBLAGES IN FRUITING AND FLOWERING TREES
The gathering of birds of many species to feed on trees bearing
fruit or flowers are a spectacular feature of the tropics. Several species
of New Guinea parrots, pigeons, and honeyeaters are mainly to be
found at these ephemeral food sources. In 1964 ‘Terborgh and I made
a study of these feeding assemblages which has been reported in detail
elsewhere (Terborgh and Diamond, 1970), but will be summarized
briefly here along with some subsequent observations.
Our method was to station ourselves at a fruiting or flowering tree
and, at two-minute intervals, to record the number of individuals of
each species in the tree. One individual recorded in one two-minute
period was taken as an arbitrary unit of “bird-usage” of the tree.
In the individual species accounts it occasionally will be stated that a
given species accounted for a certain percentage of the bird-usage
80
FEATURES OF THE EASTERN HIGHLANDS
of a particular tree; this refers to the bird-usage units of that species
as a percentage of the total bird-usage units of all species in that tree.
Usage of flowering trees was dominated by honeyeaters (notably of
the genera Oedistoma, Myzomela, and Meliphaga) and by lorikeets
(Chalcopsitta, Pseudeos, Trichoglossus, Domicella, Charmosyna, Oreo-
psittacus, and Neopsittacus). ‘Ten to 16 species were regularly seen in
one tree. Most of these species presumably were obtaining nectar and
insects from the flowers, but the parrot Trichoglossus haematodus ate
the flower receptacles as well. At a given locality essentially the same
eroup of bird species reappeared in different species of flowering trees.
Within a given tree there was some vertical segregation of the honey-
eaters, with Oedistoma pygmaeum and the several species of Myzomela
mainly in the canopy and the species of Meliphaga mainly in the lower
branches. While the honeyeaters moved in and out as individuals, the
lorikeets often arrived and departed as flocks. Several of the lories
have crimson upperparts, and of those which have green upperparts,
several have a crimson or blue rump. ‘This striking dorsal coloration
must make them conspicuous when viewed from above against the
green jungle, and may serve to attract the attention of other members
of the species to a flock flying towards a tree which it has located.
Trees bearing small fruits (up to 5 mm in diameter) were utilized
by representatives of more bird families (Columbidae, Campephagidae,
Pachycephalinae, Sturnidae, Paradisaeidae, Meliphagidae, Dicaeidae,
a few Psittacidae) than were flowering trees. Nine to 16 species were
regularly seen in one tree. Trees with larger fruits (greater than 10 mm
in diameter) were virtually monopolized by pigeons, particularly those
of the genus Ptilinopus. Ficus trees with large fruits several inches in
diameter surrounded by a woody rind were monopolized initially by
parrots (Trichoglossus haematodus, Psittaculirostris desmarestit), which
chipped holes in the rind with their strong bills. Later, members of
other families, particularly Campephagidae, Paradisaeidae, and long-
billed species of Meliphagidae, inserted their weak bills into the hole
to obtain pulp which the shorter-billed parrots had been unable to
reach.
These transient assemblages of birds that gather in fruiting or
flowering trees should not be confused with the mixed-species, itinerant,
mainly insectivorous flocks which forage through tropical forests in
many parts of the world. It is worth mentioning explicitly that I never
encountered such itinerant flocks in the Eastern Highlands. Further-
more, when I brought four of my Eastern Highlands assistants with
me to the North Coastal Range in 1966 and we met itinerant flocks
for the first time, they related to me their amazement and were
emphatic that their own birds at home didn’t behave that way! In
contrast, Gilliard (in Gilliard and LeCroy, 1968) observed mixed
flocks in the Schrader Range of the Eastern Highlands; Schodde and
Hitchcock (1968, p. 40) mentioned their presence at Lake Kutubu;
81
FEATURES OF THE EASTERN HIGHLANDS
Rand (Mayr and Rand, 1937, pp» 114, 192, 159; 161; Anehibold, Rand
and Brass, 1942, p. 246) encountered them in western New Guinea
and southeastern New Guinea; and I found them on Mt. Albert-
Edward, and at certain localities and altitudes but not at others in the
North Coastal Range. I have no explanation for this apparent un-
predictability.
RAINFALL
Rainfall patterns in the Eastern Highlands are relevant to the
problem of bird breeding seasons discussed in the next section.
Tables 2 and 3 give rainfall records for Okapa Patrol Post and for
Karimui Patrol Post, respectively, kindly supplied by the patrol officers
at these stations. Rainfall records for some other stations, including
Lufa and Goroka, have been published by Brass (1964), and records
for the Kubor Range and Lake Kutubu have been summarized by
Hitchcock (1964) and by Schodde and Hitchcock (1968).
For much of the Eastern Highlands the annual rainfall averages
between 70 and 180 inches, and there is a weakly differentiated dry
season between May and September and rainy season between Novem-
ber and April. However, the distinctness of wet and dry seasons varies
from year to year and varies among neighboring localities in the same
year.
At Okapa the annual rainfall averages 91 inches and varies in dif-
ferent years between 65 and 136 inches. ‘he wettest month, March, has
on the average 3.4 times as much rain as the driest month, June. In at
least five of the eight years up to 1965 for which there are records,
May, June, July, August, and September were drier than the average
month of that year, and December, February, March and April were
wetter than the average month. ‘The June-July dry season was quite
pronounced in 1956, 1964, and 1965. However, normally dry September
was the wettest month in 1963, and May was the third wettest month
in 1962, while normally wet February was the third driest month in
1963 (drier than June-October) and December was the third driest
month in 1964.
At Karimui the pattern is qualitatively similar but wetter overall
and less distinct. Annual rainfall averages 131 inches and varies from
113 to 158 inches, with January, February, March, April, and Septem-
ber on the average the wettest months and June, July, August, and
November the driest months. ‘The June-August dry season was fairly
pronounced in 1964, and more pronounced in 1965. However, the
wettest month, March, has on the average only 2.3 times as much
rain as the driest month, June. In 1961 normally dry July was the
wettest month. These statements are based on rainfall records taken
at Karimui Patrol Post. The situation on Mt. Karimui, for which no
records are available, is patently different. The mountain is usually
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FEATURES OF THE EASTERN HIGHLANDS
shrouded in cloud even when it is fair at Karimui Patrol Post. I was
told after descending from Mt. Karimui in August 1965 that the
month I spent on the mountain had been the driest period on record
since a rain gauge was set up at Karimui Patrol Post. On Mt. Karimui
this irreducible minimum of precipitation meant that it rained day
and night without interruption for half of the days on the mountain,
and only seven of the days we spent on it were without any rain.
At Lake Kutubu (Schodde and Hitchcock, 1968) May to October
is the wettest season, rather than the driest season as in the Wahgi
Valley and my study areas.
‘To summarize, during the particular months and years of my studies
(June-September 1964 and 1965) the particular localities I visited were
experiencing a relatively dry season. This would have been true in
these months at these localities in most but not all other years, and at
many other but not all localities in the same years.
BREEDING SEASONS IN THE EASTERN HIGHLANDS
Breeding patterns in the Eastern Highlands are complex in that
they differ among birds of different habitats, vertical strata, altitudes,
or food habits. Hence, analysis of breeding activity in separate seg-
ments of the avifauna is more informative than analysis of the avi-
fauna as a whole. The following categories, which overlap to some
extent, will be considered: grassland birds; birds of second-growth and
the forest edge; hawks; birds that frequent fruiting trees; birds that
frequent flowering trees; meliphagids generally; forest birds of the
ground and lowerstory; forest warblers; other forest species. Since my
records are most detailed for the Karimui area (Karimui, Bomai,
Soliabeda, Mt. Karimui), all remarks refer to this area unless it is
specifically stated otherwise. Breeding patterns on Mt. Karimui dif-
fered from those at lower elevations and will be cited separately, but
those from the three stations at lower elevations (Karimui, Bomai,
Soliabeda) were similar to each other and will be referred to without
distinction simply as the pattern at Karimui. Most information is
based on examination of gonad size, but some is derived from actual
nests or nestlings. It will be recalled that this information comes from
the relatively dry months of June, July, and August of 1964 and 1965.
Although several patterns emerge from my results, they must be con-
sidered preliminary for three reasons: (a) they are based on extrapola-
tion from a fraction of the year; (b) a condition of enlarged testis size
may be maintained for longer than the actual nesting seasons; and
(c) I have no information on peak seasons of fruit, flower, seed, and
insect production.
Grassland birds——Adequate information to judge breeding condi-
tion was obtained at Karimui for three species that live in grassland:
the rail Amaurornis olivaceus, the wren-warbler Malurus alboscapu-
84
FEATURES OF THE EASTERN HIGHLANDS
latus, and the mannikin Lonchura spectabilis. All adults collected of
all three species had gonads in breeding condition, and nestlings of
all three species were obtained. ‘Thus, there was a highly synchronous
breeding peak in the dry season. In addition, the cuckoo Cacomantts
variolosus, which lives in garden trees rather than in the grass itself
but uses Malurus abolscapulatus as its principal foster parent, was
breeding. Information was obtained for more grassland species at
Okapa, and all were breeding: Malurus alboscapulatus and Lonchura
spectabilis, plus the quail Synoicus ypsilophorus, the rail Porzana
tabuensis, the grass-warbler Megalurus timoriensis, and the shrike
Lanius schach. Lanius schach actually appears to leave the Okapa re-
gion in wet seasons.
Birds of second-growth, the forest edge, and open spaces (excepting
hawks).—All adults taken at Karimui of 11 species in this category
were found not to be in breeding condition: the crested swift Hemi-
procne mystacea, the swifts Collocalia hirundinacea and C. esculenta,
the kingfisher Dacelo gaudichaud, the dollarbird Eurystomus orien-
talis, the flycatchers Peltops blainvillii and P. montanus, the oriole
Oriolus szalayi, the drongo Dicrurus hottentottus, the starling Mino
dumontii, and the sunbird Nectarinia sericea. Some but not all adults
of the flycatchers Rhipidura leucothorax and Pachycephala rufiventris
and the butcherbird Cracticus cassicus were breeding. ‘here was some
breeding in the wood swallow Artamus maximus, and most individuals
of the honeyeater Meliphaga auga were breeding, in 1964, but there
was no breeding in either species in 1965, which was drier. Evidently
breeding of most forest edge species ceases completely when it becomes
sufficiently dry. The situation is thus exactly opposite to the case of
grassland birds.
Hawks.—No individual in breeding condition was encountered
among the six species of hawks whose gonads were examined (Accipi-
ter novaehollandiae and Falco berigora, in open country; Aviceda sub-
cristata, Accipiter poliocephalus, and Megatriorchis doriae, in lowland
forest; and Accipiter melanochlamys, in montane forest).
Frugivores (except meliphagids)—Under this category are lumped
42 forest species that feed to a large extent on fruit: all species of
arboreal pigeons about which evidence was obtained, totaling 12; the
parrot Psittaculirostris desmarestit; the hornbill Aceros plicatus; all
species of campephagids, totaling seven; the whistler Pachycephala
leucostigma; all species of birds of paradise and bowerbirds, 14 in all;
and all dicaeids, six in all. Of the 29 frugivores at Karimui, there were
19 species in which all individuals were uniformly in nonbreeding con-
dition; five species in which some individuals were breeding in 1964
but none in 1965 (the pigeon Ducula zoeae, the parrot Psittaculirostris
desmarestii, the birds of paradise Manucodia chalybatus and Lopho-
rina (Diphyllodes) magnifica, and the bowerbird Ailuroedus buc-
coides); two species in which some but not all individuals were breed-
85
FEATURES OF THE EASTERN HIGHLANDS
ing in 1965 (the bird of paradise Lophorina (Ptiloris) magnifica and
the flowerpecker Dicaeum geelvinkianum); one species which was not
breeding in July and August 1965 but entered breeding condition in
September 1965 (the pigeon Megaloprepia magnifica); and two species
in which virtually all individuals were breeding (the pigeon Ptilinopus
superbus and the bird of paradise Lophorina (Cicinnurus) regia). The
situation was quite different on Mt. Karimui: out of 15 frugivores,
only five species showed no signs of breeding, while in ten species
some (but generally not all) individuals were breeding. In the case of
two species, the pigeon Macropygia nigrirostris and the bird of para-
dise Lophorina (Diphyllodes) magnifica, present both on Mt. Karimui
and in the Karimui Basin, the montane population was breeding, but
the lowland population was not, although numerous adults were
present.
‘The simplest interpretation to cover most of these facts is that breed-
ing is almost entirely confined to times of local wet conditions in most
fruit-eating birds. ‘The systematic difference between Mt. Karimui and
the lower elevations is to be expected from the fact that conditions
were considerably drier at the lower elevations during my studies. ‘The
fewer breeding records from 1965 than 1964 reflect the differences in
rainfall.
Birds that depend wpon flowering trees—Thirteen species feed
mainly or almost exclusively in flowering trees: seven parrots of the
subfamily Loriinae, and six honeyeaters (Oedistoma iliolophum, O.
pygmaeum, and four species of Myzomela). Of the nine species at
Karimui, five were not breeding, one (Trichoglossus haematodus) was
breeding in 1964 but not 1965, and some but not all individuals were
breeding in three cases (Charmosyna placentis, Myzomela cruentata,
Oedistoma iliolophum). Of the four species on Mt. Karimui, two
(Oreopsittacus arfaki and Myzomela rosenbergit) were breeding, and
two (Charmosyna papou and C. pulchella) were not. At Okapa natives
and Europeans said that the several species of lories there had young
in the wet months from November on.
Honeyeaters (Meliphagidae).—There are 18 additional species of
meliphagids which are not so strikingly tied to flowering trees, though
most visit flowering and/or fruiting trees. Of the 12 species at Karimui,
eight were uniformly not breeding, two were represented by some
breeders in 1964 but none in 1965, and two had breeding individuals
in both 1964 and 1965 but fewer in 1965. ‘Thus, breeding is sup-
pressed partly or completely in all these species in a dry spell. In the
wetter conditions on Mt. Karimui only one species (Meliphaga orien-
talis) was uniformly nonbreeding, four species had some or most in-
dividuals breeding, and in two species (Melidectes rufocrissalis and
Ptiloprora guisei) all males had greatly enlarged testes.
Forest birds of the ground and lowerstory.—This category consists
of nine forest species that feed on the ground (four pigeons of the
86
FEATURES OF THE EASTERN HIGHLANDS
genera Chalcophaps, Henicophaps, and Gallicolumba, the pitta Pitta
erythrogaster, the thrushes Amalocichla incerta and Drymodes super-
ciliaris, and two logrunners of the genus Eupetes), plus 18 forest war-
blers, flycatchers, whistlers, and honeyeaters which forage largely or
exclusively in the lower story. At Karimui many, but not all, individ-
uals of eight species were breeding, virtually all individuals of one (the
flycatcher Poecilodryas placens) were breeding, three species had some
breeding individuals in 1964 but not in 1965, and three were not
breeding. On Mt. Karimui there were breeding individuals of six
species but none of eight species. In two of these six species (the war-
bler Sericornis nowhuysi and the flycatcher Peneothello cyanus) all or
most individuals were breeding. ‘wo species present both on Mt.
Karimui and at lower elevations (the warbler Crateroscelis murina
and the whistler Collwricincla megarhyncha) had breeding individuals
at low elevations but not on the mountain. ‘Thus, the pattern in forest
lowerstory birds is the opposite of the pattern among arboreal fruit
eaters, in that there is more breeding activity at Karimui than at
higher elevations. Evidently there is some tendency in lowerstory birds
to nest in the dry season, and more were therefore nesting at Karimui,
where the dry period was more pronounced.
Forest warblers.—Eleven species of forest warblers occur at Karimui
or on Mt. Karimui (Clytomyias insignis, five species of Sericornis, two
species of Crateroscelis, and three species of Gerygone). Most, but not
all, individuals of four species were in breeding condition, all indi-
viduals of five species were breeding, Clytomyias insignis was not
breeding, and Crateroscelis murina was breeding at Karimui but not
on Mt. Karimui. ‘T'wo of these 11 species (Crateroscelis) were confined
to the lowerstory, while most of the others are often but not exclu-
sively found in the lowerstory. ‘This might suggest that the breeding
of warblers is just another expression of the tendency of lowerstory
species to be breeding in June-August. However, the warbler Gery-
gone palpebrosa, which I have never seen in the lowerstory, was breed-
ing at Karimui; and six out of the eight warblers on Mt. Karimui
were breeding, although the majority of lowerstory species generally
on Mt. Karimui were not breeding. ‘Thus, lowerstory habits are not
necessarily the explanation for breeding patterns in forest warblers.
Other forest species——Among forest flycatchers (Muscicapinae and
Pachycephalinae) not confined to the lower story, breeding individuals
were encountered among seven of 14 species at Karimui and among
three of 10 species at Mt. Karimui. In only two cases (Microeca flavo-
virescens at Karimui, and Tregellasia leucops on Mt. Karimui) was
there evidence that the whole adult population had come uniformly
into breeding condition. Finally, one may consider a “wastebasket”
category consisting of all forest species not belonging to the categories
considered so far—three parrots, five cuckoos, four nocturnal species,
three kingfishers, one logrunner, one drongo, one crow, and one man-
87
FEATURES OF THE EASTERN HIGHLANDS
nikin. Only two of the 15 species at Karimui in this “wastebasket”
category were breeding, and only two of six on Mt. Karimui.
Summary of breeding patterns.—During the dry months of July-
August 1964 and 1965 at Karimui, breeding was taking place among
all grassland birds, most lowerstory forest birds, and most warblers,
but was not occurring among most second-growth birds, frugivores,
flower feeders, meliphagids, or most other middlestory and upperstory
forest species. On Mt. Karimui during these same months, where con-
ditions were wetter, breeding was taking place among most frugivores,
meliphagids, and warblers, but not among most other species. “Breed-
ing’ in these statements is taken to mean that a significant fraction
of individuals of a given species were in breeding condition.
In addition, it is worth considering in which species there appeared
to be a highly synchronized pattern, such that virtually all individuals
either were or were not breeding. Synchronous breeding peaks were
found among all species of grassland birds, two frugivores at Karimui,
three meliphagids on Mt. Karimui, one lowerstory species at Karimui
and two on Mt. Karimui, two warblers at Karimui and three on Mt.
Karimui, and one flycatcher at Karimui and one on Mt. Karimui.
Breeding was completely suppressed in all hawks, most second-growth
species, most frugivores, flower feeders, and meliphagids at Karimui,
and many other forest species of the middle- and upperstories.
‘There were three species (the pigeon Ptilinopus superbus, the war-
bler Sericornis papuensis, and the honeyeater Myzomela rosenbergit)
of which most or all specimens collected proved to be males, all in
breeding condition. All nine specimens collected of the pigeon Ptili-
nopus pulchellus proved to be males and all were not breeding. Only
males of the swiftlet Collocalia hirundinacea were collected in early
July, only females in late July. ‘The explanation for these unbalanced
sex ratios is unclear.
For comparison with the Karimui results, incidental information
about breeding patterns has been obtained in four other parts of New
Guinea. ‘The peak breeding months for most species are September-
November in the Wharton Range of southeastern New Guinea (Mayr
and Rand, 1937; Schodde, cited in Schodde and Hitchcock, 1968, p. 17;
my experience on Mt. Albert-Edward), November-March in northwest-
ern New Guinea (Rand, 1942b; Ripley (1964), however, reports May-
October), October-January in the western Papuan islands (Ripley,
1964), and September-October at Lake Kutubu (Schodde and Hitch-
cock, 1968). These peaks fall in the austral spring and summer. At these
localities, as at Karimui, the rainy season is November to April and
the dry season June to August, except at Lake Kutubu, where May-
October are the wettest months.
Factors related to breeding patterns —Rainfall patterns probably
have a significant effect on breeding, with dry weather suppressing
breeding in most species but eliciting synchronized breeding peaks in
88
FEATURES OF THE EASTERN HIGHLANDS
grassland and forest understory or ground-dwelling birds. ‘The sup-
pressing effect of very dry weather is illustrated by a comparison of
the 1964-1965 results. The following ten species were in breeding con-
dition at Karimui in July-August 1964 (July 30 to August 17) but
were not breeding (although resident) in the much drier months July-
August 1965 (July 1 to August 6): Trichoglossus haematodus, Psittac-
ulirostris desmarestii, Monarcha guttula, Pachycephalopsis poliosoma,
Artamus maximus, Manucodia chalybatus, Lophorina (Diphyllodes)
magnifica, Ailuroedus buccoides, Meliphaga flaviventer, and Meli-
phaga auga. In the cases of four additional species, breeding occurred
in July-August of both years, but the percentage of the local popula-
tion breeding was considerably higher in 1964: Ducula zoeae, Colluri-
cincla megarhyncha, Toxorhamphus poliopterus, and Melilestes me-
garhynchus. Similarly, a suspension of breeding by some bird species
in the dry season and a sudden resumption with the onset of rains has
been described in Trinidad by Snow and Snow (1964) and in Vene-
zuela by Gilliard (1959a).
Grassland birds evidently reverse the normal trend, wait for a dry
spell, and then come into breeding condition rapidly and synchro-
nously. A possible explanation is that nests in grassland, whether on
the ground or in grass, receive nearly the full force of a downpour and
are more susceptible to damage by rain than are nests in trees or the
forest. Similar considerations may apply to ground-dwelling forest
species. An analogy is provided by the duck Salvadorina waigiuensis,
which breeds in the Karimui area at the height of the dry season: it
nests on low boulders beside streams subject to flash floods after heavy
rains in the foothills, and such nests would have little chance of success
in wet periods.
It is interesting that the peak breeding season is the austral spring
and summer (October-March) in New Guinea and at most southern
hemisphere tropical as well as temperate localities, even within a few
degrees of the equator, while the peak breeding season at tropical as
well as temperate localities north of the equator is generally in the
boreal spring and summer, March-June (Moreau, 1966; Snow and
Snow, 1964). Part of the reason may be that the months of peak rain
correspondingly reverse near the equator. However, local rainfall pat-
terns may not be the sole determinant of breeding, since peak breed-
ing still occurs in the austral spring at Lake Kutubu even though the
rainfall pattern is locally reversed there, the spring being the end of
the rainy season rather than the beginning, as in much of the rest of
New Guinea. In the New World, Snow and Snow (1964, p. 32) tenta-
tively interpret a similar situation to mean that “natural selection
has led to complexes of closely knit seasonal changes, affecting both
animal and plant life, which may prevail over large areas and be
adapted to the general climatic conditions of the region but not per-
fectly adapted to special local conditions.”
89
FEATURES OF THE EASTERN HIGHLANDS
Factors which one might a priori postulate to integrate breeding
seasons over large areas at the same latitude despite local differences
in rainfall patterns include the annual cycles of daylength, light in-
tensity, and temperature. Another consideration is whether resident
birds time their breeding to avoid competition from the influx of
wintering and postbreeding birds from Australia, the palearctic zone,
and the New Guinea lowlands. As discussed on p. 00, these visitors
are strictly confined to open habitats and seem unlikely to influence
the breeding of forest species. Only four resident species of open coun-
try in the Eastern Highlands appear to face an influx of conspecific
or congeneric Australian or palearctic visitors: citing the resident first
and the vistor second, these are the falcons Falco berigora novae-
guineae and F. cenchroides cenchroides, the cuckoos Cacomantis pyr-
rhophanus excitus and C. p. prionurus, the dollarbirds Eurystomus
orientalis waigiouensis and E. 0. pacificus, and the swallows Hirundo
tahitica frontalis and H. nigricans nigricans. All four of these visiting
species are uncommon or local in the Eastern Highlands. The resident
Eurystomus and Falco were not breeding during the season of the
visitors’ presence in my study areas, while I have no information con-
cerning the breeding season in the other two cases.
Further progress in understanding breeding seasons of New Guinea
birds will require information about seasonal variations in food abun-
dance, combined with year-round records of bird cycles based on nests
and juveniles rather than on gonad condition.
NATIVE CLASSIFICATIONS AND KNOWLEDGE OF BIRDS
My introduction to the systems of zoological classification developed
by New Guinea natives came during the first week of my first season in
the Eastern Highlands, when I was staying at the Fore village of
Miarosa and was trying to learn some words of the Fore language. As
I was washing at the village spring one evening and a frog began
calling nearby, a small Fore boy explained, “Dakwo wanipindi min-
die.” I had already learned that “wanipindi” was the word for “water”
in the locative case, and that “mindie’’ was the form of the verb “to
be” used for living things, so that I assumed that the sentence meant
“There’s a frog in the water” and that “dakwo” meant “frog.” The
next day a man caught and brought me a frog while we were in the
forest, and I pointed to it and said “dakwo” to practice the new word.
“No, no,” I was told, “This isn’t dakwo. Dakwo is another fellow.
This one is ibisaraya.”” Evidently different species of frogs had different
names.
Thereafter I made a systematic effort to learn Fore names for ani-
mals, by asking individual natives to identify collected specimens, to
name bird calls we heard, to name birds we saw in the field, and to
describe animals they knew about but had not yet been able to show
90
FEATURES OF THE EASTERN HIGHLANDS
me. In this way I obtained Fore names and identifications or descrip-
tions of 110 birds, 15 small mammals, 20 large mammals, 16 frogs,
and 17 lizards and snakes. These names have been discussed elsewhere
(Diamond, 1966). Detailed analyses of zoological classification by the
Karam people of the Schrader Range have been published by Bulmer
(1967a, 1968) and Bulmer and ‘Tyler (1968). I made no attempt to
get names for plants but am sure that these were also classified, since
Paran and Esa responded to an idle remark I once made about mush-
rooms with a long lecture naming and describing 31 kinds of mush-
rooms in the vicinity of Awande, whether each was edible, and on
what each was likely to be found growing.
It turned out that the Fore had a name for every bird that occurred
regularly in the area. Usually each species had a separate name, and
even sibling species as similar as two Sericornis warblers or the two
Macropygia cuckoo-doves were distinguished. In a few cases (Gery-
gone warblers, Halcyon kingfishers, Melanocharis berrypeckers) related
species were lumped under the same name but might then be distin-
guished by an added epithet. For instance, the kingfisher Halcyon
sancta was named “‘patoroba bilong place” (the “patéroba” of settled
areas), while Halcyon megarhyncha was “‘patoroba bilong bush” (the
“patoroba” of the forest). In the cases of some birds of paradise and
bowerbirds with marked sexual dimorphism, such as Lophorina su-
perba and Amblyornis macgregoriae, the name for the adult male
differed from that for the female or immature male.
‘Towards the end of my first season, Paran, my best Fore informant
at Awande village, gave me names and descriptions of 30 birds which
he knew but which I had not yet collected. All of these were eventually
identified tentatively or definitely in 1965 and 1966. Many of the birds
he described were not only rare but small and undistinctive (e.g.,
Crateroscelis nigrorufa or the “fuintara”, Cisticola exilis or the “ikonan-
tube”, Clytomyias insignis or the “tabugiri’). Some of the birds he
discussed had been seen only once or twice in recent memory at
Awande. For instance, Paran had seen the “da” (Talegalla sp.) only
twice; the “topa” (apparently a white heron) had last been seen in
1964, when one individual appeared at Awande; and no one had seen
the “kwi-kwipa” (Gallinula tenebrosa or Porphyrio porphyrio) since
one had been captured 10 years earlier during Paran’s boyhood. These
names and descriptions proved useful in tracking down missing species.
Correspondingly, Fore knowledge of local birds was sufficiently ex-
haustive that in cases where I showed the Fore a specimen of a bird
and they stated that it did not occur at Awande, I feel that this infor-
mation can be relied upon.
The purpose behind Fore animal names appears to be utilitarian.
The only large animals to serve as sources of meat are the pig, the
cassowary, and (formerly) man, so that even the smallest birds are
hunted and eaten, as are mice, lizards, bats, and beetles. In addition,
91
FEATURES OF THE EASTERN HIGHLANDS
the feathers of many birds (birds of paradise, bowerbirds, and some
parrots and hawks) are prized for decoration. As a result, many Fore,
particularly boys and men, possess an incredibly detailed knowledge
of the habits and voices of birds and other animals in their area. It
should be realized that Fore natives vary individually in their knowl-
edge, reliability, and powers of observation, just as do Europeans.
Some of the younger generation of Fore, who are going to schools,
have acquired little knowledge of the local fauna. It should also be
realized that linguistic difficulties, misunderstandings, and inadequate
interviewing techniques can easily result in wrong information when
one is quizzing natives. Information elicited from primitive peoples
is systematically discounted by some ornithologists who have found
such information unreliable in their experience, as a result of work-
ing with poorly informed groups or poorly informed individuals, or
employing inadequate interviewing techniques. However, an undif-
ferentiated rejection or acceptance of all information supplied by
natives is no more justifiable than would be the undifferentiated re-
jection or acceptance of all information supplied by professional orni-
thologists. Some observations on bird distribution, behavior, or song
by Paran and several of my other New Guinea associates will be dis-
cussed in this monograph under the species accounts, because I found
these individuals equal to the best European observers I have known
in their powers of species recognition, knowledge of birds, and care
in separating fact from opinion or hearsay.
‘The species accounts in the remainder of this book give the Fore
name for each bird wherever identified with certainty. In some cases
the names in the North Fore dialect spoken at Awande and the South
Fore dialect spoken at Miarosa differ and are given separately. I have
also listed some names obtained from my Gimi assistants. The Gimi
and Fore languages are related but mutually unintelligible, and Gimi
and Fore names occasionally coincide but usually differ. Finally, I
have given the Daribi names given by my assistants from the Karimui
basin. Iwo supposedly unrelated languages, Daribi and ‘Tudawhe, are
spoken at Karimui, and comparison of lists of names from a Daribi
village (Bomai), a Tudawhe village (Soliabeda), and a border village
(Yudo) shows that there have been some word borrowings, so that some
Daribi names I cite may actually be derived from ‘Tudawhe. Many
more species of birds occur at Kari imui than at Awande, and I found
that the Daribi apply the same name to several related birds more
often than do the Fore.
SUMMARY
1. Paichiness of distribution—The distribution of some bird spe-
cies on the Central Range is discontinuous, in that they are absent
from the Eastern Highlands but present on other parts of the chain
92
FEATURES OF THE EASTERN HIGHLANDS
farther west and farther east. Numerous intermediate cases show that
speciation in the mountains of New Guinea usually involves the for-
mation of western and eastern isolates on the Central Range, which
then reinvade each other's ranges. The formation of such isolates on
a continuous mountain chain is made possible by distributional dis-
continuities resulting from local extinctions. ‘This distributional patch-
iness in New Guinea forest birds is probably due to low dispersal
rates and high species diversity, and also expresses itself in differences
between the avifaunas of nearby mountains and in fragmented, relict-
like distributions of some species.
2. Ecological sorting mechanims.—Completed speciation requires
not only the development of reproductive isolation but also of eco-
logical segregation. ‘The principal ecological sorting mechanism among
congeners in the mountains of New Guinea, and usually the first one
to develop, is altitudinal, such that two to five bird species replace
each other abruptly with altitude. ‘he sharpness of these transitions
is due to competition, as shown by compression of altitudinal ranges
during speciation and reinvasion and by expansion when one species
is locally eliminated. Congeners may also sort out spatially by occupy-
ing different habitat types; by foraging at different vertical levels in
the vegetational column; by allopatry; and by “checkerboard allo-
patry.’”” Nonspatial sorting related to food and foraging differences
frequently depends upon a difference in body size. ‘The ratio of body
weights of two congeners which sort out by size averages 1.90 and
usually falls between 1.5 and 2.5, due to a balance of selective pres-
sures. Instances of temporal segregation are rare in New Guinea.
3. Zoogeography—The Central Range may be divided into three
weakly defined regions, based on presence of endemic forms and ab-
sences of otherwise widespread forms. Most geographic variation in
montane birds on the Central Range is east-to-west and results simply
from distance. In birds confined to elevations below 5,000 ft variation
is mainly north-south because of the barrier posed by the Central
Range.
4. Avifauna of the Karimui Basin—Karimui, a flat mountain-
ringed basin at 3,500 ft, has an avifauna typical of the sea level low-
lands rather than of the hill slopes. Several lowlands birds isolated in
this tropical “island” within the highlands have differentiated to yield
melanistic endemic forms.
5. Altitudinal distribution.—Analysis of the altitudinal ranges of
166 forest bird species on Mt. Karimui, derived from altitudinal cen-
suses along the west ridge from 1,350 ft to the summit at 8,165 ft,
shows that species diversity decreases with increasing altitude; and
that definition of altitudinal montane “zones” of bird distribution
would be arbitrary, since species turnover is a fairly smooth function
of altitude except for the local anomalies introduced by the flat floor
of the Karimui Basin. The population structure of most species
93
FEATURES OF THE EASTERN HIGHLANDS
changes in a characteristic way with altitude: at the bottom of the
altitudinal range are immatures, nonbreeding adults appear somewhat
higher, breeding adults still higher, and another band of immatures
may exist at the upper limit of the altitudinal range. This population
structure 1s present to an exaggerated degree in birds of paradise.
6. The breeding nonforest avifauna—tIn the mountains of New
Guinea the breeding nonforest avifauna is limited in variety because
there is little natural nonforest habitat in the mountains. Some of
these nonforest species evolved in specialized habitats, at the forest
edge, or in forest; others have recently begun to spread upward from
the lowlands as native agriculture creates increasing expanses of open
country; and still others may have differentiated rapidly and recently.
7. Migrants.—Palearctic and Australian wintering migrants to the
Eastern Highlands are absolutely confined to nonforest habitats. ‘There
are a few cases of local migration of New Guinea species in the Eastern
Highlands, involving postbreeding lowlands birds, species depending
upon fruiting and flowering trees, and species escaping wet or dry
conditions.
8. Patterns of breeding activity.—In the Eastern Highlands the pat-
terns of breeding activity are very complex and vary among different
groups of birds, at different localities, at different elevations, and in
different years. In general, grassland species have a highly synchro-
nized breeding peak in the dry season; birds of second-growth, meliph-
agids, and other species dependent upon fruiting and flowering trees
tend to breed in the wet season; and lowerstory birds and warblers
tend to breed in the dry season.
9. Assemblages in fruiting and flowering trees, rainfall, and native
classification.—Bird feeding assemblages in fruiting and flowering
trees, rainfall patterns, and classifications of birds by New Guinea
natives are summarized.
94
IIl
SPECIES ACCOUNTS
Measurements and weights——All measurements are given in milli-
meters, and all weights in grams. Wing measurements are of the wing
straightened against a ruler, while tail measurements are from the in-
sertion of the central pair of rectrices to the tip of the longest rectrix.
In cases of large series I omit individual values and cite instead the
range, average value, and standard deviation in the form illustrated
by the example “8 4: 15.2-19.4 (17.4 + 1.2),” where 15.2-19.4 is the
range, 17.4 the average value, and 1.2 the standard deviation. In the
lists of specimens, a number followed by a question mark (e.g., “2 ¢,
1 9, 1imm. 4, 1 ?”) always means that the sex was not determined
and does not mean that the species identification is in question.
Specimens examined.—Under “specimens examined” I list not only
birds prepared and brought back as skins, skeletons, or in for-
malin, but also a large number of birds examined and often mea-
sured, weighed, and sexed in the field but not retained as specimens.
This latter category includes some specimens collected and offered for
sale by local natives, and some mistnetted birds that were subsequently
released, and consists principally of individuals of excessively common
species and of protected species.
Localities —The names and elevations of my collecting localities
may be summarized again here for convenient reference. Okapa area:
Awande (6,000 ft), Miarosa (5,800 ft), Okasa (3,550-4,250 ft). Mt. M1-
chael area: Lufa (6,300 ft), Mt. Michael (7,000-12,300 ft), Mengino
(4,600-6,150 ft). Karimui area: Karimui (3,650 ft), Bomai (3,250 ft),
Soliabeda (1,350-2,000 ft), Sena River (4,500 ft); Mt. Karimui Zone 1
(4,000-4,200 ft), Zone 2 (4,400-4,750 ft), Zone 3 (4,750-5,390 ft), Zone 4
(5,390-5,960 ft), Zone 5 (5,960-6,500 ft), Zone 6 (6,500-7,080 ft), Zone 7
(7,080-7,620 ft), Zone 8 (7,620-8,165 ft).
Stomach contents, voice, and abundance.—Because I did not exam-
ine stomach contents of my Eastern Highlands specimens, the stomach
contents analyses reported in the text are all from specimens I col-
lected on Mt. Albert-Edward in southeastern New Guinea in 1969.
All other remarks about specimens, behavior, and voice are based on
the Eastern Highlands unless specifically stated otherwise.
Figures 10-15, 17-35, and 37-42 depict vocalizations in simplified,
diagrammatic fashion by representing relative pitch vertically and rel-
ative duration horizontally, with syllables and sound quality specified
in some cases.
Estimates of relative abundances, obtained by the method described
on p. 10, are frequently stated, using the expression “At locality X,
species Y accounted for Z°%, of the local avifauna.”
95
SPECIES ACCOUNTS
Frequently cited references——The major previous collections in the
Eastern Highlands are cited so frequently in the text that full refer-
ences are omitted to save space. The relevant references are: Mayr
and Gilliard (1954), for Gilliard’s collections; Gyldenstolpe (1955), for
Gyldenstolpe’s collections; Sims (1956), for Shaw-Mayer’s collections;
Schodde and Hitchcock (1968), for Schodde’s collection at Lake Ku-
tubu; and unpublished manuscripts of R.N.H. Bulmer (1962, 196'7a)
summarizing his studies in Kyaka territory and in the Schrader Range.
“N.G.B.S. Newsletter” will be used as an abbreviation for the New
Guinea Bird Society Newsletter, published monthly at Port Moresby
since 1965.
Laxonomy and nomenclature. Scientific nomenclature in general
follows that used by Rand and Gilliard (1967) in their Handbook of
New Guinea Birds. Most of the deviations involve overdue or gen-
erally accepted lumping of monotypic or small genera.
Most vernacular names follow the current practice of the New
Guinea Bird Society, viz., to use current Australian names (An Index
of Australian Bird Names, CSIRO Division of Wildlife Research,
‘Technical Paper No. 20, 1969) for those New Guinea birds also found
in Australia, and to use names proposed by Rand and Gilliard (1967)
for other species. For species of New Guinea origin with a restricted
distribution in Australia, I have preferred the name proposed by Rand
and Gilliard if it differs from the Australian name and if the Austra-
lian name is less appropriate to the New Guinea population. For
example, the Australian name of Ptilinopus suwperbus, Purple-crowned
Pigeon, would be confusing in New Guinea, where there are several
additional species of pigeons with purple crowns, whereas the name
used by Rand and Gilliard, Superb Fruit Dove, had already been
applied to this species in several previous books and monographs
on New Guinea birds or southwest Pacific birds. Some names
proposed by Rand and Gilliard present problems of consistency
or suitability. For instance, Eugerygone rubra, Red-backed Warbler,
is no longer considered a warbler; /frita kowaldi, Blue-capped Bab-
bler, and the three species of Crateroscelis, Mouse-babblers, are
no longer considered babblers; Gerygone ruficollis, ‘Tree-fern Gery-
gone Warbler, is much more characteristic of other habitats than
of treefern fields; and Acanthiza murina, De Vis ‘Tree Warbler,
is the sole New Guinea representative of a genus of which the other
twelve species live in Australia and are all called thornbills (Western
Thornbills, Tasmanian Thornbill, Brown ‘Thornbill, etc.). In these
cases I have either modified the name proposed by Rand and Gilliard
or else substituted a name proposed by Schodde and Hitchcock (1968
and pers. comm.).
Seven new races based on my collections have been described previ-
ously (Diamond, 1967a), and no new names are introduced in this
monograph.
96
SPECIES ACCOUNTS
In evaluating the validity of proposed subspecies, I used as a stan-
dard the admirably thorough, sound, and consistent taxonomic analy-
ses of Mayr and Rand (1935, 1937) and of Rand (1936, 1938a, 1938b,
1938c, 1941a, 1941b, 1942a, 1942b) based on the collections of the first
three Archbold expeditions. Of the 99 New Guinea bird forms de-
scribed by Rand, there is no instance in which I was unable to discern
the characters cited in the diagnoses in the Archbold reports. Con-
versely, I have found no population of which Rand had adequate
material and in which I could detect reliable taxonomic characters
warranting naming but which Rand had refrained from naming. A
uniform and consistent taxonomic treatment for different groups of
birds and for different parts of New Guinea is an obvious prerequisite
for drawing undistorted conclusions about evolution, zoogeography,
and other biological problems of general interest, and the opportunity
to extract such conclusions seems to me to furnish the only significant
justification for concerning oneself with subspecies. In reviewing races
proposed since 1942, I have, accordingly, synonymized those races
which would not be considered distinct by the standards of the Arch-
bold reports.
CASUARIIDAE: CASSOWARIES
Casuarius bennettti shawmayerit Rothschild
Dwarf Cassowary
NATIVE NAMES. Fore: dmanani. Gimi: amananic. Daribi: képi. Pidgin En-
glish: moruk.
TAXONOMY. Apparently only one specimen of this species
known to have originated from a definite locality in the Eastern High-
lands has been taxonomically analysed. This was a female from the
Hagen Range, assigned by Sims to shawmayeri, a race originally de-
scribed from the Kratke Mountains at the eastern end of the Eastern
Highlands.
BREEDING. Gimi natives told me that eggs are found in March
and April, suggesting breeding towards the end of the rainy season.
DISCUSSION. Dwarf Cassowaries live in virgin forest of hills and
mountains up to about 7,000 ft, apparently in low numbers. ‘They are
shy and rarely seen but betray their presence by their droppings, con-
sisting of pits of large fruit which evidently represent their principal
food. When cornered, they defend themselves with powerful kicks:
Paran’s dog had been killed by a cassowary, and Gilliard (Mayr &
Gilliard, 1954) saw a human attacked and nearly killed. From native
accounts the adults are solitary and may be accompanied by one
young bird. ‘The meat is dark and said to be not unlike human flesh;
the large green eggs are eaten; and the feathers are used as brooms
97
SPECIES ACCOUNTS
and as decorations at native sing-sings. Captured young birds are kept
as pets in villages until large enough to eat.
Casuarius casuarius subsp.
Double-wattled Cassowary
Native informants at Karimui stated that two cassowaries occur
there. One was said to be a smaller bird, living on the mountain
slopes; I have seen captives kept by natives, and it is Casuarius ben-
nettii, the widespread montane cassowary. he cassowary on the floor
of the Karimui Basin itself was said to be much larger (about the
height of a man); this description was confirmed by a European resi-
dent who saw one that had been killed. This second species can only
be C. caswarius, the large cassowary of the lowlands, whose altitudinal
range generally appears to exclude that of C. bennetti. If confirmed,
the occurrence at Karimui would represent an altitudinal record for
this species, but it would not be surprising in view of the many other
lowlands species that have colonized the flat basin floor.
PODICIPEDIDAE: GREBES
Podiceps novaehollandiae novaehollandiae Stephens
Little Grebe
NATIVE NAME. Fore: nosho.
DISCUSSION. Gyldenstolpe collected one specimen on the Wahgi
River; Bulmer saw a pair and inspected a nest in the Baiyer Valley;
and natives report grebes as present in the Fore area.
PHALACROCORACIDAE: CORMORANTS
Phalacrocorax sulcirostris (Brandt)
Little Black Cormorant
Observed by Schodde and Hitchcock at Lake Kutubu, and reported
from Baiyer River (N.G.B.S. Newsletter, No. 54, p. 1, June 1970).
Phalacrocorax melanoleucos melanoleucos (Vieillot)
Little Pied Cormorant
Observed by Schodde and Hitchcock at Lake Kutubu, and found
breeding at Baiyer River (3,900 ft) by Bulmer.
98
SPECIES ACCOUNTS
ANHINGIDAE: ANHINGAS
Anhinga rufa papua Rand
Australian Darter
Observed by Schodde and Hitchcock at Lake Kutubu.
ARDEIDAE: HERONS and BIT’TERNS
Notophoyx picata (Gould)
Pied Heron
SPECIMENS EXAMINED. Karimui: | 2 (26 Aug. 1965).
WING. 224.
EXPOSED CULMEN. 64.
DISCUSSION. ‘The above specimen was collected by a native, who
said that he had shot it out of the top of a tree. A single bird stayed
for 10 days around a tea plantation at Mt. Hagen in June 1967
(N-G2B.5.. Newsletter, No: 23, ps 2, Sept. I9b7):
Egretta alba modesta (Gray)
Great White Heron
Bulmer collected one at 3,900 ft in the Baiyer Valley (18 Jan. 1956),
and Schodde and Hitchcock observed individuals at Lake Kutubu.
Egretta intermedia plumifera (Gould)
Plumed Egret
Observed at Lake Kutubu by Schodde and Hitchcock.
Nycticorax caledonicus hilli Mathews
Nankeen Night Heron
Gyldenstolpe obtained two from a swampy patch in the Wahgi
Valley. Schodde and Hitchcock found the species common at Lake
Kutubu.
Zonerodius heliosylus (Lesson)
Forest Bittern
Bulmer collected one at 4,700 ft on the Lai River (23 Nov. 1955).
99
SPECIES ACCOUNTS
Dupetor flavicollis gouldi (Bonaparte)
Black Bittern
Shaw-Mayer secured one at 7,300 ft on Mt. Giluwe.
THRESKIORNITHIDAE: IBISES and SPOONBILLS
Platalea regia Gould
Royal Spoonbill
One was collected by Gyldenstolpe in a boggy area near Nondugl.
ANATIDAE: GEESE and DUCKS
Anas superciliosa rogerst Mathews < A. s. pelewensis Hartlaub
and Finsch
Black Duck
In the Eastern Highlands the Black Duck has been collected in the
Wahgi River and near Mt. Giluwe, and observed at Lake Kutubu
and in a marsh near Okapa. ‘The two New Guinea races differ only in
size; Gilliard’s specimen agreed with the smaller pelewensis, while
specimens taken by Gyldenstolpe and Shaw-Mayer agreed with rogersi.
Anas querquedu la Linnaeus
Garganey ‘Teal
One female of this rare palaearctic visitor is cited as having been
collected at Lake Kandep (N'G. Bis, Newsletter, INo. WZ, p. 2, Och
1966).
Anas gibberifrons gracilis Buller
Gray Teal
Gyldenstolpe collected one female in the Wahgi River.
Salvadorina waigiuensis Rothschild and Hartert
Salvadori’s ‘Teal
NATIVE NAMES. Fore: noshé. Gimi: nosd. Daribi: gédi.
SPECIMENS EXAMINED. Soliabeda: 2 6,1 9 (24-28 July 1965).
WING. 2 4: 186, 189. 1 9: 180.
EXPOSED CULMEN, 2 4:-36; 38, 2 9: 35:
STOMACH CONTENTS. Insects and much gravel.
TAXONOMY. The species may increase in size with altitude. Six
100
SPECIES ACCOUNTS
males that I measured, collected at altitudes above 10,000 ft on Mt.
Wilhelm, in the Snow Mountains, and in southeastern New Guinea,
have wings of 195-201 (197 + 3) mm, larger than either of the Soli-
abeda males (from 1,350 ft, at the lower altitudinal limit of the
species).
BREEDING. Both males had enlarged testes. One of the males
was shot with the female, which contained an egg nearly ready to lay
as well as several other enlarged ova. On 27 July 1965 Paran found
a nest, a depression in a rock next to the Wi River, lined with erass
and containing three pale coffee-colored eggs. ‘hus, Salvadorina’s
breeding at Soliabeda falls in the dry season, which is teleologically
understandable: after rains in their watersheds, small highlands
streams are subject to sudden flash floods, which would immediately
destroy a nest like the one Paran found. Van Deusen (pers. comm.)
discovered the first known nest, in vegetation near the alpine lakes on
Mt. Wilhelm.
DISCUSSION. Of the 13 species of Anatidae reported from New
Guinea, this is the only endemic species, and the only one that fails
to descend to the lowlands. Most of the remaining 12 species are
largely or entirely confined to the lowlands. The poor representation
of Anatidae in the mountains is what one would expect from the very
small number and size of New Guinea’s montane lakes.
Single individuals of Salvadorina were seen several times on the Wi
River (1,350 ft) and Sena River (4,500 ft), which are rocky, turbulent,
and narrow mountain streams. In some cases the bird was perched on
boulders in the middle of the torrent, while in other cases it was first
noticed as it took flight, taxiing low over the surface of the water
instead of bounding straight up with a spring (cf., also, Mayr and
Rand, 1937, p. 11). To judge from reports of native informants, Salva-
dorina is a regular but uncommon inhabitant of rushing torrents on
mountain slopes in the Eastern Highlands. ‘The unusually low eleva-
tion of Soliabeda records is probably attributable to the fact that the
Wi River was still turbulent and in mountainous terrain. In addition,
Salvadorina occurs in another quite different habitat, namely, the
alpine lakes between 10,000 and 13,000 ft in the Snow Mountains,
Eastern Highlands, and southeastern New Guinea. Gilliard and Van
Deusen both observed it in the twin alpine lakes on Mt. Wilhelm.
Netlapus coromandelianus coromandelianus (Gmelin)
White-quilled Pygmy Goose
H. L. Bell (pers. comm.) states that L. W. C. Filewood found this
species abundant at Lake Kandep (7,000 ft).
101
SPECIES ACCOUNTS
ACCIPITRIDAE: HAWKS and EAGLES
Elanus caeruleus wahgiensis Mayr and Gilliard
Black-winged Kite
On 29 June 1965 I observed one individual over grassland at Tarabo
airstrip (6,000 ft) near Okapa. It flew back and forth 10-30 ft above
the ground, periodically stopping to hover in one spot. The manner
of hovering was distinctive: the axis of the body was inclined to the
horizontal at about 45°, head upwards, and the wing strokes were
entirely in a plane above that of the body. This behavior is essen-
tially the same as that of Elanus leucurus (sometimes considered con-
specific with E. caeruleus) which I have observed in South America.
Gilliard collected the type specimen and observed others in the
Wahgi Valley. There are also sight records by Bulmer and Bell at
Baiyer River (3,600 ft) and from Pureni in the western part of the
Eastern Highlands (N.G.B.S. Newsletter, No. 23, p. 4, Sept. 1967).
The expanses of the Eastern Highlands cleared by man seem ideal
habitat for this species, and it is surprising that it should be so local
and uncommon.
Aviceda subcristata stenozona (Gray)
Crested Lizard Hawk
SPECIMENS EXAMINED. Soliabeda: 1 ¢ (26 July 1965). Karimui: 1 ¢@ (4
Aug. 1965). Mt. Karimui zone 1: 1 ¢@,1 9 (11 Aug. 1965).
WING. 3 3G 284,294 308. 1 9% S07.
GULMEN FROM CERE. 34: 21, 22, 22.
STOMACH CONTENTS. Insects.
TAXONOMY. ‘The races stenozona of western New Guinea and
megala of eastern New Guinea differ only in size. Wing measurements
of comparative material yielded 272-305 (284 + 14) for five stenozona
males, 306 and 311 for two megala males. Values given by Rand and
Gilliard (1968) are: stenozona, g 290-303, 2 296-314; megala, ¢ 298-
316, 9 314-334. The present material is thus closer to stenozona. A
single immature female taken at Lake Kutubu by Schodde and Hitch-
cock had a wing of 294 mm and was also assigned to stenozona. ‘The
barring on the underparts of my four specimens is darker than in
most other material, nearly charcoal.
BREEDING. Although the male and female from Mt. Karimui
were together as a pair, the gonads of these four specimens were small.
DISCUSSION. Individuals or groups of up to five were seen soar-
ing high or else perched in tall trees overlooking open spaces. Once I
saw a bird attacked and driven off by the flycatcher Peltops montanus.
102
SPECIES ACCOUNTS
The two specimens from the lower slopes of Mt. Karimui may con-
stitute an altitudinal record (4,100 ft) for this lowland species.
Henicopernis longicauda longicauda (Garnot)
Long-tailed Buzzard
While hawks in general are not numerous in the Eastern Highlands,
this species is among the less uncommon. Until one learns to recog-
nize it in flight, however, Henicopernis is easy to overlook entirely,
since it is usually seen soaring rather high. ‘The wings are long,
rounded, and broad, the head is small, and the tail is long, rounded,
and slender, so that the silhouette suggests three long loops and one
short loop attached at a point and radiating at right angles. ‘he pat-
tern of the undersurface is also distinctive, the head, upper breast,
wings, and tail appearing dark, the lower breast and belly light. ‘Vhe
primaries are often held outstretched. With practice the combination
of the distinctive shape and undersurface pattern enables one to recog-
nize Henicopernis at a moderate distance. It is rarely seen sufficiently
close for the barring of the wings and tail to be visible. I saw it regu-
larly in all parts of the Eastern Highlands over both cleared country
and forest, usually soaring without flapping, sometimes alternately
soaring and flapping. About half of my observations were of pairs or
of groups of three birds. One pair was seen to chase off an individual
of the kite Haliastur indus.
Milvus migrans affinis Gould
Black Kite
NATIVE NAME. Fore: kékepa.
DISCUSSION. In the Eastern Highlands this species is widespread
in districts with large expanses of man-made grasslands (but is absent
at Karimui, where grassland is limited and probably recent in origin).
It gathers in large numbers at the leeward side of grassland fires in
wait for fleeing grasshoppers, and searches the ground for burnt prey
once the fire has passed. Elsewhere in New Guinea Milvus migrans is
a lowlands species and has not been recorded even from the extensive
man-made grasslands of the Baliem Valley of western New Guinea. I
do not know whether it breeds in the Eastern Highlands or whether
it is a seasonal nonbreeding visitor from the lowlands. The birds I
watched were always soaring low over the ground in the manner of a
harrier (Circus). ‘The indented tail, unique among New Guinea hawks,
facilitates identification in silhouette.
103
SPECIES ACCOUNTS
Haliastur indus girrenera (Vieillot)
Brahminy Kite or Red-backed Sea Eagle
NATIVE NAMES. North Fore: pirindmu. South Fore: pilam.
SPECIMENS EXAMINED. Karimui: 1 imm. ¢? (17 July 1965).
WING. 357.
EXPOSED CULMEN. 24.
STOMACH CONTENTS. Grasshoppers.
DISCUSSION. ‘This species, Henicopernis longicauda, and Milvus
migrans are perhaps the commonest hawks of the Eastern Highlands.
‘The chestnut and white adults are unmistakable, while the imma-
tures may be recognized in flight by the light “window” in the distal
half of the wing. I found Haliastur indus widespread (Okapa, Lufa,
Karimui) up to 6,600 ft, and Bulmer found a nest at 4,000 ft. It was
usually seen soaring over open country, generally singly but sometimes
in groups of up to four. About once a month it came to eat spiders
out of their webs near the European houses at Okapa, and one indi-
vidual was seen with a rat, but much of the diet is insects caught on
the wing. A soaring individual was chased off by a pair of Henicoper-
nis longicauda. While this is mainly a lowlands species, the ‘Third
Archbold Expedition (Rand, 1942b) and Ripley (1964) found that it
has also colonized the man-made grassland of the Baliem Valley of
western New Guinea.
VOICE. A weak, somewhat nasal, high-pitched downslur, sugges-
tive of the bleating of a lamb and incongruously faint for so large a
bird.
Accipiter buergerst (Reichenow)
Buerger’s Goshawk
SPECIMENS EXAMINED. Karimui: 1 ? (probably Q from measurements) (13
July 1965).
WING. 316.
TATE, 228,
CULMEN FROM CERE. 26.
SOFT PARTS. Iris: light yellow. Legs: light yellow-green. Cere: light green.
Bill: dark gray.
TAXONOMY. ‘The feathers of the back have black centers and
chestnut edges, the impression of black predominating on the upper
back and of chestnut on the lower back. ‘The head is black, with a
few white feathers; the upper tail charcoal with nine thin lighter
bars; and the upper wings charcoal brown, with faint light bars on
the inner edges of the primaries. The underparts from the chin
through the undertail coverts are white unevenly washed with light
chestnut, the chestnut being darker on the flanks and shanks and ab-
sent on the throat. There are small black streaks on the throat, bold
black streaks on the breast, flanks, and shanks, and black spots on the
104
SPECIES ACCOUNTS
lower belly and undertail coverts. The axillaries are rufous with black
centers, and the undersurface of the primaries and tail alternately
barred black and white.
Only about seven other specimens exist, of which two were available
for comparison: the type of Accipiter eudiabolus Rothschild and Har-
tert, an unsexed adult (presumed male from its measurements) from
Babooni at 3,000 ft in the mountains of southeastern New Guinea; and
an immature, also presumed to be male, from the Hydrographer
Range of southeastern New Guinea. ‘The Babooni adult has the head
and upperparts black, the underparts white with bold black streaks,
and the feathers of the upperwing coverts black with broad chestnut
edges; wing 295, tail 201, exposed culmen 22 mm. ‘Vhe Hydrographer
Range immature has the head, upperparts, axillaries, and underparts
of the body mainly rich chestnut, with bold black streaks below and
black centers to the feathers of the upperparts; wing 292, tail 206, ex-
posed culmen 21 mm. In both of these specimens, as in the Karimui
bird, the undersurface of the wings and tail is barred black and white.
The Karimui specimen is thus not fully adult but more nearly so
than the Hydrographer Range specimen, from which it differs in that
the chestnut wash of the axillaries and underparts is paler, the ground
color of the throat and a few patches on the breast and undertail
coverts are already pure white, and the black centers of the back
feathers have increased at the expense of the chestnut peripheries,
especially on the upper back.
DISCUSSION. Accipiter buergersi, the rarest species of hawk en-
demic to New Guinea, is apparently confined to low altitudes (1,900-
5,200 ft) in the mountains of eastern New Guinea. Besides Karimui
and two or three localities in southeastern New Guinea, the other
localities where it has been taken are Junzaing on the Huon Peninsula,
the type locality of Maeanderberg and the Lordberg in the mountains
south of the Sepik River, and the Sau Valley of the Eastern Highlands,
where Bulmer collected one specimen at 5,200 ft (5 Dec. 1955).
In the upper part of the altitudinal range of Accipiter buergersi
there is no other forest Accipiter, but in the lower part of its range it
is one of three forest species in its genus. The three species differ in
size, each being about 50% heavier than the next: A. buergersi > A.
novaehollandiae > A. poliocephalus.
Accipiter novaehollandiae leuwcosomus (Sharpe)
Gray Goshawk or White Goshawk
NATIVE NAMES. Daribi: kemorage (melanistic phase), pendgo (white phase).
SPECIMENS EXAMINED. Karimui: one specimen preserved in alcohol (pre-
sumed ¢ from measurements) (28 July 1964); 1 ¢ (3 Aug. 1965).
WING. Tg: 2i7. lL Grr 215,
TATE, Ver 162. Lars 158)
105
SPECIES ACCOUNTS
CULMEN FROM CERE. 14: 17. 12?: 17.
STOMACH CONTENTS. A small bird.
TAXONOMY. ‘These specimens have been described previously
(Diamond, 1967a). Accipiter novaehollandiae has been known to exist
in two color phases, an all-white phase and a colored (russet and gray)
phase, whose frequencies vary geographically. Thus, the white phase
is characteristic of ‘Tasmania; only the colored phase occurs in the
D’Entrecasteaux Archipelago (subspecies pallidimas) and the Louisi-
ade Archipeiego (subspecies misulae); and both phases occur on New
Guinea in roughly equal proportions or with the colored bird slightly
more common. In New Guinea Accipiter novaehollandiae is mainly a
lowland species, although it does reach the Baliem Valley of the west-
ern New Guinea mountains. In the Eastern Highlands it is not known
to occur outside of the Karimui Basin; it was even absent at Soliabeda
(2,000 ft) nine miles southeast of Karimui Patrol Post but beyond the
mountains ringing the basin. ‘This isolated Karimui population is re-
markable in that both of the above specimens, and all individuals seen
by me in 1964 and 1965, were melanistic, i.e., a uniform dark gray-
brown. The only other record of melanism in this species is a single
female from the Hydrographer Mountains of southeastern New
Guinea. According to Karimui natives, the dark phase (‘“kemorage’’)
is resident at Karimui throughout the year and nests at the tops of
very tall trees, while individuals of the white phase (“penago’”’) appear
only infrequently and disappear after a few days. ‘(he occurrence of
geographically localized melanistic, white, and colored phases is of
interest, as is the extremely limited range of the melanistic form in
an ecological island surrounded by areas where only the other two
forms occur.
BREEDING. ‘The gonads of the 1965 males were small.
DISCUSSION. I found solitary individuals at Karimui and Bomai
in trees overlooking gardens and cleared areas.
VOICE. (see Fig. 10). An unhurried series of 8-10 thin, high,
upslurred notes, surprisingly weak for a bird its size, and similar to
the call described for the species in Australia (Cayley, 1959).
Accipiter novaehollandiae:
ee ee ee eae, ies Fee
, 2 sec
Fic. 10. Voice of Accipiter novaehollandiae.
106
SPECIES ACCOUNTS
Accipiter fasciatus polycryptus Rothschild and Hartert
Australian Goshawk
SPECIMENS EXAMINED. Karimui: 19 (2 Aug. 1965). Bomai: 1 imm. 9 (7
July 1965).
WING. 1 9: 260. 1 imm. @: 204.
TATE... 1 9: 199. Loimm:. 9: 142.
CULMEN FROM CERE. 1 @: 18. 1 imm. 9: 17.
TAXONOMY. ‘This material agrees with the eastern New Guinea
race polycryptus. ‘The race dogwa of the Fly River bulge of southern
New Guinea differs in being considerably paler below, especially on
the abdomen, and in having the brown bars on the breast narrower
than the intervening white areas (vice versa in polycryptus).
BREEDING. ‘The Karimui adult female had two ovaries.
DISCUSSION. At Karimui the habitat preference and behavior
of Accipiter fasciatus appeared similar to those of Accipiter novaehol-
landiae, i.e., solitary individuals were seen perched in trees overlooking
native gardens and open spaces. A perched immature was repeatedly
attacked by the flycatcher Rhipidura leucophrys and finally flew off
with the flycatcher in pursuit. Other Eastern Highlands records are
from the Wahgi Valley, where Gilliard and Gyldenstolpe found it
“not common” to “fairly rare,’ and from Kyaka territory, where Bul-
mer reported it as common.
VOICE. A series of about a dozen or more thin notes, progres-
sively rising in pitch.
Accipiter poliocephalus Gray
New Guinea Gray-headed Goshawk
SPECIMENS EXAMINED. Karimui: 1 9 (13 July 1965). Bomai: 1 9 (8 July
1965). Soliabeda: 1 ¢, 1 9 (24 and 30 July 1965). Mt. Karimui Zone 3: 1 9? (20
Aug. 1965).
WING. 1 @: 195: 4 9: 209: 215, 221, 227.
GULMEN FROM CERE. 1 ¢: i7, 4 97: 19, 20, 20; 21.
STOMACH CONTENTS. A lizard.
BREEDING. ‘The gonads of all specimens were small.
DISCUSSION. My few observations of this lowlands hawk at Kari-
mui and Bomai were of solitary individuals perched in trees in native
gardens, a habitat description which could apply equally well to the
other two Accipiter species in the Karimui Basin (A. novachollandiae
and A. fasciatus). However, A. poliocephalus also lives within the
forest: the Mt. Karimui (5,000 ft) and Soliabeda females were both
taken while diving for netted or wounded flycatchers in the forest. In
addition to its greater preference for the forest interior, A. polioce ph-
107
SPECIES ACCOUNTS
alus may sort out ecologically from its congeners A. novachollandiae
and A. fasciatus on the basis of its smaller size (the two congeners
weigh 50%, more), suggesting different prey.
Accipiter melanochlamys schistacinus (Rothschild and Hartert)
Black-mantled Goshawk
NATIVE NAME. Fore: amaipdana.
SPECIMENS EXAMINED. Awande: 1 ¢@ (15 June 1965). Mt. Karimui Zone 6:
I g © Sept. 1965).
WING, 2°42 2137-220:
CULMEN FROM CERE. 2 @: 16, 17.
STOMACH CONTENTS. A small bird.
TAXONOMY. The single available specimen of the nominate
race, which is confined to the Vogelkop, is blacker above, especially on
the head, and darker below than my two specimens or other schista-
cinus material.
BREEDING. ‘Testes of both specimens were small.
DISCUSSION. Both specimens were obtained in the interior of
virgin montane forest, remote from human disturbance. The Awande
specimen was shot while perched at the top of a tree; the Mt. Karimui
specimen (ca. 6,800 ft) was driven into a mistnet by my Fore assistants
while it was trying to eat a small dicaeid Melanocharis versteri trapped
in the net. Another specimen that subsequently escaped was netted
at 8,500 ft inside the forest on Mt. Michael, where it too had presum-
ably been after a netted bird and become trapped of its own efforts.
One individual was surprised on the ground in the forest and flew
into the treetops. The habit of preying on other birds is responsible
for its Fore name, “amaipana’”’, which means “killer of one’s brother.”
This species has generally been considered rare although widespread,
and less than 30 specimens exist. It nevertheless occurs regularly in the
Eastern Highlands, even if at a low density, from at least 5,500 ft to
8,700 ft wherever there are large expanses of forest undisturbed by
man. Thus, I obtained one specimen each at my three midmontane
stations; and Gilliard, Gyldenstolpe, Shaw-Mayer, and Bulmer each
collected one to three specimens in the Wahgi Mountains, Kubor
Mountains, Schrader Range, Mt. Hagen, and Mt. Giluwe. It is the
only forest Accipiter in this altitudinal range and is replaced in the
forest at lower elevations by A. poliocephalus and A. novaehollandiae
(and the very rare and geographically limited A. buergers?). On Mt.
Karimui a vertical distance of 1,800 ft separated the sites where A.
melanochlamys and A. poliocephalus were netted. Records elsewhere
also suggest that altitudinal segregation of A. melanochlamys trom its
forest congeners is complete.
108
SPECIES ACCOUNTS
Megatriorchis doriae Salvadori and D’Albertis
Doria’s Hawk
SPECIMENS EXAMINED. Soliabeda: 1 ¢ (29 July 1965).
WING. 287.
TAME. — 250;
CULMEN FROM CERE. 25.
BREEDING. ‘The testes were small.
DISCUSSION. ‘The monotypic genus Megatriorchis is one of the
rarest endemic New Guinea hawks. The above specimen was brought
in by a Soliabeda native, who spotted it perched: ima thee inside the
forest. He hid in the undergrowth, made noises imitating a small bird,
and shot the hawk with bow and arrow when it came down to investi-
gate. The previous day I had seen probably the same individual
perched inside the forest in the crown of a flowering tree, which had
attracted many small birds and butterflies. During the 15 minutes that
I watched the hawk, it remained stationary. Megatriorchis doriae has
been recorded from forest at altitudes up to’ 3,000 Ir atea number of
localities scattered over New Guinea.
Harpyopsis novaeguineae Salvadori
New Guinea Eagle
NATIVE NAMES. Fore: tiyé6. Gimi: luipa.
DISCUSSION. Harpyopsis, the largest endemic New Guinea hawk,
is quite uncommon but widely distributed in the Eastern Highlands,
inhabiting forest up to about 9,500 ft. The individuals I observed
were either soaring just over the top of the forest or else perched
inconspicuously inside the forest in the crowns of trees. ‘The most re-
markable feature of this eagle’s habits is that it hunts by night as well
as by day, and at one camp the calls of a pair of Harpyopsis awakened
me repeatedly throughout the night. In life the feathers around the
eye form a distinct facial disc, as in owls, and may play some role in
night vision or hearing. ‘he prey consists of large marsupials and
birds and small pigs.
VOICE. A single note with a quality like the sound made by a
taut bowstring being released, followed by a hollow note repeated one
to four times and like the foraging call of a domestic chicken: bung!
—buk-buk-buk-buk.
Circus spilonotus spilothorax Salvadori and D’Albertis
Spotted Marsh Harrier
My only observation was of a black-and-white male soaring low over
109
SPECIES ACCOUNTS
grassland near Okasa (4,250 ft) in June 1965. The other Eastern High-
lands records are from grassland at Nondugl, where Gilliard and
Gyldenstolpe each saw it a few times, from grassland near Mt. Giluwe,
where Shaw-Mayer observed it often, and from gardens and grassland
in the Baiyer Valley and Schrader Range, where it was found by Bul-
mer. Its distribution is evidently quite local, and I interpret my failure
to find it in the grasslands at Okapa, Lufa, and Karimui as probably
indicating its being actually absent from those localities, since the
distinctive male is easy to recognize even at a distance.
Circus approximans gouldi Bonaparte
Swamp Harrier
SPECIMENS EXAMINED. Karimui: 1 ? (12 July 1965).
WING, 400:
TAME. 254.
CULMEN FROM CERE. 24.
SOFT PARTS. Iris: brown. Legs: yellow. Cere: yellow. Bill: black.
TAXONOMY. ‘This specimen is a nearly solid and uniform dark
brown below, except for a paler chin and some inconspicuous dark
shaft streaks. ‘The upperparts are also uniform dark brown, except
that there are a few white patches on the lower back, the nape is largely
white, and the forehead and area around the base of the bill light gray.
DISCUSSION. ‘This Australian and New Zealand race was pre-
viously known in New Guinea mainly or only from sea level on the
southern coast, and provides yet another example of a lowland species
reaching the Karimui Basin. The Second Archbold Expedition (Rand,
1941a) found both this form and Circus spilonotus spilothorax at Lake
Daviumbu in the same month, strengthening the case for considering
them as separate species. At Merauke C. approximans is present onlv
in the dry season and C. spilonotus only in the wet season (Hoogerwerf,
1964).
FALCONIDAE: FALCONS
Falco peregrinus ernestt Sharpe
Peregrine Falcon
DISCUSSION. Few Eastern Highlands records exist: a male col-
lected by Shaw-Mayer at 7,300 ft on Mt. Giluwe; one seen by Van
Deusen (Sixth Archbold Expedition) at 11,400 ft over the summit grass-
land of Mt. Michael; and possible sight records by Bulmer in Kyaka
territory.
VOICE. A fairly high-pitched, slightly hoarse, weak, upslurred
screech “ka-ah,”’ repeated about seven times in 5 sec.
110
SPECIES ACCOUNTS
Falco severus papuanus Meyer and Wiglesworth
Oriental Hobby
NATIVE NAME. Fore: akik.
SPECIMENS EXAMINED. Soliabeda: 1 9 (25 July 1965).
WING. 233.
‘EAIE, 0%.
CULMEN FROM CERE. 15.
SOFT PARTS. Ivis: brown. Legs: yellow. Cere: yellow. Bill: black.
BREEDING. ‘The specimen had two ovaries.
DISCUSSION. ‘This is apparently the only specimen collected to
date in the Eastern Highlands and was one of a pair that circled over
the village of Soliabeda. The following day one individual, presumably
the surviving mate, appeared over the village. In addition, on several
occasions at Miarosa (5,800 ft) in June 1964 I observed a pair repeatedly
alight in the top of a tall dead tree, take off, circle, and return to the
same perch. Finally, I saw one individual in Gimi territory (5,000 ft)
in July, 1964. Thus, Falco severus may be regarded as local and un-
common in the Eastern Highlands in open country with tall trees.
VOICE. A rapid series of typically falcon-like notes, descending in
pitch.
Falco cenchroides cenchroides Vigors and Horsfield
Nankeen Kestrel
SPECIMENS EXAMINED. Karimui: 5 9 (1-17 July 1965). Bomai: 1 9 (7 July
1965).
WING. 240-264 (255 + 9).
Merb 145-160 (54) = (6):
CULMEN FROM CERE. 14.5-15.0 (14.9 + 0.2).
WEIGHT. 154-189 (167 + 16).
SOFT PARTS. Iris: brown. Legs: yellow. Bill: light grey.
DISCUSSION. Falco c. cenchroides is an Australian bird for which
there were no New Guinea records until recent years, when it began
to turn up as a winter visitor (March to September) in southern New
Guinea, (N.G,B.S. Newsletter, No. 10) pe 2; Aug, 1966; No, 14, p. 3,
Dec. 1966; No. 43, p: 1, May 1969; No. 44, p. 1) July 1969; van den
Assem, 1960). As discussed previously (Diamond, 1967a), my specimens
were collected in June and July of 1965, when I frequently saw this
falcon flying over the Karimui and Bomai airstrips. In addition, I saw
an individual over Karimui airstrip in June 1966 and in April 1969,
and a photograph of a falcon shot by the patrol officer at Karimui in
1964 establishes it as this species. ‘he fact that all specimens were
females with small ovaries suggests that they were nonbreeding
vagrants. Stresemann (1914a) pointed out earlier that the records of
Falco c. cenchroides outside of Australia (from New Zealand, Java,
Babbar, Ceram, and the Aru Islands) are nearly all females.
111
SPECIES ACCOUNTS
Falco berigora novaeguineae (Meyer)
Brown Hawk or Brown Falcon
SPECIMENS EXAMINED. Karimui: 1 3 (13 Sept. 1965).
WING. 310-1.
(WALES Tso:
CULMEN FROM CERE. 21.
STOMACH CONTENTS. Hair and jaws of a small mammal.
SORT VRARGES a Lins: light brown. Legs: whitish. Cere: lemon to white. Bill:
eray.
BREEDING. ‘The testes were small.
DISCUSSION. In addition to the specimen collected, I observed
this falcon several times at Karimui and Iogoramalu in the Karimui
Basin, near Okapa, and in Gimi territory. Individuals were seen
perched in trees in gardens or open country, and twice I saw pairs
chasing each other, diving and twisting rapidly and calling. Gylden-
stolpe and Bulmer also collected specimens. ‘Thus, this falcon may be
widespread, but uncommon, in the Eastern Highlands in open country
(never in the forest interior).
VOICE. The pairs chasing each other uttered a very rapid series
of harsh notes on one pitch delivered at machine-gun rate; this could
alternatively be described as a crackling scream. ‘This series was some-
times punctuated by two more emphatic notes.
MEGAPODIIDAE: MEGAPODES or
BRUSH TURKEYS
Megapodius freycinet subsp.
Common Scrub Hen or Scrub Fowl
NATIVE NAMES. Fore: katba. Gimi: oedéc. Daribi: neyé.
STOMACH CONTENTS. Insects, centipedes, snails, pebbles, and sand.
DISCUSSION. According to native reports, Megapodius freycinet
is at Soliabeda (2,000 ft), Karimui (3,650 ft), on Mt. Karimui itself
(above 4,000 ft), and at Awande (6,000 ft), where Paran collected it.
It also occurs in Kyaka territory, where natives brought eggs to
Bulmer on several occasions. In much of its range this is a lowland
species, but the Third Archbold Expedition (Rand, 1942b) found it
up to 4,000 ft in the Snow Mountains, and Greenway (1935) recorded
it up to 5,700 ft in the Herzog Mountains.
Talegalla sp.
(Talegalla jobiensis, Brown-collared Brush ‘Turkey, and/or
Talegalla fuscirostris, Black-billed Brush ‘Turkey)
NATIVE NAMES. Fore: 6a. Gimi: alofya. Daribi: wadi.
112
SPECIES ACCOUNTS
DISCUSSION. Some species of Talegalla is widespread in the
Eastern Highlands. On Mt. Karimui, between 5,000 and 6,500 ft, I
heard calls virtually identical to the calls of Talegalla jobiensis that I
heard on the north coast. My native assistants reported Talegalla in
the Okapa area below 6,000 ft, and down to at least 2,000 ft in the
Karimui area. At Lake Kutubu Schodde and Hitchcock observed a
Talegalla which they assumed was T. fuscirostris, but they did not
obtain specimens.
The species of Talegalla involved is uncertain, and its identification
will answer a question of considerable interest. ‘There are three species
of Talegalla, which are very similar morphologically and nearly but
not quite allopatric: T. cuvieri of far western New Guinea, T. fusci-
rostris of southern New Guinea, and T. jobiensis of northern, north-
eastern, and southeastern New Guinea. T. fuscirostris is a lowland
species, apparently not recorded at elevations higher than a few
hundred feet, whereas T. cuviert and T. jobiensis occur up to 3,000-
5,000 ft. T. fuscirostris and T. cuviert overlap for a distance of at
least 150 miles in southwestern New Guinea around long. 136° E, and
in this area they appear to segregate altitudinally, with 7. cuvierl con-
fined to higher elevations. T. fuscirostris and T. jobiensis overlap for
at least 50, perhaps 250, miles in southeastern New Guinea around
long. 147° E. I would guess that the Talegalla at Okapa and Karimui
will prove to be T. jobiensis rather than T. fuscirostris, because the
altitude would be typical of the former and unprecedented for the
latter, and because the calls of the Karimui Talegalla were indis-
tinguishable from those of 7. jobiensis. ‘The evolutionary history im-
plied by these distributions is that genus Talegalla once consisted of a
superspecies ring of three allopatric forms; and that the range of the
southern form, T. fuscirostris, is being reinvaded by T. cuviert from
the west and by T. jobiensis from the east, with the forms segregating
altitudinally in each overlap zone.
VOICE. A nasal, unhurried, and very loud call which may be
rendered “wa-wa-wa’’, suggestive of a child wailing but more nasal
and deeper, and consisting of two detached upslurs followed by a
steady note (Fig. 11).
Aepypodius arfakianus arfakianus (Salvadori)
Wattled Brush ‘Turkey
NATIVE NAMES. Fore: iya. Daribi: dasiari.
STOMACH CONTENTS. A broken fruit pit, and much gravel.
DISCUSSION. ‘This species of megapode appears to be generally
distributed, though quite uncommon, in primary forest between 3,000
and 9,000 ft, having been observed or collected at several localities by
Gilliard, Shaw-Mayer, and me. According to natives, Aepypodius flies
113
SPECIES ACCOUNTS
Talegalla sp. :
eg — (nasal)
wd wd wd
| = USEC }
Fic. 11. Voice of Talegalla sp.
up into trees when disturbed, whereas Talegalla prefers to remain on
the ground. ‘This is in accord with our observations: an Aepypodius
seen by Terborgh at Okasa was in a tree, and those that I saw on the
Sena River were birds which burst up from the ground at my approach
and alighted heavily in trees.
PHASIANIDAE: QUAIL and PHEASANTS
Synoicus ypstlophorus lamonti Mayr and Gilliard
Brown Quail
NATIVE NAME. Fore: yokayo.
SPECIMENS EXAMINED. Awande: 1 ¢ (20 June 1965).
WEIGHT. 83.
WING, 91.
TAXONOMY. ‘This specimen agrees with topotypical lamonti
males. Its upperparts are more chestnut and richly colored than those
of mafulu, while dogwa and plumbeus differ in the tendency towards
uniform gray coloration of the underparts.
BREEDING. The testes of the specimen (collected in June) were
enlarged. On 18 August 1964 a Fore man at Awande showed me a
nest on the ground in grass containing five eges (one 33 x 24 mm)
colored very pale tan with dark brown speckles. ‘The “yékay6” was
said to be generally nesting then at Awande, which implies that breed-
ing is in the dry season, as true of most other grassland birds as well.
DISCUSSION. I found small groups of this quail not uncommon
in grassland and native gardens at Miarosa, Awande, and Okasa. Other
collectors have also encountered it regularly in the Eastern Highlands
wherever there are substantial expanses of midmontane grassland and
gardens. In southeastern New Guinea, but not in the Eastern High-
lands, it also occurs in alpine grassland.
114
SPECIES ACCOUNTS
VOICE. A two-note nasal whistled call, the second note a queés-
tioning upslur at a higher pitch (Fig. 12).
Synoicus ypsilophorus:
eg
ze (whistled)
2 SeC
Fic. 12. Voice of Synoicus ypsilophorus.
Coturnix chinensis novaeguineae (Rand)!
Chinese Quail or King Quail
Other collectors have found this small quail to occur, but in con-
siderably lower numbers than Synoicus ypsilophorus, in the grassland
of the central part of the Eastern Highlands (Mts. Wilhelm, Hagen,
and Giluwe, and the Wahgi Valley). At Okapa and the Karimui Basin
it is apparently absent.
LURNIGIDAE: BUSTARD QUAIL
Turnix maculosa subsp. and T. m. giluwensis Sims
Black-backed Bustard Quail or Red-backed Quail
NATIVE NAME. Fore: mu.
SPECIMENS EXAMINED. Okasa: 1 @ (23 June 1965).
WEIGHT. 26.
WING. 67.
EXPOSED CULMEN. II.
TAXONOMY. The race giluwensis, described from Mt. Giluwe
(Sims, 1954), was characterized as lacking the chestnut collar of hors-
brughi, smaller than furva, and paler below than furva or horsbrughi.
Only one specimen each (both females) of giluwensis and horsbrughi
was available for comparison. ‘The Okasa male agrees with gilwwensis
in lacking the chestnut collar of horsbrughi; the orange-brown color of
the breast 1s richer than that of giluwensis but less rich than that of
horsbrughi; the color of the lower belly is pale buff as in horsbrughi,
1 Listed as Excalfactoria chinensis in Rand and Gilliard (1967).
MS
SPECIES ACCOUNTS
not white as in giluwensis. The wing length is close to that of the two
known males of giluwensis, given as 65 mm by Sims (1956). (Rand
(1942a) gives the wing of one male horsbrughi as 72 mm, and the male
of furva is unknown). This meagre material is insufficient to assign
the Okasa population racially but suggests that it is intermediate
between giluwensis and horsbrughi.
DISCUSSION. The Okasa specimen, which apparently is the fourth
known New Guinea male of this rare species, was captured by hand by
a Fore man in grassland. The other Eastern Highlands records of this
somewhat local species are from Awande, Mt. Giluwe, and Nondugl.
RALICIDAE: RATES
Rallus pectoralis captus Mayr and Gilliard
Slate-breasted Rail or Lewin Rail
NATIVE NAME. Fore: indénpa.
SPECIMENS EXAMINED. Awande. 1 ¢, 3 9 (27 Aug. 1964; 18-20 June 1965).
WEIGHT. 14:73. 2 9: 69, 77.
WING. 1 4: 100. 3 9: 95, 96, 100.
EXPOSED GUEMEN. TV @: 27. 3 93 27, 28, 31.
TAXONOMY. ‘These Awande specimens agree with topotypical
captus from Mt. Hagen in size and in the rich brown coloration of the
top of the head and neck. ‘The male specimen has the crown dark
olive-brown rather than chestnut.
DISCUSSION. All four specimens were trapped in grassland by
natives. Most other collectors in the Eastern Highlands have also
obtained trapped specimens. ‘The species is evidently fairly common
but secretive in grassland from 5,000 to 8,000 ft. It is absent at Karimui,
probably because of the low altitude.
Rallus philippensis wahgiensis Mayr and Gilliard
Banded Land Rail
Gilliard, Gyldenstolpe, Shaw-Mayer, and Bulmer all found this
larger relative of Rallus pectoralis fairly common in swampy grass-
land in the Wahgi Valley, at Mt. Giluwe, and in Kyaka territory.
Rallus pectoralis and R. philippensis differ ecologically in that the
former prefers dry grassland, the latter swampy areas. I never encoun-
tered Rallus philippensis, and Awande natives appeared not to know
it, probably because the grassland at Awande was all dry.
Porzana pusilla palustris Gould
Marsh Crake
‘Three specimens collected at Mt. Giluwe by Shaw-Mayer provide
116
SPECIES ACCOUNTS
the only Eastern Highlands record. The species has been taken at only
one other locality in New Guinea, the Wissel Lakes (Junge, 1953).
Porzana tabuensis edwardi Gyldenstolpe
Spotless Crake
NATIVE NAME. Fore: oldlinta.
SPECIMENS EXAMINED. Awande: 1 ¢,1 9 (18-20 June 1965).
WEIGHT... 14: 47. IOs. 44.
WING. 1 4: 84. 1 Q: 81.
EXPOSED CULMEN. I ¢: 20. 1 9: 47.
TAXONOMY. ‘The Awande material belongs to edwardi, a thinly
differentiated race characterized by having the upperparts slightly
darker and the bill slightly longer than in nominate tabuensis or
richardsoni.
BREEDING. ‘The gonads of both specimens were enlarged.
DISCUSSION. Both specimens were trapped in grassland by na-
tives. The species was also collected by Gilliard, Gyldenstolpe, Shaw-
Mayer, and Bulmer and is fairly common but secretive in grassland,
mainly between 5,000 and 8,000 ft. It is absent at Karimui.
Poliolimnas cinereus leucophrys (Gould)
White-browed Crake
SPECIMENS EXAMINED. Awande: 1?
WING... 9.
TAXONOMY. ‘The race minimus is considered inseparable from
leucophrys.
DISCUSSION. Poliolimnas cinereus has a spotty distribution in the
New Guinea lowlands. The Awande specimen (6,000 ft), taken by
Paran in 1967, constitutes an altitudinal record, with the next highest
record being a specimen taken at 1,350 ft in the Astrolabe Range by
‘Tate after the Second Archbold Expedition. ‘The Awande record may
indicate a straggler rather than a resident population.
VOICE. A squeaky two-note call, the second note one-half second
after the first.
Rallicula forbesi cf. steint Rothschild
Forbes’s Chestnut Rail
NATIVE NAMES. Fore: ko-atna. Daribi: kéma.
SPECIMENS EXAMINED. Awande: 1 9 (20 June 1965). Mt. Michael: 1 ¢ (6
July 1964).
WEIGHT. I 9: SI.
WING. 1 @: 105; 1 oO: 107,
ACA, Des bGe BOs ‘On.
117
SPECIES ACCOUNTS
EXPOSED CULMEN. 1 4:28. 1 ©: 23,
STOMACH CONTENTS. Insects.
‘TAXONOMY. ‘These two specimens differ from nominate forbesi
of southeastern New Guinea in the much shorter tail, and in this
respect agree with steznz, originally described on the basis of a single
female from the Weyland Mountains in far western New Guinea. The
larger series obtained by Gilliard, Gyldenstolpe, and Shaw-Mayer all
showed great variation in plumage. In addition, specimens from Mt.
Giluwe had tails as long as those of nominate forbesi, while those from
various locations in the Wahgi Valley had short tails. The suggestion
of Sims (1956) that this variability of Rallicula forbesi in the Eastern
Highlands is due to recent reestablishment of breeding contact be-
tween the western race steini and the eastern race R. f. forbesi follow-
ing a period of isolation is a plausible one, since the details of
distribution in the genus Rallicula suggest that these rails have become
isolated, differentiated, and reinvaded each other’s ranges often (Dia-
mond, 1969).
DISCUSSION. Rallicula forbest goes about on the ground in
parties of a few individuals inside primary midmontane forest between
4,200 and 9,500 ft. It is secretive and uncommon but nevertheless
widespread, and has turned up at almost all forest collecting stations
in the Eastern Highlands within these altitudinal limits.
Rallina tricolor tricolor Gray
Red-necked Rail
SPECIMENS EXAMINED. Karimui: 1 9 (12 August 1964).
WING. 146.
TAXONOMY. ‘The specimen agrees with the New Guinea race
R. t. tricolor, and differs from the Australian race robinsoni (Cape
York Peninsula) in the darker brown belly and lower back and in the
fainter barring on the abdomen.
DISCUSSION. ‘The specimen provides the sole record for the
Eastern Highlands, and also represents an altitudinal record for this
lowland species, rarely found more than a few hundred feet above
sea level.
Amaurornis olivaceus ruficrissum (Gould)
Rufous-tailed Moorhen
NATIVE NAME. Daribi: kaba.
SPECIMENS EXAMINED. Karimui: 1 ¢, 3 9, 1 juv. ? (9 and 11 Aug. 1964;
1 July-6 Aug. 1965).
WEIGHT. 2.9% 175, leo.
WING, I @: 148. 3 9: 197, 139, 199,
EXPOSED CULMEN. 1 4: 5b. 3 Q: 28, 28, 50.
STOMACH CONTENTS. Insects.
118
SPECIES ACCOUNTS
TAXONOMY. A. o. ruficrissum of northern Australia and south-
ern New Guinea differs from A. 0. moluccanus of the remainder of
New Guinea in the lighter belly and more rufous, less dull undertail
coverts, and in that the male has a red base to the culmen, lacking in
moluccanus. The Karimui material is somewhat intermediate but
nearer ruficrissum in the color of the belly and undertail coverts; the
male has a red-orange base to the culmen. ‘The juvenile is uniformly
black.
BREEDING. All four adults had enlarged gonads. ‘This fact, plus
the record of the juvenile not more than a few days old taken in early
July, implies that breeding is in the dry season, as true for most grass-
land birds.
DISCUSSION. ‘This moorhen was present at Soliabeda as well as
at Karimui and was also collected by Schodde and Hitchcock at Lake
Kutubu. In the Karimui area and in other parts of New Guinea
where I encountered it, it lived in the tall grass that had grown up
around gardens and cleared land, remote from any swamp. It was far
more often heard than seen and proved impossible to flush; it merely
remained out of sight in the grass when disturbed.
VOICE. As described in detail elsewhere (Diamond and ‘Terborgh,
1968), the commonest vocalization in this species is a duet, in which
each bird of a pair alternately utters a cat-like wail lasting about a
second and beginning just as the wail of its partner concludes. During
the performance the birds move about in the tall dense grass, several
feet apart, and remain out of sight of each other. The duet is often
given at night as well as during the day. Solo calls consisting of a low
irregular, hen-like “chuck-chuck-chuck” or “kuk-kuk” or “bk-bk-bk”’
are heard infrequently.
Gallinula tenebrosa subsp.
Dusky Moorhen
In 1951 a European resident at Nondugl showed Gyldenstolpe
colored photographs of two moorhens which had been shot near the
Wahgi River and which Gyldenstolpe identified as this species. ‘There
is no other record from the Eastern Highlands and only a few scat-
tered records from other parts of New Guinea.
Porphyrio porphyrio subsp.
Purple Swamp Hen
Gilliard and Gyldenstolpe found evidence of the presence of swamp
hens in swampy grass near the Wahgi River, and Gilliard secured one
specimen. Bulmer obtained a sight record at Wabag (6,000 ft) in
January 1956. ‘The species occurs mainly in the lowlands but has also
119
SPECIES ACCOUNTS
been found at higher altitudes in western New Guinea (Wissel Lakes,
Baliem Valley).
CHARADRIIDAE: PLOVERS
Pluvialis dominica fulva (Gmelin)
Golden Plover
Gilliard, Gyldenstolpe, and Shaw-Mayer met small flocks at Nonduel
in March and April and in September and October. Bulmer noted
large flocks present for short periods of the austral summer in Kyaka
territory, and Bell recorded the species at Baiyer River in December
1965. ‘The Golden Plover is common in other parts of New Guinea in
migration and as a winter visitor.
SCOLOPACIDAE: SANDPIPERS, CURLEW,
anc ONIPE
Except for Scolopax saturata, all members of this family recorded in
New Guinea are nonbreeding visitors from the Palearctic and Nearctic.
Numenius minutus Gould
Little Whimbrel
Bell (1968) observed a flock at Baiyer River airstrip from 12 to 16
Dec. 1965.
Tringa hypoleucos Linnaeus!
Common Sandpiper
This is one of the most widespread wintering shorebirds in the
Eastern Highlands. ‘There are records from Nondugl in April and
October by Gilliard and by Gyldenstolpe; from Lake Kutubu in
September by Schodde and Hitchcock; from Mt. Hagen district and
Baiyer River in December by Bell; from Lanim River in September
by Bulmer; and from Baiyer River in December, January, April,
August, and October by Bulmer.
Tringa brevipes (Vieillot)?
Gray-tailed ‘Tattler
Observed by Schodde and Hitchcock at Lake Kutubu after 20 Sept.
1 Listed as Actitis hypoleucos in Rand and Gilliard (1967).
2 Listed as Heteroscelus incanus brevipes in Rand and Gilliard (1967),
120
SPECIES ACCOUNTS
Gallinago megala Swinhoe!
Marsh Snipe
Gyldenstolpe found large numbers suddenly appearing in the Non-
dugl grasslands for a few days in late September 1951.
Gallinago hardwicku (Gray)?
Japanese Snipe
Gyldenstolpe collected two specimens at Nondugl in September
1951. There is one older New Guinea record (Mt. Wilhelmina: Rand,
1942b) and several more recent ones. A few sight records of snipe of
this or the preceding species have been reported from elsewhere in the
Eastern Highlands.
Scolopax saturata rosenbergi Schlegel
East Indian Woodcock
Gilliard, Gyldenstolpe, Shaw-Mayer, and Bulmer found woodcock
uncommon in deep forest at high altitudes (8,500-10,000 ft) on several
peaks (Mt. Wilhelm, Mt. Giluwe, Lamende Range, Wahgi Divide
Mountains, Schrader Range).
Calidris ruficollis (Pallas)
Red-necked Stint
One was collected by Schodde and Hitchcock at Lake Kutubu on
30 Sept.
Calidris acuminata (Horsfield)4
Sharp-tailed Sandpiper
One specimen was collected by Schodde and Hitchcock at Lake
Kutubu on 30 Sept.
GLAREOLIDAE: PRATINCOLES
Glareola isabella Vieillot®
Australian Pratincole
SPECIMENS EXAMINED. Soliabeda: 1 9 (25 July 1965).
Listed as Capella megala in Rand and Gilliard (1967).
Listed as Capella hardwickii in Rand and Gilliard (1967).
Listed as Erolia ruficollis in Rand and Gilliard (1967).
Listed as Erolia acuminata in Rand and Gilliard (1967).
Listed as Stiltia isabella in Rand and Gilliard (1967).
Bem ow Ne
oO
121
SPECIES ACCOUNTS
WEIGHT. 48.
WING. 187.
DISCUSSION. ‘The bird collected had landed on bare ground
after circling Soliabeda village shortly after sunrise. Schodde and
Hitchcock regularly saw small groups of this Australian wintering
visitor on playing fields and the airstrip at Lake Kutubu until at least
4 Oct. Bell observed 20 at Tari airstrip on 4 Aug.
LARIDAE: GULUS and TERNS
Sterna fuscata subsp.
Sooty Tern
Single individuals were picked up at Wapenamanda and Mt. Hagen
on the morning of 29 June 1967, evidently members of a flock lost
while flying across New Guinea the previous night (N.G.B.S. News-
letter No: 23,7p. 2, sept. 196))):
COLUMBIDAE: PIGEONS
Diets of New Guinea Pigeons
‘The genera of New Guinea pigeons differ in diet. Ptilinopus,
Megaloprepia, and Ducula take only fruit whose flesh is soft enough
to crush between one’s fingers, and the stomachs of these genera never
contain gravel. ‘The size of fruit eaten varies with the size of the bird,
from 6 mm in diameter for the smallest Piilinopus to 50 mm for the
largest Ducula. In Columba, Macropygia, Reinwardtoena, Chalco-
phaps, Gallicolumba, Caloenas, and Otidiphaps the stomach often or
usually contains gravel, sand, or stones (up to 10 mm in diameter in
Caloenas and Otidiphaps). Correspondingly, the diet of these genera
consists of hard fruit, nuts, and seeds, occasionally supplemented by
insects in Chalcophaps and Gallicolumba. ‘The stomach wall of Calo-
enas is especially thick and muscular.
Niche Differences in Genus Ptilinopus
In the Papuan region there occur 13 species of Piilinopus, of which
nine range virtually over the entire island of New Guinea, one is con-
fined to the north coast (P. viridis), and three fail to reach, or else
barely reach, the main island of New Guinea. All are mainly green in
color, medium-sized to small, arboreal, and live on soft fruit. ‘The
vocalizations of all 10 species with which I am familiar are a confusingly
similar series of low ‘“‘coo’s’” or “‘hoo’s”. Above 5,000 ft one encounters
only one species (P. rivoli) or occasionally two (P. rivoli plus P.
ee
SPECIES ACCOUNTS
ornatus). At lower elevations it is the rule to find several species locally
sympatric. For instance, combining results from the three prewar
Archbold Expeditions and my three expeditions, species lists were avail-
able for a total of 29 localities at elevations of 4,000 ft or less. At three
localities, six Ptilinopus species were present; at four localities, five
species; at five localities, four species; at nine localities, three species;
at seven localities, two species; and at one locality, one species. Several
species are regularly seen in a single fruit tree. ‘Vable 4 indicates the
species present at each of my eight best-studied collecting localities in
the Eastern Highlands.
Size differences are a major factor that make this degree of local
sympatry possible. ‘Thus, approximate average weights of the 10 New
Guinea mainland species are: P. nanus, 50; P. pulchellus, 75; P. coro-
miulaius, 75; i, iozonus, 110; P. superbus, 125; P. viridis, 125; FP.
aurantifrons, 140; P. rivoli, 155; P. ornatus, 165; and P. perlatus, 240.
The five species that occur regularly at Karimui (P. nanus, P. pulchel-
lus, P. superbus, P. ornatus, P. perlatus) have weights that successively
increase by a factor of about 1.5 (50, 75, 125, 165, 240). "wo small
species with the same weight, P. pulchellus and P. coronulatus, have
locally complementary distributions so that both rarely occur at the
same locality. Other ecological differences are that P. ornatus and P.
perlatus have altitudinal distributions which largely exclude each
other; that the local distributions of P. ornatus and P. superbus in hill
forest (below P. rivoli, above most of the other species) tend to be
complementary; that P. 7ozonus and P. perlatus are more characteristic
of second-growth and the forest edge than the other species; and that
P. suwperbus had a synchronized peak of breeding in my study areas at
a time when all its congeners at the same elevation were in nonbreed-
ing condition. However, much remains to be learned about ecological
sorting within this genus.
TABLE 4
LocAL DISTRIBUTION OF SPECIES OF Ptilinopus Fruir Doves!
P. es P: PF: BP. FP:
superbus pulchellus rivoli nanus ornatus perlatus
Awande (6,000 ft) x
Mt. Michael
(ca. 8,000 ft) x
Mt. Karimui
(5,000-7,000 ft) x »
Okasa (ca. 4,000 ft) x x (rare) x
Sena River (4,500 ft) x
Karimui (3,650 ft) x x x (rare) x x x
Bomai (3,250 ft) x x (rare)
Soliabeda (2,000 ft) x <x x
1 The table indicates which Ptilinopus species were present at each of my eight best-studied
collecting localities in the Eastern Highlands.
123
SPECIES ACCOUNTS
Ptilinopus swperbus superbus (Temminck)
Superb Fruit Dove
NATIVE NAME. Daribi: ai.
SPECIMENS EXAMINED. Karimui: 2 ¢ (29 July 1964; 3 July 1965). Okasa:
7 S$ (22-25 June 1965).
WEICHT. 6 @: D4 119; 122 104 128. 132°
WING. 4 @: 123, 127, 130, 135.
STOMACH CONTENTS. Fruit, 7-26 mm. in diameter.
BREEDING. All of the specimens were males, and all had greatly
enlarged gonads. One female was observed at Karimui and another at
the Sena River but none at Okasa, where males were particularly
numerous. Perhaps the females at Okasa were remaining out of sight
on nests, unless the males arrive before the females.
At the Sena River in 1964 I found a nest 3 ft off the ground in a low
bush 30 ft from the river. It was a flimsy stick construction containing
one white egg measuring 29 x 18 mm. This nest and the enlarged
testes of the specimens suggest that the peak of breeding occurred in
the dry season at Okasa, Karimui, and the Sena River both in 1964
and 1965. Most other fruit eating species were not breeding at this
time (p. 85).
DISCUSSION. Ptilinopus superbus was the commonest Ptilinopus
at Okasa and the only one on the Sena River, but was quite uncommon
at Karimui and absent at Soliabeda and on Mt. Karimui. In general,
I have found P. superbus characteristic of forest on hill slopes up to
about 4,500 ft, largely below the altitudinal range of P. rivoli. Its
main competitor on the hill slopes is P. ornatus, which is more rigidly
tied to this habitat but is usually found in localities where P. superbus
is absent or uncommon. P. superbus occurs irregularly in second-
erowth and agricultural areas up to about 5,000 ft. It also extends to
sea level, where it comes into competition with other Ptilinopus
species.
The male displays to the female while standing on a branch, by
puffing up his breast feathers and orange collar, spreading the wings,
flaring the tail to show the light terminal band, and uttering a rasping
sound.
VOICE. ‘The distinctive slow call consists of a single note with an
‘“m’” sound, followed after a 2-sec pause by a series of three to six
identical upslurs at I-sec intervals.
Ptilinopus pulchellus pulchellus (Yemminck)
Beautiful Fruit Dove
SPECIMENS EXAMINED. Karimui: 2 4, 1 ? (29 July 1964; 1 and 17 July 1965).
Bomai: 1 ¢ (7 July 1965). Soliabeda: 4 ¢ (22-27 July 1965). Mt. Karimui Zone
124
SPECIES ACCOUNTS
ol Ss (ll Aue. 1965); Zone 2:1 ¢ (14 Aug. 1965).
WEIGHT. 8 4: 55-85 (76 + 11).
WING. 9 @: 105-109 (108 + 2).
STOMACH CONTENTS. Fruit, 7-16 mm in diameter.
TAXONOMY. Of the nine specimens sexed, seven had the testes
small, while two (Mt. Karimui Zone 2 and Karimui 17 July 1965)
had the testes somewhat enlarged. It is remarkable that only males
were collected (females are indistinguishable from males in the field).
DISCUSSION. Ptilinopus pulchellus was fairly common at Karimui
and Soliabeda in trees of forest, second-growth, and gardens, and
Schodde and Hitchcock also found it at Lake Kutubu. It is generally
distributed elsewhere in New Guinea in forest up to about 2,000 ft,
occasionally (as at Karimui) somewhat higher. Its ability to coexist
with several other species of Ptilinopus in this range is probably as-
sociated with its smaller size.
One other New Guinea species of Ptilinopus, P. coronulatus, has
approximately the same size (weight and wing length) as P. pulchellus.
‘The local distributions of these two species complement each other to
a striking degree. Thus, on the southern slopes of the Eastern High-
lands and on the upper Fly River P. pulchellus is present, P. coronu-
latus absent, while on the northern slopes of the Eastern Highlands,
on the Sepik River, and on the lower Fly River P. coronulatus 1s
present, P. pulchellus absent. Out of 23 low-altitude collecting stations
of the three prewar Archbold Expeditions, 13 supported only P.
coronulatus, six only P. pulchellus, and only four supported both
species. On the Vogelkop the Denison-Crockett expedition took P.
pulchellus at three localities, took P. coronulatus at five different locali-
ties, and found no locality supporting both species, although all these
camps were within 100 miles of each other and most were within 10
miles. There must be some common denominator to the localities
supporting one species rather than the other, but I cannot discern
what it is.
Ptilinopus coronulatus subsp.
Little Coroneted Fruit Dove
In Baiyer and Lai Gorges on the northern slopes of the Eastern
Highlands, Bulmer collected three specimens of this species at 2,000-
3,000 ft in October 1955. As discussed under the previous species, its
local distribution largely complements that of the similar sized P.
pulchellus.
Ptilinopus rivoli bellus Sclater
White-breasted Fruit Dove
NATIVE NAMES. Fore: tubi. Gimi: tibi. Daribi: birisigi.
125
SPECIES ACCOUNTS
SPECIMENS EXAMINED. Awande: 1 ¢ (26 June 1964). Okasa: 1 ¢ (23 June
1965). Mit. Michael: 1 4, 2 ©, 1 imm, 6 (4-8 July 1964). Karimui: 1 imm. ¢,
1 9 (3 and 5 Aug. 1965). Mt. Karimui Zone 3: 3 9; Zone 4: 1 @,1 9 (16-28 Aug.
1965).
WEIGHT. 3. @: 134, 145, 188. 5 O: 139° 141, 142, 152) 166.
WING. 2 @: I43, 148. 4 ©: 133, 134, 136, 137:
STOMACH CONTENTS. Fruit, 6-20 mm in diameter.
BREEDING. ‘The Okasa male, and one male and one female from
Mt. Karimui Zone 4, had enlarged gonads. The nest is a flimsy stick
construction with one egg.
DISCUSSION. ‘This is the characteristic Ptilinopus of mountain
forest from about 4,500 to 8,000 ft, occasionally to 9,000 ft, and was
fairly common at all my montane collecting stations. In addition, I
obtained four records from lower elevations: one collected at 4,200 ft
at Okasa, two collected at Karimui (3,650 ft), and an adult male seen
at Bomai (3,250 ft). In most of its altitudinal range P. rivoli is the sole
species of its genus except for occasional localities where P. ornatus
is present. Only the lowest part of its normal range is shared with
P. superbus. The few individuals that straggle below 4,000 ft (e.g., to
Karimui and Bomai) are greatly outnumbered by several species of
congeners at these localities.
Single individuals or pairs frequent the middle- and upperstories
of the forest interior, infrequently venturing into trees of second-
growth. Up to four birds may be seen simultaneously in a fruit tree.
When perched motionless and silently, which is to say most of the time,
their green plumage renders them hard to locate in the leafy crowns of
tall trees.
VOICE. A series of upslurred “hoo’s,” falling in pitch and greatly
accelerating as the series progresses. Other species of Ptilinopus (P.
perlatus, P. aurantiifrons, P. iozonus) have calls to which this same
description applies, but in no other species is the acceleration so
marked. Also, softer and slower single “hoo’s” or pairs of “hoo’s,”’ each
“hoo” initially rising, then usually falling, in pitch.
Ptilinopus nanus nanus (Temminck)
Least Fruit Dove
SPECIMENS EXAMINED. Soliabeda: 1 9 (25 July 1965).
WEIGHT. 48.5.
WING. 8).
TAXONOMY. The length of the wing agrees with that of the few
other available females of P. n. nanus, the New Guinea race (84-88
mm). Females of minimus from the western Papuan islands have a
wing length of 80-81 mm.
DISCUSSION. On two occasions I observed one or several indi-
viduals in a tall fruit tree also frequented by P. ornatus and Megalo-
126
SPECIES ACCOUNTS
prepia magnifica in the forest at Soliabeda. I also saw a pair in a tall
tree at the edge of the forest at Karimui on II Sept. 1965. ‘There are
no other Eastern Highlands records. ‘his is the smallest and rarest
wide-ranging New Guinea Ptilinopus, known only from a handful of
scattered localities and specimens in the lowlands.
VOICE. A slow, repeated, disyllabic upslur “koo-uh.”
Ptilinopus ornatus kaporensis Rothschild and Hartert
Ornate Fruit Dove
NATIVE NAME. Daribi: buma.
SPECIMENS EXAMINED. Soliabeda: 3 ¢,1 Q (25 and 26 July 1965). Karimui:
4 @,1 9,1? @ and 3 Aug. 1965).
WEIGH. 7 22 142, 157, 162, 166; 169, 178,185. 2 > lol, lob. 1: 149.
WING. 6 @: 144, 145, 151, 151, 154, 155. 1 9: 146. 1 ?: 149.
STOMACH CONTENTS. Fruit, 13-19 mm in diameter.
TAXONOMY. Racial differences in populations of this species
from the main body of New Guinea (exclusive of the Vogelkop) form
a cline around the periphery of the island, broken in the west at the
Vogelkop, where the nominate race ornatus is quite distinct in pos-
sessing a purple crown. The opposite extremes in this cline fall in
northern New Guinea (assigned to P. 0. gestrot) and in southwestern
New Guinea (P. 0. kaporensis), birds from the southeast (topotypical
gestrol) being intermediate but closer to the northern populations.
The distinguishing characters are the color of the breast (darker and
browner in kaporensis, lighter and more orange in gestrot) and the
color of the band on the upper back and neck (darker, browner, and
more richly colored, nearly chocolate, in kaporensis; lighter and more
olive suffused with orange in gestrot). Material available for compari-
son included gestroi from the northern slope of the Snow Mountains,
topotypical gestroi from southeastern New Guinea, and kaporensis
from the upper Fly River and from Mt. Goliath (southern slope of
Snow Mountains) but no topotypical kaporensis (Onin Peninsula,
southwestern New Guinea). ‘The specimens from the southern slope
of the Eastern Highlands, which geographically are intermediate be-
tween southeastern New Guinea and the Fly River or Mt. Goliath,
have the breast darker and browner than any of the comparative ma-
terial, and this probably means darker and browner even than the
type of kaporensis, which Rand (1942b) found to compare well with
the Fly River specimen and most of the Mt. Goliath specimens. In the
color of the band on the back and neck, however, the Eastern High-
lands birds agree well with Mt. Goliath and Fly River kaporensis.
The afhnities of the Eastern Highlands birds are thus with kaporensis
to the west, not with gestroz to the east.
BREEDING. ‘The gonads of all specimens were small.
DISCUSSION. ‘The characteristic habitat of P. ornatwus is forest
rad
SPECIES ACCOUNTS
of the hill slopes up to 4,500 ft, ranging down occasionally to the base
of the hills and infrequently reaching the flat country at sea level. The
slightly smaller P. swperbus is also characteristic of hill forest but, in
contrast to P. ornatus, is also found frequently in the flat lowlands.
The distributions of P. superbus and P. ornatus in the hill forest may
to some extent complement each other, as do those of the smaller P.
pulchellus and P. coronulatus in the lowlands. For instance, P. ornatus
was common at Soliabeda and Karimui, quite uncommon at Okasa,
and absent at the Sena River, while P. superbus was absent at Solia-
beda, quite uncommon at Karimui, and common at the Sena River
and Okasa.
In addition, I saw P. ornatus once at 6,500 ft in moss forest on Mt.
Karimui, separated by 3,000 vertical feet from the population on the
floor of the Karimui Basin. There are at least five other instances of
detached populations of P. ornatus at altitudes well above its usual
upper limit (ca. 4,500 ft): 7,100 ft in heavily mossed forest on the
northern slope of the Snow Mountains (Rand, 1942b), 5,200 ft in moss
forest on Mt. Menawa in the North Coastal Range (my 1966 expedi-
tion), Bijenkorf on the southern slope of the Snow Mountains (Junge,
1937), and the camps of Lord Rothschild’s collector, A. S. Meek, on Mt.
Goliath in the Snow Mountains and on the upper Angabunga River
in southeastern New Guinea. ‘The only other species of Ptilinopus at
all six of these high-altitude localities was P. rivoli, which weighs ap-
proximately the same as P. ornatus. How P. ornatus and P. rivoli sort
out ecologically in these high-altitude areas of local sympatry is un-
known.
VOICE. A low, repeated, soft “mm.”
Ptilinopus perlatus zonurus Salvadori
Pink-spotted Fruit Dove
SPECIMENS EXAMINED. Karimui: 4 ¢, 3 9 (2-5 Aug. 1965).
WEIGHT. All specimens weighed more than 210 g.
WING. 3°: 154% 157, 157. 3 9: 150; 152; 155.
BREEDING. All specimens had small gonads.
DISCUSSION. At Karimui this species and its smaller relative P.
ornatus were the two commonest species of Ptilinopus and fed together
in the same fruit trees. The two species are very similar in pattern and
are more closely related to each other than either is to any other
species. However, their typical altitudinal and habitat preferences are
totally distinct, and their occurrence together at Karimui is excep-
tional. Whereas P. ornatus is usually found on the hill slopes and rarely
reaches sea level, P. perlatus is usually confined to the lowlands, and
Karimui represents an altitudinal record for it. At the lower elevation
of Lake Kutubu (2,450 ft) Schodde and Hitchcock found P. perlatus
128
SPECIES ACCOUNTS
very common but did not obtain P. ornatus. In my experience, P.
perlatus is characteristic of second-growth trees and the forest edge
rather than the interior of primary forest. Its occurrence in the low-
lands is quite local, and it must be considered the least common wide-
spread New Guinea Ptilinopus, except for P. nanus.
VOICE. A series of low, downslurred “hoo’s,”’ delivered at a rate
slightly less than one per second. The rate is either constant, or else
accelerates slightly but by much less than does the similar call of P,
rivoli. Also, a very soft single “hoo” dropping in pitch at the end, with
or without an initial rise in pitch.
Megaloprepia magnifica interposita Hartert
Magnificent Fruit Dove or Wompoo Pigeon
NATIVE NAME. Daribi: somagoi.
SPECIMENS EXAMINED. Karimui: 5 ¢, 5 9Q (10 July, 15 July, 3-5 Aug., 11
Sept. 1965); 1 ? (15 Aug. 1964). Soliabeda: 1 ¢, 1 9 (22 and 28 July 1965).
WEIGHT. 6 4: 180, 192, 201, +210, +210, +210. 5 9: 175, 177, 182,
189. 210.
WING. 6 4: 163-175 (167 +5). 6 Q: 158-171 (163 + 4).
STOMACH CONTENTS. Fruit, 9-22 mm in diameter.
TAXONOMY. ‘The races to which this Eastern Highlands material
is related are poliura (southeastern New Guinea), assimilis (Cape York
Peninsula, Australia), and interposita (type locality the Wandammen
Peninsula of northwestern New Guinea, also southwestern New Guinea
from the Onin Peninsula to the Fly River). In the deep, bright yellow
color of the undertail coverts the Eastern Highlands specimens re-
semble the type of interposita and Fly River birds, whereas the under-
tail coverts of poliwra from southeastern New Guinea are paler and
duller. The wing spots of poliwra are brighter yellow, those of inter-
posita from the Wandammen Peninsula, Weyland Mountains, and
southern slopes of the Snow Mountains are more whitish; Fly River
and Eastern Highlands birds are intermediate in this respect. ‘The
breast is lightest and reddest in poliura, deeper and more purple in
interposita from southwestern New Guinea and the type of interposita,
and still deeper and more purple in Fly River and Eastern Highlands
birds. In Australia Megaloprepia magnifica becomes more purple
ventrally and larger as one proceeds from the Cape York Peninsula
(assimilis) through northern Queensland (ker?) to southern Queensland
and New South Wales (magnifica). The race assimilis is close to Fly
River and Eastern Highlands birds in the purple color of the breast
but is larger. My measurements of the wing yield 4 ¢ 172-188
(is2 == 7), 9 9 I/G-X6l (E79 = 3) tor assimilis, whereas Fly River
birds are the same size as Eastern Highlands birds: 5 g 162-173
C6225), 5 § 158-162 (162 26 4),
In explanation of these characters, it may be recalled that lower sea
levels during parts of the Pleistocene created a land bridge, now
129
SPECIES ACCOUNTS
covered by the shallow Torres Straits, between the Cape York Peninsula
of Australia and the Fly River bulge of southern New Guinea. The
distributions of numerous New Guinea birds and mammals attest to
the earlier existence of this bridge (Tate, 1952; Keast, 1961). Across
this bridge the New Guinea and Australian populations of Megalo-
prepia would have been continuous, and the Fly River bulge would
have been a meeting point not only of poliwra and interposita but of
assimilis as well. It seems unwise to dismember these clines further,
and the Eastern Highlands population is therefore best assigned to
interposita, since it agrees most closely with this race. The details are
however of zoogeographical interest and may be briefly summarized:
Fly River and Eastern Highlands birds are rather different from the
southeastern race poliwra and stand closest to the western race inter-
posita, from which they deviate in having a darker, more purple breast
(assimilis genes) and yellower wing spots (poliwra genes).
BREEDING. ‘T'wo specimens taken on 11 Sept. 1965, both males,
had enlarged testes, and one female, taken on 4 Aug. 1965, had some-
what enlarged ovaries. ‘The other nine specimens sexed (five taken in
July, four in early August 1965) had small gonads. This seems to
suggest breeding recommencing at the end of the dry season. ‘The nest
is a flimsy stick construction on a branch 6-15 ft above the ground,
with one white egg.
DISCUSSION. At Karimui and Soliabeda Megaloprepia was less
common than the commoner species of Ptilinopus, with which it
shared use of fruit trees in the shaded forest interior while avoiding
trees of the forest edge and second-growth. It remained within the
shaded lower half of the crowns, whereas species of Ptilinopus often
chose exposed perches at the tops of trees after feeding. In addition,
we heard it once on the lowest part of Mt. Karimui (4,000 ft). It is
basically a lowland species, and the only other Eastern Highlands
records are from Lake Kutubu (2,450 ft).
VOICE. ‘Three deep, equally-spaced notes on virtually the same
pitch, the whole call lasting about | sec. So well are the sounds ap-
proximated by the syllables “hoo-wa-hoo” that I initially assumed that
a man was calling in the forest, and it did not even occur to me to
consider the possibility that it might be a bird.
Ducula pinon jobiensis (Schlegel)
Pinon Imperial Pigeon
DISCUSSION. Bulmer collected one specimen of this lowland
species at 2,000-3,000 ft in the Baiyer Gorge.
VOICE. Very low-pitched notes, in either of the two patterns de-
picted in Figure 13 (p. 133).
130
SPECIES ACCOUNTS
Ducula rufigaster rufigaster (Quoy and Gaimard)
Purple-tailed Imperial Pigeon
SPECIMENS EXAMINED. Karimui: 1 ? (30 July 1964). Soliabeda: 1 4 (30
July 1965).
WING. Tas 198, «es 202;
STOMACH CONTENTS, Fruit.
TAXONOMY. The race pallida was separated on the grounds
that “birds from the south slopes of the Oranje Mountains differ
from all other populations by the decidedly paler breast and abdomen”
(Junge, 1952, p. 248) and was also characterized as larger than nomi-
nate rufigaster (Rand and Gilliard, 1968, p. 173). Five specimens col-
lected by Meek on the Setekwa River, 50 miles west of the type
locality of pallida (Noord River), do not appear to me paler ventrally
than other New Guinea populations. The average wing lengths of 11
populations from various parts of New Guinea, including the popula-
tion from the Setekwa River, all fall between 192 and 199 mm, with
a spread of 10 to 25 mm in each population. ‘Thus, the distinctness of
pallida from rufigaster seems doubtful.
BREEDING. ‘The testes of the male were small.
DISCUSSION. At Karimui and Soliabeda this pigeon was quite
uncommon, solitary, and usually seen perched motionless in the mid-
dlestory of the shaded forest interior, occasionally at the forest edge.
Ducula chalconota smaragdina Mayr
Red-breasted Imperial Pigeon
NATIVE NAMES. North Fore: kintanamu. South Fore: mimpi. Gimi: mipi.
SPECIMENS EXAMINED. Miarosa: 1 9 (17 June 1964). Awande: 1 ¢ (19
June 1965). Mt. Michael: 1 ¢,2 Q (2-12 July 1964).
WHING e224 3) OOO R 2 0b e 20:
STOMACH CONTENTS. Fruit.
TAXONOMY. ‘These specimens agree with the race smaragdina
from the main body of New Guinea, which differs from D. c. chal-
conota of the Vogelkop in having a longer wing and a greener, less
purple back. ‘There is some individual variation in the back color,
which is particularly green in the Miarosa female, more purple in the
Awande male.
DISCUSSION. Ducula chalconota, the largest montane pigeon, is
the high altitude representative of D. rufigaster, occurring uncom-
monly and as solitary individuals in the middle- and upperstory of
forest at about 6,000-8,500 ft. I observed it on Mt. Michael and in the
Okapa area, while Gilliard and Bulmer collected it on Mt. Hagen. It
was absent on Mt. Karimul.
VOICE. A low quiet, unhurried note “hmmm,” either constant in
131
SPECIES ACCOUNTS
pitch or else upward inflected, given either twice in succession or just
once.
Ducula zoeae (Lesson)
Zoe Imperial Pigeon
NATIVE NAME. Daribi: hee.
SPECIMENS EXAMINED. Karimui: 6 ¢, 3 9 (11 and 13 Aug. 1964; 9 July-5
Aug. 1965). Soliabeda: 1 ¢,1 Q (21 July 1965).
WING. © (62213, 216,218, 220; 7220; 221. 49-1214, 290. 993 295,
STOMACH CONTENTS. Fruit, 19-45 mm in diameter.
BREEDING. The 11 Aug. and 13 Aug. males had enlarged testes,
and the 5 Aug. female had an enlarged ovary, while the July specimens
had small gonads. ‘Thus, breeding may have been commencing towards
the end of the dry season, as with Megaloprepia magnifica.
DISCUSSION. At all my low-altitude stations up to a ceiling vary-
ing locally between 4,000 and 4,700 ft, Ducula zoeae was a common
species whose characteristic calls were heard daily. Most of its altitudi-
nal range, which extends down to sea level, is shared with its less
common congener D. rufigaster, but D. rufigaster does not extend to
quite so high altitudes as does D. zoeae. Other records of D. zoeae in
the Eastern Highlands are at Lake Kutubu (2,450 ft) and at 4,000 ft
in the Baiyer Valley.
D. zoeae was usually seen singly or in small numbers, from the
middlestory up to the crowns of the tallest trees. Unlike D. rufigaster
and D. chalconota it seemed equally at home at the edge of the forest
as in the forest interior. ‘The flight is heavy, and the wing beat slow.
VOICE. A low but resonant series of broken “hoo’s.” Each “hoo”
rises slightly in pitch, lasts somewhat more than I sec, and is broken
into usually five separate notes: hoo, h-h-h-h-hoo, h-h-h-h-hoo, h-b-h-h-
hoo etc. (rie. 13):
Gymnophaps albertisi albertisi Salvadori
D’Albertis’ Pigeon
NATIVE NAME. Fore: pupunte. Daribi: buruma.
SPECIMENS EXAMINED. Okasa: 3 ¢ (23 and 24 June 1965). Mt. Michael: 1
Q (4 July 1964). Karimui: 1 ¢,1 9 (6 Aug. 1965). Soliabeda: 1 9 (22 July 1965).
WING. 4. 4: 180, 187, 197, 204 1 9: 19L
BREEDING. The Okasa specimens had enlarged gonads, while
those from other localities did not.
DISCUSSION. D’Albertis’ Pigeon is widely but erratically dis-
tributed in the Eastern Highlands, and was present at all my collecting
stations except Mengino and Bomai, ranging up to 11,000 ft in stunted
moss forest on the summit ridge of Mt. Michael. In the vicinity of
mountains it sometimes descends to sea level, but it is usually absent
N54
SPECIES ACCOUNTS
Ducula zoeae:
—_ — _ —— ———= etc.
hoo — h-h-h-h-hoo:_h-b-h-h-hoo h-h-h-h-hoo:_b-h-h-h-hoo
2 sec
Ducula pinon:
: era
Fic. 13. Voices of Ducula zoeae and D. pinon.
from the flat lowlands. In contrast to most other pigeons in the hills
and mountains of New Guinea, this is a social bird par excellence. I
rarely saw a solitary individual, occasionally met pairs, but usually
found it in groups of three to 30, the average size being about a
dozen. Flocks perch in tall trees of the forest interior, and if disturbed,
swiftly circle high over the forest before realighting. In fruit trees
they keep to themselves and do not mix with other species of pigeons.
A characteristic habit or display is for flocks to fly down a mountain
at high speeds along the crest of a steep ridge, dropping thousands of
feet in a single dive.
VOIGE. LUnever heard a vocalization.
Columba vitiensis halmaheira (Bonaparte)
White-throated Pigeon
In New Guinea this appears to be a scarce species; the only Eastern
Highlands record comes from Nondugl, where Gyldenstolpe collected
one specimen from a small flock.
Macropygia amboinensis kerstingi Reichenow
= Cc
Amboina Cuckoo-dove
NATIVE NAMES. Fore: kubétya. Gimi: kubdfa. Daribi: bube.
SPECIMENS EXAMINED. Awande: 2 9 (16 and 18 June 1965). Okasa: 3 @, 3
133
SPECIES ACCOUNTS
Q (23-25 June 1965). Mt. Michael: 1 @ (13 July 1964). Karimui: 3 ¢,1 9? (9
July-3 Aug. 1965). Soliabeda: 1 4,2 ©, 1? (21-25 July 1965). Mt. Karimui Zone
2 6 (O-t) Aus. 1965),
WEIGHT. 9 4: 131-179 (149+ 17). 8 9: 107-158 (136 + 17).
WING. 7 ¢: 158-170 (167+ 4). 7 Q: 143-171 (161 + 10).
STOMACH CONTENTS. Gravel and ground-up fruit.
TAXONOMY. As previous observers have noted, series of this
species from the same locality show great individual variation. ‘The
legs are bright purple, bright red-purple, bright carmine, or bright
pink in most adult males, dull dark brick-brown in one adult male
and in immature males, and dull purple, dull brick-red, or dull dark
brick-brown in females. The bill is pale flesh, light horn-colored, dark
horn-colored, gray-brown, dark gray, or completely black, and this
variation does not correlate with sex or other characters. ‘Ihe amount
of barring on the back, neck, cheeks, and breast of adult females and
immature males is variable. ‘There is no consistent variation in size or
in weight with altitude. This large and inadequately understood in-
dividual variability reinforces the doubts expressed by previous au-
thors (Gyldenstolpe, 1955; Gilliard and LeCroy, 1961) whether the
race kerstingi can be maintained as distinct from cinereiceps. When
comparison is restricted to adult males, specimens from northern New
Guinea (kerstingi) have slightly darker backs than material from the
D’Entrecasteaux Archipelago, the type locality of cinereiceps. My East-
ern Highlands series is nearer kerstingt, less dark dorsally, but darker
ventrally, as Rand (1942a) noted for Fly River kersting:. Southeastern
New Guinea specimens are nearer cinereiceps.
BREEDING. Gonads were small in most specimens, except that
they were slightly enlarged in two females and one male from Okasa,
and in one male each from Karimui and Mt. Karimui.
DISCUSSION. Macropygia amboinensis occurred at all of my
Eastern Highlands collecting stations except Bomai and the higher
elevations on Mts. Michael and Karimui, and has also been collected
by most other workers in the Eastern Highlands. ‘Vhe ecological dif-
ferenecs between M. amboinensis and M. nigrirostris are complex. Both
species may be found from the mountains down to sea level, and the
habitat of both has been recorded as forest or second-growth, depend-
ing upon the locality. ‘Their local distributions tend to be, but are not
strictly, complementary, such that only one of the two is common at a
particular location. Detailed consideration of my distributional records
and the published records of others suggests the following correlations:
(1) The most clear-cut difference is that there are practically no records
of M. amboinensis in deep primary forest above about 5,000 ft,
whereas M. nigrirostris occurs in forest up to altitudes varying between
7,000 and 9,000 ft, frequently into the mossy zone. For instance, M.
nigrirostris is the only species in forest above 5,000 ft on Mt. Karimui
in my experience and on Mts. Wilhelm, Kubor, and Hagen (Mayr and
134
SPECIES ACCOUNTS
Gilliard, 1954). (2) M. amboinensis is the dominant species in trees of
open country cleared by man (native gardens, isolated casuarina stands
in grassland, edge of the forest) above 5,000 ft. ‘This is the case at
Miarosa and the base of Mt. Michael (present study) in the Wahgi
Valley of the Eastern Highlands (Mayr and Gilliard, 1954), and in the
Baliem Valley of western New Guinea (Rand, 1942b). ‘hus, like many
other lowland species, M. amboinensis has been able to spread upward
into areas cleared by man. (3) Below 5,000 ft M. amboinensis is the
dominant species of the forest, as at Okasa, Soliabeda, the lower slopes
of Mt. Karimui up to 4,500 ft, and many other localities. ‘Vhe exclusion
of M. nigrirostris from the forest at low altitudes is certainly not com-
plete, but where M. nigrirostris is common below 5,000 ft, it seems
generally to be in second-growth, as around Karimui station (3,650 ft)
and at Lake Kutubu (2,450 ft). The size difference may play a role in
making possible the local sympatry at lower elevations, since M. am-
boinensis is about 1.7 times as heavy as M. nigrirostris.
M. amboinensis was generally seen as solitary individuals or pairs
in the middlestory of trees, not in the crown. Some practice is re-
quired to distinguish it in the field from M. nigrirostris. ‘The green-
naped, light-breasted males of M. amboinensis are easily identified in
good light. Females and immatures are more difficult to recognize, but
M. nigrirostris is smaller and its brown of a richer hue. The vocaliza-
tions of the two species are very distinct.
VOICE. A series of doubled-noted hoots delivered at the rate of
one to two per second. ‘The two notes of each hoot are connected;
the second note is about a major third in pitch higher than the first
and is accented. Successive hoots do not vary at all in pitch (Fig. 14).
Some calls of Reinwardtoena reinwardtsi are nearly indistinguishable.
Macropygia nigrirostris nigrivostris Salvadori
Black-billed Cuckoo-dove
NATIVE NAMES. Fore: piripiri. Daribi: bube.
SPECIMENS EXAMINED. Awande: 3 6, 2 9 (16-18 June 1965). Okasa: 1 9
(24 June 1965). Karimui: 7 ¢, 3 9 (3 July-5 Aug. 1965). Mt. Karimui Zone 1: 2
6 (i! and I2 Aug. 1965); Zone 2: 1 4 (5 Sept. 1965).
WEIGHT. 13 ¢: 68-104 (91411). 6 2: 66-104 (88 + 14).
WING. 8 4: 139-153 (145-45). 4 9: 144, 145 (3).
STOMACH CONTENTS. Gravel and ground-up fruit.
TAXONOMY. While both Macropygia amboinensis and M. nigri-
rostris have quite variable plumage, two features by which they may
be distinguished in any plumage are that the uppertail is always barred
in M. nigrirostris, never in M. amboinensis; and that the bill of M.
nigrirostris is stouter.
In the present series the pattern of variation is as follows. Adult
males have the dorsal surface of the tail barred, the lower back some-
135
SPECIES ACCOUNTS
Macropygia amboinensis:
AP Sl lt See
ae sec
Fic. 14. Voice of Macropygia amboinensis, and one of the two common calls of
Reinwardtoena reinwardtsi (virtually indistinguishable from call of Macropygia
amboinensis).
times faintly barred, the remainder of the plumage totally unbarred.
Adult females have not only the uppertail but also the whole back
and breast barred. Barring in immatures of both sexes is distributed
as in the adult female but is heavier, and Gilliard and LeCroy, (1967,
p. 194) have pointed out detailed differences in barring patterns be-
tween adult females and immatures. The legs are bright pink or bright
reddish purple in adult males, dull brick-red in adult females and
immatures. ‘The bill is black in adult males, a horn-color of varying
depth in adult females and immatures.
BREEDING. ‘The gonads were enlarged in two of the three Mt.
Karimui specimens but in no other material. ~
DISCUSSION. ‘The ecological differences between this species and
M. amboinensis have been discussed under M. amboinensis. In the
ecologically disturbed area around Karimui Patrol Post (3,650 ft)
M. nigrirostris was commoner than M. amboinensis. It was absent
from the primary forest of Mt. Karimui between 4,500 and 6,500 ft
but reappeared in the moss forest at 6,500-7,600 ft.
VOICE. Quite different from M. amboinensis, and more reminis-
cent of Ptilinopus: a quiet series of about a dozen muted “‘coo’s’’ at
the rate of three or four per second, initially rising and then dropping
in pitch, slightly decreasing in volume, and with no change in tempo
(unlike most species of Ptilinopus, which accelerate).
Reinwardtoena reinwardatsi gviseotincta Hartert
Great Cuckoo-dove
NATIVE NAMES. Fore: ébiye. Gimi: yukiba. Daribi: yuguri.
SPECIMENS EXAMINED. Karimui: 1 ? (12 July 1965). Bomai: 1 9 (6 July
1965). Mt. Karimui Zone 1: 1 9 (9 Aug. 1965); Zone 3: 1 g (16 Aug. 1965).
WEIGHT. 1.208. 1 4,1 oO, Torr S210;
WING. 1 4: 242. 2 9: 214, 236. 1 ?: 228.
BREEDING. ‘The gonads of all specimens were small. A male
was collected on a nest of sticks 15 ft above the ground, containing a
single white egg measuring 40 25 mm.
136
SPECIES ACCOUNTS
DISCUSSION. ‘This species is fairly widespread at almost all ele-
vations in the Eastern Highlands but is everywhere uncommon. A
sighting in Zone 6 of Mt. Karimui was in moss forest, and Gilliard
found Reinwardtoena up to 11,000 {ft in moss forest on Mt. Hagen.
All my sightings were of solitary individuals 10-40 ft above the
ground; some were in deep forest, some at the edge of forest, and one
in second-growth. The long tail and notably slow wing beat make the
flight of this graceful pigeon a joy to watch.
VOICE. There are two distinct calls: a series of disyllabic hoots
virtually identical to the call of Macropygia amboinensis (Fig. 14,
p. 136); and a single note followed by a rapid series of a dozen notes
dropping in pitch, suggesting laughter.
Chalcophaps stephani stephani Pucheran
Stephan’s Ground Dove
SPECIMENS EXAMINED. Karimui: 1 ¢,1 @ (16 July and 3 Aug. 1965).
WRIGHIIY. “1 @2 126. 1 9-120,
WING, «1 age 144. 1 2138:
STOMACH CONTENTS. Fruit, seeds, and gravel.
TAXONOMY. ‘These two specimens are darker, more lavender-
gray, and less brown on the breast than other New Guinea material.
The darkness is in conformity with the general trend at Karimui.
BREEDING. ‘The gonads of the male were much enlarged. A nest
was located 30 ft above the ground in a fork of a tree.
DISCUSSION. I have no other records than the two specimens.
At Lake Kutubu (2,450 ft) Schodde and Hitchcock found this species
to be the common ground dove of second-growth and the forest edge,
while Gallicolumba jobiensis was common in primary forest. Else-
where in New Guinea Chalcophaps stephani mainly occurs near sea
level.
VOICE. A series of faint notes delivered at a rate of 3-4 notes per
second and lasting 8-15 sec. ‘Towards the end of the series the notes
rise slightly in pitch and become slightly louder.
Gallicolumba rufigula alaris Rand and G. r. septentrionalis Rand
Red-throated Ground Dove
NATIVE NAME. Daribi: pémo.
SPECIMENS EXAMINED. Karimui: 1 ¢ (11 Aug. 1964); 1 9 (6 Aug. 1965).
WEIGHT. 1 9: 121.
WING, 1 @: 156. 1 Oe 192.
STOMACH CONTENTS. Fruit and some insects.
TAXONOMY. When compared with alaris from the Fly River
(including the type) and Setekwa River, the two Karimui specimens
differ somewhat in being darker on the back and especially darker on
137
SPECIES ACCOUNTS
the nape, but they share the most distinctive character of the race, the
absence of the gray area on the side of the head. The gray edges of the
upperwing coverts are comparable to those in Setekwa River birds,
and broader and grayer than in Fly River birds. The races orientalis
(southeastern New Guinea), septentrionalis (northern New Guinea),
and nominate rufigula (Vogelkop) all differ in possessing a gray area
on the head. In addition, the gray edges of the wing coverts are nar-
rower in septentrionalis, broader in nominate rufigula, and duller and
less distinct in orientalis than in the Karimui birds. The race helvi-
ventris of the Aru Islands agrees in lacking the head patch but has
the covert edgings much more vinaceous. Northern watershed speci-
mens from the Jimmi Valley were assigned to the northern race sep-
tentrionalis by Gilliard and Gyldenstolpe, and Gyldenstolpe (1955)
specifically mentioned the presence of a well-defined gray area on the
side of the head.
When examined in March 1966, the two Karimui specimens still
had the breast quite distinctly washed with yellow-orange, but this
wash was completely absent in prewar material. Rand (194la) com-
mented previously on the rapid postmortem fading of this wash.
BREEDING. ‘The gonads of the male were much enlarged.
DISCUSSION. Both specimens were collected while on the ground
in forest. Karimui natives are familiar with the species, but it is evi-
dently quite uncommon and shy. Like G. beccarit, G. rufigula becomes
locally abundant when wild bamboo produces seeds.
VOICE. A very faint, upwards inflected “br-r-r-r-r,” easily over-
looked or mistaken for a frog.
Gallicolumba beccarw beccarit (Salvador)
Beccari’s Ground Dove
NATIVE NAME. Fore: manumu.
SPECIMENS EXAMINED. Awande: 2 ? (21 June 1964, and 19 June 1965). Men-
gino: 1 g (16 July 1964). Mt. Karimui Zone 4: 1 9 (30 Aug. 1965); Zone 8: 1 ¢
(10 Sept. 1965).
WEIGHT. 1 9: 59, I 2: 56;
WING: I 9: 105. 2 72 105, 109,
STOMACH CONTENTS. Gravel and seeds.
DISCUSSION. ‘This species is widespread but usually uncommon
in montane forest between about 5,000 and 9,000 ft, and is generally
the sole species of ground dove at these altitudes. It lives on the ground
in the forest and is quite shy (all my specimens were netted). Paran
told me that about once a decade, when the wild bamboo produces
seeds, G. beccarii appears in enormous numbers at Awande to eat the
seeds and to nest.
138
SPECIES ACCOUNTS
Gallicolumba jobiensis jobiensis (Meyer)
White-breasted Ground Dove
SPECIMENS EXAMINED. Karimui. 2 ¢,2 9,12 (3 July-5 Aug. 1965).
WEIGHT. 2 4: 182, 158. 2 9: 126,181. 1 ?: 154,
WING. 2 @: 147, 149. 2 9: 140, 141.
STOMACH CONTENTS. Fruit, gravel, and insects.
BREEDING. The testes of the two males were slightly enlarged.
DISCUSSION. All the specimens were solitary and on the ground
inside the forest.
The distribution of this species seems highly erratic. ‘The five speci-
mens from Karimui suggest that it was uncommon but not rare there.
At Lake Kutubu Schodde and Hitchcock found this species to be the
common ground dove of the forest interior. Bulmer collected one
specimen at 6,700 ft, the Fore said that it occurred at Awande, and
a single trapped specimen was brought to Gilliard at Kup at some
elevation about 4,000 ft. Most other New Guinea records are from low
elevations.
Henicophaps albifrons albifrons Gray
White-capped Ground Pigeon
NATIVE NAME. Daribi: odu.
SPECIMENS EXAMINED. Karimui: 1 ¢ (3 Aug. 1965), 2 9 (14 Aug. 1964).
Soliebeda: 1 9 (23 July 1965).
NVUIN Ge Le ae eSiien 22 1S el8b, liso.
BREEDING. One of the two 1964 females had an enlarged ovary,
but the gonads of the other specimens were small.
DISCUSSION. ‘The two Karimui females were collected together.
The Soliabeda female was alone at the top of a tree, and the Karimui
male was one of a pair on the ground. My only experience with this
uncommon species in the field was with a solitary individual at
Soliabeda, flying 10 ft above the ground in partly cut forest with tall
trees.
VOICE. ‘The call, delivered from a perch in a tree, is a rapid series
of “hoo’s” at the rate of 5 per second for 20 sec. The series slightly
rises in pitch, and has a hollow quality similar to the voice of
Macropygia nigrirostris or Centropus menbehki.
Otidiphaps nobilis cervicalis Ramsay
Magnificent Ground Pigeon
NATIVE NAMES. Fore: waibo. Gimi: kwéyo. Daribi: wagari.
SPECIMENS EXAMINED. Awande: 1 9 (19 Oct. 1967).
WING. 199.
STOMACH CONTENTS. Ground-up fruit, and pebbles 10 mm in diameter.
139
SPECIES ACCOUNTS
TAXONOMY. This specimen agrees with cervicalis of southeast-
ern New Guinea, and differs from nominate nobilis of western New
Guinea, in the light gray rather than iridescent green-purple nape,
the dark green rather than dull purple rump, and the deep green
rather than deep purple underparts. A native-produced specimen from
the Jimmi Valley on the northern slope of the Eastern Highlands was
also assigned to cervicalis by Mayr and Gilliard (1954).
DISCUSSION. ‘This is a rare and local inhabitant of the forested
hill slopes in the Eastern Highlands, ranging occasionally up to 6,000
ft. My only field record was of a call heard at 5,100 ft on Mt. Karimui.
VOICE. A plaintive tremulous, descending whistle lasting 2 sec.
Goura scheepmakeri subsp.
Scheepmaker’s Crowned Pigeon
Crowned Pigeons, presumably of this semispecies, were reported to
Schodde and Hitchcock as occurring in the Lake Kutubu district.
Goura victoria beccarii Salvadori
Victoria Crowned Pigeon
DISCUSSION. Gyldenstolpe mentions individuals of this northern
Goura semispecies at Nondugl aviary, captured somewhere on the
northern slopes of the Eastern Highlands. Both semispecies of Goura
are to be expected only at low elevations.
VOICE. A very low “buk-buk-buk.”
PSI? PAGCIDAK: PAROS
Chalcopsitta scintillata chloroptera (Salvadori)
Yellow-streaked Lory
SPECIMENS EXAMINED. Soliabeda: 1 ¢,1 9 (25 July 1965).
WEIGHT. 1 $:>210. 1 9: 195.
WING. “Use l705 i Ox Gs;
TAXONOMY. Both specimens lack the yellow area in the under-
wing which is always present in nominate scintillata and often reduced
or lacking in chloroptera. The underwing coverts are entirely green in
the Soliabeda male, while in the female they are green for the proxi-
mal two-thirds and red for the distal third. This character also places
the Soliabeda birds with chloroptera and differentiates them from
nominate scintillata, in which the underwing coverts are largely red.
BREEDING. ‘The testes of the adult male were slightly enlarged.
DISCUSSION. I encountered no individuals of this lowland spe-
140
SPECIES ACCOUNTS
cies besides this pair, and there are no other records from the Eastern
Highlands.
The fact that the Soliabeda birds are chloroptera reinforces earlier
indications that chloroptera, originally described from the southeast-
ern coast, may extend in the foothills far west of the easternmost
coastal population of C. s. scintillata (originally described from the
southwest). ‘Phe localities from which these two races have now been
recorded are as follows: nominate scintillata, along the southwestern
coast from ‘Triton Bay (long. 134°E) east to at least the mouth of the
Fly River (long. 143°E); chloroptera, on the southeastern coast be-
tween long. 146°E and 148°E, and apparently extending westward in
the foothills and on the upper reaches of the south coast rivers to
Soliabeda (upper Purari drainage, long. 145°E), upper Fly River (one
specimen collected by the Second Archbold Expedition, long. 142°F),
upper Eilanden River (ca. long. 139°E), and upper Noord River (ca.
long. 138°E). Both races show much individual variation (e.g., variable
amounts of yellow in the primaries of chloroptera, of red in the pre-
dominantly green underwing coverts of chloroptera, and of green in
the predominantly red underwing coverts of nominate scintillata).
The Noord River population includes typical chloroptera, typical
nominate scintillata, and intermediates. Probably hybridization of the
two races is now occurring after a prolonged break in contact at the
Fly River, as also seems to be the case with Pstttaculirostris desmarestit.
‘This would be in keeping with the obvious evolutionary history of the
genus Chalcopsitta as a superspecies ring in the New Guinea lowlands,
with contact between populations having been broken and _ reestab-
lished a number of times. ‘The genus consists of three virtually allo-
patric semispecies, whose distributions now form an incomplete circle
(broken in the east between Astrolabe Bay and the Kemp Welch
River) around the periphery of New Guinea: C. duivenbodei on the
north coast, C. atra in the far west, and C. scintillata on the south coast
and Aru Islands. The unique type of C. atra spectabilis (intermediate
between C. a. insignis and C. s. scintillata) also suggests hybridization
between former isolates at at least one other point in the ring besides
the Fly River.
Pseudeos fuscata incondita (Meyer)
Dusky-orange Lory
NATIVE NAMES. Fore: paila. Daribi: bu-kinikini.
SPECIMENS EXAMINED. Karimui: 1 ¢,1 92 (5 Aug. and 11 Sept. 1965).
WHRIGHT. Ll as 143.
WING, 1 as 159; 1 Osho,
TAXONOMY. ‘The male is in the red phase, while the female is
in an intermediate red-orange phase.
BREEDING. ‘The testes of the male were enlarged.
141
SPECIES ACCOUNTS
DISCUSSION. ‘The erratic distribution of this lory in the Eastern
Highlands suggests seasonal migration within New Guinea. On none
of my visits to the Okapa area (June and August 1964; June 1965;
June 1966) did I see it in the field. Yet the Fore were familiar with the
“paila” and said that it was numerous in the rainy season (November
onward). Several Europeans at Okapa had captive specimens in both
the orange and the yellow phases, which the Fore had captured as
nestlings. Similarly, the Daribi were familiar with the Dusky Lory un-
der the name of “bu-kinikini,’ but I found it rare at Karimui and
absent at Soliabeda, Bomai, and Mt. Karimui. In the Wahgi Valley,
Gilliard never encountered it on his two visits (both March or April
to August), and Gyldenstolpe between August and early November col-
lected only one specimen (late October). Apparently the Eastern High-
lands population breeds during the rains and then leaves before the
onset of the dry season, perhaps retreating to the coastal lowlands.
Other Eastern Highlands records include Lake Kutubu, the Schrader
Range, and Kyaka territory. This lory gathers in flocks in flowering
tees:
VOICE. A loud, shrill cry quite similar to that of Trichoglossus
haematodus, but shorter, hoarser, and even more erating.
Trichoglossus haematodus intermedius Rothschild and Hartert
Rainbow Lory
NATIVE NAMES. Fore: k’re-’k’re. Gimi: k’re-k’re. Daribi: kinikini.
SPECIMENS EXAMINED. Mengino: 1 ¢, 2 9 (15 July 1964). Karimui: 6 ¢,
6 9 4-15 Aug. 1964; 3 July-11 Sept. 1965). Soliabeda: 2 ¢@, 2 9 (25-29 July 1965).
WEIGHT. 9 42°27, 129) 1355, lov, Wa 4 OIG, 20 Io, aa
WING. 6 ¢: 143, 143, 144, 144, 145, 146. 7 9: 135, 136, 137, 138, 138, 142, 144.
STOMACH CONTENTS. Flowers.
TAXONOMY. ‘This is a variable species, in which each local pop-
ulation shows some distinct features (Cain, 1955). My specimens are
closest to intermedius from the Wahgi Valley except that the upper
belly is less dark green. ‘Vhe race berawensis from western New Guinea
differs in the more extensive blue on the face and forehead; berauensis
from the Fly River, in having the dark green of the upper belly spot-
tier; and micropteryx of southeastern New Guinea, in the paler breast
and back, browner nape and cheeks, and narrower green edges to the
red of the breast. T. h. caeruleiceps of the lower Fly River is very
distinct in its much darker green belly, pronounced orange wash on
the abdomen, reduced green edges to the red of the breast, and more
extensive blue on the forehead and sides of the head. ‘The material of
Gyldenstolpe and Gilliard from the Wahgi Valley was also assigned to
intermedius, while Schodde and Hitchcock placed one specimen from
Lake Kutubu with micropteryx.
BREEDING. The testes of both males collected at Karimui in
142
SPECIES ACCOUNTS
1964 were enlarged. In 1965 the gonads were enlarged only in a
Soliabeda male. According to the Fore this lory, like many others,
breeds at Okapa during the rainy season.
DISCUSSION. I encountered Trichoglossus haematodus, one of
the most widespread and conspicuous lories in the Eastern Highlands,
at all my collecting stations except Mt. Michael, up to 4,500-5,000 ft
in primary forest and up to 6,500 ft in the casuarina groves and trees
of villages and open country. It usually occurs in small, noisy groups
of up to half a dozen individuals perched in the middlestory and
canopy of trees or calling loudly in flight. Unlike Pseudeos fuscata, to
which it is otherwise rather similar in voice and habits, 7. haematodus
does not leave the Eastern Highlands during the dry season.
Pairs of birds that were perched facing each other 1 ft apart dis-
played by opening and rapidly fluttering the wings to reveal the
strikingly colored pattern of the underwing. Many species of lories
have patterned underwings that may function in species-specific dis-
plays.
Terborgh’s observations on feeding trees indicate that the main
food source of T. haemotodus is flowers and that it takes fruit less fre-
quently. At Karimui it was the second commonest parrot in flowering
trees, accounting for about 10% of the bird-usage, and it actually fed
on the blossoms by biting off the receptacles and allowing the petals
to fall to the ground. However, ‘Verborgh also encountered an unusual
relation between this species and honeyeaters in Ficus trees at Karimui
and Miarosa. ‘The fruits of these cus were about 8 cm long and 4
cm in diameter and protected by a thick, woody pericarp which pre-
cluded any direct assault on the soft pulp inside. Underneath the trees
were lying fallen fruits with neatly cut holes about 1 cm in diameter
at the blossom ends. Observation of the feeding birds showed that
these holes were being made by Trichoglossus haematodus and an-
other parrot, Psittaculirostris desmarestti, which tossed aside chips
with a shake of their heads and gained access to the pulp within five
minutes. Not even these parrots, though, could penetrate the side walls
of the pericarp, and they always had to abandon the fruit with most
of the pulp remaining because of their short bills. Honeyeaters such
as Melilestes megarhynchus, Melidectes torquatus, Meliphaga flavi-
venter, and several other species of Meliphaga then inserted their
long, delicate bills, which would have been useless for opening the
fruit, and ate more of the pulp. Initially, when most of the fruits were
still unopened, 99%, of the bird-usage in these Ficus trees was by
Trichoglossus and Psittaculirostris, and 1% by honeyeaters, Later,
when most of the fruits had been opened and the parrots had ob-
tained as much pulp as their bills could reach, honeyeaters accounted
for up to 93% of the usage, and parrots only for 7%.
VOICE. A high, short, very grating shriek, louder than that of
143
SPECIES ACCOUNTS
Neopsittacus musschenbroekii but less hoarse and grating than that of
Pseudeos fuscata.
Psitteuteles goldiet (Sharpe)
Red-capped Streaked Lory
NATIVE NAME. Fore: yani.
DISCUSSION. This is the least common of the midmontane lories,
known from eight scattered localities in the Eastern Highlands be-
tween about 5,500 and 8,500 ft. My only field observation was of a
noisy flock flying over the forest at 6,900 ft above Miarosa. According
to the Fore it breeds at Okapa during the rainy season as do other
lories, and several caged birds which had been brought in as nestlings
were kept as pets by Europeans at Okapa. In Kyaka territory Bulmer
commonly noted flocks of up to 40 in December and January but only
occasionally saw the species during the rest of the year, implying
local migration.
Domicella lory somu Diamond and D. I. erythrothorax (Salvadori)
Western Black-capped Lory
NATIVE NAMES. Fore: korié. Gimi: korié. Daribi: sému.
SPECIMENS EXAMINED. D. Il. somu: Karimui, 1 4, 2 9; Bomai, 2 4, 2 9;
Soliabeda, 2 g, 2 9; Mt. Karimui Zone 1, 1 ¢ (7 July-11 Aug. 1965). D. 1.
erythrothorax: Sena River, 1 ¢ (26 July 1964).
WING. D. lI. erythrothorax; 1 8, 167 (see Diamond, 1967a, for weights and
wing measurements of D. 1. som).
TAXONOMY. As discussed elsewhere (Diamond, 1967a), the race
somu, with a restricted range in the Karimui Basin and Purari River
drainage, is unique in lacking the neck band present in all other races
of D. lory but absent in D. hypoinochrous. Surprisingly, the single
specimen which ‘Verborgh and I collected 15 miles east of Karimui at
the Sena River, on the slopes of the mountain walls ringing the Kari-
mui Basin but still within the basin, definitely belongs to the south-
eastern New Guinea race erythrothorax because of the presence of the
neck band (and slightly larger size). Schodde and Hitchcock assigned
the Kutubu population to D. 1. rubiensis, but their description implies
that it actually belongs to D. 1. somu, since they mention the absence
of the neck band.
BREEDING. ‘The gonads were small in 11 specimens, slightly en-
larged in the two others, suggesting that breeding was suspended in
the dry season, as with other lories.
DISCUSSION. Domicella lory is generally distributed along the
southern edge of the Eastern Highlands up to about 4,500 ft, being
uncommon above 3,000 ft but numerous at Soliabeda (2,000 ft). In
virgin forest the lories Domicella lory, Trichoglossus haematodus, and
144
SPECIES ACCOUNTS
Pseudeos ftiscata all extend up to nearly the same altitude (the latter
two consistently several hundred feet higher), but Domicella lory has
failed entirely to follow the other two species in colonizing the man-
made second-growth habitats of the Eastern Highlands at 5,000-6,500
ft. The social behavior is also different. Trichoglossus haematodus and
Pseudeos fuscata are often seen in small flocks, both when in flight
and when in feeding trees. All my sightings of Domicella lory in flight
were of individuals or pairs, and the largest number that I saw to-
gether in a feeding tree was three. ‘To judge from the trees it fre-
quents, it feeds principally on flowers, secondarily on fruits.
VOICE. More varied and more musical than that of Trichoglossus
haematodus or of Pseudeos fuscata. The commonest vocalization con-
sists of two identical squeals in immediate succession, with a quality as
of sleigh-bells when heard in the distance; this double squeal cannot
be confused with the voice of other New Guinea parrots. In addition,
a variety of high piercing squeaks and loud whistles may be emitted,
particularly when the birds are perched.
| Domiceta hypoinochroa devittatus (Hartert) |
Eastern Black-capped Lory
DISCUSSION. ‘This species has never been found west of Hall
Sound in southeastern New Guinea. Mayr and Gilliard (1954) list it
from the Eastern Highlands on the basis of a captive bird photo-
graphed by Gilliard at Kup and said to have been caught south of the
Kubor Range. ‘This captive was probably a misidentified individual of
Domicella lory somu, the common Domicella of the Eastern High-
lands. At the time when Gilliard examined the Kup pet in 1952, the
race D. 1. somu was still unknown, and the best way of distinguishing
D. hypoinochroa and D. lory was thought to be the neck band present
in the former and absent in the latter. However, somu lacks the neck
band, and must be distinguished from D. hypoinochroa by subtler
points easily overlooked or invisible in a photograph.
Charmosyna papou goliathina Rothschild and Hartert
Papuan Lory
NATIVE NAMES. Fore: waiya. Gimi: waiya. Daribi: hade.
SPECIMENS EXAMINED. Mt. Michael: 1 9 (4 July 1964). Mt. Karimui Zone
gl 6.2 2 (27-28 Aue, 1965).
WING. 1 @: 150. 3 Q: 145 (2), 147.
STOMACH CONTENTS. A paste of flowers.
TAXONOMY. ‘The Mt. Michael female and Mt. Karimui male
are in the black phase, while the Mt. Karimui females are in the red
phase. ‘The black phase seems several times more numerous than the
red phase in the Eastern Highlands. ‘The specimens agree with golia-
145
SPECIES ACCOUNTS
thina of western New Guinea and differ from stellae of southeastern
New Guinea in that the longest rectrices are yellow-green rather than
reddish yellow, and in that the red of the longest uppertail coverts is
tipped with green.
BREEDING. The Mt. Karimui male and female were a pair with
enlarged gonads.
DISCUSSION. Charmosyna papou is a regular inhabitant of the
forest interior from 5,500 to about 9,000 ft and has turned up at vir-
tually all collecting stations in this habitat. It is locally common, but
its abundance depends upon the presence of the flowering trees on
which it feeds. Unlike some other lories, C. papou remains within the
forest, and I never saw it in open country.
The altitudinal range of C. papow lies above those of all its con-
geners, except that C. pulchella occasionally overlaps C. papou. C.
papow shares its range with four other lories, but these five high-
altitude lories sort out largely on the basis of size: Oreopsittacus arfaki,
ca. 20 g; Neopsittacus pullicauda, 38 g; Neopsittacus musschenbroeku,
50 g; Psitteuteles goldier, 60 g; Charmosyna papou, 85 g.
VOICE. A shrill, somewhat grating cry, more grating than that
of Oreopsittacus arfaki or Neopsittacus pullicauda but much weaker
than that of Trichoglosslsus haematodus. A loud wing beat which I
heard frequently on Mt. Karimui was attributed to this species by
the Fore:
Charmosyna pulchella bella (De Vis)
Little Red Lory
NATIVE NAMES. Fore: stshuke. Gimi: ani.
SPECIMENS EXAMINED. Mt. Karimui Zone 2: 2 ¢,1 9; Zone 3: 1 4,2 Q
(14-17 Aug. 1965).
WEIGHT. 2) 6237; 3 O's S10, 32:8, 34;
WING. 3 4: 93, 94,95. 8 Q: 87, 91, 92.
TAXONOMY. The specimens are indistinguishable from bella
of southeastern New Guinea and ‘Telefolmin.
BREEDING. ‘The testes were slightly enlarged.
DISCUSSION. ‘This lory is sparsely distributed in hill forest along
the slopes of the Eastern Highlands between about 2,000 and 5,800 ft.
Its altitudinal range lies largely but not entirely below that of C.
papou. Elsewhere in New Guinea its range lies entirely above that of
C. placentis, but in the Karimui area, where C. placentis extended up
to 4,750 ft, C. pulchella and C. placentis overlapped considerably and
fed in the same flowering trees at the Sena River and on Mt. Karimui.
C. pulchella is one of several hill forest species with a discontinuous
altitudinal range at Karimui because the tropical flat basin floor di-
vides its normal range on the hill slopes (p. 55). On Mt. Karimui all
146
SPECIES ACCOUNTS
the specimens and observations of C. pulchella fell between 4,400 and
5,290 ft. The species was also present at the Sena River, (4,500 ft) on
the eastern wall of the Karimui Basin. I never encountered it on the
flat basin floor (3,650 ft), and Yudo natives confirmed that it was ab-
sent there and to be found only on the mountain. A few individuals
were seen again when we went down to Soliabeda (2,000 ft) beyond the
basin walls.
Like other lories, C. pulchella congregates noisily in flowering trees.
Like C. papou and unlike C. placentis or Trichoglossus haematodus,
it remains within the forest and rarely ventures into open country.
VOICE. A short note similar to that of C. placentis but sweeter,
less shrill or staccato.
Charmosyna wilhelminae (Meyer)
Pygmy Streaked Lory
One specimen collected by Bulmer at 4,500 ft in the Lanim Valley
is the sole Eastern Highlands record of this rare lory. A flock that I
observed at 5,000 ft on Mt. Albert-Edward in southeastern New Guinea
was feeding at flowers of the oak Castanopsis.
Charmosyna placentis placentis (Yemminck)
Yellow-fronted Blue-eared Lory
NATIVE NAME. Daribi: abubage.
SPECIMENS EXAMINED. Karimui: 4 ¢ (2-10 July 1965). Soliabeda: 4 @, 5
© (23-30 July 1965). Mt. Karimui Zone 2: 2 ¢, 1 9 (14-15 Aug. 1965).
WEIGHT. 9 4: Soliabeda, 33.0, 36.5, 39.0, 42.0; Karimui, 31.0, 33.0, 40.0; Mt.
Karim, 30.5; 62.0) (overall average 35.5 3-4.5). 8 92 Soliabeday 26.5, 290, 29.3;
30.3, 33.5; Karimui, 33.0, 36.3; Mt. Karimui, 35.7 (overall average 31.7 + 3.5).
WING. 9 ¢: Soliabeda, 84, 86, 86, 87; Karimui, 86, 86, 90; Mt. Karimui, 88, 91
(overall average 87 + 2). 11 9: Soliabeda, 81, 82, 83, 84, 84; Karimui, 85, 86, 88,
89; Mt. Karimui, 88, 91 (overall average 86 + 3).
STOMACH CONTENTS. Flowers, seeds, and a paste of flowers.
TAXONOMY. ‘These specimens have a blue rump patch of the
same size as in nominate placentis from the Fly and Oriomo Rivers,
Kei Islands, and Aru Islands (the patch is much larger in ornata and
absent in swbplacens of southeastern New Guinea). My females aver-
age slightly darker above, and my males deeper red on the flanks,
than the comparative material of placentis, in line with the general
trend towards darkness at Karimui. ‘he measurements (wings of males
and females, and weights of females but not of males) suggest an in-
crease in size with altitude.
BREEDING. ‘The gonads were considerably enlarged in one male
and one female from Karimui and in two males from Soliabeda, and
slightly enlarged in both males from Mt. Karimui.
147
SPECIES ACCOUNTS
DISCUSSION. ‘This species has hitherto been considered a char-
acteristic bird of the New Guinea lowlands at elevations near sea level.
Its presence in abundance at Karimui (3,650 ft) may constitute an alti-
tudinal record. It remained numerous on the lower slopes of the basin
walls and finally disappeared at 4,750 ft on Mt. Karimui. At Soliabeda
(2,000 ft) it was even more abundant. There are no other Eastern
Highlands records, and its distribution or area of abundance in the
Eastern Highlands may prove to be confined to the Karimui area.
C. placentis congregated to feed in flowering trees, where it was the
most numerous species in the Karimui area and provided from 15%
to as much as 65% of the bird-usage. We had no records of it in the
many fruiting trees kept under observation. In the flowering trees it
distributed itself uniformly throughout the crown. At any time a given
tree might hold up to 25 individuals of C. placentis, which came and
went in groups of two to 10, kept up an incessant chatter, and often
paused from feeding to chase each other.
VOICE. A single short note “tsss’’—crisp, shrill, high, and staccato,
and considerably less substantial than the notes of Neopsittacus mus-
schenbroeku, Trichoglossus haematodus, or Pseudeos fuscata.
Oreopsittacus arfaki grandis Ogilvie-Grant
Plum-faced Mountain Lory
NATIVE NAMES. Fore: tushuke. Gimi: gigi.
SPECIMENS EXAMINED. Mt. Michael: 2 6, 3 9 (I-11 July 1964). Mt.
Karimui Zone 4: 1g, 1 9= Zones: 1 6,2 93 Zone 6: 1 > (28 Aug.-5 Sept. 1965):
WRIGHT. 2 622233, 225. 4.07 217, 21:8) 220; 225.
WING. 2 4:81, 81. 5 oO: 81,82 82. 84 ee
STOMACH CONTENTS. Flower paste.
TAXONOMY. Eastern Highlands birds lack the red abdominal
spot present in major and nominate arfaki.
BREEDING. ‘The testes were enlarged in birds from Mt. Karimui.
DISCUSSION. ‘This small, high-altitude lory is characteristic of
moss forest, and its vertical distribution on a given mountain seems
to correlate with the distribution of this vegetational type. On Mt.
Karimui, where the cloud level is lower than in most other parts of
the Eastern Highlands, Oreopsittacus descended to 5,600 ft. On Mt.
Michael, where the trees were heavily mossed only above 8,700 ft,
Oreopsittacus was found only above this altitude and occurred up to
about 10,500 ft in the moss forest. At Okapa I collected up to 7,500 ft
and found neither heavy moss nor this parrot, but the Fore were
familiar with the “tushuke” and said that it lived somewhere on top. It
occurs in flocks of two to six and frequents flowering trees.
VOICE. A short, weak, unvoiced, repeated note “ts,” weaker than
that of other montane lories and more suggestive of a warbler than of
a parrot.
148
SPECIES ACCOUNTS
Neopsittacus musschenbroeku major Neumann
Yellow-billed Mountain Lory
NATIVE NAMES. Fore: kasa. ’ Gimi: kasa.
SPECIMENS EXAMINED. Awande: 7 ¢,5 9 (14-20 June 1965). Mengino: 1
@ (15 July 1965). Mt. Karimui Zone 3: 2 9 (17 Aug. 1965).
WEIGHT. 6 @:-50-62 G22). 7 Ox 40-55 (00 4 I).
WING. 8 4: 109-117 (13 +53). 7 9: 111-116 (113 = 2).
STOMACH CONTENTS. Dry and granular flower remains,
BREEDING. All specimens, except one Awande female, had small
gonads. According to the Fore, the “kasa” breeds in the rainy season.
DISCUSSION. Neopsittacus musschenbroeku is widespread at
about 5,000-8,500 ft throughout the Eastern Highlands. It is one of
the few midmontane forest birds that have profited greatly from hu-
man activities, most of the other species in man-made habitats in the
Eastern Highlands being originally lowland birds. For example, it
was numerous in partly cut forest around Okapa, Lufa, and Mengino,
and was the commonest parrot at Miarosa village (5,800 ft), congre-
gating in casuarina stands along the gardens and roads in flocks of
up to 50. On the other hand, it seems to require at least the close
proximity of forest and is absent in areas (such as Goroka) where the
forest has been largely or completely eliminated for miles around to
leave just isolated groves. N. musschenbroekw is invariably seen in
pairs or larger groups, not as individuals.
VOICE. A shrill! “ss”, “ks’, or “ts”, somewhat staccato and hoarse,
and moderately high in pitch. In comparison with the notes of other
lories that may be found in its altitudinal range, the note of N.
musschenbroeki is harsher, shorter, and more grating than that of
N. pullicauda, more staccato and considerably stronger than that of
Oreopsittacus arfaki, and less harsh and somewhat weaker than the
notes of Pseudeos fuscata or Trichoglossus haematodus. The note is
heard frequently whether the birds are perched or in flight.
Neopsittacus pullicauda pullicauda Hartert
Orange-billed Mountain Lory
SPECIMENS EXAMINED. Mt. Michael: 1 ¢ (5 July 1964).
WING. 100.
STOMACH CONTENTS. Dry and granular flower remains. This species and
N. musschenbroekii may actually eat flowers, whereas Oreopsittacus arfaki and
Charmosyna papou, whose stomachs contain soft paste, perhaps only take nectar.
BREEDING. ‘The gonads were very small.
DISCUSSION. ‘To my surprise, the specimen was caught in a mist-
net a few feet above the ground in tall forest on level ground at 8,000
ft (Neopsittacus generally stays in the canopy).
This is the high-altitude Neopsittacus species, living in moss forest
149
SPECIES ACCOUNTS
throughout the Eastern Highlands at about 7,500-10,000 ft. Its alti-
tudinal range overlaps that of N. musschenbroekii by 500 or 1,000 ft,
and in the intermediate zone both species can be found in the same
flowering tree. This overlap may be made possible by the fact that
N. musschenbroeku is about 40% heavier than N. pullicauda. ‘The
main plumage differences between these two sibling species are that
N. musschenbroekii has a yellow bill, yellow undertail, and light shaft
streaks on the crown, while N. pullicauda has a more orange bill, olive
undertail, and lacks light shaft streaks on the crown.
VOICE. A gentle sibilant note, harsher than that of Oreopsittacus
arfaki but weaker and less shrill than that of Neopsittacus musschen-
broekit.
Psittaculirostris desmarestti cervicalis (Salvadori and D’Albertis)
Large Fig Parrot
NATIVE NAME. Daribi: shuashta.
SPECIMENS EXAMINED. Karimui: 4 ¢ (12 Aug. 1964; 4-5 Aug. 1965). Bomai:
1 $ (9 July 1965). Mt. Karimui Zone 1: 1 @ (11 Aug. 1965).
WHICH. 2.02 l0s us. “ho 26:
WING, 4 6: 12,16, MGA doe Ail:
TAXONOMY. When compared with specimens of cervicalis (the
southeastern New Guinea race) from the Brown River, Kumusi River,
Hydrographer Mountains, and Holnicote Bay, Karimui specimens dif-
fer only in that the throat and cheeks are paler, more yellow and less
orange, and in that the ear coverts are even deeper red-orange. From
Fly River material collected by the Second Archbold Expedition and
classified as godmani cervicalis (Rand, 1942a), Karimui specimens
differ in that the breast band is deeper blue and in that the ear coverts
are darker and deeper orange, but they agree in the color of the throat
and cheeks. ‘The Karimui population is thus intermediate between
southeastern New Guinea birds and the variable Fly River popula-
tion but is nearer the former.
BREEDING. ‘The gonads of the 1964 male were considerably en-
larged, but those of the 1965 males were only slightly enlarged.
DISCUSSION. ‘The three semispecies of genus Psittaculirostris
constitute a superspecies ring in the lowlands around the periphery of
New Guinea. As discussed by Rothschild (1920), Fly River birds ap-
pear to be hybrids between the distinct races P. d. godmani of the
southwest and P. d. cervicalis of the southeast. Evidently the Fly River
was one of four major points (the other three being Huon Gulf, Hum-
boldt Bay, and Geelvink Bay) where distribution in the Psittaculiros-
tris ring was formerly broken, but reestablishment of contact at the
Fly River between populations on either side of the break evidently
occurred before differences had developed to the point of reproductive
150
SPECIES ACCOUNTS
isolation. The evolutionary history of the genus Chalcopsitta must
have been similar.
The presence at Karimui (3,250-4,200 ft) constitutes an altitudinal
record for this lowland genus, and other records of P. desmarestti tor
the Eastern Highlands are lacking. As in the case of several other low-
land species, so for P. desmarestii the Karimui population represents
an isolated island of distribution, since I did not find the species at
Soliabeda (2,000 ft) immediately to the south of the Karimui Basin,
nor did Schodde and Hitchcock report it from Lake Kutubu (2,450 ft).
At Karimui this parrot was uncommon, was seen in groups of two to
six, and occurred much more often in fruit trees than in flowering
trees, where it might be seen at any height from the lowest to the high-
est branches. ‘Together with Trichoglossus haematodus it cut holes
with its heavy bill through the woody pericarp of Ficus fruits, most
of whose pulp was then eaten out by long-billed honeyeaters (see
p. 143). One pair was seen coming out of a hole in a tree.
VOICE. A high, thin, downslurred note quite unlike that of the
other parrots in feeding trees at Karimui.
Psittaculirostris edwardsit (Oustalet)
Edward's Fig Parrot
DISCUSSION. Mayr and Gilliard recorded two specimens col-
lected by N. B. Blood in the Jimmi River region, and another speci-
men at Nondugl aviary from unknown locality. Gyldenstolpe men-
tioned four further Nondugl aviary birds originating from the Jimmi
River. This is the Pstttaculirostris form of northeastern New Guinea
and presumably confined in the Eastern Highlands to low elevations
on the northern slopes.
VOICE. A very short, metallic note “ks” with a sharp attack, as
the sound of coins falling on concrete.
Opopsitta gulielmitertit swavissima (Sclater)
King William’s Fig Parrot
SPECIMENS EXAMINED. Bomai: 2 ¢ (a and b),1 9 (6 July 1965).
WEIGHT. 2 4: 33 (a), 27 (b). 1 9: 33.
WING. 2 4: 77 (a), 70 (b). 1 9: 79.
TAXONOMY. All three specimens agree with swavisstma of south-
eastern New Guinea in having the forehead blue, whereas it is brown
in fuscifrons, the race of southwestern New Guinea from the Fly River
westward. The inner edges of the innermost primaries are yellow, a
character shared by both these races and by melanogenia of the Aru
Islands. ‘The small male labelled ‘“b” resembles the adult female in
151
SPECIES ACCOUNTS
plumage and is presumably a second-year bird (Mayr and Rand, 1037)
DISCUSSION. In addition to collecting this lowland species at
Bomai, I observed it at Karimui and Soliabeda as well, but there are
no other Eastern Highlands records outside of the Karimui area. It
was uncommon and was seen in groups of two to four, containing birds
of both sexes. We had a few observations of it in flowering trees.
VOICE. ‘There are two call notes: a short, unmusical, staccato
ks”; and a short, more musical, less staccato “‘ts’’ with a sweet quality.
Both notes are less sharp, shrill, or grating than those of Charmosyna
placentis.
Opopsitta diophthalma festetichi (Madarasz)
‘T'wo-eyed Fig Parrot
NATIVE NAMES. Fore: ninti or isoateo.
SPECIMENS EXAMINED. Okasa: 1 9 (24 June 1965).
WEIGHT. 56.
WING. 94.
TAXONOMY. ‘The races of this parrot have been reviewed re-
cently by Forshaw (1967). I reassessed the New Guinea mainland and
Aru Islands populations in the light of 51 specimens at the American
Museum of Natural History: 11 from southeastern New Guinea Chee
> 9, limm. ¢), four fom the Kasterm Fhehlands (1 45 2 5 lam:
@), one ( g ) from the Huon Peninsula, one (¢ ) from the Ramu River,
two (¢) from Astrolabe Bay, five from the North Coastal Range (2 4,
3 @), 19 from western New Guinea and the western Papuan islands
(lO"3, 7 2; 2mm. ¢), ones) trom the Bly Raver, and seven <,
2 ¢,2imm. ¢) from the Aru Islands. ‘The material from eastern and
northeastern New Guinea (Eastern Highlands, Huon Peninsula, North
Coastal Range, Huon Peninsula, Ramu River, Astrolabe Bay) differs
from the southestern and western populations in having a wider yellow
band across the crown of the male and in that the backs of the females
are slightly darker. ‘The name festetichi (type locality Astrolable Bay)
is available for this eastern and northeastern population to distinguish
it from nominate diophthalma (type locality ‘Triton Bay, western New
Guinea). In both the male and female characters the southeastern
population is not separable from the western population and is sep-
arable from festetichi, so that the race coccineifrons from southeastern
New Guinea must be considered synonymous with nominate dioph-
thalma (not with festetichi, as Forshaw had suspected). Neither For-
shaw nor I could detect the alleged variation in the depth of the red
color of the forehead and cheeks, sometimes cited as a racial character.
In the race aruensis the yellow crown band of the male is either lack-
ing (Aru Islands) or very narrow (Fly River), and the female of aru-
ensis is quite distinct.
152
SPECIES ACCOUNTS
BREEDING. ‘The Okasa female was one of a pair coming out of
a hole in a tree and may have been preparing to breed, as suggested
by the enlargement of the ovary.
DISCUSSION. ‘This fig parrot has a very spotty distribution at
lower elevations (i.e., up to about 5,000 ft), and apparently lives in
second-growth trees, in partly cut forest, and at the forest edge rather
than in primary forest. In 1964 ‘Terborgh saw the species several times
at Okasa (3,500-4,250 ft) in stranglers bearing fruits about 5 mm in
diameter. Gilliard reported it as “not uncommon up to 5,000 feet in
the riverine forests near Kup” (Mayr and Gilliard, 1954, p. 339); Gyl-
denstolpe collected 10 specimens near Nondugl but met it nowhere
else; Bulmer obtained one specimen about 6,000 ft in Kyaka territory;
and there are sight observations from Baiyer Valley (N.G.B.S. News-
letter, No. 54, p. 1, June 1970). Birds that I saw were usually solitary,
and perched at the tops of dead trees or else probed bark while hang-
ing upside down.
VOICE. A sharp, sweet, somewhat metallic “ks,” similar to the
call of Psittaculirostris edwardsit but weaker and with a less sharp
attack.
Micropsitta bruijni bruijni Salvadori
Mountain Pygmy Parrot
This locally distributed species of mountain forest is known in the
Eastern Highlands from one specimen taken at 7,500 ft in the Kubor
Mountains by Gilliard, six taken on the Schraderberg by Burgers, one
collected at 6,700 ft at Minj by N. B. Blood, sightings by Bulmer in
the Jimmi Valley, and sightings at 5,500 ft in the Hagen area (N.G.B.S.
Newsletter, No. 54, p. 1, June 1970). It works along and around
branches in nuthatch-like fashion. The very similar Micropsitta pusio
may turn up at lower elevations.
Probosciger aterrimus goliath (Kuhl)
Palm Cockatoo
NATIVE NAMES. South Fore: pakitéra. Daribi: boriau.
SPECIMENS EXAMINED. Okasa: 1 ? (26 June 1965).
WING. 390.
TAXONOMY. From the large size I presume that this specimen
belongs to the race goliath, inhabiting the foothills of southern New
Guinea, rather than to nominate aterrimus, an Australian race which
reaches the coastal lowlands of southern New Guinea. Schodde and
Hitchcock also identified the Lake Kutubu population as goliath. The
race stenolophus of northern New Guinea has much narrower crown
feathers.
153
SPECIES ACCOUNTS
DISCUSSION. ‘The specimen was collected while eating fruit at
the top of a tree in the Okasa forest at about 4,200 ft. According to
Fore natives living near this forest, the “pakitora” is quite uncommon
there. My Daribi assistants also knew the “boriai’” but said that it
appears only very infrequently at Karimui (3,650 ft).
VOICE. A rapid series of alternate upslurs and downslurs “whik-
whik-whik .. .” at a rate of 4 per second. Occasionally, a melodious,
whistled, jumbled call with a quality similar to Domicella lory.
Cacatua galerita triton ‘VYemminck
Sulphur-crested Cockatoo
NATIVE NAMES. Fore: wai. Gimi: é6de. Daribi: na4ra. Pidgin English: koki.
DISCUSSION. In the areas of the Eastern Highlands which I vis-
ited and which are still little disturbed by man, C. galerita is a ubiqui-
tous and conspicuous inhabitant of primary forest up to about 5,000
ft, locally (Awande, Mt. Michael, Mt. Karimui) up to 7,000 ft, and
was present at all my Eastern Highlands collecting stations. In areas
with moderate human population densities, particularly those where
shotguns are permitted, this cockatoo tends to disappear because its
yellow crest plumes are prized for decoration. Gyldenstolpe and Gilli-
ard did not meet cockatoos in the central parts of the Eastern High-
lands in the wild state. ‘Their studies were confined to altitudes above
4,000 ft, largely above 5,000 ft, but at these elevations cockatoos should
merely be uncommon rather than absent, and I suspect that the species
has already been extermniated by the higher human population densi-
ties in these areas.
The Sulphur-crested Cockatoo was usually seen in small groups of
from a few to a dozen individuals. It is most conspicuous when soar-
ing, somewhat hawk-like, high over the forest and calling loudly. ‘This
behavior seems to constitute a social display, for a few birds will land
screaming in a tree, to be followed by several others which can be
seen and heard as they come flying in from a mile away. A perched
bird that is alarmed bends its head and sways, raising its yellow crest,
to observe a human intruder from various angles.
VOICE. The call is the loudest of any New Guinea bird: a fa-
miliar, ear-splitting, harsh scream, delivered perched or in flight. A
very different and infrequent vocalization, given only when perched,
consists of a virtually continuous sound which goes on for several
minutes, interrupted briefly every 114 sec while the bird catches its
breath. The pitch is nearly constant but rises slightly before each
catching of breath. The sound is not loud and is difficult to localize
if one does not happen to see the singer. ‘The quality is totally un-
musical, somewhat rasping, invites confusion with an insect, but most
closely resembles the voice of a dog whose vocal cords have been dam-
aged.
woe
SPECIES ACCOUNTS
Psittrichas fulgidus (Lesson)
Vulturine Parrot
NATIVE NAMES. Fore, Gimi, and Daribi: kabare.
DISCUSSION. My sole field observation of this remarkable parrot
in the Eastern Highlands came at Soliabeda, in an area of very low
human population density, where one bird perched at the top of a tall
tree in a native garden and then flew off. ‘he distinctive flight con-
sists of alternate rapid flaps and soaring, reminiscent of the Black
Vulture Coragyps atratus of the New World. ‘To judge from native
reports, it is present but rare at lower elevations in Fore, Gimi, Daribi,
and ‘Tudawhe areas. Schodde and Hitchcock saw groups of up to 20
at Lake Kutubu.
The red feathers of the “kabare” are prized by natives far more than
the plumage of any other bird, including any bird of paradise. Even
in a poor area like Karimui, a Psittrichas commanded the relatively
enormous sum of twenty dollars, equal to the price of a large pig and
not much less than the price of a wife. Despite this popularity I saw
or heard of only three captive Psittrichas in the Karimui area, an in-
dication of its rareness even in areas with sparse human population.
This is one of the few montane species whose existence is threatened
directly (i.e., as opposed to being threatened indirectly through de-
struction of habitat) by man.
VOICE. Very harsh, rasping, drawn-out screams. Often the screams
are in pairs, the first at constant pitch and hoarse, the second up-
slurred, nasal, and squeaky as the sound of a rusty gate.
Larius roratus pectoralis (Muller)
Red-sided Eclectus Parrot
NATIVE NAMES. Gimi: néra. Daribi: anari (male), tégali (female).
SPECIMENS EXAMINED. Karimui: 1 ¢ (12 Aug. 1964). Mt. Karimui Zone 1:
1 g (11 Aug. 1965).
WING. 2 4: 256, 263.
BREEDING. ‘The testes of the Mt. Karimui specimen were large,
while those of the Karimui bird were very small.
DISCUSSION. ‘The Red-sided Eclectus Parrot was found up to
4,800 ft throughout the Karimui area, where my census figures showed
that it was consistently about one-third as numerous as Cacatua
galerita. In the lowlands this species is more numerous and sometimes
assembles in considerable numbers, circling conspicuously high over-
head at sunset like the Nestor parrots of New Zealand. However, in
the Karimui area it was invariably solitary. It is probably widespread
at the lower elevations in the Eastern Highlands, since my Gimi as-
sistants were familiar with it and my Awande assistants knew it from
low elevations in Fore territory. It was found common at Lake Kutubu
155
SPECIES ACCOUNTS
(2,450 ft) by Schodde and Hitchcock and up to at least 5,000 ft in
Kyaka territory by Bulmer.
VOICE. A loud harsh scream, quite similar to that of Cacatua
galerita, but with a more croaking quality and less deafening.
Geoffroyus geoffroyi aruensis (Gray)
Red-cheeked Parrot
SPECIMENS EXAMINED. Soliabeda: 1 9 (22 July 1965).
WEIGHT. | 169.
WING. 155.
TAXONOMY. The single specimen agrees with aruensis, e.g., in
that the rump is green rather than red.
DISCUSSION. At Soliabeda (2,000 ft) this was the fourth com-
monest parrot, and Schodde and Hitchcock found it widespread at
Lake Kutubu (2,450 ft), but I saw it only a few times at Karimui
(3,650 ft). This is a lowland species with an altitudinal range only
marginally overlapping that of its congener G. simplex, which was
absent at Soliabeda but common at Karimui and higher elevations.
G. geoffroyi occurred singly or in groups of a few birds, and most often
attracted attention when calling loudly in flight.
VOICE. A series of identical notes repeated at the rate of two to
three per second for 5 to 10 sec. The notes are high and fairly loud,
and have a bright, piercing, ringing quality as of a metal surface being
struck. Each note is upslurred, virtually disyllabic. Only two other
species have calls with which the call of Geoffroyus geoffroy: can be
confused. At a great distance and when not heard clearly, the call of
G. simplex may sound similar. Some calls of the drongo Dicrurus hot-
tentottus, which also consist of a series of ringing, metallic, disyllabic
notes, are very similar and at times virtually indistinguishable. With
practice the drongo calls can usually be distinguished by the facts that
the series in the drongo usually consists of fewer notes; that the
drongo’s notes are sometimes downslurred, whereas they are always
upslurred in G. geoffroyi in the Eastern Highlands (though not in
other parts of New Guinea); and in that the call is usually delivered
in flight by G. geoffroyi but when perched by the drongo. ‘Vhe quality
of the voice of G. geoffroyi shows considerable geographical variation.
Geoffroyus simplex subsp.
Blue-collared Parrot
NATIVE NAMES. Fore: k’ri-k’ro. Daribi: sibunane.
BREEDING. Said by natives to breed in the rainy season at
Okapa.
DISCUSSION. Although I found this parrot widespread and fairly
156
SPECIES ACCOUNTS
common throughout the Eastern Highlands at 3,000-6,500 fi 1t 1680
incredibly shy that a specimen with locality data has yet to be col-
lected. Usually I saw it in flocks flying overhead at a great height and
calling, and neither I nor my native assistants were able to approach it
more closely than 200 feet when it was perched in flowering trees, with-
out its panicking and bolting. It is said to eat acorns of the oak Casta-
nopsis. A nestling reared in captivity learned to speak a few words.
VOICE. In pattern, similar to the voice of G. geoffroyi, 1.e., a series
of piercing high notes delivered at the rate of about two per second.
The successive notes are at nearly, but not exactly, the same pitch and
vary slightly in quality, whereas successive notes of G. geoffroyi are
identical. The sounds are well represented by a repetition of the Fore
name: k’ri-k’ro-k’ri-k’vo- etc. As usually heard, i.e., at a great height,
the quality is appealing and somewhat musical, as of sleigh-bells in
the distance or of a rusty gate swinging back and forth. Under these
circumstances confusion with the voice of G. geoffroyi is possible. ‘The
charm of the call disappears when heard nearby, and it becomes loud,
harsh, piercing, and unmusical.
Alisterus chloropterus callopterus (D’Albertis and Salvadori)
Papuan King Parrot
NATIVE NAMES. Fore: awe. Gimi: kafe. Daribi: tanabo.
SPECIMENS EXAMINED. Awande: 1 ¢, 2 9 (16 June 1965). Okasa: 2 9,1?
(22-23 June 1965). Karimui: 1 ¢, 1 imm. ¢ (5 Aug. 1965).
WEIGHT. 2 4: 167,173. 2 9: 155,170. 1 ?: 148.
WIENG. 84,0195. 2) 0 2190) 97.
TAXONOMY. In the narrowness of the blue band on the upper
back, the two adult males agree with callopterus, the form of central
New Guinea, rather than with nominate chloropterus, the southeast-
ern New Guinea race. ‘The band is slightly broader in the male from
Awande than in the male from Karimui (100 miles west of Awande),
possibly suggesting an approach to nominate chloropterus.
BREEDING. ‘The gonads were enlarged in the Okasa specimens,
small in Karimui specimens.
DISCUSSION. ‘The Papuan King Parrot was present in low num-
bers at all my forested collecting localities below 6,500 ft in the south-
ern parts of the Eastern Highlands and probably occurs down to the
foot of the hills. Apart from a Nondugl aviary bird and a pet bird of
uncertain origin recorded by Gilliard and Gyldenstolpe, and probably
taken at lower elevations in the Jimmi Valley, records of this forest
species from the central parts of the Eastern Highlands seem to be
lacking, possibly because of the extensive deforestation there below
6,500 ft. It is virtually confined to the shaded forest interior, occurs
singly or in groups of two or three, remains unobtrusively in the lower
branches and middlestory, moves deliberately, and flies only short dis-
157
SPECIES ACCOUNTS
tances and without calling when disturbed. Because of its retiring be-
havior it is heard more often than seen.
VOICE. ‘Two distinctive calls: (a), a single, low-pitched, fairly
loud, hoarse, harsh, very short and staccato note. (b) ‘(he commoner
call is a series of five or six similar notes at 11% sec intervals, at medium
pitch, with a somewhat piercing quality, and each note being slightly
downslurred. The pitch usually drops slightly with successive notes
im the series (ip. 15),
Alisterus chioropterus:
Fic. 15. Voice of Alisterus chloropterus.
Niche Differences in the Genus Psittacella
The four species of this genus apparently sort out ecologically in a
simple fashion. ‘There are two large species, P. brehmii and P. picta,
and two small species, P. madaraszi and P. modesta. At a given altitude
one may find one large species and one small species, but the two large
species have nearly mutually exclusive altitudinal ranges, as do the
two small species. Details of plumage and distribution suggest that,
of the three speciations in this genus, the first was the split between a
large form and a small form, the second produced the second large
species, and the most recent produced the second small species (Fig.
16).
Psittacella brehmu pallida Meyer
Brehm’s ‘Tiger Parrot
NATIVE NAMES. Fore: tange. Gimi: akoi.
SPECIMENS EXAMINED. Mt. Michael: 2 9 (1 and 4 July 1964). Mt. Karimui
Zone 4:1 @ (31 Aug. 1965).
WEIGHT. I-34. 92.
WING. I @: 116. 1 9: J21.
TAM. TOs 8;
STOMACH CONTENTS. Small seeds and ground-up fruit.
TAXONOMY. This species shows much individual variation in
plumage, but within the limits of this variation Eastern Highlands
158
SPECIES ACCOUNTS
P. picta P. brehmii P. modesta P, madaraszi
large size large size small size small size
higher altitudes lower altitudes higher altitudes lower altitudes
Fic. 16. Reconstructed evolutionary history of Psittacella. An ancestral form
speciated to yield one large form and one small form, each of which again speciated
to yield a pair of similar-sized forms with mutually exclusive altitudinal ranges.
material is closest to the southeastern New Guinea race pallida. ‘Vhe
Mt. Karimui male has the head and upper breast darker than in any
other specimen available for comparison, but this character is sub-
ject to postmortem fading (Rand, 1942b, p. 452) and varies individ-
ually. The western race intermixta differs in that it is considerably
larger, while harterti females (Huon Peninsula) have a slightly paler
head and less extensive barring on the breast and flanks. ‘The race
buergersi from the Schraderberg is probably synonymous with pallida
(Gilliard and LeCroy, 1968).
BREEDING. ‘The testes of the Mt. Karimui male were slightly en-
larged. .
DISCUSSION. Psittacella brehmi is an uncommon and solitary
but widespread inhabitant of montane forest between about 5,300 ft
and 9,000 ft, and has been encountered sparingly by most collectors
at most localities within this zone (the claim of Gyldenstolpe, 1955,
p. 57, that “it inhabited grassland and shrub country” appears unique).
The individuals that I saw were 10 to 50 ft above the ground, slowly
walked along a branch supporting themselves by the legs and bill with-
out using the wings, and periodically stopped to bite at something.
‘They spent at least 10 minutes in the same tree and often remained
motionless for several minutes.
Records from the Eastern Highlands and other parts of New Guinea
indicate that the altitudinal ranges of P. brehmii and its similar-sized
high-altitude relative P. picta are nearly mutually exclusive, with the
transition altitude varying locally between 8,000 and 10,000 ft. On the
159
SPECIES ACCOUNTS
Huon Peninsula, where P. picta is absent, P. brehmii may extend up
to 3,800 meters (= 12,500 ft), as suggested by altitudes on labels of
specimens collected by Keysser and examined in the American Mu-
seum of Natural History. In plumage P. brehmii and P. picta are more
similar to each other than to their smaller relatives P. modesta and
P. madaraszi but the two large species differ more from each other
than do the two small species, suggesting that the former speciation
occurred earlier. In addition, the races of P. picta are quite distinct,
also indicating that it is a relatively old species.
VOICE. P. brehmii is generally silent, but on a single occasion I
saw it give a two-note call (Fig. 17). The first note was short and
harsh, and was followed at a 1/4-sec interval by a nasal note which first
rose and then fell in pitch.
Psittacella brehmii
ee
(harsh) (nasal)
} 1 sec ,
Fic. 17. Voice of Psittacella brehmii.
Psittacella picta excelsa Mayr and Gilliard
Painted ‘Tiger Parrot
NATIVE NAMES. Fore: tange.
SPECIMENS EXAMINED. Mt. Michael: 1 ? (9 July 1964).
WING. 109.
TAIL. 70:
STOMACH CONTENTS. Fruit and granular vegetable matter.
TAXONOMY. The Mt. Michael specimen has the crown, breast,
and edges of the primaries somewhat lighter and more brightly colored
than in excelsa from Mts. Hagen, Wilhelm, and Kubor, but these dif-
ferences are minor. It differs most obviously from nominate picta in
the olive-brown (instead of red-brown) crown and from lorentzi in
the olive-brown (instead of blue-green) throat and in the red (instead
of barred yellow) upper tail coverts.
160
SPECIES ACCOUNTS
DISCUSSION. ‘The specimen was netted at 10,200 ft in stunted
moss forest. The two individuals that I saw in the field were perched
15 or 20 ft above the ground in small trees. Paran said that this
“tdnge” (as opposed to P. brehmii, the other “tange’’) was absent from
the mountain behind Awande (about 7,500 ft high) but was present on
a higher mountain behind Okapa (somewhat over 8,000 ft high). Com-
bining this information with the collections of Gilliard and of Shaw-
Mayer, one can say that this parrot is a regular inhabitant of forest on
high mountains from elevations of about 8,000 ft up to at least 11,000
ft, and probably higher.
Relations of Psittacella madaraszi and P. modesta
Psittacella madaraszi and P. modesta were formerly considered con-
specific. Since the available evidence indicates that they are distinct
and since some specimens have been misidentified in the literature
of the last 15 years, the material in the American Museum of Natural
History and the British Museum (Natural History) was reviewed. ‘Vable
5 summarizes measurements.
The morphological differences are as follow. P. modesta is in general
larger (Table 5), but P. madaraszi increases in size from east to west,
so that the westernmost population, P. madaraszi major, 1s as large as
P. modesta. P. madaraszi has a stouter bill (culmen from cere 15 mm,
as opposed to 13 mm in P. modesta). The female plumages are distinct:
P. modesta females have orange-and-black barred breasts and brown
heads, while P. madaraszi females have no or only obscure barring on
the breast and have a green head with a blue forehead and a barred
crown and hindneck. Adult males of the two species are very similar
and have brown heads with yellow centers to the feathers and unbarred
breasts. A yellow collar is present in adult males of P. modesta except
for the nominate race, and is lacking in adult males of P. madaraszt.
In each species the immature male resembles the adult female. ‘There
are no morphologically intergrading specimens, although an intergrade
would be easy to recognize in the female.
Regarding recently collected material, all four of my Eastern High-
lands specimens are P. madarszi hallstromi. Gilliard collected four
specimens in the Eastern Highlands which he described as “P. modesta
hallstromv” (= P. madaraszi hallstromi). Examination of these speci-
mens shows that three of them, including the type of hallstromz, belong
to P. madarasz. ‘The fourth, an immature male collected at 7,000 ft
on Mt. Hagen and mentioned by Mayr and Gilliard (1954, p. 340), is
unmistakably P. modesta and has a barred breast, green-washed brown
head without yellow centers to the feathers, long wing (typical of
P. modesta, longer than any known specimen of P. madaraszi), and
lacks a yellow collar (present in adult males but lacking in females
and immature males of P. modesta). At ‘Velefolmin, Gilliard collected
161
SPECIES ACCOUNTS
TABLE 5
MEASUREMENTS OF Psittacella modesta AND Psittacella madaraszi
Wing (2)
P. modesta modesta
Vogelkop (28, 29)
P. modesta collaris
95; 96
Snow Mountains,
southern slopes [94, 96, 97, 98(3), 99, 102]a
P. modesta subcollaris
Snow Mountains,
northern slopes
(10g, 10Q)b [94-100]
Telefolmin (1 @) 95
P. modesta subsp.
Eastern Highlands
Go. bo)
P. madaraszi madaraszi
98, 99, 100, 101
Southeastern New
Guinea (18g, 69) 87(4), 88(2), 89, 20(2)5 913);
92(2), 93(3), 94
P. madaraszi huonensis
Huon Peninsula
(1g, 19Q)b 93
P. madaraszi hallstromi
Eastern Highlands
(4g,49Q)b
P. madaraszi subsp.
Telefolmin (29)
P. madaraszi_ major
91, [91 Ja, 92, [93], 94(2), [95], 96
Snow Mountains,
northern slopes
(19)
Weyland Mountains
(24,29) 797
* From published measurements of other authors.
b Including type.
Wing (©)
9395
[98, 98]a
[92-102]a
92
88; 93; 96
95, 98
95
[90], 94, 97
Altitude (ft)
5,500-5,900
5,500-8,000
5,900-9,200
7,300
7,000-9,500
6,000-7 900
4,000-5 ,000
5,200-8,000
5,850
5,300
4,300-5,600
one male and two females, which he listed as P. modesta subcollaris
(Gilliard and LeCroy, 1961, p. 41). Examination shows that the male
is P. modesta subcollaris and that the females are P. madaraszi. In the
Wahgi Valley Gyldenstolpe collected three males which I have not
examined but which were evidently P. madaraszi hallstromi. At 8,000-
9,500 ft in the Eastern Highlands Shaw-Mayer collected five specimens
which Sims (1956, p. 404) assigned to hallstromi. Examination of these
specimens in the British Museum (Natural History) shows that one is
a juvenile P. madaraszi hallstromi (4, wing 91, collected at 8,000 ft in
the Kubor Range, specimen number B.M. 1953.17.88) and that the
162
SPECIES ACCOUNTS
other four are P. modesta (é, wing 100, 8,500 ft, Mt. Giluwe, B.M.
1953.17.85; ¢, wing 99, 8,000 ft, Mt. Giluwe, B.M. 1953.17.86; ¢@, wing
102, 8,000 ft, Mt. Giluwe, B.M. 1953.17.87; ¢, wing 101, 9,500 ft,
Hagen Range, B.M. 1953.17.89). Shaw-Mayer collected an additional,
unreported specimen of P. madaraszi at 7,500 ft at Baiyanka on the
Purari-Ramu Divide (4, wing 94, B.M. 1942.1.6.7).
The distributional details are as follows. P. modesta, consisting of
the races collaris, subcollaris, and nominate modesta, occurs from the
Vogelkop in the extreme west to long. 145°E in the Eastern Highlands
but is absent farther east in the Herzog Mountains, southeastern New
Guinea, and the Huon Peninsula. P. madaraszi, consisting of the races
huonensis, hallstromi, major, and nominate madaraszi, occurs from
eastern New Guinea (southeastern New Guinea, Herzog Mountains,
Huon Peninsula) west along the Central Range to the Weyland Moun-
tains but is absent on the Vogelkop in the far west. The two species
are sympatric for a distance of at least 400 miles on the Central Range.
Their altitudinal ranges appear to be mutually exclusive, with P.
modesta living at higher altitudes than P. madaraszi. ‘Thus, in the
Snow Mountains the Third Archbold Expedition took P. modesta at
5,900-9,200 ft, P. madaraszi at 5,300 ft. At Telefolmin P. modesta was
taken at 7,300 ft, P. madaraszi at 5,850 ft. In the Eastern Highlands
P. modesta has been taken at 7,000-9,500 ft, P. madaraszi at 5,200-
8,000 ft.
‘The reconstructed evolutionary history is that P. modesta arose as
a western isolate, P. madaraszi as an eastern isolate, of what was
originally one small-sized Psittacella species. Each form has reinvaded
most of the other’s geographical range, with altitudinal segregation
as the ecological sorting mechanism, but P. modesta has not yet reached
the eastern tip of New Guinea, and P. madaraszi has not yet reached
the western tip. The birds of paradise Epimachus fastosus and E.
meyert have the same distribution and presumably the same history.
Psittacella modesta subsp.
Modest ‘Tiger Parrot
As discussed above, this species was taken at 7,000 ft on Mt. Hagen
by Gilliard, and at 8,000-9,500 ft on Mt. Giluwe and the Hagen Range
by Shaw-Mayer.
Psittacella madaraszi hallstromi Mayr and Gilliard
Madarasz’s ‘Tiger Parrot
NATIVE NAME. Fore: anko.
SPECIMENS EXAMINED. Awande: 2 ¢,1 9 (16-20 June 1965). Mt. Karimui
zone 5: 1 9 (30 Aug. 1965).
WRIGHT. 2 @: 415; 460, 2 9: 35,3, 36.0.
WING. 2 4: 94,96. 2 9: 89, 92.
163
SPECIES ACCOUNTS
CULMEN FROM CERE. 2 ¢: 15, 15. 1 9: 14.
STOMACH CONTENTS. Granular vegetable remains.
TAXONOMY. The Eastern Highlands race hallstromi differs from
nominate madaraszi of southeastern New Guinea in its slightly larger
size, in the slightly darker brown head and throat of the male, and in
the slightly darker green back and brighter red and darker black on
the nape of the female.
BREEDING. ‘The gonads were small in all specimens.
DISCUSSION. I have no observations of this parrot in life. All
were taken singly in forest between 6,000 and 6,500 ft, and one was
netted. As mentioned previously, Gilliard and Gyldenstolpe each ob-
tained three specimens, and Shaw-Mayer two.
Loriculus aurantufrons meeki Hartert
Bat Lorikeet
STOMACH CONTENTS. <A white paste or powder (sap?), and fruit.
DISCUSSION. ‘The Eastern Highlands records for this rare little
parrot are one female taken at Lake Kutubu (2,450 ft) and assigned to
meeki by Schodde and Hitchcock; one specimen collected at 4,000 ft
in the Lanim Valley by Bulmer; two Nondugl aviary birds examined
by Gyldenstolpe, said to be from the Jimmi River; and possible sight-
ings by ‘TVerborgh at Karimui in August 1964. ‘he wing beat is rapid,
fluttering, and atypical for a parrot.
CUCULIDAE: GCUCKOOS
Cuculus saturatus horsfieldi Horsfield and Moore
Oriental Cuckoo
Gilliard collected an adult male of this palearctic migrant at 5,200 ft
near Nondugl on 10 May, Bulmer had a possible sighting in Kyaka
territory at 4,600 ft in October, and there is one sighting from Baiyer
River (N.G.B.S. Newsletter, No. on pp» t, jume 1970):
Cacomantis vartiolosus oreophilus Hartert
Gray-breasted Brush Cuckoo
NATIVE NAMES. Fore: petetébeye. Daribi: sadewe bilong place (= “sadewe of
the village’: cf. Cacomantis castaneiventris).
SPECIMENS EXAMINED. Awande: 1 ¢ (17 June 1965). Karimui: 6 ¢,2 9,
limm. ¢, 1 imm. 9, 1 imm. ? (11 Aug. 1964; 3 July-4 Aug. 1965). Bomai: 1 ¢ (6
July 1965).
WEIGHT. 7 @: 32, 33, 35 (3), 36°); 2 QO: 35,40. bimm, ¢ 35. Jammy ?:
Sm. i amo. 2: 30.
WING. 7 4: 117 (2), 120, 121 @), 122, 125. 2 9: 2, 121. Timm, g: i2i. |
im. O: 117. 1 amin, re Ilo.
164
SPECIES ACCOUNTS
EXPOSED GULMEN,. 6 4: 15.5, 16:5, 17:0, 175, 18:5, 19.0; 2 @: 18:0; 19.0. 1
imm. ¢: 185. 1 imma. ? 14.5.
STOMACH CONTENTS. Insects; less often, insects and fruit.
TAXONOMY. These specimens agree with orcophilus in that the
bill is stouter than in infaustus. Of the three immature Karimui speci-
mens, the male is in nearly adult plumage, with barring remaining
only on the belly; the female is more immature, being entirely barred
below and barred in a few patches above; while the specimen of in-
determinate sex is entirely barred above and below.
BREEDING. Of the eight adult males, the Bomai specimen, the
Awande specimen, and three Karimui specimens had small testes, while
the other three Karimui specimens had the testes slightly enlarged.
Both adult females (from Karimui) contained large eggs nearly ready
to lay.
Paran shot the immature Karimui specimen of indeterminate sex
while it was being fed by two wren-warblers Malurus alboscapulatus,
which were also collected and proved to be two immature males. He
said that at Awande, too, “asasaba’s’ (Malurus alboscapulatus) were
the species that raised “petetébeye’s” (Cacomantis cuckoos). Malurus
alboscapulatus was clearly breeding in the dry season both at Awande
and at Karimui (p. 216). ‘The two laying females and the young im-
mature suggest that Cacomantis variolosus also breeds in the dry
season and has synchronized its breeding to that of its principal host.
DISCUSSION. ‘This cuckoo is conspicuous because of its song,
fairly common, and nearly ubiquitous in gardens, second-growth, trees
in open country, and towns, ranging up to about 6,300 ft in this
habitat. It has surely increased greatly in numbers and spread upward
somewhat as a result of clearing of the forest in the Eastern Highlands.
The preferred perches of C. variolosus are in the middlestory of trees
overlooking open spaces. It is most frequently seen at heights of
15-60 ft above the ground, rarely down to 10 ft or up into the crowns
of tall trees.
Ecological relationships among the three species of genus Cacomantis
in New Guinea seem to me simple. C. variolosus inhabits open country
from sea level up to about 4,000-6,300 ft, the upper limit depending
upon the extent of ecological disturbance caused by man. C. castanei-
ventris inhabits the forest interior over much of this same altitudinal
range and is found regularly at the upper part of this range in the
absence of man but is rare at sea level. C. pyrrhophanus inhabits both
second-growth and the forest interior from about 5,000 ft upward and
in effect takes over the niches of both its congeners at higher eleva-
tions. In undisturbed areas altitudinal exclusion of the other two
species by C. pyrrhophanus seems complete, though there may be
marginal overlap in disturbed areas.
VOICE. ‘There are two songs, both unmistakable (Fig. 18). ‘The
first is a series of a half dozen similar three-note phrases, the series
165
SPECIES ACCOUNTS
progressively rising in pitch, accelerating, and becoming louder, as if
the singer were becoming progressively more excited. Each three-note
phrase is in dotted rhythm and can be represented by the syllables
“pe: - -te-to”, giving rise to the Fore name “petetdbeye’. The whole
seriés consists of four to six repetitions of this basic phrase and lasts
up to 5 sec. The other song, which is similar in form to the songs of
several species of Chrysococcyx, has the same quality but consists of a
series of nearly identical notes delivered at the rate of one per second,
each note dropping in pitch at the end. The series may remain at
constant volume and on constant pitch. It may also progressively drop
slightly in pitch and grow louder without accelerating. A bird deliver-
ing the second song may break into the “pe-- -te-to” song and then re-
turn to the second song, giving the impression of an idling engine
occasionally being set into motion. Both songs have the same whistled,
shrill quality, carry well in open country, and are among the most
characteristic sounds of Karimui station. Once I heard the “pe- - -te-to”
song from a bird still in immature plumage.
Cacomantis pyrrhophanus prionurus (Lichtenstein)
and C. p. excitus Rothschild and Hartert
Fan-tailed Brush Cuckoo
SPECIMENS EXAMINED. Awande: 1 ¢@ (17 June 1965).
WEIGHT. 50.
WING. 138.
TAIL. 140.
STOMACH CONTENTS. Insects.
TAXONOMY. ‘The Awande specimen belongs to the Australian
race prionurus, recognized by its pale cinnamon underparts and dull
gray back. It had a small ovary and must have been a winter visitor.
‘Three specimens collected at Nondugl by Gyldenstolpe in August
and September 1950 provided the first, and only other, record of this
race from New Guinea. It has also been found on migration to the
Aru Islands. ‘The resident New Guinea race excitus is quite distinct
in its dark rufous-brown underparts and dark shining slaty dorsal
coloration.
BREEDING. Bulmer recorded one egg in a nest of the wren-
warbler Malurus alboscapulatus, which is the principal host of Caco-
mantis variolosus at lower elevations; and I observed a juvenile being
fed by the warbler Sericornis nouhuysi.
DISCUSSION. Gilliard, Gyldenstolpe, Shaw-Mayer, and Bulmer
each collected one or two specimens of C. p. excitus, and Gilliard re-
ported it “very common,” but this is probably true only very locally,
as I did not find it either on Mt. Michael or on Mt. Karimui and
other observers have generally found it rare. In the Eastern Highlands
its altitudinal range is about 5,000 to 9,500 ft, largely above that of the
166
Cacomantis variolosus:
(qd) <n. 7 eer
pe teto pe teto etc.
Cacomantis castaneiventris (7):
rae
a
3 SEC
a
Eudynamis scolopacea :
2 Séc
Fic. 18. Voices of three New Guinea cuckoos,
SPECIES ACCOUNTS
other two Cacomantis species. Individuals that I observed were solitary
and perched either on the ground or on stumps in forest clearings,
or else on large branches up to 50 ft above the ground in open forest.
Most of the time they were motionless and perched upright in a
thrush-like posture, but one hovered for a few seconds to pluck some-
thing from the foliage.
VOICE. A rapid, plaintive trill, descending slightly in pitch, and
similar to some calls of the kingfishers Halcyon megarhyncha and H.
torotoro but weaker.
Cacomantis castaneiventris weisket Reichenow
Chestnut-breasted Brush Cuckoo
NATIVE NAMES. Fore: petetédbeye. Gimi: fétote. Daribi: sidewe bilong bush
(= “sadewe of the forest”: cf. Cacomantis variolosus).
SPECIMENS EXAMINED. Mengino: 1 @ (15 July 1964). Karimui: 3 ¢, 3 9,
limm. ¢ (2 Aug. 1964; 3 July-4 Aug. 1965). Soliabeda: 1 ¢, 1 imm. 9 (24 and 30
July 1965). Mt. Karimui Zone 1: 1 ¢ (12 Aug. 1965).
WRHRIGEIE. 4 ° 4:32, 36, 37, 58. 3 9: 95, 37, 38, Vamim: 2227, Timm. O34
WING.” 4s 10, 1 7, 119. SO 07, 2s WAS Sosa 0 a mot
Qs UGH
TAXONOMY. ‘The wing measurements place this series with
wetskei, which differs from arfakianus only in its slightly larger size.
BREEDING. ‘The gonads were small except in the Soliabeda male
and a Karimui male and female, which had the gonads slightly en-
larged.
DISCUSSION. All 11 specimens were obtained inside the forest,
and seven of them were netted, suggesting that this cuckoo spends
much time in the understory. One immature male and subadult female
were found together in the same net. I never saw the species in life,
but song records and reports of natives indicate that it is uncommon,
local, and shy, and lives in forest up to about 6,000 ft.
VOICE. Natives attribute a mournful, slow, three-note song at
medium-low pitch to this species. ‘he pattern is distinctive: the first
and third notes are leisurely and upslurred, the second note shorter,
and the first note is higher in pitch and separated by a longer time in-
terval from the second and third (Fig. 18, p. 167). The identification is
consistent with the three-note songs described for this species by Stein
(1936, p. 49) and by Ripley (1964, p. 36). On the north coast of New
Guinea I heard the same song differing only in that the final note was
trilled, so that the identification may still be compatible with Rand’s
description (Mayr and Rand, 1937, p. 63) of the song of C. castanei-
ventris as a “full rich trill.”
168
SPECIES ACCOUNTS
Genus Chrysococcyx!
The three resident New Guinea species form an altitudinal sequence
in forest, with C. ruficollis at the highest elevations, C. meyerii at
middle elevations, and C. malayanus (not yet reported from the Eastern
Highlands) at the lowest elevations. An additional species, C. lucidus,
occurs in open country in the Eastern Highlands as a winter visitor
from Australia.
Chrysococcyx lucidus plagosus (Latham)
Golden Bronze Cuckoo
SPECIMEN EXAMINED. Lufa: 1 9 (29 June 1964).
DISCUSSION. My sole specimen of this Australian winter visitor
was shot from a perch 20 ft above the ground in a lawn casuarina at
Lufa Patrol Post (6,300 ft). A modest number of specimens have been
obtained by others (four at Nondugl by Gyldenstolpe, one each at
Nondugl and in the Kubor Mountains by Gilliard, four in Kyaka
territory by Bulmer), suggesting that the species’ appearances in the
Eastern Highlands are regular.
Chrysococcyx ruficollis (Salvadori)
Mountain Bronze Cuckoo
I did not encounter this cuckoo, which appears to be rare and local
in mountain forest between 6,000 and 8,500 ft. Ihe known records in
the Eastern Highlands are two specimens collected in the Kubor Range
(one each by Gilliard at 7,000 ft and by Hitchcock at 6,400 ft), one on
Mt. Wilhelm at 8,500 ft by Gilliard, and one at 6,000 ft on the north
slope of Mt. Hagen by Bulmer.
Chrysococcyx meyerit Salvadori
Meyer’s Bronze Cuckoo
SPECIMENS EXAMINED. Awande: 1 9 (19 June 1965). Soliabeda: 1 @ (23
July 1965).
WEIGHT. 1 3: 19.3. 1 9: 20.7.
WING. 1 4:90. 1 9: 92.
DISCUSSION. Meyer's Bronze Cuckoo is solitary and uncommon
but widespread up to 5,000 ft, rarely up to 6,000 ft, and has been
encountered by most collectors. It is found in the forest or at the forest
edge but seems to prefer the latter. In the field this cuckoo suggests a
sluggish flycatcher in its behavior. It moves slowly in the middlestory,
usually 15 to 40 ft above the ground, catching insects.
1 Listed as Chalcites in Rand and Gilliard (1967).
169
SPECIES ACCOUNTS
VOICE. A series of six or seven identical notes at intervals of
about | sec or less, each note dropping slightly in pitch at the end.
The ‘pitch is high, the quality plaintive and clear, the volume soft. In
pattern this resembles the second song of Cacomantis variolosus but is
much weaker in volume and slightly more leisurely in tempo, pre-
cluding the possibility of confusion. The song of Chrysococcyx lucidus
lucidus, which I heard in New Zealand, is similar in pattern. Published
descriptions of the songs of C. ruficollis (Stein, 1936), C. basalis (Cayley,
1959), and C. lucidus plagosus of Australia (Cayley, 1959) are also
similar, but that of C. malayanus (Cayley, 1959) is apparently quite
different.
Eudynamis parva parva Salvadori!
Black-capped Koel
SPECIMENS EXAMINED. Mengino: 1 imm. ¢ (15 July 1964). Okasa: 1 ¢,19
(a) (24 June 1965). Karimui: 1 @ (b) (4 Aug. 1965).
WEIGHT. 1 4:40. 2 @: 49 (a), 40 (b).
WING. 1 62102, 2 92 104); 100\(b)., 1 amam:. 4; 105;
STOMACH CONTENTS. Fruit.
‘TAXONOMY. ‘The coloration of these birds is not gray but in-
stead is rufous, placing them with the race nominate parva (grisescens
is restricted to northern New Guinea). The iris was red. ‘The immature
male from Mengino has the throat already orange-brown, as in the
adult male, rather than gray, as in the female and immature male;
the mustache stripe is charcoal but not yet the steely blue of the adult
male; the gray stripe between the eye and mustache is already
ochraceous posteriorly; the crown is still dark brown, with just a few
scattered steely blue feathers.
BREEDING. ‘The Okasa male and female, a pair shot from the
same tree, both had enlarged gonads.
DISCUSSION. ‘The above specimens constitute the sole Eastern
Highlands records of this very uncommon cuckoo. ‘The Mengino
specimen was shot 50 ft above the ground in a fruit tree at 4,600 ft in
open second-growth forest near a river, at the upper limit of the
species’ altitudinal range. In addition to these specimens, a pair was
seen in the middlestory of a tree at Karimui.
VOICE. ‘The song, delivered from a treetop perch, is similar to the
common song of Eudynamis scolopacea (Fig. 18, p. 167) but weaker: a
series of pairs of notes, each pair louder and on a higher pitch than
the previous pair. ‘The elements are sometimes three repeated notes
or else single downslurs rather than a pair of notes. Another variant
is a series of identical upslurs at 114-sec intervals, similar to the voice
of Cacomantis variolosus in quality.
1 Listed as Microdynamis parva in Rand and Gilliard (1967); see Mayr (1944).
170
SPECIES ACCOUNTS
Eudynamis scolopacea rufiwenter (Lesson)
Indian Koel
SPECIMENS EXAMINED. Karimui: 1 ¢ (11 Sept. 1965).
WING. 185.
STOMACH CONTENTS. Fruit, 9-41 mm in diameter.
TAXONOMY. The small size places the specimen with rufiventer,
the breeding race of most of New Guinea, rather than with swbcyano-
cephala, which is in southern New Guinea as a wintering visitor from
Australia and as a resident (Rand, 1941). Females of these two races
are quite distinct in plumage. Cuckoos agreeing closely with the female
of rufiventer were seen but not collected at Karimui.
BREEDING. ‘The testes were somewhat enlarged.
DISCUSSION. ‘The Indian Koel was heard or seen nine times at
Karimui in August 1964. In 1965 the only records were of the specimen
(11 Sept.) and of a song heard on 15 Sept. I was particularly interested
in tracing the unmistakable song and am certain that I did not hear
it in the dry months of June, July, or August 1965. Probably local
migration within New Guinea is involved. All Koels seen were solitary,
and were either feeding in the middlestory of fruit trees, or else sing-
ing from commanding perches at the tops of very tall trees overlooking
open spaces. Eastern Highlands records of this tropical species are
lacking except at Karimui.
VOICE. ‘There are four types of vocalizations, all loud. ‘The com-
mon song is a series of about a dozen pairs of notes, the first note of
each pair on the same pitch as the second but shorter (Fig. 18, p. 167).
Successive pairs rise in pitch and increase slightly in volume. The whole
song lasts about 5 sec. Another song consists of three upslurs, the
second following 2 sec after the first and on the same pitch, the third
1 sec after the second and on a slightly higher pitch (Fig. 18). Each
upslur has a sense of forward momentum and sounds as if it came from
a bird much larger than the Koel. A single such slur is sometimes given
as a call. ‘he common alarm note is a rapidly repeated, musical,
emphatic, staccato note “whik-whik-whik .. .”” reminescent of Para-
disaea minor. An alternative alarm note is a disagreeable upslur.
Caliechthrus leucolophus (Muller)
White-crowned Koel
NATIVE NAME: Daribi: poiba.
SPECIMENS EXAMINED: Karimui: 1 ¢ (17 July 1965). Mt. Karimui Zone
4:1 ¢ (28 Aug., 1965).
WEIGHT: 26% 113, 125.
WING...2 @: 169, 175.
STOMACH CONTENTS. Large insects.
BREEDING. Both specimens had small testes.
171
SPECIES ACCOUNTS
DISCUSSION. ‘These two specimens provide the sole records for
this low-altitude treetop cuckoo in the Eastern Highlands. Elsewhere
in New Guinea, too, the species is local, being common in a few areas
and absent in others. The Karimui bird was perched alone in a tree
at the edge of a clearing. The Mt. Karimui bird, one of two in ad-
Jacent trees in primary forest, was collected at 5,700 ft, which is
unusually high for the species. Daribi natives said that it occurs
regularly in the Karimui area.
VOICE. According to natives, three or four loud rapid notes on
the same pitch.
Centropus menbeki menbeki Lesson
Greater Coucal
NATIVE NAME. Daribi: pini.
SPECIMENS EXAMINED. Karimui: 2 ¢, 1 9 (13 and 14 July 1965). Bomai:
1 2 (7 July 1965). Soliabeda: 1 ? (22 July 1965).
WING. 2 @: 212, 243. 2 9: 230, 235.
TAIL. 2 @5 343,366, 2°02 340, 345.
EXPOSED CULMEN. 2 @: 46,47. 2 9: 49, 51.
CULMEN FROM BASE. 2 ¢@: 49,52. 2 9:57, 58.
STOMACH CONTENTS. Insects.
TAXONOMY. All specimens have the plumage unbarred. ‘The
iris was red, and the bill whitish.
BREEDING. Both males had two testes, which were small.
DISCUSSION. ‘There are no Eastern Highlands records for the
Greater Coucal other than from the Karimui area, where it was
uncommon. The presence at Karimui (3,650 ft) and on the lower slopes
of Mt. Karimui (up to 4,200 ft) is somewhat above the normal altitudi-
nal range of the species. It ranges from the ground to about 50 ft
above the ground, and has a characteristic habit of clumsily hopping
up the trunk of a small tree, switching its long tail from side to side
as it goes. The Soliabeda specimen was surprised while trying to eat
two small birds out of our mistnets.
Although C. menbeki and C. phasianinus both may be found in
second-growth, their average habitat preferences are different: C.
phasianinus is mainly found in grassland, while C. menbeki (but never
C. phasianinus) occurs in the forest. In addition to these differences in
habitat preference, it is surprising how infrequently the two species
even occur in the same locality.
VOICE. The common call is a series of about four hollow, boom-
ing, sometimes disyllabic notes “hoo” or “coo-a”, similar in quality to
the calls of Macropygia nigrirostris or species of Ptilinopus. This call
may be heard for some time after sunset. Less frequently heard is a
peculiar and totally unmusical call consisting of a staccato upslurred
172
SPECIES ACCOUNTS
grunt like the croak of a bullfrog, followed by a dry, rapid, rattled
series of notes rising in pitch.
Centropus phasianinus propinguus Mayr
Pheasant Coucal
DISCUSSION. ‘The only definite Eastern Highlands records of this
grassland coucal are from the grassland of the Baiyer Valley, where
Gilliard and Bulmer collected it and Bell found it common up to
about 5,000 ft. In addition, native informants told Bulmer that it
occurs in the Kaironk Valley of the Schrader Range up to 5,500 ft and
near Banz. Elsewhere in New Guinea C. phasianinus is confined to
altitudes generally below 2,500 ft, and it has not colonized the Baliem
Valley grasslands of western New Guinea. However, the large expanses
of grassland in the Eastern Highlands appear to be ideal habitat for
it, and it will be interesting to see whether it succeeds in colonizing
other parts of the Eastern Highlands besides the Baiyer Valley.
VOICE. A series of dull, hollow, booming notes which accelerate
in tempo, similar in quality to the call of C. menbeki.
TYTONIDAE: BARN OWLS
Niche Differences in the Genus Tyto
‘The three species of Tyto in the Eastern Highlands sort out by
habitat: 7. tenebricosa in forest (and alpine grassland), 7. alba in
disturbed habitats, T. capensis in midmontane grassland.
Tyto alba meeki (Rothschild and Hartert)
Barn Owl
NATIVE NAME. Fore: waiyimpémpo.
SPECIMENS EXAMINED. Awande: 2 6 (1 and 4 July 1967).
WING. 276, 278.
DISCUSSION. ‘The specimens were collected from second-growth
trees in the Okapa area where I also saw an individual at sunset flying
at the edge of the forest and uttering monosyllabic screams. The Barn
Owl is generally distributed in low numbers through the Eastern High-
lands (other records by Gilliard, Gyldenstolpe, and Bulmer).
Tyto tenebricosa arfaki (Schlegel)
Sooty Owl
NATIVE NAME. Fore: ési.
STOMACH CONTENTS. A ball of hair with skulls of small mammals.
173
SPECIES ACCOUNTS
DISCUSSION. The Fore knew of only one pair of this little-known
forest barn owl living within a wide radius of Okapa. Shaw-Mayer col-
lected one individual each at 8,500 ft in the Hagen Range and the
Lamende Range, Gilliard collected a young bird at 7,000 ft in the
Kubor Range, and there are records of two birds brought into Nondugl
aviaries. On Mt. Albert-Edward I found an individual sleeping during
the day in a hole under the roots of a forest tree.
Van Deusen (pers. comm.) analysed bones in the regurgitated pellets
of two individuals of Tyto tenebricosa and kindly provided the follow-
ing details, which offer valuable insight into its diet. The first set of
pellets was found under a boulder in alpine grassland on Mt. Wilhelm
at 12,000 ft. The three major prey species were the rats Rattus niobe
and Pogonomelomys sp., and, to a lesser extent, the bandicoot Pero-
ryctes longicauda. All three live in alpine grassland. ‘There were also
a few bones of the small marsupial Eudromicia caudata from the ad-
jacent subalpine forest, the rats Neohydromys sp. and Pogonomys sp.,
the large wallaby Thylogale bruijni (probably taken as carrion from
wild dog kills), and an unidentified bird. The second set of pellets came
from a boulder roost at the edge between forest and grassland at 7,800
ft in the Cromwell Mountains. ‘The dominant prey species proved to
be the common arboreal forest marsupial Psewdocheirus forbes. Other
prey included the giant rats Hyomys sp. and Mallomys sp., the small
forest rat Anisomys sp., the rat Rattus exulans, three forest marsupials
(the bandicoot Peroryctes raffrayanus, the flying phalanger Petaurus
breviceps, the wallaby Dorcopsulus vanheurnz), the large wallaby Thy-
logale bruijni (probably taken as carrion), the forest bats Nyctimene
sp. and Syconycteris sp., and unidentified birds.
VOICE. According to natives, a long, high-pitched, slightly down-
slurred note, suggestive of the neighing of a horse.
Tyto capensis papuensis Hartert
Grass Owl
Of the three barn owls occurring in the Eastern Highlands, this
seems to be the least uncommon. Gilliard, Gyldenstolpe, and Shaw-
Mayer all observed it over grassland and obtained two specimens each.
It has also been discovered in western New Guinea in the grassland
of the Baliem Valley by Ripley (1964).
SE RIGIDAK: OW Ls
Ninox theomacha theomacha (Bonaparte)
Papuan Boobook Owl
NATIVE NAME. Fore: atoku.
SPECIMENS EXAMINED. Karimui: 1 @ (12 July 1965).
174
SPECIES ACCOUNTS
WING. 186.
STOMACH CONTENTS. Insects 25 mm long, and a few mammal hairs.
BREEDING. The testes were small.
DISCUSSION. The specimen was caught in a mistnet inside the
forest at Karimui. I heard the call of this owl, which is the commonest
nocturnal bird call in New Guinea forests, from the forest and forest
edge up to 6,450 ft on Mt. Karimui. Other Eastern Highlands records
are from Lake Kutubu (Schodde and Hitchcock), the Baiyer Valley
(Bulmer), Kup (Gilliard), and 8,200 ft in the Hagen Range (Shaw-
Mayer).
VOICE. ‘Two identical downslurred notes at an interval of slightly
less than | sec, resembling the call of Ninox novaeseelandiae which I
heard in New Zealand. On different occasions I recorded the call as
“kyew kyew” or “hyew-ew hyew-ew” or “kru kru.”
PODARGIDAE: FROGMOUTHS
Podargus papuensis Quoy and Gaimard
Great Papuan Frogmouth
NATIVE NAMES. Fore: yasa. Daribi: uni.
DISCUSSION. Podargus papuensis is rarely seen but is widespread
through the Eastern Highlands in low numbers in forest and in 1so-
lated trees up to about 7,000 ft. My sole observation was in daylight
at Karimui, when my Fore assistants pointed out to me a bird which
had been disturbed by their approach. It was perched motionless and
upright, with its head turned towards us, 30 ft above the ground in a
tall tree at the edge of a clearing. ‘This frogmouth sometimes sits with
its mouth wide open for long periods, with a sticky substance on its
palate that might serve to attract or trap insects.
Podargus ocellatus ocellatus Quoy and Gaimard
Little Papuan Frogmouth
NATIVE NAME. Fore: atoku.
SPECIMENS EXAMINED. Okasa: 1 ¢ (22 June 1965). Karimui: 4 9 (15 Aug.
1964; 3-16 July 1965). Bomai: 1 9 (9 July 1965).
WEIGHT. 2 9: 140, 141.
WING, I 4: 84. boc 1s, 17/6, 278; 179) 187.
TAXONOMY. All specimens are in the red-brown phase.
BREEDING. All had small gonads, suggesting that breeding is
limited to the wet season.
DISCUSSION. ‘Two of the specimens were netted in the forest; one
was caught by hand while sleeping on a branch during the day; and
three were shot also while sleeping on branches during the day, one
175
SPECIES ACCOUNTS
of these being in a group of three. I never observed this frogmouth
in the field, but, to judge from the number of specimens, it may be
the least uncommon nocturnal bird in forest up to about 5,000 ft.
Bulmer and Gyldenstolpe secured one specimen each.
AEGOTHELIDAE: OWLET-NIGHTJARS
Aegotheles bennettii terborghi Diamond
Collared Owlet-nightjar
SPECIMENS EXAMINED. Karimui: 1 ¢ (16 Aug. 1964).
WING. 154.
TAME. 22:
DISCUSSION. ‘The unique type of this very distinct race was
caught by hand by a Karimui native, who found it sleeping in a tree
during the day. It differs from other races most notably in its larger
size (wing and tail 25% longer than in nominate bennetti, the other
southern New Guinea race) and in its much darker, blacker upper-
parts (see Diamond, 1967a, p. 6 for diagnosis). It represents an altitudi-
nal record for this lowland species, and is of interest in illustrating
evolutionary divergence in lowland birds isolated in the Karimui
Basin as well as the tendency of these populations towards darker
plumage (p. 56).
Aegotheles wallacii wallacii Gray
Wallace’s Owlet-nightjar
SPECIMENS EXAMINED. Karimui: 1 @ (1 July 1965).
WEIGHT. 48.5.
WING. 122.
TAL. 169.
TAXONOMY. Comparative measurements of all 23 specimens
known of this species have been summarized elsewhere (Diamond,
1969). The Karimui specimen completely lacks a light collar on the
hindneck, of which there are suggestions in some gigas and manni
specimens. The light buffy V-shaped mark on the forehead is about as
distinct as in the one available specimen of nominate wallacii from
the Vogelkop and more distinct than in the one available wallaci
specimen from the Aru Islands. ‘The lower belly is palest and has the
fewest markings in the Vogelkop specimen. ‘The loose arrangement of
white dots on the undersurface of the tail into bands is least distinct
in the Karimui specimen.
DISCUSSION. ‘This is a rare species with a very local distribution,
recorded near sea level in the Aru Islands, southwestern New Guinea,
and the Vogelkop and at medium elevations in the Weyland Moun-
176
SPECIES ACCOUNTS
tains, North Coastal Range, and Karimui. ‘The Karimui specunen
represents a range extension to the east.
Aegotheles albertisti salvadoriu Hartert
Mountain Owlet-nightjar
NATIVE NAMES. Fore: atéku. Gimi: mautla.
SPECIMENS EXAMINED. Awande: 1 9, 1 juv. ? (15 and 18 June 1965). Okasa:
2 6 (22 and 25 June 1965).
WEIGHT. 2 4: 36, 39. 1 9: 37. 1 juv. ?: 25.
WING, 2 a2 12, Te 1 oy 123. 1 jov. 7-106.
PAR. 202 80 + 96.. L yay. 72246;
STOMACH CONTENTS. Insects.
TAXONOMY. The type of Aegotheles archboldi differs in its
richer dorsal coloration, coarser white spots on the back, and coarser
and darker breast markings. Since both A. albertisi: and A. archboldi
exist in two color phases and show much individual variation and
since the complex plumage patterns defy simple description, identifica-
tion in the absence of comparative material is impossible.
BREEDING. ‘The testes were large in one Okasa male, small in the
other.
DISCUSSION. ‘This aegothelid was also collected by Gilliard, Gyl-
denstolpe, and Shaw-Meyer, and is common but secretive in forest
from about 4,500 to 8,500 ft. ‘Iwo of the adults were netted, two others
were seen flying out of holes in trees when disturbed during the day,
and the juvenile was caught by hand in a garden.
The relations among the small midmontane forest aegothelids of the
albertisit-archboldi complex remain to be clarified. A. archboldi was
originally described as a race of A. albertisii. Proceeding from west to
east in the mountains of New Guinea, the distribution of the popula-
tions is as follows: A. albertisi albertisti, Vogelkop; A. albertisti
wondiwoi, Wandammen Peninsula; A. albertisii salvadoriit, Weyland
Mountains to Wissel Lakes; A. archboldi, Wissel Lakes to Mt. Wil-
helmina; A. albertisi salvadoru, Mt. Goliath; A. archboldi, Tele-
folmin; and A. albertisit salvadori, Eastern Highlands, Huon Penin-
sula, and southeast New Guinea. The only locality where both A.
albertissi and A. archboldi have been taken is the Wissel Lakes. As
Ripley (1964) points out, the altitudinal range of A. albertisti (ca.
3,800-8,600 ft, combining various records in the literature) is on the
average lower than that of A. archboldi (ca. 7,000-12,000 ft, except for
a native-procured ‘Telefolmin specimen listed (Gilliard and LeCroy,
1961, p. 43) as from 4,800 ft but probably from an unknown eleva-
tion. Thus, the geographical distributions are disjunct and largely
but not completely complementary (“checkerboard allopatry,” p. 39),
and the altitudinal ranges are possibly complementary in the areas of
local overlap.
SPECIES ACCOUNTS
Aegotheles insignis insignis Salvadori
Large Owlet-nightjar
NATIVE NAMES. Fore: atéku. Gimi: maula.
SPECIMENS EXAMINED. Mt. Karimui Zone 3: 1 03 Zone 4: 1 ss-Zone 81 As
I Q (17 Aug.-6 Sept. 1965).
WEIGHT. 243 75,79. 2 On Ge.
WING, 24: 157, 163; 2 ©: 160, 165.
STOMACH CONTENTS. Insects.
TAXONOMY. Three of the specimens are in the dark phase,
while the Zone 8 male is intermediate between the dark phase and
light phase. Gilliard and LeCroy (1961, p. 43) point out that dark
phase plumage probably succeeds light phase plumage in the same
individual.
BREEDING. Both males had the testes slightly enlarged.
DISCUSSION. All four specimens were caught in mistnets between
5,000 and 7,900 ft. Other collectors in the Eastern Highlands have also
found the species at most collecting stations in forest between about
4,000 and 8,500 ft, a range which d. insignis shares with A. albertisit.
Niche differences probably are related to size, since A. insignis is twice
as heavy as A. albertisit.
CAPRIMULGIDAE: GOATSUCKERS
Caprimulgus macrurus yorki Mathews
White-tailed Nightjar
NATIVE NAMES. Fore: k’ru-k’runta.
STOMACH CONTENTS. Insects.
BREEDING. Three “nests” were found: sets of two coffee-colored
eges with sparse, small, ight grey-brown blotches, measuring 30-34
22-23 mm, laid directly on the ground.
DISCUSSION. ‘This goatsucker was heard calling from grassland
and gardens at Miarosa (5,800 ft), Okapa (6,600 ft), and Mengino
(6,150 ft), but not at my other stations. In particular, despite the vast
expanses of grassland at Okasa, and despite the number of lowland
species that have colonized Karimui, I found it at neither locality and
presume it to be absent because its distinctive, frequently-given call
could not have been missed. ‘The experience of other observers in the
Eastern Highlands indicates an equally spotty distribution. ‘The clear-
ing of the forest has permitted this lowland species to spread up to the
newly created midmontane grassland not only in the Eastern High-
lands but also at Telefolmin (Gilliard and LeCroy, 1961, p. 43) and in
the Baliem Valley of western New Guinea (Rand, 1942b, p. 457;
Ripley, 1964, p. 40). It emerges just after sunset to hawk for insects in
cleared areas.
178
SPECIES ACCOUNTS
VOICE. Several (usually two or three) identical metallic notes
“tuck” separated by intervals of slightly less than 1 sec. The quality
may be compared to a human clicking the tongue against the palate,
or to an anvil being struck with a hammer. ‘The call carries a half mile
across the grassland on a still night.
Eurostopodus mystacalis mystacalis (Temminck)!
White-throated Nightjar
SPECIMENS EXAMINED. Soliabeda: 1 ¢ (21 July 1965).
WEIGHT. 145.
WING. 234.
TAIL. 148.
BREEDING. ‘The testes were small.
DISCUSSION. ‘The specimen was shot while hawking over Solia-
beda village at dusk. The only other Eastern Highlands record of this
Australian migrant is of a single individual taken at Kup (5,000 ft)
on 15 May 1952 (Mayr and Gilliard, 1954). All New Guinea records
are from the eastern half of the island except for one bird taken at
the Idenburg River by the Third Archbold Expedition. At Lake
Kutubu Schodde and Hitchcock observed Eurostopodus nightjars with-
out large white marks in the tail or wing and tentatively identified
them as E. papuensis, but this description might also fit E. mystacalis.
Eurostopodus archboldi (Mayr and Rand)
Archbold’s Nightjar
STOMACH CONTENTS. Large insects, including a cicada.
DISCUSSION. Four specimens of this rare high-altitude nightjar
have been taken in the Eastern Highlands: one at 10,000 ft in moss
forest on Mt. Hagen (Mayr and Gilliard, 1954, p. 342), one at 8,000 ft
on Mt. Hagen, and two at 7,300-7,500 ft on Mt. Giluwe (Sims, 1956,
407). In 1964 the Seventh Archbold Expedition obtained a single
specimen on the Huon Peninsula, whence the species had not pre-
viously been recorded. Only eight other specimens have been reported:
the type series of three from 7,900 ft on Mt. Tafa, southeastern New
Guinea (Mayr and Rand, 1935, p. 4); a specimen I collected at 8,200 ft
on Mt. Albert-Edward, southeastern New Guinea; three from about
10,600 ft in the Lake Habbema area of western New Guinea (Rand,
1942b, p. 457); and one from 7,500 ft in the Ilaga Valley of western
New Guinea (Ripley, 1964, p. 40). The geographical range is thus
extensive, but the distribution is extremely spotty. My Mt. Albert-
Edward specimen was one of a pair hawking silently in a forest clearing
at dusk, flying from just above the treetops to about 30 ft above the
ground and periodically perching on a dead tree.
1 Listed as Eurostopodus albogularis in Rand and Gilliard (1967).
179
SPECIES ACCOUNTS
APODIDAE: SWIFTS
[Chaetura novaeguineae subsp. |
New Guinea Spine-tailed Swift
Gyldenstolpe (1955, p. 69) included this species in the Eastern High-
lands avifauna on the strength of seeing “numbers of large swifts—
presumably representing the above-mentioned species” along with the
smaller Collocalia esculenta at Nondugl. Chaetwra novaeguineae has
apparently been collected only in the lowlands, whereas another large
swift, Collocalia whiteheadi, has been collected in the Wahgi Valley.
The sight record of Chaetura novaeguineae based on size is therefore
presumed to be a case of misidentification.
Collocalia esculenta esculenta (Linnaeus)
Glossy Swiftlet
NATIVE NAME. Fore: kisabe.
SPECIMENS EXAMINED. Awande (6,000 ft): 1 g (25 Aug., 1964). Sena River
(4,500 ft): 1 Q (27 July 1964). Soliabeda (1,350 ft): 1 ? (26 July 1965). Mt. Karimui
Zone 5 (6,400 ft): 1 ¢ (28, Aug. 1965).
WEIGHT. 14 (6400 ft)8.0) 1 (sb 0tt): 47-3:
WING. 2 4: 106 (6,000 ft), 109 (6,400 ft). 1 9: 102 (4,500 ft). 1 ?: 98 (1,500 ft).
STOMACH CONTENTS. Insects.
‘TAXONOMY. ‘This small series illustrates the clinal increase in
size with altitude which emerges more clearly from larger series (e.g.,
Archbold, Rand, and Brass, 1942, p. 285).
BREEDING. ‘The gonads were small in these specimens. Three
nests were found under overhanging boulders beside streams. ‘The nests
were crude cups of moss 10 cm in diameter and 8 cm deep, and con-
tained two young. An adult gathered moss for one of these nests by
repeatedly hovering to pick it off the bark of a tree.
DISCUSSION. ‘The habitat preferences of this swiftlet depend
upon altitude. Above 5,000 ft C. esculenta may be seen over almost any
kind of habitat (midmontane grassland, gardens, second-growth, pri-
mary forest, alpine grassland) up to 12,000 ft. At these higher eleva-
tions it sometimes forms mixed flocks with C. hirundinacea, but on
the average C. esculenta forages nearer the ground and closer to
vegetation, C. hirundinacea usually higher and more in the open,
though under misty conditions GC. hirundinacea may also forage within
a few feet of the ground. C. esculenta generally flies below the level of
the treetops, occasionally even inside the forest in more open areas,
and circles and skims close to the foliage. In the hill forest below 4,500
ft in the parts of the Eastern Highlands I studied, C. esculenta is strik-
ingly and absolutely confined to river gorges. ‘hus, in the vicinity of
Karimui (3,650 ft) and Bomai (3,250 ft) airstrips C. hirwndinacea was
present daily in flocks of dozens, sometimes over a hundred birds, at
180
SPECIES ACCOUNTS
any height down to nearly ground level, but not once did I see C.
esculenta in these flat cleared areas. A few hundred feet away C. escu-
lenta reappeared as the only swift in the river gorges at the edge of
these two airstrips. Similarly, C. esculenta was the only swift at our
4,500 ft camp on the Sena River, in the river gorges between Karimut
and Soliabeda, and in the Wi River gorge at 1,350 ft below Soliabeda,
where the one Soliabeda specimen was collected, but the species was
absent from the rolling terrain 500 ft above the Wi River, where
Soliabeda village itself was located. At Lake Kutubu (2,450 ft) Schodde
and Hitchcock (1968, p. 35) also noted ecological segregation between
C. esculenta and the other swiftlet present (C. vanikorensis or C.
hirundinacea), with the former confined to a river and nearby forest
glades, the latter over cleared areas distant from primary forest.
Collocalia hirundinacea hirundinacea Stresemann
Mountain Swiftlet
NATIVE NAME. Daribi: hawaiyabo.
SPECIMENS EXAMINED. Karimui: 7 ¢, 10 9, 4 ? (2-16 July 1965). Bomai:
I 3 (7 July 1965);
WEIGHT. 7 @: 8.7, 9.0 (3), 9.3 (2), 10.0. 9 9: 8.0, 9.0 (7), 9.3. 2 ?: 9.0 (2).
WING. “8 @: 105-113: (109 a= 3). LOMO TOS= NE (LOS) =. 2). 4k OS aS
113.
STOMACH CONTENTS. Insects.
TAXONOMY. Collocalia hirundinacea and C. vantkorensis are im-
possible to distinguish in the field and very difficult in the hand. Char-
acters that I found useful are that C. vanikorensis has a slightly larger
(wider and longer) bill; most but not all C. hirundinacea have feathered
tarsi, and most but not all C. vanikorensis have unfeathered tarsi; C.
vanikorensis tends to be more brown on the uppersurface and under-
surface, while C. hirundinacea tends to have more of a blue sheen
above (especially on the head) and to be more gray and paler below;
and C. hirundinacea shows more white at the base of the dorsal feathers
when the back is ruffled. All specimens collected at Karimui and Bomai
proved to be C. hirundinacea.
DISCUSSION. ‘The sex ratio pee progressively throughout
July 1965. Only males were taken on 2, 3, 4, and 7 July. On 10 July
one male and three females were ce on 11 July one male and one
unsexed bird. Only females were taken on 12, 14, 15, and 16 July.
Flocks of brown swiftlets which must have been either this species
or C. vanikorensis (as opposed to the blue C. esculenta) were observed
daily over airstrips, grassland, and cleared land from 2,000 ft up to
alpine grassland at 11,000 ft. In general, C. hirundinacea is found at
higher elevations, C. vanikorensis at lower elevations. Since my Karimui
and Bomai specimens were all C. hirundinacea, 1 presume that those
from the higher stations, where no specimens were taken, were also
C. hirundinacea. As discussed under the preceding species, habitat
181
SPECIES ACCOUNTS
segregation between C. hirundinacea and C. esculenta was strict at
lower elevations, but mixed flocks occurred at higher elevations.
VOICE. A high-pitched twittering.
Collocalia whiteheadi nuditarsus Salomonsen
Whitehead’s Swiftlet
The only Eastern Highlands records are four specimens collected by
Gilliard at 5,000-6,000 ft in the Kubor Mountains, plus breeding rec-
ords at 6,000 ft in the Kaironk Valley of the Schrader Range (Bulmer,
pers. comm.). Somadikarta (1967) was the first to recognize that the
Idenburg River swiftlets, formerly placed in the same subspecies as
the populations now known as nuditarsus, actually belong to a distinct
species, C. papuensis. Whether nuditarsus is actually conspecific with
C. whiteheadi of the Philippines is uncertain.
BEMIPROGCNIDAE- CRESTED: SWIPES
Hemiprocne mystacea mystacea (Lesson)
Mustached Swift
NATIVE NAMES. Gimi: arakada. Daribi: gébisige.
SPECIMENS EXAMINED. Karimui: 1 9 (4 Aug. 1965). Bomai: 2 ¢,2 92 (6-8
July 1965).
WEIGHT. 2°62 69273; 93°03 72, 73; 74
WING. 2 4: 225, 240. 3 9: 229, 232, 234.
STOMACH CONTENTS. Insects.
BREEDING. All the Bomai specimens (collected in July) were in
nonbreeding condition, while the Karimui female (collected in August)
had enlarged ovaries.
DISCUSSION. I found this striking bird only in the Karimui
region, up to 4,500 ft (Soliabeda, Bomai, Karimui, Sena River), where
its distribution seemed local—i.e., it was uncommon and seen ir-
regularly except at Bomai, where several were seen every day in a
particular garden. Group of two to four birds, occasionally solitary
individuals, occupied conspicuous perches in open areas in the crowns
of tall trees with little foliage, from which they sallied out, circled
briefly with strong, falcon-like flight, and returned to the same perch.
The Soliabeda, Bomai, and Karimui birds were in highly disturbed
habitats (gardens and other areas which had been extensively cleared
but had a few tall trees remaining). ‘The Sena River record was in
undisturbed forest, where a pair used a tall tree on the bank overlook-
ing the open space over the river as a perch. Gyldenstolpe (1955,
p. 72) once saw one individual of H. mystacea over Nondugl (5,200 ft),
and Bulmer mentions it as present below 5,000 ft in the Schrader
Range.
182
SPECIES ACCOUNTS
H. mystacea is worth citing as an example of a low-elevation species
that has failed to colonize Eastern Highlands garden areas above 5,000
ft, although the nature of the habitat seems to correspond well to its
normal lowland habitat and although the colonization would have
necessitated only a small step above its normal altitudinal range, which
extends to ca. 4,000 or 4,500 ft.
VOICE. The call is a high downslur, nearly a squeal.
ALCEDINIDAE: KINGFISHERS
Ceyx azureus lessonii (Cassin) and C. a. ochrogaster (Reichenow)!
Azure Kingfisher
SPECIMENS EXAMINED. Sena River: 1 ¢@, 1 9 (26 and 27 July 1964).
WING. 1 ¢: 73.
STOMACH CONTENTS. Insects.
TAXONOMY. ‘The darker ochraecous wash of the underparts
places the specimens with the southern New Guinea race lessonii, to
which the Lake Kutubu population also belongs. Sims (1956) assigned
a female taken at Minj (5,000 ft) in the Wahgi Valley by Shaw-Mayer
to the northern New Guinea race ochrogaster.
DISCUSSION. ‘This kingfisher lives near small streams, including
brooks as little as 2 ft across, or in the understory of the forest. All
individuals that I saw were solitary and perched within 5 ft of the
ground. Periodically a perched bird quickly raises and lowers its whole
body while holding its legs in place, an action comparable to a man on
horseback “posting” in the saddle when the horse is in the trotting gait,
and similar to the “posting” of Salpinctes obsoletus (Troglodytidae) of
North America and Xenicus gilviventris (Acanthisittidae) of New
Zealand. ‘The purpose of the action is probably to assist vision by
obtaining a different perspective or estimating distance by parallax.
Ceyx lepidus solitarius ‘Temminck
Dwarf Kingfisher
SPECIMENS EXAMINED. Karimui: 1 ¢, 2 ? (6 Aug. 1964; 9 July-3 Aug. 1965).
Soliabeda: 1 4,1 9Q (25 July, 1965).
WHIGHT. 2 $213.6, 145. 1.9% 140. J
WING. @. 45102, DowNos fOs Do. LPs Be,
STOMACH CONTENTS. Insects.
Ps Los0}
BREEDING. ‘he gonads were small in all specimens, implying
that breeding is not in the dry season.
DISCUSSION. ‘There seem to be no other Eastern Highlands
records yet of this tiny kingfisher, but it will undoubtedly turn up
1 Listed as Alcyone azurea in Rand in Gilliard (1967).
183
SPECIES ACCOUNTS
elsewhere below 4,000 ft. In the Karimui region it was uncommon
and solitary in the understory of the forest interior and forest edge, up
to about 15 ft above the ground. Often it occurs away from water, but
I saw one pair perched on a branch above a small brook, periodically
plunging into the water directly beneath them and immediately re-
turning to the same perch. Like Ceyx azureus, C. lepidus periodically
“posts” up and down on its legs.
Melidora macrorhina macrorhina (Lesson)
Hook-billed Kingfisher
NATIVE NAME. Daribi: sugaba.
SPECIMENS EXAMINED. Karimui: 2 ¢, 1 ? (6 and 8 Aug. 1964; 3 July 1965).
Bomai: 1 ¢,1 9 (6 and 7 July 1965). Soliabeda: 2 ? (26 and 29 July 1965).
WEIGHTY. 2 f= 91, l02. 3° 0:93, 97. 110.
WING. 3 4: 116,118,119. 3 @: 111, 119, 120.
TAXONOMY. The light highlights on the black crown of the
female place this series with nominate macrorhina, as expected on
geographical grounds, rather than with the north coast form jobiensis.
‘The 29 July Soliabeda female, but not the other specimens, is buffy
below, a character of immatures and some adults.
BREEDING. ‘The gonads were slightly enlarged in the 26 July
Soliabeda female, not enlarged in the other specimens.
DISCUSSION. Five of the seven specimens were netted. In the
Karimui area this is evidently a not uncommon species in the lower-
and middlestory of the forest interior. Elsewhere in New Guinea M.
macrorhina is regularly encountered only at elevations below 3,000 or
2,000 ft, and the only other Eastern Highlands record is a single female
taken at Lake Kutubu. This kingfisher is nocturnal, and is unobtru-
sive, solitary, and silent during the day, so that my sole sighting was
of a bird 20 ft above the ground in dense foliage.
VOICE. ‘The call (Fig. 19, p. 186) is heard only at night. It consists
usually of two whistled notes, the second conspicuously shorter than
the first, a half-tone higher, sometimes trilled, and following the first
by slightly less than 1 sec. Occasionally two additional shorter notes
are added at a higher pitch. The kingfishers Melidora macrorhina,
Clytoceyx rex, Halcyon torotoro, H. megarhyncha, H. macleayi, and
Tanysiptera galatea betray their relationship in the similar whistled
qualities of their calls.
Clytoceyx rex rex Sharpe
Shovel-billed Kingfisher
NATIVE NAMES. Fore: userépo. Gimi: mégofo. Daribi: idadandgo.
SPECIMENS EXAMINED. Awande: 1 ¢ (11 July 1967).
WING. 165.
184
SPECIES ACCOUNTS
EXPOSED CULMEN. 37.
WHOLE CULMEN. 46.
TAXONOMY. In dorsal coloration the specimen is darker than
Fly River, Bernhard Camp, and Jimmi River specimens, comparable
to Mt. ‘Tafa specimens, and paler than ‘Velefolmin specimens. Since
variation in darkness follows a checkerboard geographical pattern, the
race septentrionalis is not recognized.
DISCUSSION. ‘The only other Eastern Highlands specimens of this
uncommon and sparsely distributed species are three from the Jimmi
River presented to Gilliard, and one shot at 6,000 ft on Mt. Wilhelm
by Shaw-Mayer. I heard its song at eight localities between 2,000 and
6,500 ft, from the forest or forest edge. The specimen was surprised on
the ground while eating a snake. According to Paran, Clytoceyx is
solitary, and most in evidence at Awande in the rainy season, when
it perches on trees or fences overlooking gardens and flies down to the
ground to locate its food of large insects, grubs, and small mice.
VOICE. An unmistakable, clear, liquid, ventriloquial three- or
four-note, whistled call that is delivered from the crown of a tall tree
and carries well. ‘The whole call takes not quite | sec. ‘The commonest
pattern is of three notes in descending pitch, the first two separated in
pitch by an interval of a second or minor third, the second two by a
fourth or major third (Fig. 19). Less often there is a fourth note, giving
rise onomatopoetically to the name “u-se-ré-po”. At each note of the
call the tail is jerked. Another call consists of three or four louder,
rapid, muscal notes on the same pitch. I have heard the call only in
the morning, and most often shortly after dawn.
Dacelo gaudichaud Quoy and Gaimard
Gaudichaud’s Kookaburra
NATIVE NAME. Daribi: anuai.
SPECIMENS EXAMINED. Karimui: 2 4,2 9 (13-18: July, 1965). Bomai: 1 @
(8 July 1965). Soliabeda: 1 ¢,4 9 (22-29 July 1965).
WEIGHT. 4 ¢: 132, 138, 139, 161. 6 9: 144, 146, 146, 146, 149, 156.
WONG. ©4120) 134 isbe 37, 6-O 2 193). 137, 137, 138.188, 139.
STOMACH CONTENTS. Insects, often large ones such as grasshoppers.
BREEDING. All specimens had small gonads, hence breeding is
not in the dry season.
DISCUSSION. ‘The Karimui population of this noisy, conspicuous,
and belligerent lowland kingfisher was considerably above its normal
altitudinal ceiling, and elsewhere in the Eastern Highlands it has
been recorded only at Lake Kutubu (2,450 ft). It was usually seen
perched singly in the middlestory, occasionally in the understory or
crown, of second-growth, gardens, forest edge, and occasionally forest.
The tail is often cocked up when perched or when fighting. I have
observed fights with others of its species and with Coracina papuensis.
185
Melidora macrorhina:
1sec
Clytoceyx rex:
or “
u se re po
| 1 sec |
Dacelo gaudichaud:
(a) — =
‘2 sec
Halcyon megarhyncha and H. torotoro:
pore ce or eee *,
Oley gaia se eae Sty cots
2 sec
Tanysiptera galatea:
oe or is le ie
Fic. 19. Voices of six New Guinea kingfishers.
186
SPECIES ACCOUNTS
VOICE. Loud, and exercised from conspicuous perches. Most
vocalizations consist of a series of a half dozen or more separate notes
at intervals of 1 sec. The quality is either: (a) clear barking notes as
of a small but noisy dog, all barks being on the same pitch; (b) more
squealing, downslurred barking notes, all again being on the same
pitch; or (c) barking notes, each of which is trilled; in this case suc-
cessive notes are on lower pitches (Fig. 19, p. 186). As with Dacelo
leachii and D, novaeguineae, two nearby individuals of D. gaudichaud
often bark simultaneously; this may represent male-female duetting.
Halcyon torotoro meeki Rothschild and Hartert
Lowland Yellow-billed Kingfisher
SPECIMENS EXAMINED. Karimui: 1 9 (14 July 1965).
WEIGHT. 38.
WING. 73.
CULMEN FROM ANTERIOR EDGE OF NOSTRIL. 33.
STOMACH CONTENTS. Insects.
TAXONOMY. ‘The underparts of the Karimui specimen are as
pale as in meeki and brevirostris, and paler than in pseustes. ‘The
length of the culmen from the anterior edge of the nostril to the tip
was measured in four females each of four races: tentelare, 32, 33, 33,
DAS PsCuUstes; 30). 924925, 04; DREUITOSLTIS, 28,29) Dlx: O15) MEER, ly 38,
34, 34. The short bill of brevirostris eliminates it, and the Karimui
specimen must be assigned to meeki, a slight westward range extension.
The ranges of southern New Guinea races given by Rand (1938, pp.
11-12) may thus be reformulated as follows: meeki, southeastern New
Guinea, west to Karimui (Purari drainage, long. 145°E); pseustes,
southern New Guinea from the mouth of the Fly River (long. 143°E)
west to the Mimika River (long. 136°E) except for the range of brevi-
rostris; nominate torotoro, northern and western New Guinea, on the
south coast east to Etna Bay (long. 135°E); brevirostris, near the coast
at long. 142°E in the bulge of southern New Guinea.
BREEDING. ‘The ovary was small.
DISCUSSION. ‘This was quite an uncommon species in the Kari-
mui Basin, and has not been recorded elsewhere in the Eastern High-
lands. On the three occasions I saw it, solitary individuals were perched
2-15 ft above the ground in open second-growth, and jerked or cocked
the tail at frequent intervals. The altitudinal range of H. torotoro
excludes that of its higher-altitude sibling H. megarhyncha. In the
Karimui Basin the transition takes place at some altitude between
3,650 ft, the highest H. torotoro record, and 4,400 ft, the lowest H.
megarhyncha record (the transition usually takes place somewhat be-
low 3,000 ft in the rest of New Guinea). Although H. torotoro and
H. megarhyncha are very similar in appearance as well as in voice,
they are evidently distinct species: their sizes do not overlap if indi-
187
SPECIES ACCOUNTS
viduals of the same sex are compared, the bill is generally pure yellow
in adult H. torotoro but has some black in adult H. megarhyncha of
all races except sellamontis, neither species approaches the other
morphologically with altitude, and the two forms show independent
geographic variation in subspecific traits.
VOICE. A lengthy “police whistle’ identical to the call of H.
megarhyncha (Fig. 19, p. 186), as also noted by Stein (1936, p. 47). The
whole body shakes as the bird calls.
Halcyon m egarhyncha subsp.
Mountain Yellow-billed Kingfisher
NATIVE NAMES. Fore: patorédba bilong bush (= “patordéba of the forest’: cf.
Halcyon sancta). Gimi: guralo. Daribi: wiyokoldlo.
SPECIMENS EXAMINED. Awande: 1 9? (19 June 1965).
WEIGHTY. 63.
WING. 90.
EXPOSED CULMEN. ‘To margin of lateral feathering, 38; to margin of medial
dorsal feathering, 41.
TAXONOMY. ‘The Eastern Highlands population may eventually
require separation as a new race nearest nominate megarhyncha but
larger, but the material available at present is too limited to warrant
doing so now. Only two other Eastern Highlands specimens have been
measured: a male, wing given as 96 mm by Mayr and Gilliard (1954,
p. 343); and male, wing given as 89 mm by Gyldenstolpe (1955, p. 72).
My measurements of female specimens in the American Museum of
Natural History yield: nominate megarhyncha, 8 9° from southeastern
New Guinea, wing 81-88 (83 + 2; seven values fall between 81 and
83), exposed culmen to margin of medial feathering 36-40 (38.5);
wellsi, 2 9 from the Weyland Mountains, wing 86 and 87. The Awande
female thus has a longer wing than any other female measured, and
this is probably also true of the male listed by Mayr and Gilliard. ‘The
bill of the Awande specimen is yellow except that the upper surface of
the upper mandible is black. The bills of H. m. megarhyncha and
wellst are similar, but sellamontis of the Huon Peninsula has the
whole bill yellow. The race wells: differs from nominate megarhyncha
in the buffier, less white underparts and the darker (bluer) back, in
which respects the Awande bird agrees with megarhyncha.
BREEDING. ‘The gonads were small, as also true of my sole H.
torotoro specimen, so that these two resident species presumably breed
when the migrants H. sancta and H. macleayii are gone.
DISCUSSION. Halcyon megarhyncha is found strictly in the forest
interior, from about 4,000 to 6,500 ft. Its altitudinal range is com-
pletely exclusive of that of its low-altitude sibling H. torotoro, From
April to September the Australian migrants H. sancta and H. macleayi
share its altitudinal range but are confined to open country, so that
188
SPECIES ACCOUNTS
habitat exclusion is complete. It would be interesting to know ge
megarhyncha comes out to the forest edge and second-growth when
the migrant flood departs.
H. megarhyncha is locally not uncommon, and calls often and loudly
while perched in the middlestory of forest trees. It must remain motion-
less inside the foliage, for I found the singers exasperatingly impos-
sible to see, and Stein (1936, p. 48) and Rand (1942b, p. 460) report
similar experience. The minimal number of specimens collected in
the Eastern Highlands reflects this invisibility rather than true rare-
ness.
VOICE. A long call identical to the call of H. torotoro, closely
similar to a police whistle in quality, and consisting of a series of rapid
notes lasting a total of several seconds. The pitch may continuously
drop throughout the series; may initially rise before dropping; or may
drop, then rise, and sometimes drop again (Fig. 19, p. 186). How /.
torotoro and H. megarhyncha, which are so similar both in appearance
and in voice, distinguish each other near the transitional altitude is
unknown.
Halcyon macleayii incincta Gould and H. m. elisabeth (Heine)
Forest Kingfisher
SPECIMENS EXAMINED. Karimui: 4 ¢,1 9 (3 July-3 Aug. 1965). Soliabeda:
26,1 9, (20-29 fully 1965).
WEIGHT. 6 ¢: 41, 41, 42, 42, 43,48. 2 9: 41, 48.
WING. 6 4:°86;,89, 90; 90, 91, 9. 2 O87, 88:
STOMACH CONTENTS. Insects and a frog.
TAXONOMY. All eight specimens have the blue-green backs of
the race incincta, a migrant from eastern Australia. At Nondugl,
Gyldenstolpe took two specimens, one of imcincta and one of the resi-
dent eastern New Guinea race elisabeth. In addition, I saw but did
not collect the species at Okasa (4,250 ft), and Schodde and Hitchcock
met incincta at Lake Kutubu (2,450 ft).
BREEDING. All specimens had small gonads.
DISCUSSION. Of the two Halcyon species that migrate to New
Guinea during the Australian winter, H. sancta is common up to
7,500 ft, whereas H. macleayit becomes uncommon above 5,000 ft and
apparently disappears above 5,500 ft. Below 5,000 ft both may be
found at the same locality, but H. sancta is up to twice as common. I
can discern no differences in wintering habitat preferences where they
overlap at these lower altitudes. Both species choose open habitats
with trees in which they perch 5-40 ft above the ground, most often in
native gardens, and eschew the forest and even the forest edge. In
flight I find the white wing patches of H. macleayii a useful field mark.
VOICE. ‘The migrant incincta was silent during the time I ob-
served it in the Eastern Highlands. ‘The call of the resident elisabeth
189
SPECIES ACCOUNTS
IS a short. descending, musical, rattled trill, like a shorter, slower,
louder version of the call of H. torotoro and H. megarhyncha.
Halcyon sancta sancta Vigors and Horsfield
Sacred Kingfisher
NATIVE NAMES. Fore: patéroba bilong place (= “patdéroba of the village”:
cf. Halcyon megarhyncha). Gimi: guralo.
a eacele ae Serge ces hin ies ie a 2 es Aug. 1964; 15-20 en ee
arimul: ce o, 1964: ae o, 1965). ai: , 965).
Soliabeda: io os July 1965). PR TEM UNDE acer eater
WEIGHT. 12 ¢: 39-50 (44+ 3). 3 9: 37, 45, 49.
WING. 12 4: 89-98 (92 +3). 3 9: 85, 92, 92.
STOMACH CONTENTS. Insects, including large grasshoppers.
BREEDING. All specimens had small gonads.
DISCUSSION. ‘The sex ratio of the specimens (15. 4/2 2) as quite
disparate in this Australian wintering visitor. Other collections have
often shown equally disparate ratios. For instances, Gyldenstolpe (1955,
p. 75) obtained 0 ¢, 8 ¢@ at Nondugl; the Second Archbold Expedi-
tion (Rand, 1942a, p. 321) had a total of 0 4,6 @ at three localities,
2 4,0 Q ata fourth; the Third Archbold Expedition (Rand, 1942b,
p. 461) had 10 ¢, 3 9. It appears that males and females may have
different local preferences in their wintering grounds.
Of the migrants that visit the Eastern Highlands during the
Australian winter, this is the commonest species above 5,000 ft, and
one of the commonest below this elevation. All observers have found
it at most localities in suitable habitats up to 7,500 ft from April
through October. As stated under H. macleayii, it is to be found in
open habitats with trees, such as native gardens, around human habita-
tions, and in casuarina stands, never inside the forest. Much smaller
numbers must have reached the Eastern Highlands before the spread
of native agriculture. Insects are caught by pouncing, occasionally in
midair by sallying, and are killed by being pounded against a surface.
VOICE. Usually silent; occasionally, a soft rasped note.
Tanysiptera galatea subsp.
Common Paradise Kingfisher
STOMACH CONTENTS. Insects.
DISCUSSION. Dr. Bulmer has kindly informed me that natives
reported racket-tailed kingfishers, probably of this species, as not un-
common in the Jimmi Valley, that he examined plumes procured by
natives, and that N. Camps collected one specimen. ‘The species is to
be expected at the lowest elevations along the edges of the Eastern
Highlands.
New Guinea has two common lowland forest kingfishers which are
190
SPECIES ACCOUNTS
in related genera and ecologically somewhat similar: Tanysiptera
galatea and Halcyon torotoro. They differ ecologically in size (average
weight of the former 56 g, the latter 41 g), vertical preference (the
former more confined to the understory), and upper altitudinal limit
(the latter extending to higher altitudes).
VOICE. The call is a slow, soft, upslurred, mournful whistle last-
ing 2-3 sec and repeated at intervals. The song is a loud, rising, ac-
celerating, whistled trill usually preceded by a longer, more nasal note
(Bie: 19, p. 1c6):
MEROPIDAE: BEE-EATERS
Merops ornatus Latham
Rainbow Bee-eater
NATIVE NAME. Daribi: sibudili.
SPECIMENS EXAMINED. Okasa: 1 ¢,2 9 (24 June 1965). Karimui: 1 ¢,4 9
(3 July-4 Aug. 1965). Bomai: 2 9, 1 ? (5-7 July 1965).
WEIGHT. 2 4: 82, 38. 7 9: 26,27 (2), 28 (2); 29 (2). 122 29.
WING. 2 4: 104, 109. 7 Q: 104, 105 (4), 107 (2). 1 ?: 106.
STOMACH CONTENTS. Insects.
TAXONOMY. Five specimens have black throats (an adult char-
acter), five do not (an immature character); four have elongated tail
feathers (an adult character), six do not (an immature character).
There seems to be no fixed sequence for acquisition of the black throat
and long tail: four specimens with black throats have short tails, three
with long tails lack black throats, one has both a black throat and
long tail, and two have neither.
BREEDING. All specimens had small gonads.
DISCUSSION. As in the case of Halcyon sancta, the sex ratio is
quite disparate (2 ¢,7 ?). This has also been true for the specimens
collected by some other expeditions in New Guinea, suggesting some
segregation of the sexes on the wintering grounds.
This Australian wintering visitor has also been recorded from April
through at least August by Gilliard in the Wahgi Valley and by Bulmer
in Kyaka territory and in the Schrader Range. Schodde and Hitchcock
found it appearing at Lake Kutubu in early October and following the
Soro River, evidently at the beginning of the return migration to
Australia. In suitable habitats M. ornatus is not uncommon below
4,500 ft and locally up to 5,200 ft, above which altitude it seems rare
or absent. It apparently may gather in roosts, since flocks regularly
fly in one direction just after sunrise and return in the opposite direc-
tion towards sunset. Its habitat preference is large expanses of open
country with a scattering of trees to provide exposed perches (never
in the forest interior). From these exposed perches it sallies out like a
flycatcher, or soars and floats like a swallow, in a short flight, before
19]
SPECIES ACCOUNTS
returning to the perch. The flights in a smooth arc remind me of the
motion of a model airplane attached to a fixed length of cord. Merops
ornatus sometimes circles in a breeze with spread wings for several
seconds without flapping, and may be one of the smallest birds that
actually soars.
VOICE. -A: single musical note (‘bk’); or three or four such notes
run together in very rapid succession.
CORACIIDAE: ROLLERS
Eurystomus orientalis pacificus (Latham) and E. 0. waigiouensis Elliot
Dollarbird
SPECIMENS EXAMINED. E. 0. pacificus: Karimui: 1 @ (12 July 1965);
etter es A er eae E. o. waigiouensis: Karimui: 5 ¢,6 9 (11-12 Aug.
WEIGHT. E. 0. pacificus: 2 9: 132, 135. E. 0. waigiouensis: 4 6: 148, 153, 167,
1905 5. OS 157, 163, 166, 074, 192:
WING. E. 0. pacificus: 2 9: 190, 194. E. 0. waigiowensis: 5 @: 191, 192, 192, 196,
201; 5 2: 190, 193, 196, 203, 204.
STOMACH CONTENTS. Insects up to several centimeters in length.
TAXONOMY. At Karimui both the resident breeding form waigi-
ouensis and the Australian wintering visitor pacificus occurred, with
the former far more numerous. The sole specimen obtained at Solia-
beda was pacificus, as was the sole bird Schodde and Hitchcock col-
lected at Lake Kutubu. Bulmer collected both races in Kyaka territory.
With practice these two races can be distinguished in the field by the
darker and richer coloration, bluer underparts, and greener upperparts
of waigiouensis and by the paler, browner color of pacificus. ‘Vhe larger
size of waigiouensis appears most clearly in the weights.
BREEDING. All specimens of both races had small gonads. As dis-
cussed by Mayr (1942, p. 253), crossing of these races is prevented
because pacificus remains in nonbreeding condition while in New
Guinea. The small gonads of all my waigrouensis specimens as well
suggest that waigiouensis is also in nonbreeding condition (like most
species of second-growth in the Karimui area) when pacificus arrives,
and breeds in the rainy season after the latter’s departure. A Novem-
ber-December breeding season of waigiouensis in southeastern New
Guinea is also suggested by the records of the First Archbold Expedi-
tion (Mayr and Rand, 1937, p. 88). In addition, although both races
do sometimes occur at the same locality, usually one or the other
race vastly predominates.
DISCUSSION. ‘The Dollarbird was generally solitary and selected
exposed perches in the tops of tall, dead or leafless trees overlooking
open spaces, in partly cleared forest or near the edge of the forest, in
the manner of a giant flycatcher. From these perches it periodically
192
SPECIES ACCOUNTS
sallied out and circled around with a floppy, caprimulgid-like flight
before returning to the perch. Once a display was observed between
two birds perched in a tree, which bowed to each other rhythmically
for a minute before flying off together. Circling flights observed in the
morning at considerable heights may also constitute a display.
VOICE. A short, low-pitched, monosyllabic croak or snapping
sound reminiscent of a frog or insect, and somewhat suggestive in
quality of the disyllabic call of Mino dumonti.
BUCEROTIDAE: HORNBILLS
Aceros plicatus ruficollis (Vieillot)
Papuan Hornbill
NATIVE NAMES. Fore: ane. Gimi: ahana. Daribi: weézi.
SPECIMENS EXAMINED. Karimui: 1 ¢@ (a) (15 Aug. 1964). Soliabeda: 1 ¢
(b) (24 July 1965).
WING. 2 @: 487 (a), 415 (b).
TAIL, 1-4: 232%(b):
TAXONOMY. ‘The race junge: from eastern New Guinea was sep-
arated from ruficollis of western New Guinea on the basis of larger
average size apparent in long series, but there is a good deal of overlap
(Mayr, 1937, p. 13). Since the wing length of males averages 424 mm
in ruficollis but 442 mm in jungei, the present scanty material (aver-
age 426 mm) must be assigned to the smaller ruficollis. A male from
Lake Kutubu had a wing of 420 mm and was also assigned to ruficollis
by Schodde and Hitchcock.
BREEDING. Both specimens had small testes.
DISCUSSION. In the Karimui area this species was uncommon
and was usually seen singly or in pairs in the forest. Hornbills prob-
ably are widely distributed at low elevations (below ca. 3,500 ft) at the
edges of the Eastern Highlands.
VOICE. A series of grunts, “uh----uh----uh----uh”; in flight, a thump-
ing or chugging sound made with the wings at each beat, as of a steam
locomotive slowly starting up.
PIT TiIDAR: PIPTAS
Pitta erythrogaster macklotit Temminck
Blue-breasted Pitta
NATIVE NAME. Daribi: towaga.
SPECIMENS EXAMINED. Okasa: 1 9, 1 imm. Q (24 and 25 June 1965).
Karimui: 1 ¢,4 9,2?,1imm. ¢, 1 imm. 9, 1 imm. ? (3-12 Aug. 1964; 3 July-3
Aug. 1965). Bomai: 1 imm. g (7 July 1965). Soliabeda: 1 ¢,1 Q (28 July 1965).
WiEIGIII. “Weg revsn Soe. 780.80, cu, 95, 1 rs 86, 2 im, fo 76, 85. 1
193
SPECIES ACCOUNTS
immu. 9: 72.
WING. 1 G2 106, 6 ©3105 0s, 106) 107) 107) 108) 2 am), G2 105, 108: 2
imm. 9: 104,108. 1 imm. ?: 111.
STOMACH CONTENTS. Insects.
‘TAXONOMY. ‘The New Guinea races are quite distinct, and the
present series agrees well with macklotii from southeastern New
Guinea and the Fly River. The race habenichti differs in the brighter,
redder, less brown nape; loriae in the very dull brown nape which
hardly contrasts with the darker brown of the crown; and oblita in the
paler, less bright nape and the bluer, less green back. The five young
birds have the belly lighter and more rose-orange in color, the throat
and breast mottled brown, and only scattered bright feathers on the
nape.
BREEDING, ‘The gonads were enlarged in the Soliabeda pair but
small in all other specimens. This fact, combined with the number of
young birds, suggests that (except at Soliabeda) breeding was recently
completed, i.e., before the dry season.
DISCUSSION. Pitta erythrogaster is widely distributed in forest
on the hill slopes up to 3,500 or 4,000 ft. Natives said that it lived on
the ground in the forest interior, and Paran encountered it in small
parties of about three birds on the forest floor.
VOICE. According to Daribi natives, the call consists of two low-
pitched, slow, tremulous whistles, the first rising in pitch, the second
falling in pitch.
Pitta sordida novaeguineae Miller and Schlegel
Black-headed Pitta
This pitta, whose altitudinal spread upwards from the lowlands is
much more limited than that of P. erythrogaster, is known in the
Eastern Highlands only from the Jimmi Valley of the northern water-
shed, where Mayr and Gilliard (1954, p. 343) recorded one specimen
and Bulmer (pers. comm.) collected three more.
ALAUDIDAE: LARKS
Mirafra javanica subsp.
Singing Bush Lark
DISCUSSION. ‘This grassland species was collected in the Batyer
Valley by J. Kikkawa. It has been recorded at only four other localities
in New Guinea.
VOICE. The call note has the quality of the sound produced by
running one’s finger along the teeth of a comb, as does the call of
Peltops montanus.
194
SPECIES ACCOUNTS
HIRUNDINIDAE: SWALLOWS
Hirundo tahitica frontalis Quoy and Gaimard
Pacific Swallow
NATIVE NAME. Fore: onuguteyabi.
STOMACH CONTENTS. Insects.
DISCUSSION. ‘This swallow is common in open country, particu-
larly around villages and settlements, but there may be major gaps 1
its distribution within the Eastern Highlands (e.g., it was absent from
the entire Karimui area).
Hirundo nigricans nigricans Vieillot
‘Tree Martin
The sole Eastern Highlands records to date are sight records at ‘Tari
Airstrip on 4 August 1965, by H. L. Bell, and in the Hagen area in
March 1970 (N.G.B.S. Newsletter, No. 54, p. 2, June 1970).
CAMPEPHAGIDAE: CUCKOO-SHRIKES
This whole family appears to share a common breeding cycle, since
all or almost all individuals of all species I met in the Eastern High-
lands were in nonbreeding condition at the times of my visits.
Campochaera sloeti flaviceps Salvadori
Golden Cuckoo-shrike
NATIVE NAME. Daribi: yommu.
SPECIMENS EXAMINED. Karimui: 1 ¢ (6 July 1965). Bomai: 1 ¢,1 9,1?
(6 July 1965).
WEIGHT. 2 4: 41 402 1:92 41, IP 36.
WONG. 2G = LOSS IOS. oe Os ee.
TAXONOMY. ‘These specimens differ from nominate sloeti: of
the Vogelkop and Idenburg River in having the posterior half or two-
thirds of the crown washed with yellow-green.
BREEDING. ‘The gonads were small in all specimens, suggesting
breeding in the rainy season, as with most other campephagids.
DISCUSSION. Campochaera, a monotypic and endemic New
Guinea genus, has a very spotty distribution on the lower hill slopes
and near the foot of the hills. ‘he population in the Karimui area is
at present the only one known in the Eastern Highlands, and is about
1,000 ft higher than the usual altitudinal ceiling of the species. Be-
sides collecting it at Karimui and Bomai, I observed it at Soliabeda.
Its habitat in the Karimui area was tall trees in forest and at the edge
of forest, especially trees bearing fruit, while the duetting displays
195
SPECIES ACCOUNTS
were carried out in the crowns of the tallest trees emerging above the
canopy. Ogilvie-Grant (1915, p. 121) remarked that “it appeared to
have the habits of a Flycatcher, darting out to catch some passing in-
sect and returning again to the same perch”, but this description could
well have been based on the conspicuous duetting display and it may
not have been catching insects at all.
VOICE. A duet which has been described in detail elsewhere
(Diamond and Terborgh, 1968). Briefly, one individual delivers a
rapid series of a half dozen high, musical notes, all on the same pitch.
Simultaneously with the last several notes, a second individual gives
a whistled upslur, downslur, or two or three slurs. In groups of two
to four the brightly colored singers move through the canopy, repeat-
ing the performance from the tops of trees successively several hundred
feet apart. This display is similar to that of Coracina montana and
C. schisticeps.
Lalage atrovirens subsp.
Black-browed ‘Triller
STOMACH CONTENTS. Fruit up to 4 mm in diameter, and insects.
DISCUSSION. Bulmer collected two specimens in the Jimmi Val-
ley. The species will probably turn up elsewhere at low elevations on
the northern watershed. It lives singly or paired in the treetops, more
often in light forest or at the forest edge than in primary forest.
VOICE. Emphatic upslurs or groups of 5-7 notes reminiscent of
laughter, repeated 3-9 times (Fig. 20, p. 207).
Lalage leucomela polygrammica (Gray)
Varied ‘Triller
SPECIMENS EXAMINED. Karimui: 1 ¢, 1 ? (1 and 3 July 1965). Bomai: 1 9
(8 July 1965). Soliabeda: 1 ¢, 2 9 (22-27 July 1965).
WEIGHT. 2 4: 28, 29. 3 9: 25, 26,27. 1 ?: 24.
WING, |) 2 6593, 94. (3 Ol: 88, 88, 94,
TAXONOMY. Rand (1942a, p. 324) has discussed individual
variation in this race. My two adult males resemble southeastern New
Guinea birds in their gray, distinctly barred underparts and are
slightly more barred than one topotypical polygrammica male from
the Aru Islands. Eleven out of 14 males from the Fly River area are
nearly whitish below with the barring very obscure, and the remain-
ing ones are comparable to southeastern New Guinea males.
BREEDING. ‘The gonads were small in all specimens.
DISCUSSION. ‘This is the triller of the southern watershed, occur-
ring up to 4,500 ft. Besides collecting it in the Karimui area, I ob-
196
SPECIES ACCOUNTS
served it once at Okasa. Its habitat was the middle- and upperstories
(20-100 ft above the ground) of trees at the edge of forest, 1m partly
cut forest, and in isolated tall trees, not in the well-shaded forest 1-
terior. It was usually seen in pairs and moved about in leisurely fash-
ion.
VOICE. ‘The only sound I heard was, on one occasion, a scolding
downslurred note similar to that of the honeyeater Melilestes mega-
rhynchus,
Niche Differences in the Genus Coracina
This large genus is represented in New Guinea by I1 species, of
which 10 have been reported from the Eastern Highlands. All live in
the middle- and upperstories (I have yet to take any member of the
genus in a mistnet). All species take both insects and fruit, but C.
schisticeps, C. montana, C. boyeri, and perhaps C. lineata take mainly
fruit, while C. tenuirostris, C. caeruleogrisea, C. papuensis, C. longi-
cauda, and perhaps C. melaena and C. morio, take mainly insects.
Three of the species are large (weights 95-150). One of these, C.
novaehollandiae, is an Australian winter visitor of which only strag-
glers reach the Eastern Highlands and remain in open habitats. ‘The
other two are forest species whose altitudinal ranges are nearly mutu-
ally exclusive: C. longicauda at higher elevations, where it encounters
only the somewhat larger C. caeruleogrisea and the considerably
smaller C. montana near the lower limit of its range; and C. caeruleo-
grisea at lower elevations, where it overlaps several congeners half its
size.
The remaining seven species are small (50-70). Two of these (C.
tenuirostris and C. papuensis) are low-altitude species which are usu-
ally found in open and forest edge habitats. Of the five small forest
species (C. montana, C. schisticeps, C. morio, C. melaena, C. boyert)
two have mutually exclusive altitudinal ranges: C. montana at higher
elevations, above not only C. schisticeps but also above the three other
small forest species; and C. schisticeps at lower elevations. ‘The dis-
tributions of C. schisticeps and C. morio in the hill forests (ca. 1,500-
4,500 ft) tend to be locally complementary, and both are uncommon
at sea level, where the common forest species are C. melaena in the
middlestory and C. boyeri in the crowns. C. melaena and C. boyeri
have altitudinal ceilings lower than the ceilings of C. schisticeps and
C. morio but overlap these two hill forest species to a considerable
extent. Unsolved problems are how these four small forest species sort
out on the lower hill slopes, and what habitat differences C. schisticeps
and C. morio recognize to achieve partly complementary local dis-
tributions.
SPECIES ACCOUNTS
Coracina melaena melaena (Lesson)
Black Graybird
SPECIMENS EXAMINED. Karimui: 1 ¢, 2 9 (1 July-4 Aug. 1965). Bomai:
3 $,1 2 ? (69 July 1965). Soliabeda: 1 ¢,1 9, 1 imm. ¢ (24-27 July 1965).
WEIGHT. 4 @: 56, 57, 60, 63. 2 O15); 57, . | © #255; Tamim, 4s
aoe DO. 123) 12% 124) 126,126. 3 on lis, tis. 120) 1 OF Ps 6, im:
STOMACH CONTENTS. Insects and fruit.
TAXONOMY. The races C. m. melaena of western New Guinea and
meeki of eastern New Guinea differ in darkness of the female and in-
tergrade clinally, melaena being the darker race. The backs of my fe-
males are quite richly colored and compare well with nominate me-
laena from the Vogelkop and Idenburg River. The undersurfaces of
meeki females from southeastern New Guinea are paler, more cinna-
mon, less richly colored than in nominate melaena. In this respect Fly
River birds (assigned to meeki) are intermediate; my females are on
the average more richly colored than the Fly River birds and are
nearly as richly colored as Vogelkop and Idenburg River females. This
gives a checkerboard distribution as one proceeds from west to east
on the south coast: melaena from the Vogelkop to the Eilanden River,
meeki at the Fly River, melaena at Karimui, and meeki in the south-
Casts
BREEDING. One Karimui female had enlarged ovaries, but the
gonads were small in all other specimens.
DISCUSSION. In the Karimui region Coracina melaena is some-
what above its usual altitudinal ceiling and was nearly as common as
its much noisier close relative Coracina schisticeps, with which it
sometimes occurred in the same tree. C. schisticeps and C. melaena
differ somewhat in altitudinal preference, C. schisticeps having a
higher ceiling, and C. melaena being common in the flat lowlands
where C. schisticeps is uncommon or absent. At Lake Kutubu (2,450
ft) Schodde and Hitchcock found C. melaena common and did not
obtain C. schisticeps. C. melaena spends more time in the middlestory
(15-40 ft above the ground), C. schisticeps and C. boyeri in the crowns.
Coracina schisticeps poliopsa (Sharpe)
Gray’s Graybird
NATIVE NAME. Daribi: désin.
SPECIMENS EXAMINED. Karimui: 2 ¢ (12 July 1965). Soliabeda: 7 ¢,4 9,
limm. ¢ (21-29 July 1965).
WERIGHO. 9 4: 48-57 O33). 4°90 40) 52, 50,58. Jimmy Oy 26.
WING. 9 At 110-120 (ib = 3). 4 OF 108, 102, 113, Tia 1 mom. “es 109,
TAXONOMY. Subspecific differences are largely confined to the
female. The races C. s. schisticeps, reichenowi, and moskowski of
western and northern New Guinea differ from my material in that the
198
SPECIES ACCOUNTS
female’s chin is rufous rather than gray; in addition, in reichenowt
the male’s chin is darker. Compared to poliopsa females from south-
eastern New Guinea, the Fly River, and the southern slope of the
Snow Mountains, Soliabeda females differ in the deeper and richer
rufous of the underparts, and in the greater extent of gray on the
chin. It seems desirable to await more material from the Eastern High-
lands to decide whether the chin color is sufficiently consistent to
warrant racial separation. ‘The deep rufous underparts of Soliabeda
females are unlikely to be a racially significant character, since com-
parison of recent and older collected material from the same locality
shows that this ventral color is subject to postmortem fading.
BREEDING. ‘The gonads were small in all specimens.
DISCUSSION. Coracina schisticeps has not yet been recorded from
the northern watershed of the Eastern Highlands and is “inexplicably”
missing from the Huon Peninsula and most of the southeastern New
Guinea. At Karimui and Soliabeda it was common and conspicuous
in the treetops, accounting for about 5% of bird individuals.
VOICE. As described elsewhere (Diamond and ‘Terborgh, 1968)
groups of two to four C. schisticeps carry out treetop displays similar
to those of C. montana and Campochaera sloetit, except that the fe-
male does not reply to the male’s vocalizations and there is therefore
no duet. The male’s call is a series of loud, nasal, whistled slurs similar
to those of C. montana males, the first one upslurred, later notes down-
slurred. ‘his conspicuous performance is repeated from the tops of
different tall trees.
Coracina tenuirostris subsp.
Cicada Bird
STOMACH CONTENTS. Insects in most stomachs, fruit in half the stomachs
examined.
DISCUSSION. Bulmer collected two specimens (one each at 4,200
and 5,000 ft) in the Baiyer Valley and one at 5,000 ft in the Jimmi Val-
ley. This species lives at the forest edge and in gardens and second-
growth.
VOICE. ‘The song is a series of identical clear notes repeated at 3
times per second for 10-30 sec and crescendoing slightly. One or two
other birds may fly into a treetop with a singer and give musical chirps,
but the group display is less ritualized than in the case of C. schisticeps
and C. montana.
Coracina morio incerta (Meyer)
Miuller’s Graybird
SPECIMENS EXAMINED. Okasa: 1 9 (22 June 1965). Mengino: 1 ? (15 July
1964), Bomai: 1 ¢, 1 @ (6 July 1965).
199
SPECIES ACCOUNTS
WEIGHT. 1 4:57. 2 9: 53, 60.
WING, 1 4:18 26-118 114 “ie.
EXPOSED CULMEN. 1 ¢:17. 2 9: 16,17. 12:17.
cal
TAXONOMY. The Okasa female, presumably a young bird, has
the axillaries nearly white with gray barring, a condition that can be
matched in some specimens available for comparison. It also has very
faint suggestions of barring on the belly.
BREEDING. All specimens had small gonads.
DISCUSSION. There is some tendency, though not a strict one,
for the local distributions of Coracina morio and C. schisticeps in the
hill forest to complement each other, just as the distributions of C.
montana and C. schisticeps complement each other altitudinally. ‘This
is reflected in the fact that I collected no C. schisticeps at the three
localities where I took C. morio, and no C. morio at the two localities
where I took C. schisticeps.
VOICE. C. morio is apparently the author of a song consisting of
a buzzy note repeated at 5 times per second for about 4 sec, suggestive
of the song of C. tenuirostris.
Coracina montana montana (Meyer)
Black-bellied Graybird
NATIVE NAMES. Fore: onténti. Gimi: inane.
SPECIMENS EXAMINED. Awande: 1 ? (18 June, 1965). Okasa: 1 imm. ¢ (23
jume 1965). Mt. Michael: 2° ¢ J and) 13 Jruly, 1968); Mii. Kanu Zone 2: 3 i>
2 Os Zones: 50.4 95 Zone 4nd 6.70 9 (16 Aug.) Sept. 1965):
WEIGHT. 10 6259-70" 66-24). “9°02 57-70(G324) 2) ammo, he DO er 256:
WING, 10 4% 127-135 (Isla23)2 16.92 124) 126, 1260127, 127, 131. eam 73
123.. W 22 W224.
STOMACH CONTENTS. Fruit 2-3 mm in diameter (four stomachs); fruit and
insects (two stomachs).
TAXONOMY. Birds from eastern New Guinea were formerly sep-
arated in the race minus from those of western New Guinea on the
basis Of thei’ shorter wine (ca, 126-135 3, I2sloly o> vs: 13a la0e es,
127-137 ¢). The present series belongs to the smaller eastern popula-
tion, but minus is now not recognized because size decreases clinally
from west to east on the Central Range (Diamond, 1969; Rand and
Gilliard, 1967). The two largest males (wing 135) came from 8,000 ft
on Mt. Michael, whereas all other measured males came from altitudes
of 4,000-6,000 ft. Other than this I cannot discern an increase in size
with altitude. In the Snow Mountains, Rand (1942b, p. 463) also found
that birds from 9,200 ft were distinctly larger but that there was no
correlation between size and altitude between 4,000 and 7,200 ft. The
Okasa immature male, which has the shortest wing of the series and
a low weight, has the tail, primaries, and lores, but not the chin,
throat, or breast, black; there are scattered black and white barred
feathers (dark subterminally, white terminally) on the head, cheeks,
200
SPECIES ACCOUNTS
and belly, and the tips of the secondaries and undertail coverts are
barred black and white. The unsexed Awande specimen, which also
has a short wing and low weight, is similar except that the tail in addi-
tion has white tips and the barring on the body is more limited. Rand
(1942b, p. 463) describes an immature male in similar plumage, and
the American Museum of Natural History has a few other similar
specimens.
BREEDING. ‘The gonads were small in most specimens but were
slightly enlarged in four males and one female from Mt. Karimut.
DISCUSSION. ‘This is a conspicuous and widespread species in
mountain forest between about 4,000 and 9,000 ft, living in the crown
of the forest (not in second-growth). The lower limit to its range is
fixed by competition with its low-altitude congener, C. schisticeps:
the altitudinal ranges of these two species are mutually exclusive, and
I have not detected either species penetrating the other’s altitudinal
zone by more than 100 vertical feet. Thus, in the Karimui area C.
schisticeps was common up to 4,200 ft, above which it disappeared
completely, while C. montana was common above 4,230 ft, with a
single observation at 4,140 ft. In contrast to the sharp lower limit, C.
montana becomes gradually less common at higher altitudes de-
pending locally on the elevations at which the forest becomes stunted
and tall trees disappear. On Mt. Karimui, where the forest became
heavily mossed at 6,500 ft and progressively more stunted thereafter,
C. montana accounted for about 10% of all bird individuals at 4,200-
5,800 ft, about 5% at 5,800-7,280 ft, and was absent above 7,280 ft.
On Mt. Michael, where heavy moss and stunting set in only at 8,700 ft,
C. montana was still common around our camp at 8,000 ft. but was
only encountered once above 8,700 ft.
VOICE. A loud and precisely synchronized duet which is delivered
in the treetops by parties of two to four birds and has been described
elsewhere (Diamond and ‘Terborgh, 1968). The female delivers several
harsh notes on the same pitch in synchrony with slurred whistles from
the male. Each day the singers follow apparently the same route, se-
lecting display trees several hundred feet apart.
Coracina boyeri subalaris (Sharpe)
Rufous-underwing Graybird
SPECIMENS EXAMINED. Karimui: 7 ¢, 8 9 (11 Aug. 1964; 2 July-5 Aug.
1965). Soliabeda: 2 ¢,1 9 (22-26 July 1965).
WEIGHT. 9 ¢: 62-74 (68+4). 6 Q: 61-68 (64 + 3).
WING Sr Gebel sock by Os I20elab (lee S25).
STOMACH CONTENTS. Fruit.
TAXONOMY. ‘These agree with subalaris of southern New
Guinea in the gray (rather than white) lores of the female, in the pale
rufous underwing coverts, and in the long wing.
201
SPECIES ACCOUNTS
BREEDING. All specimens had small gonads.
DISCUSSION. This graybird lives in small parties of three to five
birds in the treetops. The Karimui population may represent an alti-
tudinal record for this lowland species, and the only other Eastern
Highlands record is provided by one specimen from Lake Kutubu
(25450 it).
VOICE. C. boyeri has a display similar to that of C. montana and
C. schisticeps, in which a group of birds flies among the tops of tall
trees, stopping in each to call. The display vocalizations consist of
single or paired, high, sweet, whistled slurs “tsyew,’ more often up-
slurred than downslurred, repeated at intervals of several seconds.
Other quieter calls are a low clucking, a cat-like “mew,” and repeated
chirping notes “chuck.”
Coracina caeruleogrisea strenua (Schlegel) and C. c. adamsoni
(Mayr and Rand)
Stout-billed Graybird
NATIVE NAMES. North Fore: kabagége. South Fore: kaiyowafyowa. Gimi:
kudiye.
SPECIMENS EXAMINED. Awande: 2 6, 3 9 (26 June 1964; 17-19 June 1965).
Okasa: 1 9 (22 June 1965). Karimui: 5 ¢,1 9, 1 imm. @ (11-15 Aug. 1964; 10
July-4 Aug. 1965). Bomai: 1 ¢, 1 9 (6 July 1965). Mt. Karimui Zone 1: 2 9;
Loewe t &, 9; Zone 3: 1 70) (O21 Aue. 1965):
WEIGHT. Okapa area (Awande and Okasa): 1 ¢@, 151; 4 9, 120, 126, 130, 136.
Karimui area (Karimui, Bomai, Mt. Karimui): 5 ¢@, 127, 138, 144, 152, 158; 4 9,
lso,0037, 139, 1425 i imm. 35 Lb:
WING. Okapa area: 2 ¢, 173, 182 (178 226); 2 OQ, 167, 170) (169 2-2), Kazim
area: 5 g, 168, 169, 171, 173, 174 (171 + 3); 5 9, 158, 165, 167, 167, 168 (165 + 4);
Ioan a 72s
CULMEN FROM NOSTRIL. Okapa area: 1 4, 29; 2 9, 26, 27. Karimui area:
Dade 204 275 Bile Boies OD, 205 215 27 2oy ols, aim as 26.
STOMACH CONTENTS. Large insects (two stomachs); 17 mm fruits (one stom-
ach). ;
TAXONOMY. ‘The race adamsoni of southeastern New Guinea
differs from strenua of western New Guinea in its larger average size,
slightly darker ochraceous axillaries, and slightly paler gray general
coloration. My measurements of wing length give: for adamsoni from
southeastern New Guinea, 5 g, 171-180 (177 + 4), 49, 163-170 (167 =
3); strenua from the Vogelkop to Telefolomys 8° 45 165-179 (17225);
6 9, 161-166 (165 + 2); nominate cacruleogrisea from the Fly River,
Z &, 169 and 170: (170 22 1), 2 95, o0"and 164 (162 325). Miy limaited
material suggests that the Awande-Okasa population averages as large
as adamsoni while the material from the Karimui area (100 miles to
the west) is as small as strenua. ‘The Awande-Okasa material ap-
proaches strenua in the paleness of the underwing coverts. Gilliard
and Gyldenstolpe assigned their Wahgi Valley material to strenua.
202
SPECIES ACCOUNTS
Evidently the populations of this species are nearer adamsoni in the
easternmost part of the Eastern Highlands and are nearer slrenwa
farther west. C. c. caeruleogrisea of the Fly River and Aru Islands 1s
generally paler.
BREEDING. All specimens had the gonads small.
DISCUSSION. ‘This large cuckoo-shrike is widespread but not nu-
merous throughout the Eastern Highlands from about 3,000 ft to a
ceiling varying locally between 5,200 ft (Mt. Karimui) and 8,000 ft
(Mt. Michael), and has been met by most observers. ‘Vhe other large
New Guinea cuckoo-shrike, C. longicauda, has an altitudinal range ly-
ing mostly above that of C. caeruleogrisea but not completely exclusive
of it. Like other Coracina species, C. caeruleogrisea lives in the upper-
story of the forest in small groups of about three birds. It walks along
tree limbs in a manner reminiscent of a crow.
VOICE. Varied, with both quiet and louder calls: a harsh, grating
scream, fairly but not very loud; a soft “chirp”; and a soft slurred
note first dropping then rising again in pitch, with a mewing quality,
like the call of the bowerbird Ailuroedus crassirostris.
Coracina novaehollandiae melanops (Latham)
Australian Graybird
SPECIMENS EXAMINED. Awande: 1 ¢@ (16 June 1965). Karimui: 3 ¢, 1 imm.
é6,2imm. 9 (14 Aug. 1964; 9-11 July 1965).
\WARIGIBIIE, Sy aS IPA Wee Nee, oP NING), IL aieaam, AS VAD), Il anaoham, GS ils
NVING. 3 7 196) 1972 2007 too 199) i immo 192. 2mm OP Sl 196:
CULMEN EROM BASE. 3 ¢°°27,.28, 295 1° 9:30. 1 imm: -¢? 29). 2 amm.
On Z83, 29.08
TAXONOMY. ‘These Australian migrants belong to the eastern
and southern Australian race melanops, not to the ‘Tasmanian race
novaehollandiae or the northern Australian race didima, which have
also been recorded from New Guinea. C. n. didima differs in its
shorter wing and paler color, C. n. novaehollandiae in its shorter bill.
BREEDING. ‘The gonads were small in all specimens.
DISCUSSION. ‘This is the first record of this Australian migrant
for the Eastern Highlands or for such high altitudes anywhere in New
Guinea. Small flocks of two to six birds were seen on six occasions at
Awande, Karimui, and Bomai in isolated tall trees in open country,
i.e., in gardens and near Karimui airstrip, a habitat in which no spe-
cies of Coracina resident at Karimui or Awande was ever seen. These
may have been itinerant groups which remained for short periods,
since all my sightings in the Karimui Basin fell in the periods 4-14
Aug. 1964, and 8-11 July 1965.
203
SPECIES ACCOUNTS
Coracina papuensis subsp.
Papuan Graybird
STOMACH CONTENTS. Insects (no fruit).
DISCUSSION. ‘This non-forest campephagid is generally a low-
land species in New Guinea but has colonized two widely separated
areas of the Eastern Highlands. At Lake Kutubu (2,450 ft) Schodde
and Hitchcock found it common in flocks at the forest edge and in
second-growth along the lake. Bulmer found it breeding commonly
in gardens and open country of the Baiyer Valley up to 5,500 ft. The
flight is distinctively undulating; the bird alternately flaps to gain alti-
tude, then glides down with the wings held stiffly vertically down-
wards.
VOICE. A weak, high-pitched, disyllabic note “chee-yu” or “whe-
chew,” the first note higher than the second, with a squealing or breezy
quality more like the voice of a parrot (e.g., Domicella lory) than of
most graybirds.
Coracina longicauda grisea Junge
Black-hooded Graybird
SPECIMENS EXAMINED. Agotu (Gimi territory): 1 ¢ (22 July 1964).
WING. 176.
STOMACH CONTENTS. Large insects (four stomachs); insects and fruit (five).
TAXONOMY. The races grisea of western New Guinea and nom-
inate longicauda of southeastern New Guinea differ only in size. I
measured wings of males of both races, with the following results,
proceeding from west to east: Snow Mountains (grisea), 171, 172, 173;
Telefolmin, 170, 173; Mts. Hagen and Wilhelm, 172, 173, 174; Agotu,
176; Huon Peninsula, 176; southeastern New Guinea (C. J. longi-
cauda), 177, 179, 180, 180, 186 (180 a= 3). There is obviously a climal
increase in size from west to east, as shown previously by the more
extensive measurements of Gilliard and LeCroy (1961, p. 46). Five
males from the Wahgi region, slightly farther west than Agotu, mea-
sured by Gilliard and LeCroy (1961, p. 47) and by Gyldenstolpe (1955,
Dp 77), gave 169; 172, 172,174, 076, yielcine am averaseror N7omter all
six Eastern Highlands males. On this basis I refer the whole Eastern
Highlands population to grisea. More specimens would be required
to decide if birds at the eastern end of the Eastern Highlands really are
nearer nominate longicauda.
DISCUSSION. My sole encounter with this species was in forest
near Agotu, where a tongue of moss forest descended to about 5,800 ft,
giving this unusually low record for the species. ‘Mhe specimen was one
of a flock of six hopping slowly along branches in the tree crowns. Most
other observers in the Eastern Highlands have encountered C. longi-
204
SPECIES ACCOUNTS
cauda but at altitudes above 7,000 ft and ranging in the moss forest
up to 10,500 or even 12,000 ft.
VOICE. A high parrot-like slur similar to the note of Coracina
papuensis, and a short low-pitched chirp.
MOTACILLIDAE: PIPITS and WAGTAILS
Motacilla cinerea caspica (Gmelin)
Gray Wagtail
Gyldenstolpe met this palearctic wintering visitor a few times in
August in boggy grassland at Nondugl. Bulmer observed and collected
it near streams in Kyaka territory during December through February
at 4,000-5,500 ft and also commonly in the Schrader Range up to
7,500 ft.
Anthus gutturalis rhododendri Mayr
Alpine Pipit
SPECIMENS EXAMINED. Mt. Michael: 3 ? (10 July 1964).
WING. 2 7: 92, 95.
STOMACH CONTENTS. Insects (five stomachs), grass seeds (two), insects and
seeds (three).
TAXONOMY. ‘The Mt. Michael specimens are smaller than nomi-
nate gutturalis of southeastern New Guinea and have the abdomen
slightly darker; wollastoni of western New Guinea has the black mark-
ings on the side of the throat reduced or absent.
DISCUSSION. ‘This high-altitude pipit was met only in alpine
grassland at 12,000 ft near the summit of Mt. Michael where, during
our brief experience in this habitat, it was the only conspicuous bird.
It walked on the ground with a waddling gait, and perched in bushes
in the grassland and in 40-ft trees at the forest edge. Alpine grassland
has now been surveyed ornithologically on four high peaks of the
Eastern Highlands (Mts. Michael, Wilhelm, Hagen, and Giluwe), and
Anthus gutturalis has been found on all four.
VOICE. ‘The varied song consists of high, thin, clear notes, each
repeated several times before repeating a different type of note. ‘The
call is a faint, high-pitched “tsip” or “‘tsee,’’ sometimes repeated.
Anthus novaeseelandiae exiguus Greenway
Richard’s Pipit
NATIVE NAME. Fore: iboroéto.
DISCUSSION. I met this pipit in twos and threes on the lawns of
European houses at Okapa (6,600 ft) and on the nearby grass airstrip
205
SPECIES ACCOUNTS
at Tarabo (6,000 ft), but nowhere else. Other observers have found it
numerous at a number of Eastern Highlands localities between 4,000
and 7,000 ft in very close-cropped grass, such as on sheep pastures,
football fields, airstrips, and lawns. The erassland on grown-over native
gardens seems too tall for it. Elsewhere in New Guinea it has been
recorded only near Wau and Port Moresby.
MUSCICAPIDAE: OLD WORLD INSECT EATERS
‘TURDINAE: ‘THRUSHES
Drymodes superciliaris brevirostris De Vis
Northern Scrub Robin
NATIVE NAME. Daribi: gisonabo.
SPECIMENS EXAMINED. Karimui: 1 $?, 3 @, 1 ? (4-7 Aug. 1964; 2 July-6
Aug. 1965). Bomai: 1 @ (7 July 1965).
WHIGEIE) I 42 58.. 1 492 52) 2O% 26, 41.
WING. 1 g: 95. 1 $?: 90. 3 Q: 80, 82, 82.
CULEMEN FROM NOSTRIL. 1 ¢@: 13.0. 1 A?: 11.5. 3 9: 95, 95, 100:
TAXONOMY. ‘These specimens are closest to brevtrostris, from
which they differ only in the darker, blacker crown (tendency towards
nigriceps). In the pale color of the underparts they are closer to brevi-
rostris from southeastern New Guinea than from the Fly River; the
latter are browner below. ‘The races nigriceps and beccarii have duller
and darker backs, and nigriceps in addition is darker and grayer below.
DISCUSSION. D. superciliaris is a shy and uncommon, but wide-
spread, inhabitant of the forest floor from about 1,800 to 4,200 ft. I
never saw it in life but heard its song several dozen times at Okasa,
Soliabeda, Karimui, and on the lower slopes of Mt. Karimui.
VOICE. An unmistakable series of clear, very slow, chromatically
descending, somewhat plaintive, whistled notes. The quality and the
medium pitch are similar to those of Crateroscelis murina. In the
Eastern Highlands the song consists typically of five notes, each lasting
11% sec and separated by an equal pause from the next mote; so that
the whole song lasts about 15 sec. ‘he first note is initially down-
slurred. Successive notes each remain on constant pitch, and each is
about a half-tone lower than the next (Fig. 20). ‘he downslurred first
note is sometimes given alone as a call, and takes more practice to
identify but is still diagnostic.
Saxicola caprata wahgiensis Mayr and Gilliard
Pied Chat
NATIVE NAMES. Fore: pobogile. Gimi: médi. Daribi: dédi.
SPECIMENS EXAMINED. Awande: 1 ? (17 June 1965).
206
SPECIES ACCOUNTS
Lalage atrovirens:
eee HO ag or or Poe Oye ee 6) es
(each phrase is repeated 3 to 9 times )
Drymodes superciliaris:
—————
5 sec
Fic. 20. Voices of Lalage atrovirens and Drymodes superciliaris.
WEIGHT. 19:5:
WING. 78.
DISCUSSION. ‘This is a common species in cleared country (native
gardens, or grassland with perches such as fences or trees, but not pure
grassland) almost anywhere in the Eastern Highlands. Solitary in-
dividuals or pairs perched on fences, dead branches, the ground, and
at the tops of isolated trees, whence they periodically sallied out in
flycatcher fashion. One was seen with a berry in its bill. Males showed
territorial behavior, frequently singing from conspicuous perches gen-
erally 10 to 30 ft, sometimes as much as 70 ft, high.
VOICE. <A weak, somewhat hoarse, high-pitched, whistled song of
four to eight notes in irregular rhythm. Each singer has several pat-
terns, each of which is delivered several times in succession (with
moderate pauses between repetitions) before proceeding to the next
pattern. One of these four-note patterns is the source of the Fore name
“pobogile.”
Zoothera dauma papuensis (Seebohm)!
White’s Ground ‘Thrush
SPECIMENS EXAMINED. Okasa: 1 @ ? (a) (23 June 1965). Sena River: 1 ? (b)
(28 July 1964). Mt. Karimui Zone 3: 1 ¢ (c) (19 Aug. 1965).
WING. I S(er Wa Tos Gye 117.
DISCUSSION. ‘This thrush is one of the rarest species of the New
1 Listed as Oreocincla dauma in Rand and Gilliard (1967).
207
SPECIES ACCOUNTS
Guinea hill forest and has been taken in New Guinea only in five
areas. ‘he 12 specimens known to me are as follows: Huon Peninsula,
one each collected by Wahnes and Keysser on the Sattelberg; Adelbert
Mountains, one taken by Ziegler (3,800 ft); southeastern New Guinea,
one collected by the First Archbold Expedition at Mafulu (4,000 ft),
three collected at 4,000 ft on the Aroa River by Weiske, and the type
specimen taken by Goldie; southern slope of the Eastern Highlands,
my three specimens; southern slope of the Snow Mountains, one taken
by Meek at 4,000 ft. All three of my specimens came from primary
forest: the Okasa specimen was shot on the ground at 3,550 ft, the Sena
River one netted at 4,500 ft, and the Mt. Karimui one netted at 5,100
ft. ‘he species has a wide extralimital range (Asia to the Solomon
Islands) and is among the minority of hill forest species that “hopped”
directly into the New Guinea mountains from the outside, instead of
evolving from a lowland ancestor.
Amalocichla incerta olivascentior Hartert
Lesser New Guinea Thrush
NATIVE NAME. Gimi: kabiya-agéra.
SPECIMENS EXAMINED. Mt. Karimui Zone 4: 1 ¢ (30 Aug. 1965).
WEIGHT. 30.7.
WING. 76.
TAME. ST.
EXPOSED CULMEN. 16.
STOMACH CONTENTS. Insects.
TAXONOMY. The Mt. Karimui specimen was compared with
brevicauda of southeastern New Guinea, olivascentior of the Snow
Mountains and Weyland Mountains and North Coastal Range, nomi-
nate incerta of the Vogelkop, and one Mt. Goliath specimen referred
to brevicauda. ‘Vhe color of the underparts is grayest and dingiest, least
brown, in olivascentior from the Weyland Mountains and North
Coastal Range, followed by Snow Mountains olivascentior; south-
eastern New Guinea brevicauda is browner, and Vogelkop A. 1. incerta
brownest, least gray. The Mt. Karimui specimen is intermediate be-
tween Weyland Mountains and Snow Mountains birds. The Mt.
Goliath specimen is slightly browner than the Mt. Karimui specimen
but still much less brown than southeastern Drevicauda. ‘The white
feathers on the side of the throat are conspicuously tipped with dark
in Weyland Mountains and Snow Mountains birds and the Mt. Kari-
mui specimen but not in southeastern birds or the Mt. Goliath speci-
men. The Mt. Karimui specimen is therefore assigned to olivascentior
on the basis of the ventral coloration and throat feathers. Its wing and
tail lengths are shorter than those of 10 male olivascentior measured
(wing 77-84, tail 55-61). Gyldenstolpe assigned his seven specimens from
the Wahgi Valley to brevicauda, apparently on geographical grounds
and without comparison of material. He specifically mentions, however,
208
SPECIES ACCOUNTS
the dark margins of the throat feathers, so that they undoubtedly
belong to olivascentior. The measurements he reports (e.g., wing 79-82
in males) are comparable to Snow Mountains and Weyland Mountains
olivascentior. ‘Vhe allocation of the single Mt. Goliath adult is uncer-
tain. In his check list Mayr (1941) includes the Sepik Mountains in the
range of brevicauda, presumably on the basis of one specimen which
Biirgers collected on the Schraderberg and which I was not able to
examine.
The revised ranges of the subspecies are therefore: brevicauda, south-
eastern New Guinea, Huon Peninsula, and (requires confirmation)
Schraderberg; olivascentior, Eastern Highlands, North Coastal Range,
Snow Mountains, and Weyland Mountains; Mt. Goliath (eastern ex-
tremity of Snow Mountains), allocation uncertain; and nominate
incerta, Vogelkop.
BREEDING. ‘The testes of the Karimui specimen were greatly
enlarged.
DISCUSSION. My specimen was netted at 5,960 ft, while Gylden-
stolpe’s specimens were all trapped at 8,000 ft. The only other Eastern
Highlands records are from the Schrader Mountains (specimens ob-
tained by Biirgers and Bulmer). This is evidently a rare and local
thrush of the forest floor.
Turdus poliocephalus erebus Mayr and Gilliard
Island ‘Thrush or New Guinea Blackbird
STOMACH CONTENTS. Fruit (two stomachs), insects (one), fruit and insects
(one).
The Island ‘Vhrush was found in alpine grassland above 1,000 ft on
the summits of Mts. Hagen, Wilhelm, and Giluwe by Gilliard and by
Shaw-Mayer. On Mt. Albert-Edward I found it characteristic of the
edge between forest and alpine grassland, hopping on the ground and
perching in low bushes or up to 40 ft high in forest edge trees, some-
times feeding in grassland several hundred feet from the forest edge
but retreating to the forest when alarmed.
VOICE. ‘The call-notes are typical for the genus Turdus: a re-
peated buzzy alarm note with the tail flicked at each note, a squawk,
, ee 99
and a weak, high, slightly downslurred “‘tsr” or “‘sss.
ORTHONYCHINAE: LOGRUNNERS
As constituted by Deignan (1964) in Peters’ Check-list, Vol. 10, this
family consists of nine genera, restricted to Australia and New Guinea
except for the species Eupetes macrocerus from Malaya, Sumatra, and
Borneo. ‘The members of this family were formerly considered to be
babblers (Timalimae). ‘The genera Drymodes and Crateroscelis, also
previously considered babblers, have been transferred to ‘Vurdinae
209
SPECIES ACCOUNTS
and Malurinae, respectively, so that the Timaliinae are now considered
to be represented in New Guinea only by Pomatostomus isidori and
P. temporalis of the lowlands.
Eupetes castanonotus pulcher Sharpe
Midmountain Eupetes
NATIVE NAMES. Gimi: liyéta. Daribi: giwibo.
SPECIMENS EXAMINED. Sena River: 1 ¢ (29 July 1964). Karimui: 3 ¢, 1
- 2 alee fea Me cheers 1965). Bomai: ' é 9 July 1965). Soliabeda:
en, QO (41-29 July Dy. fit; Karimut Zone i: 2 9, 1 ami: g (iei2 Ame:
WEIGHT.” 3 @: 70) 72,74, "2 93 70,74. Imm. 4:61, ers 58:
WING. (6°42 94, 96,97, 975,99) 99. 4° O 2895/89: 9193. 1 imm 496) r= 8o:
TAXONOMY. In two of my females the eye stripe is completely
blue, while in the other two and in one of the unsexed Karimui speci-
mens the stripe becomes paler or whitish blue posterior to the eye.
T'wo of the females have white spots on the undertail coverts, while
this cannot be checked in the other two due to loss of feathers. ‘he
immature male has scattered chestnut feathers on the blue of the
lower back and rump.
Material available for comparison consisted of nominate castano-
notus from the Vogelkop, saturatus from the southern slope of the
Snow Mountains, wropygialis from the northern slope of the Snow
Mountains, specimens from the Adelbert Mountains assigned to buer-
gersi, two males of uncertain affiliation (perhaps bwergersi or par) from
Keku in the Finisterre Mountains of the Huon Peninsula, and pulcher
from southeastern New Guinea. ‘Topotypical material of bwergersi
(Sepik Mountains) and par (Saruwaged Mountains) was lacking.
Females of the races E. c. castanonotus, saturatus, uropygialis, and from
the Adelberts differ from mine in having the eye stripe blue in all speci-
mens. The general color of E. c. castanonotus is quite similar, but its
blue is slightly paler and less purple, and the chestnut of its back is
very slightly paler. £. c. satwratus differs conspicuously in its much
deeper blue ventral coloration and its slightly darker chestnut back.
E. c. wopygialis differs conspicuously in that the lower back of the
female is blue, not chestnut, and differs slightly in the generally paler
coloration. The Adelbert series and the Keku males differ in the
lighter tone both of their chestnut and blue parts. E. c, pulcher is
quite ‘close ini its general coloration; the blue of pulcher averages
slightly lighter and less purple, but many specimens can be matched
with mine; the chestnut averages lighter in pulcher. Females of pulcher
have the eye stripe pale blue or whitish, often mixed with rusty (none
of my females has this rusty). From the description (Mayr, 1931, p. 691),
buergersi is eliminated by the blue eye stripe of all three known
females and perhaps by its darker general coloration. E. c. par (Meise,
1930, p. 17) apparently agrees with my specimens in that the eye stripe
210
SPECIES ACCOUNTS
of the female may be either solid blue, or else pale blue behind the
eye; it apparently differs in its longer wing (the original description
cited 2 ¢ 101, 101; 2 9 98, 99); Meise said that males of par were
indistinguishable from saturatus, which would make them much darker
blue than my specimens; Mayr said that females of par had a lighter
chestnut back than E. c. castanonotus, which would make them lighter
chestnut than my specimens.
The Eastern Highlands series thus shares the variably light eye stripe
of females of the eastern populations pulcher and par. ‘Vhe small dit-
ferences in general coloration between it and the slightly lighter
pulcher to the east are what one would expect of a clinal east-to-west
darkening along the southern watershed that would culminate in
saturatus of the Snow Mountains. ‘The incidence of light female eye
stripes seems also to decrease clinally east-to-west on the southern
watershed from pulcher through the Eastern Highlands to saturatus.
The Eastern Highlands population is therefore best assigned to the
most similar end of the cline, pulcher.
BREEDING. The gonads were enlarged in all adult males and
half the females. Evidently this ground nesting species (Rand, 1942a,
9° OF
p. 327) breeds in the dry season.
DISCUSSION. FE. castanonotus lives strictly in the forest interior
on the ground. Single birds or pairs were occasionally seen walking
with bobbing head and fairly long, leisurely strides on the forest floor.
When disturbed, the birds flew just above (i.e., less than 1 ft above)
the ground and realighted 10-20 ft away, in a manner reminiscent of
South American tapaculos (Pteroptochos megapodius, Rhinocryptidae).
‘They were much more often heard than seen.
While E. castanonotus has yet to be recorded away from the Karimui
area, it 1s likely to turn up at most stations in hill forest between about
1,500 and 4,500 ft. The genus Eupetes provides an instance of perfect
triple exclusion, in that the altitudinal range of E. castanonotus lies
strictly above that of E. caerulescens (transition at ca. 1,000-2,000 ft,
E. caerulescens from there down to sea level) and strictly below that of
E. leucostictus (transition at ca. 4,100-5,000 ft). The evolution of the
genus is particularly easy to trace, since each species shows consider-
able subspecific variation in color pattern and some race of each species
can be found which approaches in its color pattern some race of the
species above or below it.
VOICE. ‘The song (Fig. 2la) consists of three to six clear, bell-like,
whistled notes at moderate pitch, all on the same pitch and identical
except for dramatically increasing in volume, and concluded by a very
loud downslurred “chew!” Sometimes a short, unmusical, fainter,
downslurred “ksew” comes between the whistled notes and the “chew!”
The whole song takes about 2 sec. On paper this looks like the song
of several Pachycephala species (P. soror, P. hyperythra, P. rufiventris),
Zi
SPECIES ACCOUNTS
but in practice I felt no danger of confusion, possibly because the
final “chew!” is so distinctive. Heard more often is the common call
(Fig. 21b), three very loud, identical, explosive notes “chew! chew!
chew!” in rapid succession. When heard at a distance, this call can
easily be confused with a similar call of the whistler Pachycare flavo-
grisea, but that of Eupetes castanonotus is louder, somewhat more
rapid, and comes from the ground, not from the trees.
Eupetes castanonotus:
a CHEW ! a ksew CHEW !
whistled, explosive
bell-like
2 sec
2 sec
(b) 7a
CHEW! CHEW! CHEW!
pass ae
Fic. 21. Voice of Eupetes castanonotus.
Eupetes leucostictus loriae Salvadori
High Mountain Eupetes
NATIVE NAME. Fore: iré.
SPECIMENS EXAMINED. Miarosa: 1 ? (17 June 164). Mt. Michael: 1 9, 1 ?
(4 and 7 July 1964). Mt. Karimui Zone 4: 1 ¢,1 9 (30 Aug. and 1 Sept. 1965).
WHRIGHYT. 1 @,49. 1 9, 5.
WING. 1 4,85. 8°99, 77, 83:
TAXONOMY. ‘This series is closest to loriae of southeastern New
Guinea and amabilis of the Huon Peninsula. The latter is blue below
with very little olive, while the former has an olive wash on the breast
and flanks. My specimens have a lesser extent of olive wash than the
average amount for loriae but can be matched by some southeastern
New Guinea birds and are therefore assigned to loriae, as Mayr and
Gilliard (1954, p. 345) similarly concluded for their Wahgi Valley
series.
BREEDING. ‘The male had somewhat enlarged gonads.
DISCUSSION. ‘This eupetes is inconspicuous and uncommon but
212
SPECIES ACCOUNTS
nevertheless widespread on the forest floor from ca. 5,000 to 8,500 ft,
having been encountered by most collectors in the Eastern Highlands.
Its habits are generally similar to those of £. castanonolus, 1.€., 1t walks
on the ground singly or in pairs, but it differs conspicuously in not
possessing or frequently exercising a loud voice.
VOICE. The only sound that I heard was, on one occasion, a faint
“tsip” from one bird of a pair, like the call of a young chick.
Melampitta lugubris longicauda Mayr and Gilliard
Lesser Melampitta
STOMACH CONTENTS. Insects (one stomach), insects and seeds (one).
DISCUSSION. In the central and northern parts of the Eastern
Highlands Gilliard, Gyldenstolpe, Shaw-Mayer, Hitchcock, and Bulmer
recorded this species in dense undergrowth on the forest floor at higher
elevations (ca. 7,000-10,000 ft). Despite intensive netting in apparently
suitable habitats at Okapa, Mt. Michael, and Mt. Karimui I never
obtained it.
Ifrita khowaldi kowaldi (DeVis)
Blue-capped Ifrit
SPECIMENS EXAMINED. Miarosa: 2 ¢ (17 and 24 June 1964). Mt. Michael:
1 9 (5 July 1964). Mt. Karimui Zone 5:1 ¢,1 9; Zone 6: 1 6,2 9: Zone 7: 2
asl oO; Zone 8; 1 S52 9 (28 Aug-s: Sept. 1905).
WEIGHT. 4 @: 33, 34, 35, 36. 6 9: 30, 32, 33(4).
WING. 7 4:86, 86, 87, 88, 89, 90; 92° 6 9: 82584, 86,86, 86, 87.
STOMACH CONTENTS. Insects.
TAXONOMY. These belong to the eastern race; brunnea of
western New Guinea differs by being more brown and less olive above
and by having the remiges and rectrices more rufous. ‘Vhe ear stripe is
white in males but not in females.
BREEDING. ‘The gonads were slightly enlarged in two Mt. Kari-
mui males but were small in other specimens.
DISCUSSION. Virtually all collectors in the Eastern Highlands
have found the Ifrit fairly common in primary midmontane forest up
to about 9,500 ft. Its lower limit is not more than 1,000 ft below the
lower limit of the mossy zone, at an altitude depending upon local
conditions (usually 6,500 or 7,000 ft). It feeds nuthatch-like in moss-
covered trees, probing in moss, pounding at the bark, often bending
over a branch head downwards to probe the undersurface while still
gripping the uppersurface with its legs, and bracing itself stiffly on its
tail. he tip of the tail consequently becomes abraded, as in the case
of the New Zealand warbler Mohoua ochrocephala, which has similar
habits. The Blue-capped Ifrit forages from the understory to the crowns
213
SPECIES ACCOUNTS
and is often seen in pairs or groups of three, calling frequently and
loudly.
VOICE. A rapid series of a half dozen rasped, scratchy, identical
notes, similar to the call of Machacrirhynchus nigripectus or that of
Mohoua ochrocephala.
MALURINAE: WREN WARBLERS
The warblers of New Guinea and Australia fall into four groups
whose relationships are unclear. One group, represented in New
Guinea by five species in the genera Malurus, Todopsis, Cheno-
rhamphus, and Clytomyias, is confined to Australia and New Guinea.
A second, represented in New Guinea by 19 species in the genera
Sericornis, Acanthiza, Gerygone, and perhaps Crateroscelis, is centered
on New Guinea and Australia but has a few representatives on islands
to the east and west. A third consists of grass warblers of predomi-
nantly Eurasian and African genera, of which Acrocephalus, Megalu-
rus, and Cisticola have breeding representatives (five species) in New
Guinea. The fourth consists of the Eurasian and African tree warblers,
of which only Phylloscopus trivirgatus reaches New Guinea. ‘The third
and fourth groups comprise the subfamily Sylviinae. While earlier
authors (and Rand and Gilliard, 1967) placed the second group (Ser-
cornis and its allies) with the Sylviinae and kept the first group as a
separate subfamily Malurinae, Mayr and Amadon (1951) and Keast
(1961) tentatively combine the second group with the Malurinae. ‘The
latter arrangement is arbitrarily adopted here.
Malurus alboscapulatus mafulu Mayr and Rand and
M. a. kutubu Schodde and Hitchcock
Black and White Wren Warbler
NATIVE NAMES. Fore: asasaba. Gimi: férotoro.
SPECIMENS EXAMINED. Awande: 3 ¢, 2 juv. ? (15-20 June 1965). Karimui:
bos, | O,.3 mom, 4, 1 juv. 6 (12 Aus. 19649 12-17 July 1965). Boman 1 OG
July 1965). Soliabeda: 2 6,2 9 (24-29 July 1965).
WEIGIIE, 10) 65 °933-123, (dll = 0). 4 Os 9575) 10s OS ALO 3 siamma
10,05-10:0; 11-0. Lyjuv. gs 10:5:
WING, 10° 4348-52 (50: a2). 4°O 7494). 3 mms (5 48; 495 S9F ay. a
46.
STOMACH CONTENTS. Insects.
TAXONOMY. All four females have white lores and scapulars
and are charcoal brown to black above but vary in the color of the
underparts. One, with enlarged ovaries, is white below except for a
faint breast band (Soliabeda); a second, caught on the nest, is entirely
white below (Bomai); a third has a white chin and belly but blackish
throat and breast (Karimui); and the fourth is largely charcoal brown
below with scattered white feathers and a whitish chin (Soliabeda). Of
the three immature males (all from Karimui), one has white lores,
214
SPECIES ACCOUNTS
scapulars, and underparts, and the upperparts are black except for
scattered brown feathers; a second is similar except that the upper-
parts are charcoal brown; the third is gray-brown above including the
scapulars, and dirty white below with a darker breast band. ‘The
juvenile male is charcoal brown above including the scapulars, while
the underparts and lores are white (Karimui). Both Awande juveniles
are gray-brown except for a white chin. Adult males are black except
for white scapulars; one has a few feathers of the belly white but still
has enlarged testes. The many female and immature Malurus (i.e.,
those apart from the black adult males with white scapulars) seen in
the field at Karimui were equally variable; some were pure white
below, others had a dark breast band, and still others were largely
dark below except for a light throat.
The color of the females and the large size place this series with
mafulu, a race described from the midmontane grasslands of south-
eastern New Guinea and later found in those of the Wahgi Valley.
M. a. naimii, the lowland race of southeastern New Guinea to the west
of Port Moresby, differs by its smaller size and on the average by the
smaller extent of dark areas on the flanks and breast of the female.
M. a. dogwa of the Fly River lowlands differs markedly in the much
smaller size, brown upperparts, buffy flanks, and whitish underparts
without a breast band. Only the Bomai female and the Karimui
juvenile have any buff on the flanks, and these small amounts can
be matched in mafulu. It is interesting that in the case of Malurus
alboscapulatus the newly created grasslands of the Karimui Basin
have been colonized by the midmontane race (mafulw) rather than
by the lowland races (naimii or dogwa).
The Wahgi Valley population was also assigned to mafulu by Mayr
and Gilliard. Surprisingly, Schodde and Hitchcock (1968, p. 42) state
that the population at Lake Kutubu (2,450 ft) and Mendi (5,900-7,000
ft) is very different and they describe it as a new race kutwbu, since
the female is uniformly sooty black. This pattern is shared by three
other spatially isolated populations: moretoni in the southeastern New
Guinea lowland to the east of Port Moresby, aida in the northwestern
New Guinea lowland and midmontane grasslands of the Weyland
Mountains, and randi in the midmontane grasslands at the Wissel
Lakes. The mafulu group (female white ventrally) also has an inter-
rupted distribution in the midmontane grasslands of southeastern
New Guinea and the eastern part of the Eastern Highlands (mafuluw),
the lowlands of southeastern New Guinea (naimiz), the lowlands of
northeastern New Guinea (tappenbecki), and the Vogelkop (nominate
alboscapulatus). ‘The third group, in which females are brown and
white, occurs in the lowlands of southern New Guinea (dogwa) and
southwestern New Guinea (lorentzi) and the midmontane grasslands
of the Baliem Valley (balim). An explanation for the checkerboard
distributions of the three female plumage types is lacking.
215
SPECIES ACCOUNTS
BREEDING. Natives brought in two nests, bowls of dry grass
which were said to have been on the ground in grassland. One con-
tained one fledgling, the other two. All adult males taken at all my
collecting localities had greatly enlarged testes. Evidently breeding is
concentrated in the dry season. According to Fore informants nests of
Malurus are those most often victimized by the cuckoo Cacomantis
variolosus.
DISCUSSION. Malurus alboscapulatus is probably the most abun-
dant and ubiquitous Eastern Highlands bird in grassland and gardens,
up to about 7,000 ft. At Soliabeda, where a few years before my arrival
two small gardens had been cleared in the middle of a large area of
undisturbed forest, M. alboscapulatus was already common and breed-
ing though the other characteristic species of the midmontane grass-
lands had not yet been able to colonize this tiny island of suitable
habitat. It is usually seen in small groups, perhaps family groups, of
two to six birds. ‘The birds can readily be called out of the depths of
the grassland to perches on tall stalks of grass, but I have never seen
them perched more than 6 ft above the ground. Their flights are
jerky, brief, and give every impression of being weak, despite which
they must have crossed nine miles of unbroken forest, including a
mountain ridge 1,000 ft high, to have reached Soliabeda from the
nearest grassland.
VOICE. A rapidly spitted twitter or jumble of notes full of sibi-
lants, inspiring the Fore name “a-sa-sa-ba.”
Todopsis cyanocephala bonapartii Gray
Blue Wren Warbler
DISCUSSION. Schodde and Hitchcock found this low-altitude
wren warbler at Lake Kutubu in dense second-growth and at the forest
edge.
VOICE. A rapid series of spitted notes very similar to the call of
Malurus alboscapulatus.
Clytomytas insignis oortt Rothschild and Hartert
Rufous Wren Warbler
NATIVE NAME. Fore: tabugiri, or asasaba bilong bush(= “asasaba of the
forest”: cf. Malurus alboscapulatus).
SPECIMENS EXAMINED. Awande: 1 ¢@ (a) (14 June 1965). Mt. Karimui Zone
br 1 OF Zone 6: 1 4; Zone 8: 1s.
WHICH. 2 6712.0, 1275 9295 152(a)yels0:
WING, 202 56,582 2 O59 K(a),. 57.
STOMACH CONTENTS. Insects.
TAXONOMY. These agree with oorti in the underparts being
buffier than in nominate insignis of the Vogelkop, which has a nearly
white throat.
216
SPECIES ACCOUNTS
BREEDING. The Awande female (marked ‘“‘a’’) contained two
nearly formed eggs, while the gonads of the Mt. Karimui specimens
were small.
DISCUSSION. Clytomyias insignis seems to be local and uncom-
mon in midmontane forest. All four of my specimens were netted,
indicating preference for the understory. The Fore said that Clytomyzas
is very similar in behavior to Malurus alboscapulatus and that it goes
about near the ground and around fallen trees with cocked tail.
SYLVIINAE: WARBLERS
Niche Differences in the Genus Sericornis
‘This genus presents not only difficult taxonomic problems but also
some of the most complicated altitudinal relationships in New Guinea,
including parallel four-species and two-species altitudinal sequences
(Diamond, 1969). Four of the species (S. spilodera, S. arfakianus, S.
perspicillatus, S. papuensis) are small (weight ca. 9-12), have small
bills (culmen from base 12-13), forage both in the lower- and middle-
stories, and constitute a four-species altitudinal sequence. ‘The remain-
ing two widespread species, S. virgatus (absent in the Eastern High-
lands) and S. nouhuysi, are large (weight ca. 11-18), have large bills
(culmen from base 15-16), forage mainly in the lower story, and ex-
clude each other altitudinally, but each overlaps several of the small
species. S. beccari is related to S. virgatus but is confined to the flat
lowlands of southern New Guinea and the Cape York Peninsula of
Australia, while S. nigroviridis is a mystery bird of distinct appearance,
known from one specimen.
The altitudinal ranges of the abundant four small Sericornis species
on Mt. Karimui were:
S. spilodera 1,350 ft or below—4,240 ft
S. arfakianus 4,400 ft —5},000 ft
S. perspicillatus —6,350 ft 5,100 ft
S. papuensis —summit (8,165 ft) 6,450 ft
All transitions were sharp, i.e., no individual of any species was
caught, seen, or heard in another’s altitudinal band, except for two
S. spilodera shot together at 4,950 ft, far into S. arfakianus’s band.
‘These two were young birds with short wings, low weights, and gonads
not discernible, and conform to the general rule (p. 00) that normal
altitudinal limits are most likely to be violated by young birds.
Sericornis spilodera guttatus (Sharpe)
Pale-billed Sericornis
NATIVE NAME. Daribi: saboba.
SPECIMENS EXAMINED. Karimui: 9 ¢,1 9 (31 July-17 Aug. 1964; 3-17 July
ald
SPECIES ACCOUNTS
1965). Bomai: 1 ¢ (8 July 1965). Soliabeda: 2 ¢
Karimui Zone 1: 4 ¢,2 9; Zone 2: 1 @; Zone 3: 2:
WEIGHT. 10 @: 10.3-13.2 (119+0.8). 3 9: 10.
WING. 10 @: 56-63 (60+ 2). 2 9:57, 58.
(22 and 27 July 1965). Mt.
? (10-16 Aug. 1965).
b, LO 0:
TAXONOMY. ‘This series is closest to guilatus of southeastern
New Guinea and agrees with it in size but differs in the darker throat
spots, deeper yellow color below, darker upper tail, and darker,
brighter, and more olive back. S. spilodera thus follows the general
trend at Karimui towards dark coloration. However, these differences
seem not sufficiently marked to justify naming a new race. The race
wurot of the Fly River is paler and greener below, paler and less bright
above, slightly smaller, and the throat spotting is paler; granti of the
southern slope of the Snow Mountains is paler below, paler, browner,
and less olive on the back, tail, and crown, and with paler, browner,
less gray and less clear throat spotting; nominate spilodera of northern
New Guinea is paler and less yellow below, darker on the crown, and
the throat spotting is less dark; ferrwgineus of Waigeu is much paler
above and below and has the throat spotting nearly obsolete. ‘The
whole culmen measures 13 to 14 in my specimens.
BREEDING. At Karimui both in 1964 and 1965, and at Bomai
and Soliabeda, most males and the one female obtained had greatly
enlarged gonads, suggesting breeding in the dry season for this species
of the understory. The sex ratio of specimens for those localities was
very unequal in favor of males (12:1), as in the case of S. papuensis on
Mt. Karimui (possibly because females were staying on nests?). In
contrast, all specimens obtained on Mt. Karimui had very small
gonads. These individuals near the upper limit of the altitudinal
range must be young birds or nonbreeding adults, a pattern observed
in other species as well (cf. p. 30).
DISCUSSION. _ S. spilodera is the Sericornis species with the lowest
altitudinal range and is present in hill forest on all major mountain
ranges of New Guinea, becoming rare or absent at sea level. The
altitude of 4,240-4,400 ft for the S. spilodera-S. arfakianus transition on
Mt. Karimui is somewhat higher than usually found elsewhere in New
Guinea, probably due to the effect of the tropical Karimin Basin
shifting most hill forest ranges upwards.
In the field S. spilodera is most easily distinguished from other
species of Sericornis by its pale, horn-colored bill.
Sericornis arfakianus (Salvadori)
Gray-green Sericornis
SPECIMENS EXAMINED. Okasa: 2 6, 1 ? (22-24 June 1965). Sena River: 1 ?
(26 July 1964). Mt. Karimui Zone 2:1 ¢,1 9, 2; Zone 3: I ¢ (13-18 Aug, 1965).
WEIGHT, 4 2: 9.0, 9.0, 9.0, 10.3. 1 Os wo. 2 rs 0b, 9:5)
WING. 4 @: 48, 52, 55,56. 1 9:48. 27: Ol, ol.
CULMEN FROM BASE. 4 ¢@: 120, 125, 125, 13.0. i Qe 120) «22 120; 120.
218
SPECIES ACCOUNTS
TAXONOMY. The slight geographical variation in this species
follows an irregular checkerboard that renders impossible the defini-
tion of subspecies with coherent ranges. In dorsal coloration, popula-
tions from the Weyland Mountains, Cyclops Mountains, North Coastal
Range, and southeastern New Guinea are brighter than birds from
the Idenburg slopes, Adelbert Mountains, and Telefolmin, with
Vogelkop birds still duller. ‘The underparts range from yellowish with
slight streaking to grayer with more streaking, the sequence of popula-
tions from yellower to grayer being Cyclops > Weyland > North
Coastal Range > southeastern New Guinea > Adelbert Mountains >
‘Telefolmin, Vogelkop, Idenburg slopes.
BREEDING. ‘Testes were large in both Okasa males and one of
the Mt. Karimui males.
DISCUSSION. This is the second member in the altitudinal se-
quence of small Sericornis, living above S. spilodera and below S.
perspicillatus between ca. 4,000 and 5,000 ft. On most New Guinea
mountains, in my experience and to judge from records of the First
and the Third Archbold Expeditions (Mayr & Rand, 1937; Rand,
1942b), its total vertical range is only about 1,500 ft, but even with
respect to this narrow band its Mt. Karimui range is compressed (to
600 ft).
VOICE. Dry scolding notes, and a dry call note “chip.”
Sericornis perspicillatus Salvadori
Buff-faced Sericornis
NATIVE NAME. Fore: pasagekiyabi.
SPECIMENS EXAMINED. Awande: 2 ¢, 1 9 (20 June 1964; 15 June 1965).
Mit. Michael: I 4 (2: July 1964), Mt. Karimui Zone 3: 2 ?; Zone 4; 2 4, 1
fone pez .3., 1 PG Aug-1 Sept. 1965).
WEIGEL. 95) @: 8.0; 8.8, 9.0, 100; 10.2. Bb O::-8:0;
WING. ais Oils Oto (5), 2.0% 50,51.
CULMEN FROM BASE. 7 4: 12.0 (3), 22.5, 13:0) (3).
STOMACH CONTENTS. Insects.
BREEDING. ‘Three of the four Mt. Karimui males had greatly
enlarged testes, suggesting breeding at the same time as other species
of Sericornis at Karimui.
DISCUSSION. 8S. perspicillatus and S. nouwhuysi are superficially
rather similar because of the yellowish underparts and orange face in
both species. ‘Ihe iris is generally dull brown in S. perspicillatus,
whereas it is often red-brown in S. nouhwysi.
S. perspicillatus is the third in the altitudinal sequence of small
Sericornis. As with many other birds of midmontane forest, the descent
of stunted moss forest to 6,500 ft on Mt. Karimui resulted in an
unusually low upper limit for S. perspicillatus (6,350 ft), and its upper
limit on most other mountains (e.g., Mt. Michael) is around 8,000 ft.
219
SPECIES ACCOUNTS
It flits nervously in small groups of a few to eight birds in the under-
story and in trees up to about 40 ft above the ground, not only in the
forest interior but also at the forest edge, in partly cut forest, and
even in casuarina groves which have no bushes in the understory.
VOICE. ‘The common call is a dry “chip,” recognizable with prac-
tice. A more distinctive vocalization, perhaps the song, is a rapid series
of “chip’s’” which progressively rise in pitch and grow thinner in
quality; the series may be preceded by a more substantial “chip” at a
higher pitch (Fig. 22). There are also scolding and chattering notes.
Sericornis perspicillatus:
Sericornis papuensis:
or -
or or Ve ne
ne on ae
Fic. 22. Voices of Sericornis perspicillatus and S. papuensis.
Sericornis papuensis papuensis (De Vis)
and (?) S. p. buergerst Stresemann
Papuan Sericornis
SPECIMENS EXAMINED. Mt. Michael: 1 ? (6 July 1964). Mt. Karimui Zone
5: 1 9, 1 imm. ?; Zone 6:9 g, 2 9; Zone 7: 2 4; Zone 8,9 @ (8 Aug.-8 Sept.
1965).
220
SPECIES ACCOUNTS
WEIGHT. 10 ¢: 9.8-11.7 (10.7+0.5). 3 9: 10.0, 10.3, 10.7. 1 imm, ?: 95.
WING. 10 @: 55-6! (522 1), 93 O: 52,59, 59. 1 imum, 7:52:
CULMEN FROM BASE. 10 @: 12.0-14.0 (12.8+0.5). 3 Q: 125 (8). 1 imm.
cay dere
STOMACH CONTENTS. Insects.
TAXONOMY. ‘The mark which I find most useful in the hand for
distinguishing specimens of this difficult and variable species from
other Sericornis and from Acanthiza murina is on the forehead: for
2-3 mm from the base of the bill the forehead has the same light orange
color as the face, with dark tips to the feathers; this contrasts with the
darker color of the more posterior parts of the forehead and the
crown. From 8S. perspicillatus, 8. papuensis is distinguished by this
forehead mark, the subterminal tail band, longer wing, and usually
by the browner, less olive, coloration, especially on the wings and tail.
Marks I find useful, besides the forehead, for separating the super-
ficially similar Acanthiza murina are: the color of the tail (dull gray
or gray-olive, especially near the tip, in A. murina, brownish-olive in
S. papuensis; the subterminal tail band (much more distinct in the
former); the light orange eye ring of the latter, absent in the former;
the edges of the primaries (olive in the former, often brownish-olive
in the latter); the whitish chin, speckled with black on the sides, in
A. murina; and, in life, the dull brown iris of S. papuensis, as opposed
to the whitish or pale tan iris of A. murina.
All of my males prepared as skins, and two of my three females, form
a quite uniform series, showing little variation. “The Mt. Karimui Zone
5 female from the lower limit of the altitudinal range differs in its
darker wings and underparts. The Zone 5 immature taken with this
female is quite yellow below, a feature which can be matched in young
birds from other parts of New Guinea. Elsewhere in New Guinea this
is a highly variable species, with plumage varying from brownish to
greenish. On the average the brownest series are those assigned to the
race buergerst from the Snow Mountains and Telefolmin (topotypical
buergersi from the Schraderberg were unavailable); nominate papuen-
sis from southeastern New Guinea averages more olive, and meeki
from Mt. Goliath is much greener. ‘The present series is intermediate
between nominate papuensis and buergersi but closer to papuensis,
from which it differs mainly in that papuensis averages less dark above.
Both the more brownish and the more greenish individuals from the
Snow Mountains have darker underparts than the Mt. Karimui series,
and no Mt. Karimui specimen is really brown as are many from the
Snow Mountains.
The two specimens collected by Gilliard on Mt. Wilhelm and Mt.
Hagen, which Mayr and Gilliard (1954, p. 347) assigned to buergersi,
are browner below and duller above than Mt. Karimui specimens. I
have not seen Gyldenstolpe’s five adults from the Wahgi Valley, which
he described (1955, p. 93) as being variably browner or greener above
and which he assigned to buergersi. More specimens of this variable
PPA |
SPECIES ACCOUNTS
species would be needed to decide whether Wilhelm-Hagen-Wahgi
birds really do average browner than those from Mt. Karimui 50
mules to the south.
BREEDING. The disparate sex ratio is notable (20 g, 3 9). With-
out exception all males had greatly enlarged testes. Of the two Zone 6
females, one had the ovaries enlarged, while I did not record ovary
size in the other. The two Zone 5 specimens (6,450 ft) were an im-
mature with undiscernible gonads and a female with small ovaries,
conforming to the general pattern of nonbreeding birds being found
at the extremes of the altitudinal range, From 6,550 it up tothe
summit of Mt. Karimui (8,165 ft) S. papuensis was singing often and
loudly.
DISCUSSION. S. papuensis is the fourth and highest in the alti-
tudinal sequence of small Sericornis. It foraged from near the ground
to about 30 ft above the ground in groups of up to four birds, gleaning
mainly on branches and twigs, rarely on main trunks.
When I discussed Sericornis ecology previously (Diamond, 1969, pp.
26-30), I had not had an opportunity to study S. papuensis in the
presence of Acanthiza murina. ‘The published records of the First
Archbold Expedition from Mt. Albert-Edward appeared to suggest that
interaction with A. murina altered the niche of S. papuensis so that the
latter and S. perspicillatus overlapped in altitudinal range. ‘This proves
not to be the case, as shown by more recent field work I carried out on
Mt. Albert-Edward. Despite the presence of A. murina, the ranges of
S. papuensis and S. perspicillatus there are still nearly mutually ex-
clusive, as on Mt. Karimui in the absence of A. murina, and overlap
is confined to a few immature individuals.
VOICE. ‘The song (Fig. 22, p. 220) is a loud, rapid, unmusical out-
burst with a somewhat harsh and unclear quality, consisting of five to
eight notes, all of which are on one of two (occasionally one of three)
pitches. The song is sometimes prefixed by two or three faint notes.
The call is a dry “chip” similar to that of other Sericornis species.
Sericornis nouhuysi stresemanni Mayr and
S. n. oorti Rothschild and Hartert
Large Mountain Sericornis
NATIVE NAME. Fore: mabiséna.
SPECIMENS EXAMINED. Awande: 3 6, 2 9 (20-28 June 1964; 15-20 June
1965). Mt. Michael: 2 ¢,1 9 (4-10 July 1964). Mt, Karim Zone 3: I 4, 1°;
Zone 4: 2 4; Zone 5: 2 6,2 9; Zone 6: 2 9; Zone 8: 1 ¢ (16 Aug.-7 Sept. 1965).
WEIGHT. 9 4: 13.0-17.7 (1621.7). 7 9: TE0-Ib.8 (146s: 0.0),
WING. 10 @: 61-67 (6442). 10 Q: 58-64 (61 + 2).
CULMEN FROM BASE. 10 ¢@: 14.0-17.0 (15.341.1). 10 9: 140-160 (54+
().8).
STOMACH CONTENTS. Insects.
222
SPECIES ACCOUNTS
TAXONOMY. The Awande and Mt. Karimui series differ in color,
though not in size. Awande specimens have quite olive backs, rather
yellow underparts, not dingy on the breast, and are closest to oorti of
southeastern New Guinea. Mt. Karimui specimens are more rufous
above, less yellow below, dingier on the breast, and closer to the de-
scription of stresemanni from the Schraderberg and to the series col-
lected by Gilliard in the Wahgi region and assigned to stresemannt.
The race S. n. nouhwysi of western New Guinea is even more rufous,
while Telefolmin birds were assigned to stresemanni. Evidently there
is a west-to-east color cline in S. nouhwysi, from more rufous in the
west to more olive in the east. It seems most convenient to place the
stresemanni-oort? transition between the Wahgi-Karimui area and the
Okapa area 100 miles to the east.
‘The iris is either brown, orange-brown, or red-brown in both males
and females.
BREEDING. Gonads were enlarged in only two Awande speci-
mens, but were greatly enlarged in all Mt. Karimui males and two of
the females. All five Sericornis species taken in the Karimui area were
evidently breeding in July, August, and September. At 10,200 ft on
Mt. Michael an adult was captured on its nest on 10 July 1964. The
nest was 4 ft above the ground in a bunch of moss hanging from a tree
limb, and contained two eggs.
DISCUSSION. S. nouhwysi has been found common by most col-
lectors at most localities in the Eastern Highlands from about 4,500 to
11,000 ft or more, in forest and dense second-growth. Compared to the
four smaller species of Sericornis, S. nouhuysi spends a greater fraction
of its time in the understory, forages more on main branches and
trunks than on twigs and small branches, and often works vertically
up a trunk in the manner of a creeper or nuthatch, probing at the
bark and moss. ‘The altitudinal range of S. nowhwysit takes in the entire
ranges of S. perspicillatus and S. papuensis on Mt. Karimui and over-
laps S. arfakianus at the Sena River and Acanthiza murina on Mt.
Michael. “The independence of its distribution from that of these
other small warblers confirms that it occupies a different type of niche,
associated with its more marked preference for the understory, dif-
ferent foraging technique, and larger size and longer bill (all but one
of my S. nouhuysi specimens weigh more than all specimens of all
other Sericornis species I took).
VOICE. ‘The Fore pointed out to me a song said to come from this
species: a thin, slightly sibilant, rambling warble of only modest length,
dropping in pitch towards the end. The call notes are a dry ‘ chip”,
a spitted note unlike that of S. perspicillatus, and a rasped scolding
note.
223
SPECIES ACCOUNTS
Sericornis beccarti subsp.
Beccari’s Sericornis
or
Sericornis virgatus subsp.
Perplexing Sericornis
These two species include a confusing group of populations among
which the species lines are not fully clear. As discussed elsewhere
(Diamond, 1969), S. virgatus is apparently a hill forest species living
at 2,000-4,500 ft on the Vogelkop, the northern slopes of the Central
Range west of long. 143°F, and several of the “north mountain
islands.” S. beccarit is apparently confined to the flat lowlands of
southern New Guinea and the Aru Islands near sea level, and to the
Cape York Peninsula of Australia. At Lake Kutubu (2,450 ft) Schodde
and Hitchcock (1968, p. 45) took one specimen of this complex, the
first record from the southern hill slopes of the Central Range. ‘They
tentatively related it to the population tdenburgi of northwestern
New Guinea, based on the published description of that race, but
they lacked comparative material, without which racial assignment of
Sericornis forms is practically impossible, and they recognized that the
affiliation with tdenburgi was improbable on geographical grounds.
Further speculation as to the race and species represented must await
taxonomic comparison of the specimen.
Acanthiza murina (De Vis)
New Guinea Mountain ‘Thornbill
SPECIMENS EXAMINED. Mt. Michael: 1 Q, 4 ? (10 July 1964).
WWIINKG I et fey I a (Ole
GULMEN FROM BASE. TO} 2b. Ter Tio!
STOMACH CONTENTS. Insects.
TAXONOMY. Acanthiza murina is confusingly similar super-
ficially to the smaller species of Sericornis. It has a distinct, dark, sub-
terminal band on the tail, a whitish iris, an absolutely even, dingy,
ventral coloration (no orange on the throat or cheeks), and a dull,
even, olive color above. Differences between it and Sericornis papuensis
are summarized under S. papuensis. It may be recognized in the field
by the very short tail and the whitish underparts.
DISCUSSION. Of the small warblers, Acanthiza murina has the
highest altitudinal range, being confined to moss forest above 8,000 ft,
mainly above 10,000 ft. Its altitudinal range is shared with Sericornis
papuensis and S. nouhuysi, from which it differs ecologically in that
it forages from the treetops down to 15 ft above the ground, never
lower. The ecologically equivalent warbler at lower elevations is
224
SPECIES ACCOUNTS
Gerygone cinerea. Acanthiza murina is usually seen in loose flocks of
3-10 individuals spread out over several adjacent trees, eleaning (not
sallying or hovering) on leaves and small twigs, rarely on larger
branches. It moves by short hops of several inches up to a foot, and
the flocks feed systematically for several minutes in one tree before
moving on to the next tree.
VOICE. ‘The song consists of four similar pairs of notes, the first
member of each pair being on a higher pitch than the second. Members
of flocks continually give faint, sweet, almost tinkling contact calls, by
which the flock is easily located.
Songs of the Genus Gerygone
There are five species of Gerygone whose songs I have had the op-
portunity to hear at various localities: G. ruficollis in the Eastern
Highlands and a recording from the Baliem Valley of western New
Guinea, G. chrysogaster in the Eastern Highlands and North Coastal
Range, G. magnirostris in the North Coastal Range and on Karkar,
G. olivacea in southeastern New Guinea and in Queensland (Aus-
tralia), and G. igata in New Zealand. The songs of all five are high-
pitched, have a thin and somewhat plaintive quality, are delivered
fairly fast, and are based on machine-gun patterns of about a half
dozen notes which are repeated two to nine times in immediate suc-
cession. Figure 23 illustrates the songs of G. ruficollis and G. chryso-
gaster in the Eastern Highlands, and of G. olivacea, G. magnirostris,
and G. igata for comparison. I did not hear songs from G. cinerea, G.
chloronota, or G. palpebrosa.
Gerygone cinerea Salvadori
Gray Gerygone Warbler
SPECIMENS EXAMINED. Awande: 1 ¢,1 9 (14 and 20 June 1965). Miarosa:
by 2 jone 1964). Mc. Kanmini Zone 3: 1 fs Zone 7:2 ¢ (20 Amges Sept.. 1965):
WEIGHT. 4 4: 7.0, 747, 7.7, 8.0. 1 9: 72.
WING, 4-23 51, 52). 593545 1 9252, 1 P51.
STOMACH CONTENTS. Insects.
BREEDING. All three Mt. Karimui specimens had the testes much
enlarged.
DISCUSSION. ‘This tiny, inconspicuous warbler occupies between
about 5,400 and 8,500 ft in forest the niche that Acanthiza murina
does at higher altitudes. Flocks of 2-10 individuals spread out over
several trees, forage from the treetops down to 15 ft above the eround,
gleaning leaves, twigs, and small branches. Unlike Acanthiza murina,
Gerygone cinerea occasionally hovers and makes short sallies, its be-
havior is more nervous, and its foraging less systematic. At the forest
edge it also gleans in the understory. ‘The transition altitude between
225
SPECIES ACCOUNTS
Gerygone ruficollis (Eastern Highlands):
Gerygone chrysogaster (Eastern Highlands):
eee as
Gerygone olivacea (southeast New Guinea, east Australia):
Es AuSeC
Gerygone igata (New Zealand):
Opes SSeeee. | sek Pe Sere ats SP ee aoe ee eee
Gerygone magnirostris (north New Guinea):
aA 7 Ae EA Say
Fic. 23. Voices of five Gerygone warblers.
226
SPECIES ACCOUNTS
Gerygone cinerea and Acanthiza murina varies locally between 7,800
and 9,300 ft. This is one of the few instances of sharp altitudinal re-
placement among non-congeneric species in New Guinea.
VOICE. Members of flocks give faint, sibilant contact calls.
Gerygone chrysogaster chrysogasler Grey
Yellow-bellied Gerygone Warbler
SPECIMENS EXAMINED. Soliabeda: 1 ¢ (25 July 1965).
WEIGHT. 9.5.
WING. 55.
STOMACH CONTENTS. Insects.
TAXONOMY. ‘The black bill and the brownish-olive upperparts
place this specimen with nominate chrysogaster (notata and dohertyi
have olive upperparts and a horn-colored bill).
BREEDING. ‘The greatly enlarged testes of the specimen and the
frequent songs indicate that G. chrysogaster was in breeding condition
at Soliabeda.
DISCUSSION. At Soliabeda this low-altitude warbler was common,
accounting for 69% of bird individuals. One song was heard at 2,150 ft
between Soliabeda and Karimui, but the species was absent at Karimui,
and there are no other Eastern Highlands records. Single individuals
or small groups were seen moving rapidly in forest and in second-
growth trees, generally in the middlestory but sometimes descending
to within a few feet of the ground. G. palpebrosa was also present at
Soliabeda but apparently did not descend into the lower story.
VOICE. A typical weak Gerygone song consisting of a repeated
five-note pattern introduced by three or four extra notes (Fig. 23, p.
226). If one discharges a gun and G. chrysogaster is in the vicinity, it
immediately sharts up its song. A continuous dry chirping may also be
heard occasionally.
Gerygone chloronota subsp.
Gray-headed Gerygone Warbler
I have no specimens but do have sight records of birds observed at
such close quarters as to preclude the possibility of misidentification.
Between 6 and 9 July 1965 I saw one individual daily at Bomai,
flitting 10-60 ft above the ground in a tree at the edge of a ravine. In
August 1964, ‘lerborgh studied some groups of 2-6 in a small flowering
tree at Karimui, standing at the edge of a garden several hundred yards
from forest. ‘hey were the only bird species exploiting this tree and
kept up a nearly continuous flow of soft vocalizations as they probed
between the petals. ‘The species evidently prefers trees in open habitats.
In New Guinea G. chloronota is a rare species, known from few and
scattered localities.
227
SPECIES ACCOUNTS
Gerygone palpebrosa inconspicua Ramsay
Black-headed Gerygone Warbler
SPECIMENS EXAMINED. Karimui: 2 ¢ (1 and 2 July 1965).
WEIGEHE. 2 62-8, 9.
WING. 2 @: 52, 54.
STOMACH CONTENTS. Insects.
TAXONOMY. ‘These belong to the southeastern New Guinea
race. The race tarara from coastal parts of the southern New Guinea
bulge west of the Fly River differs in the male most obviously in the
brownish tinge of the black parts of the throat and head; wahnesi
of northern and northeastern New Guinea differs in the black crown
(olive in the Karimui specimens); and nominate palpebrosa of the
Vogelkop differs in the brighter and more yellow-green back. The
dorsal coloration of museum specimens foxes with age.
BREEDING. ‘The testes were greatly enlarged in both specimens.
DISCUSSION. In the Karimui region G. palpebrosa was present
from 4,200 ft down to at least 1,350 ft. ‘There seem to be no other
Eastern Highlands records yet, but it will doubtless turn up at other
hill forest stations. Pairs or small groups flit rapidly in trees of the
forest and at the forest edge. I never netted this warbler or saw it less
than 10 ft from the ground.
Gerygone magnirostris subsp.
Swamp Gerygone Warbler
STOMACH CONTENTS. Insects.
DISCUSSION. ‘This lowland warbler has been observed in stream-
side vegetation at 3,500 ft near Baiyer River, an altitudinal record
for the species (J. Kikkawa, pers. comm.; N.G.B.S. Newsletter, No. 54,
p24, Jr: 19:70):
VOICE. A repeated four-to-six-note pattern of slurs (Fig. 23), with
a slightly nasal quality and a substantial volume, suggestive of a species
of Lalage rather than Gerygone.
Gerygone ruficollis insperata De Vis
Red-necked Gerygone Warbler
NATIVE NAME. Fore: enemesilo.
SPECIMENS EXAMINED. Awande: | juv. ? (18 June 1965). Miarosa: 1 ? (17
June 1964). Lufa: 2 ? (29 June and 13 July 1964).
WEIGHT. 1 juv. ?2 7.
WING. 1 juv. ?: 48.
STOMACH CONTENTS. Insects.
TAXONOMY. The Awande juvenile is pale lemon below, becom-
ing paler towards the lower belly, dark brown with an olive tinge on
228
SPECIES ACCOUNTS
the back, and dingy dark olive on the head. ‘The tail feathers have the
characteristic white spots.
DISCUSSION. With the possible exception of the honeyeater
Melidectes torquatus, no other midmontane species has profited so
spectacularly from native agriculture as G. ruficollis. In primary forest
it is hard to see but easy to locate by song. ‘These songs indicate that
it was not uncommon in primary forest on Mt. Karimui between 4,500
and 6,500 ft, and present but uncommon around 8,000 ft on Mt.
Michael. However, in trees in open habitats cleared by man in the
Eastern Highlands, and especially in casuarina groves, this nervous
warbler is often the most abundant bird, occurring in flocks of up to
20 and even up to 50 birds. It has similarly colonized the open habitats,
again casuarina groves in particular (Rand, 1942b, p. 476), in the
Baliem Valley of western New Guinea. In southeastern New Guinea,
but apparently not elsewhere, it is common in treeferns in alpine
grassland.
VOICE. An unmistakable, high pitched, plaintive, fairly rapid (ca.
six notes per second), and lengthy song. ‘The first 10 to 30 notes
gradually descend in pitch, eventually level out at a constant pitch,
and thereafter every fifth note is lower in pitch by an interval of a
third or a fourth, forming a five-note pattern which is the origin of
the Fore nace ‘“‘e-ne-me-si-lo”. The “enemesilo” phrase is repeated
three to nine times to conclude the song (Fig. 23, p. 226). Sometimes all
the “enemesilo” phrases are omitted, and the song terminates after the
initial 10 to 30 notes. Voice recordings from the Baliem Valley which
I have heard are essentially the same.
Ripley (1964, p. 59) gives the following description of the song of the
warbler Phylloscopus trivirgatus in the Baliem Valley: ““Uhese warblers
have a long, drawn-out series of single whistling notes on a descending
scale, rather pretty but mournful. ‘The notes remind me of the song of a
North American White-throated Sparrow, Zonotrichia.” ‘This is an
accurate description of the Gerygone ruficollis song. ‘The song of
Phylloscopus trivirgatus is also high but completely different in form
and pattern, and the attribution of the Gerygone song to Phylloscopus
is surely an error, one, however, which could arise easily, as both are
small and nervous tree warblers.
Niche Differences in the Genus Crateroscelis
This genus of three species, which was formerly placed with the
babblers but is now assigned to the warblers, illustrates the course of
altitudinal sequences of more than two congeners in New Guinea. All
three species are quite similar in appearance and habits. In a few areas
all three are present and replace each other subsequentially: C. mwrina
at low altitudes, C. nigrorufa at middle altitudes, C. robusta at higher
altitudes. In many or most areas, however, C. nigrorufa is absent, and
229
SPECIES ACCOUNTS
there is a sharp boundary between the altitudinal ranges of C. murina
and C. robusta. This “squeezing-out” of the middle species seems to be
a common result of multiple altitudinal replacement in other genera
as well (Pe 94),
Crateroscelis murina murina (Sclater)
Lowland Mouse-Warbler
NATIVE NAMES. Gimi: sikaide. Daribi: kirili or pfroni.
SPECIMENS EXAMINED. Okasa: 3 ¢ (23 and 26 June 1965). Karimui: 4 ¢,
2 9, (31 July-16 Aug. 1964; 1-14 July 1965). Soliabeda: 2 Q (23-30 July 1965).
Mt. Karimui Zone 1: 3 ¢,5 9; Zone 2: 1 4; Zone 3:1 4,1 @ (9-19 Aug. 1965).
WEIGHT. 9 : 13.0-17.0 (15.0 41.1). 8 9: 14.3-16.0 (15.0 + 0.7).
WING. 10 ¢: 59-63 (61 +1). 10 9: 55-61 (57 + 2).
TAXONOMY. The races monacha, pallida, fumosa, and capitalis
are paler, especially on the underparts, while fumosa and capitalis are
also smaller. The Eastern Highland series belongs to the dark-bodied,
dark-crowned, widespread New Guinea race C. m. murina. Birds from
the Karimui area average darker than those from Okasa, as is often
the case at Karimui.
BREEDING. Gonad condition was as follows: Okasa, testes small
in all males examined; Karimui, testes enlarged in most but not all
males, one nestling brought in on 16 July; Mt. Karimui, testes small
in three males, slightly enlarged in a fourth, ovaries small in all
females. ‘his suggests that breeding of this lowerstory species is in the
dry season at Karimui but not at Okasa or on Mt. Karimui.
DISCUSSION. C. murina is the commonest bird in the understory
of hill forest. In the flat coastal plains and near sea level it is quite
sparsely distributed, sometimes even absent. At Soliabeda (1,500-2,000
ft) it still accounted for only 1.2% of all birds. This figure had risen
to 4.9% at Karimui (3,650 ft), and thereafter C. murina became
progressively more abundant as we worked our way up Mt. Karimui,
accounting for 9.2% of all birds in Zone 3, until it abruptly disap-
peared between 5,390 and 5,400 ft, where it was replaced by C. robusta.
One of our campsites was near this transition altitude, and I had
frequent opportunity to check the strictness of the transition by noting
that C. murina songs always came from below this altitude, C. robusta
from above.
C. murina remains within the forest and is seen as solitary indi-
viduals or as small parties of two or three birds, on the ground, on
fallen logs, in low bushes, and in thickets. I have not seen it more
than 5 ft above the ground. Singers are dificult to locate, and the
species is far more often heard than seen.
VOICE. A beautiful whistled song characterized by the distinct-
ness of its notes, the simplicity of its patterns, and the very clear
quality sug egestive of a flute in the higher registers, and, like a flute,
230
SPECIES ACCOUNTS
with a “breathy” quality when heard close. Patterns consist of 2-10
notes, most commonly of either three or four notes. Frequently each
note has the same time value; sometimes a dotted rhythm is used, the
first note may be detached, or a subsequent note may be held longer
than the others. An average length for the song is 1-114 sec, but 3 sec
is not unusual. The pitch range of the whole song may be encompassed
within a fourth or major third, and intervals between successive notes
are close to half-tones, whole tones, or thirds. A given individual uses
at least eight to 10 different patterns. Each pattern is repeated three to
six, sometimes up to 16, times, with pauses between repetitions, before
the singer goes on to the next pattern. Figure 24 illustrates many of the
C. murina patterns I have heard in the Eastern Highlands (patterns |
through 10 were used successively by one singer, I] through 18 by
another). ‘Phe call is a dry “chip” or a squawk.
Crateroscelis murina:
(1) --- (2) --— () aes (2 een (61) i
2 sec 3
(oan (0 adem (S) ae (S)GaT Oh Soe See |
(M1) = (2 ao (13) = (4)= 2 Ga > 2
GS) eee hr) | aoa G3) @
Crateroscelis robusta:
Fic. 24. Song patterns of Crateroscelis murina and C. robusta.
Crateroscelis nigrorufa nigrorufa (Salvadori)
Midmountain Mouse-Warbler
NATIVE NAME. Fore: funtara.
SPECIMENS EXAMINED. Awande: 1 ¢ (12 July 1967).
WING. 61.
TAXONOMY. In the coloration of the lower flanks, belly, and
undertail coverts my specimen agrees with nominate nigrorufa of
23]
SPECIES ACCOUNTS
southeastern New Guinea rather than with blissi of the Snow Moun-
tains and Weyland Mountains, which is darker and blacker. Com-
pared to my specimen, southeastern New Guinea birds are slightly
browner, less black, dorsally, but this may be due to foxing, since
they were collected in 1903-1906.
DISCUSSION. ‘The only other Eastern Highlands specimen of this
rare species was taken by Gyldenstolpe in the Wahgi Mountains.
Crateroscelis robusta robusta (De Vis)
Mountain Mouse-Warbler
NATIVE NAME. Fore: séka.
SPECIMENS EXAMINED. Awande: 2 6, 2 9 (28 June 1964; 16-18 June 1965).
=
Mt. Michael: 2 ¢, 2 9 (4-12 July 1964). Mt. Karimui Zone 4: 1 ¢,1 9, 1 imm.
@; Zone : lg. 24 2, 1 mm. 4; Zone 7 I Oo; Zone 3: 1 & (17 Aug.-8) Sept. 1965).
WEIGHT. 5b ¢2 16.0, 16:0, W6i8, 17.3, 175. 5 9: 14.3, 16:0; 163; 6:8) USi0:
lim, fe 7.0) mm, “o. 18:0:
WING. 5 @: 60, 61, 62, 62, 63. 5 9: 56, 57, Gl, 61, 64. limm. 4:59. 1 imm.
Soe.
en CONTENTS. Insects.
TAXONOMY. As already noted by Gyldenstolpe (1955, p. 84) and
by Mayr and Gilliard (1954, p. 345), there is much individual variation
in the color of the underparts (brown or gray). ‘The present series sug-
gests that this is to some extent, though probably not entirely, a
matter of age, the brownest underparts being in the younger birds. ‘The
color of the throat also varies, and is whitest in specimens with the
least brown underparts. Gyldenstolpe’s series also suggests a correlation
with age: all nine of his gray males have longer wings than four of his
five brown males, and four of his six gray females are larger than three
of his four brown ones. However, one of his brown males had enlarged
testes. Eye color also varies individually, as noted previously by Mayr
and Rand (1937, p. 107). In adult males the iris is orange, straw-
colored, or light straw-orange; brown or orange-brown in adult females;
and dull brown or gray-brown in immatures of both sexes.
The race sanford: differs in its much buffer underparts, deficiens in
the absence of the gray breast-band.
BREEDING. ‘Testes were enlarged in all three adult males from
Mt. Karimui and in one of the two adult males from Awande.
DISCUSSION. C. robusta is nearly as ubiquitous and common in
forest at higher altitudes as C. murina is at lower altitudes. It ap-
peared abruptly on Mt. Karimui at its maximum abundance at 5,400
ft, and accounted for about 5% of the local avifauna in the zones from
5,400 to 6,250 ft, about 3°% in the zones of stunted moss forest from
6,250 to 7,280 ft, about 1% up to 7,610 ft, and was not encountered
on the top 500 ft of the mountain. On Mt. Michael it was present
from the base of the forest (7,000 ft), commonest around 8,000 ft where
the forest was still tall, and still present up through the progressively
Zou
SPECIES ACCOUNTS
more stunted moss forest until at least 10,200 ft. Its behavior is similar
to that of C. murina, i.e., it remains in the lowest story of the forest
or searches in leaf litter on the ground, singly or in pairs, and is much
more often seen than heard.
VOICE. A whistled song similar to that of C. murina but with a
less flute-like, thinner, and more tinkling quality, more irregular
rhythm, more hurried delivery, and consisting on the average of more
notes (five or six is typical). As does C. murina, C. robusta alternates
patterns, a few of which are given in Figure 24 (p. 231). Its call is a
short, dry, very buzzy note similar to that of Sericornis nouhuysi.
SYLVIINAE: WARBLERS
Acrocephalus arundinaceus subsp.
Great Reed Warbler
DISCUSSION. According to Bell (1968) this warbler is very numer-
ous in dense “pit-pit” (tall cane grass) and tall elephant grass in the
Baiyer River Valley of the northern watershed (3,600 ft) and on the
Sepik-Wahgi Divide at 6,600 ft. One specimen has been collected near
Mt. Hagen, and there is a sight record from Pureni (N.G.B.S. News-
letter, No. 28, p. 3, Feb. 1968). Its distribution throughout New Guinea
is spotty.
VOICE. A lengthy, rich song composed of single and paired notes
of varied qualities, each repeated 3-5 times before going on to the
next note. The form suggests New World mimic thrushes (Mimidae)
or the European Song Thrush (Turdus philomelos).
Megalurus timoriensis wahgiensis Mayr and Gilliard,
M. t. montanus Mayr and Gilliard,
and M. t. macrurus (Salvadori)
‘Tawny Grassbird
NATIVE NAME. Fore: kasaru.
SPECIMENS EXAMINED. Awande: 1 ¢, 2 9, 1 juv. @ (19-20 June 1965).
Miarosa: 1 ? (15 June 1964). Karimui: 2 9 (3 July 1965).
WEIGHT. 1 4: 26. 3 9: 23, 25, 27.
WING. U2 05: 4 G2 64, 66; 66,70), fr: 63, “L juv. a2 bi.
TAIL. 1 4: 11% 3 G: 97, 102, 105, 1%: 101.
STOMACH CONTENTS. Insects (three stomachs), insects and seeds (one).
TAXONOMY. ‘This species is subject to both altitudinal and local
geographical variation. ‘The first high-altitude form discovered was
alpinus from the alpine grasslands of southeastern New Guinea, which
differed from macrurus of the southeastern midmontane and_ low-
altitude grasslands in its longer wing, grayer underparts, and darker
back. The midmontane and alpine forms of the Snow Mountains dif-
233
SPECIES ACCOUNTS
fered in the same way and were also assigned to macrurus and alpinus.
Mayr and Gilliard (1951, p. 9) then described two analogous forms
from the Wahgi region of the Eastern Highlands: wahgiensis of the
midmontane grasslands, fairly close to alpinus in color but with a
longer tail and shorter wing; and montanus of the alpine grasslands,
ah the short alpinus tail but darker color. The situation subse-
quently became more complicated when Sims (1956, p. 415) analysed
Shaw-Mayer’s material from Mt. Giluwe and showed that the color
patterns there were “upside-down,” i.e., the alpine form was the paler
one, although it had the shorter tail as in the Wahgi Valley. More
recently Gilliard and LeCroy (1961, p. 48) assigned Telefolmin mid-
montane birds to neither macrurus nor wahgiensis but to alpinus on
the basis of the longer wing, and Schodde and Hitchcock assigned one
specimen each from Lake Kutubu (2,450 ft) and Mendi (6,900 ft),
between Telefolmin and the Wahgi Valley, to macrurus. Wing and
tail measurements and color may vary independently, and future
collectors should state all three; Mayr and Gilliard gave no wing
measurements in their descriptions of wahgiensis and montanus, and
Gilliard and LeCroy gave no tail measurements for the ‘Telefolmin
series.
In dorsal color the Awande and Karimui specimens are close to
wahgiensis from Mt. Hagen (Tomba) and Mt. Kubor, and differ from
each other only in the darker crown of Karimui birds. The race
macrurus of southeastern New Guinea is shorter winged, shorter tailed,
and less dark above. M. t. muscalis of the Fly River differs in its
streaked crown, shorter wing, and paler, less brown dorsal coloration.
My measurements of Hagen and Kubor wahgiensis give: Hagen, 3 9,
wing 64, 66, 68, tail 97, 95, 98; Kubor, 1 9, wing 63, tail 104. While
the number of specimens is small, there seems to be local variation in
measurements even in midmontane wahgiensis of the Eastern High-
lands. Combining my measurements of females with Gyldenstolpe's
measurements (1955, p. 90) of Nondugl females, one finds wing length
decreasing in the order Karimui > Hagen = Awande > Kubor =
Nondugl; tail length decreases in the order Awande = Kubor >
Hagen = Karimui > > Nondugl. This local variability, plus the
upside-down and checkerboard racial distributions mentioned in the
previous paragraph, makes reassessment of the New Guinea races of
this species desirable.
BREEDING. One of the Karimui females had enlarged ovaries.
This, plus the capture of the juveniles, suggests some breeding in the
dry season, but perhaps not the uniformly synchronized breeding seen
in Malurus alboscapulatus.
DISCUSSION. AIl observers have found this warbler common and
widespread in the midmontane grassland of the Eastern Highlands.
Its habitat is grass several feet high, from which it is harder to flush
than Malurus alboscapulatus. Bell (1968) found in the Baiyer Valley
AN
SPECIES ACCOUNTS
that Acrocephalus arundinaceus has colonized tall, ungrazed grass,
whereas Megalurus timoriensis is in grass grazed down by cattle to
balls about three feet high. Gilliard, Shaw-Mayer, and Bulmer also
found Megalurus timoriensis in alpine grassland. ‘This warbler is thus
of interest in having a discontinuous altitudinal range in alpine grass-
land (> 11,000 ft), man-made midmontane grassland (ca. 3,000-7,500
ft), and grassland of the lowlands.
VOICE. The call is a single dry note “tsip’” or “tsick’’ or “CRUD;
which often sounds disyllabic. ‘here is also a repeated scolding note
“buk-buk-buk . . .” like the clucking of a chicken
Cisticola exilis diminuta Mathews
Golden-headed Fantail Warbler
NATIVE NAME. Fore: ikonantube.
SPECIMENS EXAMINED. Okasa: 1 ¢, 1 @ (26 June 1965).
WEIGHT. 1 4: 85. 1 @: 82.
WING. 1 @: 50. 1 9: 43.
STOMACH CONTENTS. Insects.
BREEDING. Neither specimen had the gonads enlarged, suggest-
ing that this species may be exceptional among grassland birds in not
breeding during the dry season.
DISCUSSION. ‘The distribution of Cisticola exilis in the Eastern
Highlands is local. Besides collecting it in the grasslands at Okasa in
June 1965 and observing it there in August 1946, I was told that it
occurred at Awande, but I found it nowhere else, and it was certainly
absent at Karimui. Neither Gilliard nor Gyldenstolpe met it in the
Wahgi Valley. Bell (1968, and pers. comm.) found it near Mt. Hagen
“quite numerous but strictly localized to areas of short blady grass”;
Shaw-Mayer collected one at 7,300 ft on Mt. Giluwe; and Bulmer
reports it as common in the Baiyer Valley around 4,000 ft and in the
Kaironk Valley of the Schrader Range up to 5,000 ft.
VOICE. ‘The very distinctive song consists of two very faint notes
“hoo-hoo” 14-1 sec apart, followed by a disyllabic musical note of bell-
like absolute clarity (Fig. 25). So different are the qualities of the two
halves of the song that they give the impression of coming from two
different species. The song is given from the top of a bush, or even of
a tree, if any is in the vicinity. The dry, drawn-out call note suggest a
SMEEZE OF all DHSECE.
Phylloscopus trivirgatus giulanettit (Salvadori)
Leaf Warbler
NATIVE NAME. Fore: pasésule.
SPECIMENS EXAMINED. Miarosa: 2 ? (11 and 17 June 1964). Mt. Michael:
1 ? (13 July 1964). Mengino: 1 ¢ (15 July 1964). Mt. Karimui Zone 3: 1 ? (16
Aug. 1965).
235
SPECIES ACCOUNTS
Cisticola exilis:
ae
hoo-hoo
Fic. 25. Voice of Cisticola exilis.
WEIGHT. 1 ?: 8.3.
STOMACH CONTENTS. Insects.
TAXONOMY. ‘These specimens were prepared as skeletons or
preserved in formalin and hence cannot be analysed racially. Mayr
and Gilliard (1954, p. 347) and Gyldenstolpe (1955, p. 96) assigned
specimens from the Wahgi region to giulanettit.
DISCUSSION. P. trivirgatus is one of the half dozen commonest
forest birds in its altitudinal range (ca. 4,300-7,000 ft, in the forest
interior, forest edge, and trees of second-growth and gardens). On Mt.
Karimui it first appeared around 4,300 ft, was commonest between
5,100 and 5,900 ft where it accounted for 5-10% of all birds, and dis-
appeared around 6,100 ft. On some outlying mountain ranges of New
Guinea (e.g., the North Coastal Range and Vogelkop) it is largely con-
fined to altitudes below 5,000 ft. While it sometimes descends to within
10 ft of the ground, it usually remains in the treetops where it is more
easily identified by voice than by sight. As it gleans, it moves by short
hops of a few inches to a foot and occasional flights of several feet.
VOICE. A high, brief, patternless warble, often concluding with a
very high upslur apparently continuing to go up out of the human
auditory range.
MUSCICAPINAE: FLYCATCHERS
Peltops blainvillii (Lesson and Garnot)
Lowland Peltops Flycatcher
SPECIMENS EXAMINED. Soliabeda: 2 ¢, 2 Q (25-30 July 1965).
WEIGH. 2 62 29:7, 300; 2 Os 28:0; 3055.
WING. 2 @: 96, 96. 2.9: 93, T00.
GULMEN FROM BASE. 2 62 20, 26. 2 Q% 22,.26)
SOFT PARTS. Iris deep red or dull red.
STOMACH CONTENTS. Insects.
236
SPECIES ACCOUNTS
BREEDING. All specimens had small gonads.
DISCUSSION. ‘The genus Pellops is endemic to New Guinea, has
no close relatives, and consists of two monotypic species. P. blainvillit
is so exceedingly similar morphologically to its higher-altitude sibling
species P. montanus that it was not until 1921 that montanus was de-
scribed even as a subspecies of P. blainvillit. ‘The careful comparisons
of Stresemann (1923, p. 92) and Gilliard and LeCroy (1961, p. 52) re-
vealed only two diagnostic characters. First, P. montanus has a longer
wing and tail than P. blainvillii; this suffices to separate most but not
all specimens, because there is some overlap and also because P. mon-
tanus decreases in size with decreasing altitude (Bergmann effect) and
approaches the size of the low-altitude form P. blainvillii. ‘The second
diagnostic character is the larger white patches on the back and on
the cheek of P. montanus; this is also a good but not perfect distin-
euishing character, since there is some overlap with P. blainvillit. Both
species behave similarly and inhabit the same kinds of habitats; I
have been unable to detect ecological differences except in altitudinal
range. In the field, however, the two species are easily distinguishable
by their dissimilar and frequently given calls.
The altitudinal ranges of P. blainvillit and P. montanus are mutually
exclusive. I found P. blainvillii only between 1,500 and 2,000 ft at my
lowest-altitude locality, Soliabeda. My lowest record of P. montanus
was at 2,680 ft between Yudo and Soliabeda, leaving a gap of 680 ft in
which no Peltops was found. ‘There appeared to me to be a similar
gap in the North Coastal Range distributions, and I cannot find a
published record of P. montanus under 2,500 ft, nor of P. blainvilli
above 2,000 ft except for a specimen taken by Schodde and Hitchcock
at Lake Kutubu (2,450 ft). Hence the apparent gap could be real.
At Soliabeda P. blainvillit was uncommon. Groups of up to four
perched in the crowns of tall trees overlooking small clearings in the
forest. From these perches they sallied after insects often at considerable
distances from the crown, sometimes to within a few feet of the
ground. The territory must be large, since successive treetop perches
may be several hundred feet apart.
VOICE. Unique among New Guinea birds in that the notes are
very sharply sucked in. ‘The rhythm is usually iambic, e.g., three rapid
pairs of notes, all on the same pitch, the first member of each pair
short and unaccented (Fig. 26). At each pair of notes the head is thrown
violently up and down. Sometimes three or four sucked-in notes in
rapid succession on the same pitch are given in uniform rhythm. The
quality is totally unmusical, i.e., clicked or snapped. Once I heard a
faint call of three rapid unmusical notes on successively lower pitches,
like a weak, abbreviated fragment of the usual Peltops montanus call.
237
SPECIES ACCOUNTS
Peltops blainvillii:
Peltops montanus:
oo oC
Fic. 26. Voices of Peltops blainvillii and P. montanus.
Peltops montanus Stresemann
Mountain Peltops Flycatcher
NATIVE NAME. Gimi: ilémukokdya.
SPECIMENS EXAMINED. Okasa (4,250 ft); 2 ? (22 and 25 June 1965). Mt.
Michael (7,000 ft): 1 g¢ (13 July 1964). Mengino (4,600-5,100 ft): 1 9, 2 ? (16 July
1964). Karimui (3,650 ft): 4 ¢@, 4 9 (2 July-3 Aug. 1965). Bomai (3,250 ft): 1 ?
(6 July 1965).
WREIGIUI, 4 °¢: da.0, 240 (2), 300. 4 07 2910, 207, 30, 3210. 3) Pr: 2720) SILO:
IPs
WING. 5 42 109, 110°6), Has 4° O2 001; 10 (2), LOT Aros O77 TLOe Wis:
GULMEN FROM BASE. 5 @<,23 (4), 25; 4° O% 20; 2; 22.23. 2 P, 235, 24.
SOFT PARTS. Iris: red, orange, or orange-brown.
BREEDING. Gonads of all specimens were small.
DISCUSSION. P. montanus is erratically distributed in the East-
ern Highlands from 2,600 ft to 5,100 ft (Mt. Karimut) or to 7,200 ft (Mt.
Michael). It was present at most of my stations in this altitudinal range
but was definitely absent at Okapa (6,600 ft), Awande (6,000 ft), and
Miarosa (5,800 ft), both in my experience and in the much more ex-
tensive experience of my Fore assistants. It was not found in the
Wahgi Valley by Gilliard or Gyldenstolpe. Bulmer found it breeding
in Kyaka territory in 1955, when he collected five specimens and re-
corded one nest between 4,300 and 5,300 ft, but he did not encounter
it in the four months he spent there in 1959.
238
SPECIES ACCOUNTS
The habitat of this species may be easily summed up as tall forest
trees at the edge of open spaces. From conspicuous perches which are
rarely less than 40 ft above the ground, P. montanus sallies into the
open space to catch insects and returns to its perch or to a neighboring
tree, sitting stationary between sallies. On the Sena River the trees
chosen were on the river bank, and the open space exploited was that
over the river. In the virgin forest on Mt. Karimui the perches were
initially in tall trees emerging above the canopy or on a ridge. When
we had cleared small campsites in the forest, trees at the edge of these
were also used. At Karimui, where many gardens and roads had been
cleared, trees at the border between these clearings and the forest fur-
nished perches. It is surprising that this species, which concentrates
at natural open spaces in the forest, is not far more abundant and
widely distributed in settled areas of the Eastern Highlands. Like P.
blainvillti, P. montanus is a social species, seen usually in groups of
2-5, though sometimes singly. Once I saw a P. montanus attack a
perched hawk Aviceda subcristata and force it to take flight.
VOICE. A very rapid series of a half dozen to a dozen notes
lasting in all less than | sec, and descending slightly in pitch (Fig. 26,
p. 238). The volume is soft, and the quality quite similar to that of
the sound produced by running one’s finger along the teeth of a comb.
Infrequently one hears a hoarse upslurred “‘wheep.”
Songs and Niche Differences in the Genus Rhipidura
It is interesting to attempt to deduce species relationships within
this genus from songs, since the 12 species I have encountered fall into
three distinct groups on this basis. R. leucothorax and R. threnothorax
resemble each other in their loud, explosive songs very unlike those of
the other 10 species. ‘The harsh song of R. lewcophrys places it by itself
in another group. ‘The songs of the remaining nine species (R. atra, R.
brachyrhyncha, R. albolimbata, R. hyperythra, R. rufiventris, R. rufi-
dorsa, plus R. fuliginosa which I heard in New Zealand, Australia,
and Espiritu Santo, R. dahli of New Britain, and R. nebulosa of
Samoa) are all weak and high-pitched, and consist of either or both
of two kinds of notes: clear whistled notes and jumbled twitters. R.
albolimbata and R. hyperythra, in whose songs clear whistled notes
predominate, are particularly close to each other, and R. brachyrhyn-
cha and R. atra are close to each other in their squeaky, tinkling, jum-
bled songs. ‘The songs of R. fuliginosa and R. dahli are similar to those
of R. brachyrhyncha and R. atra. This vocal “classification” is in rea-
sonable accord with a grouping based solely on morphology.
Niche differences among the nine species of Rhipidura in the East-
ern Highlands may be summarized in oversimplified form as follows.
R. ieucophrys is confined to open country such as gardens and villages,
whereas the other species all live in the forest or dense second-growth.
239
SPECIES ACCOUNTS
‘Three species live mainly in the forest understory and have mutually
exclusive altitudinal ranges: R. threnothorax at low altitudes (to ca.
3,000 ft), R. atra at middle altitudes (3,000 to ca. 7,000 ft), R. bra-
chyrhyncha at high altitudes (above ca. 7,000 ft). ‘The latter two ven-
ture into the middlestory, which R. threnothorax rarely does. ‘The
altitudinal range and the lowerstory preference of R. leucothorax are
similar to those of its close relative R. threnothorax, but R. leuco-
thorax lives in thickets at the forest edge and in dense second-growth,
R. threnothorax in the forest interior. R. hyperythra and R. albolim-
bata are forest species that forage from the crowns to the understory,
most often in the middlestory or crown, and have mutually exclusive
altitudinal ranges, R. albolimbata living at higher elevations (above
4,500 ft) than R. hyperythra (below 4,500 ft). R. rufiventris lives in
more open parts of the forest and is commonest at the forest edge or
adjacent to open spaces (not in the thickets utililized by R. leuco-
thorax), occurs from sea level to ca. 5,000 ft, forages from the crowns
to the understory, and is larger than R. hyperythra or R. albolimbata.
R. rufidorsa is a forest species of the lower- and middlestories, confined
in the Eastern Highlands to elevations below 2,000 ft, where R. hy-
perythra is uncommon.
Rhipidura threnothorax threnothorax Miller
Sooty ‘Thicket Fantail
NATIVE NAME. Daribi: tusada.
SPECIMENS EXAMINED. Karimui: 9 ¢, 2 9, 1 nestling (1-15 Aug. 1964; 2-12
July 1965). Bomai: 2 ¢, 1 9 (6 July 1965). Soliabeda: 1 ¢, 1 Q (28-30 July
1965).
saree GO Ge tbe, 170; 8b, 19:0, 195 200. Al Oe TAL7, 1682161551910:
WING. 902 75-85) (81 s213). <3) Qs 71); 74, 75.
WAIL, 9 62 96-101 (98 S22). 45-O)39895, 925 94.
STOMACH CONTENTS. Insects.
TAXONOMY. ‘Three males and one female have pale rufous tips
to the upperwing coverts, a character subject to individual variation
in other areas as well (Rand, 1942a, p. 333; 1942b, p. 477).
BREEDING. One female at Karimui was captured on a nest from
which the nestling was secured. ‘Testes were large in both Bomai males;
enlarged in half of the Karimui males, small in the others; and small
in the sole Soliabeda male. Evidently many, but not all, individuals of
this understory species were breeding in the dry season.
DISCUSSION. R. threnothorax has not yet been recorded in the
Eastern Highlands outside the Karimui area, where it is somewhat
above its usual ceiling. It is shy, solitary, and infrequently seen but
not uncommon. Of the forest fantails this is the most sharply re-
stricted to the understory, the few that I saw being at most 3 ft above
the ground and sometimes on the ground. It favors those parts of the
forest or forest edge that have particularly dense undergrowth.
240
SPECIES ACCOUNTS
VOICE. The song is of five notes, the first upslurred, the first two
softer, the last three a loud “‘chew-chew-chew” (Fig. 27, p. 242). The
call is a loud, spitted “pik! pik! pik!” The vocalizations are reminiscent
of Ewpetes castanonotus or Pachycare flavogrisea.
Rhipidura leucothorax clamosa Diamond
White-breasted Thicket Fantail
SPECIMENS EXAMINED. Karimui: 1 ¢ (10 July 1965). Soliabeda: 2 g (25-
28 July 1965).
WEIGHT. 2. ¢: 190; 19/7.
WING... 3 42 78; 79, 79:
‘TEAL. 3 &:. 86, 89, Of.
STOMACH CONTENTS. Insects.
TAXONOMY. In accord with the general trend at Karimui this
race differs in its darker, blacker plumage (Diamond, 1967a).
BREEDING. ‘Testes were small in one Soliabeda male; enlarged in
the second Soliabeda male, which had been consistently singing from
the same thicket for several days; and enlarged in the Karimui male.
DISCUSSION. R. leucothorax is not a forest bird but remains
inside the densest and most tangled thickets and in second-growth and
at the forest edge, not necessarily in the shade of taller trees. While it
was a noisy singer and moved about within these thickets, it stayed
largely concealed and was difficult to observe. ‘The unbalanced sex ratio
in my series (3 ¢, 0 9) and in the American Museum of Natural
History’s collection (21 4, 6 ¢?) is probably because females of this
skulking bird are much more likely to escape detection than singing
males. It adopts an exaggerated fantail posture, with the tail fully
spread.
‘This fantail was uncommon and local in the Karimui area, where
it was considerably above its normal altitudinal range. In the Karimui
Basin itself my only records were of a song heard in a thicket near
Iogoramalu (3,700 ft) and of the specimen brought in at Karimui by
a native. The species was also taken by Schodde and Hitchcock at Lake
Kutubu (2,450 ft) and has been observed in the Baiyer Valley (3,500
tt; IN.G_ Bs. Newsletter, No. 54, p. 2, June 1970).
VOICE. ‘The song consists of a short series of rapid, spitted or
staccato, imitially faint notes which accelerate and rise in pitch, to
terminate in a loud, explosive, slurred note suggestive of a Pachy-
cephala (Fig. 27).
Rhipidura rufidorsa subsp.
Gray-breasted Rufous Fantail
SPECIMENS EXAMINED. Soliabeda: 1 ¢ (27 July 1965).
WEIGHT. 10.
241
SPECIES ACCOUNTS
Rhipidura threnothorax:
a
whee chew chew chew
Rhipidura leucothorax:
faint | loud,explosive
Rhipidura rufidorsa:
—_ or
—
Rhipidura brachyrhyncha:
Rhipidura hyperythra:
Rhipidura albolimbata:
Rhipidura rufiventris:
Rhipidura leucophrys:
(3 ax aera eee —
high | harsh [righ | harsh
Fic. 27. Voices of Rhipidura flycatchers.
SPECIES ACCOUNTS
WING. 66.
STOMACH CONTENTS. Insects.
TAXONOMY. The rufous of the back in Rhipidura rufidorsa
becomes gradually paler as one proceeds counter-clockwise around the
periphery of New Guinea: kumust on the north coast of southeastern
New Guinea is the brightest, kwbuna on the south coast of southeastern
New Guinea is the dullest, and nominate rufidorsa of the rest of New
Guinea is intermediate. R. r. Rubuna is also the palest form. The
Soliabeda male is somewhat brighter than kubuna, duller and darker
than nominate rufidorsa from the Fly River, and considerably darker
than rufidorsa from the Weyland Mountains and south slope of the
Snow Mountains. It is closest to Fly River rufidorsa, but a definite
racial assignment requires more material.
BREEDING. ‘The testes were enlarged.
DISCUSSION. ‘The sole specimen was netted, and no other was
seen or heard.
VOICE. ‘Three or four high, whistled notes, suggestive of Cratero-
scelis murina in quality but slightly upslurred. The notes either move
progressively down the scale, or else the last note returns to the pitch
or the first (Kio. 27, p, 242).
Rhipidura brachyrhyncha devist North
Dimorphic Rufous Fantail
SPECIMENS EXAMINED. Mt. Michael: 1 9, 1 ? (5 and 13 July 1964). Mt.
Karim Zone 6:1 9; Zone J: 1 ¢; Zone 8: 1 go 1 2 (2-7 Sept. 1965):
WEIGHT. 2 4: 10.3, 10.7. 2 9: 8.3, 9.7.
WING. 2 3: 71 7A. 3 9: 66, 67, 6S.
STOMACH CONTENTS. Insects.
TAXONOMY. Rhipidura brachyrhyncha_ exists in two color
phases, as discussed by Mayr and Rand (1937, p. 164): a darker phase
with much black in the tail (Type I of Mayr and Rand) and a paler
phase with no black in the tail (Type Ht of Mayr and Rand). All of
my specimens are ‘Type I except the Mt. Karimui Zone 7 male, which
is Type II. The five specimens which Gyldenstolpe (1955, p. 97)
collected in the Wahgi Mountains and the three which Biirgers took
in the Schrader Range (Stresemann, 1923, p. 8) were all Type I. Mayr
and Gilliard (1954) do not state the proportions in Gilliard’s speci-
mens, but four of these are present in the American Museum of
Natural History, and all are Type I. In the Snow Mountains the Third
Archbold Expedition (Rand, 1942b, p. 478) obtained 15 of Type I and
4 of Type H, Ripley (1964, p. 60) 3 and 1, and the Lorentz expedition
(Junge, 1939, p. 29) 2 and 0. In southeastern New Guinea I found
‘Type I twice as common as ‘Type II. However, Mayr and Rand (1937,
p. 167) state that the proportions are nearly equal in the Vogelkop
and the Huon Peninsula. Evidently there is some geographic variation
243
SPECIES ACCOUNTS
in the proportions, with Type l predominant in the central body of
New Guinea. The related fantail R. fuliginosa of New Zealand also
has two color phases whose relative abundances vary geographically.
BREEDING. Gonads were small.
DISCUSSION. ‘Three species of Rhipidura—R. brachyrhyncha, R.
atra, and R. albolimbata—occur in the forest interior at higher eleva-
tions and are usually not in second growth. Niche differences involve
altitudinal distribution, and vertical distribution within the forest:
As regards altitudinal distribution, all three species may coexist in
an overlap zone around 7,000 ft, where R. atra and R. brachyrhyncha
overlap by about 500 feet, but for the most part R. brachyrhyncha
lives from this altitude up to about 11,000 ft, R. atra from this altitude
down to 3,000 or 4,000 ft. These two species are much more closely
related to each other than to other New Guinea Rhipidura species,
as indicated by their songs, their lowerstory preferences, and the similar
plumage of R. atra females and R. brachyrhyncha Type 1. They may
have originated as an altitudinal pair and eventually developed
enough differences in foraging behavior (see Mayr and Rand, 1937,
p. 161) to permit modest altitudinal overlap. R. albolimbata shares
the whole altitudinal range of R. brachyrhyncha and much of the
range of R. atra, extending from 11,000 down to 4,500 ft, where it is
replaced by R. hyperythra.
As regards vertical distribution within the forest, R. albolimbata
and R. hyperythra are characteristic of the middlestory (5-60 ft above
the ground, occasionally lower or higher). R. atra and R. brachy-
rhyncha are most often seen in the understory (ca. 2-8 ft above the
ground, sometimes to 20 ft) and favor denser thickets than do R.
albolimbata and R. hyperythra. The characteristic habitat of R. brachy-
rhyncha is stunted moss forest, and it has turned up at most Eastern
Highlands collecting stations in this habitat. ‘Vhus, the lower altitudi-
nal limit of R. brachyrhyncha and the upper limit of R. atra are
strongly dependent upon local conditions. In the Okapa area, where
we ascended to 7,500 ft without reaching moss forest, R. brachyrhyncha
was absent, and R. atra went up to at least this altitude. On Mt.
Michael, where moss forest began at 8,700 ft, R. brachyrhyncha
descended to 8,000 ft, and R. atra ascended at least to 8,100 ft. On
Mt. Karimui, where moss forest began rather abruptly at 6,500 ft,
R. brachyrhyncha descended only to this altitude, and R. atra dis-
appeared at it except for two records at 7,620 tte
VOICE. ‘The high-pitched song has a tinkling quality when heard
nearby but sounds squeaky at a distance. It consists of a run-together
descending cascade of about seven fast notes, introduced by a couple
of detached, higher-pitched, slower, upslurred notes (Dig.e2 7, ip. 2a2)-
The call note is slightly upslurred, high-pitched, and squeaky.
244
SPECIES ACCOUNTS
Rhipidura atra atra Salvadori
Black Fantail
NATIVE NAME. Fore: t’re-t’re.
SPECIMENS EXAMINED. Awande: 2 4,1 9,2 imm. ¢ (19 June 1964; 15-17
June 1965). Okasa: 1 ¢ (23 Aug. 1964). Mt. Michael: 1 @ (4 July 1964) Sena
River: 1 ¢ (27 July 1965). Mt. Karimui Zone 1: 4 g, 1 9; Zone 2: 1 ¢, } es
Zone 3: 1 4; Zone 4: 2 4, 1 @; Zone 8: 1 @ (9 Aug.-7 Sept. 1965),
WEIGHT. 10 42 17.0:140 @26= 1,0). 4 OQ: 105, 10.7, 12.0, 1233. 2 1mm oi
NOMS W223.
WING. 10 @: 76-81 (79+ 2). 3 9: 73, 74, 77 (74.7). 2 imm. ¢: 73, 74,
STOMACH CONTENTS. Insects.
TAXONOMY. The two immature males are in a brown plumage
resembling that of the adult female. The other males are in a fully
adult black plumage except for a few brown feathers on one.
BREEDING. The gonads were slightly enlarged in one Mt. Kari-
mui male and one Awande male and were small in all other specimens.
DISCUSSION. Rhipidura atra was present at all my forest collect-
ing stations between about 2,800 and 8,000 ft, with the conspicuous
exception of Karimui (3,650 ft) and Bomai (3,250 ft). The possibility
that it was present but overlooked at Karimui and Bomai is negligible,
because both my Fore assistants and I knew its song, and it is readily
netted, conspicuous, and common where present. The most surprising
discovery of my walk from Karimui down to Soliabeda and vice versa
was the observation of four R. atra at 2,770 ft and of one at 3,230 ft
in hill forest on the outer side of the Karimui Basin’s mountain wall.
As we went up from the Karimui Basin onto the slopes of Mt. Karimui,
R. atra reappeared at 4,090 ft and continued up to 6,500 ft, with two
records at 7,620 ft. ‘This hill and mountain forest species thus has a
discontinuous altitudinal range at Karimui (present at 2,770-3,230 ft
and 4,090-7,620 ft, absent in between), as a result of the intrusion of
tropical conditions and flora on the basin floor. It is usually solitary
and has a typical Rhipidura behavior, displaying with fanned tail and
spinning about 180 degrees on its perch.
VOICE. ‘There are three kinds of vocalizations: (1) the call note, a
loud metallic “pink!”; (2) a very faint twittering progressing upscale;
and (3) a tinkling high-pitched jumble of a song similar to that of
R. brachyrhyncha, with a sweet and piercing quality. Once I heard
this song from an individual in brown plumage (a female or immature
male).
Rhipidura hyperythra muelleri Meyer
Chestnut-bellied Fantail
SPECIMENS EXAMINED. Okasa: 1 4, 1 9, 2 ? (23-26 June 1965). Guwasa
>
(Gimi territory): 1 Q (24 July 1964). Karimui: 1 ¢, 2 9, 1? (1 July-4 Aug. 1965).
Bomai: 1 ? (6 July 1965).
245
SPECIES ACCOUNTS
WEIGHE. 1 42 105) 292 10) 107,02 7ol0 aeons:
WING. 1 4: 73. 2 9: 66, 71. 3 ?: 72, 75, 78.
TAXONOMY. The distinguishing racial feature is the extent of
white on the tips of the outer tail feathers: 15-20 mm in castaneothorax
of southeastern New Guinea, 6-14 mm in muelleri from the remainder
of New Guinea. The figures for my series place it with mueller:
Okasa, 1 ¢ 12 mm, 2? 14 and 14 mm; Bomai, | ? 11 mm; Guwasa, 1 9
9 mm. The greater extent in the specimens from Okasa, lying farther
to the east, suggests an approach to castaneothorax. A specimen from
Lake Kutubu examined by Schodde and Hitchcock also belonged to
muellert.
BREEDING. ‘The gonads were enlarged in one Karimui male but
not in the other specimens.
DISCUSSION. R. hyperythra is the middlestory forest fantail of
lower altitudes up to about 4,500 ft and was collected or observed at
all my stations in this range, above which it is replaced by R. albo-
limbata. Its maximum abundance is in hill forest near the upper limit
of its altitudinal range: i.e., it was common at Okasa (3,500-4,250 ft),
and in Zone | of Mt. Karimui it accounted for 4% of census totals.
At Bomai and Soliabeda it was quite uncommon. ‘To judge from pub-
lished records and my own experience, it is rare and often absent in the
flat floodplains of the lowlands (Sepik Basin, southwestern New Guinea
floodplain, Fly River). Its habit of tipping forward, nervously flutter-
ing its wings and fanning its tail, then spinning around 180 degrees to
repeat the performance, resembles R. albolimbata.
VOICE. ‘The quality is as in the song of R. albolimbata, the pat-
tern is drastically abbreviated, as if R. albolimbata were sparing itself
its usual ad nauseam repetitions: two short and detached, high-pitched,
faint but clear and tinkling notes, the second following the first by
less than J sec and a minor third higher in pitch (Fig. 27, p. 242). An
even fainter twittering similar to that of R. albolimbata, R. rufiventris,
and R. atra was heard once.
Rhipidura albolimbata Salvadori
White-eared Fantail
NATIVE NAMES. Fore: ninikésu.
SPECIMENS EXAMINED. Awande: 1 ¢ (a), 3 ? (b, c, d) (28 June 1964; 14-19
June 1965). Mt. Karimui Zone 3: 1 ? (e); Zone 5: 1 ¢ (fH, 1 @ (g), 1 ? (h); Zone
6: 2 ? (4, j); Zone 7: 1 g (Kk), 1 ? (1); Zone 8: 1 ? (m) (16 Aug.) Sept. 1965).
WEHIGHT. 3 42.9.9 (@), 107_@, T4&),. 1 9: 9 (1 2 20 qh); 93 1);
9.3 (i), 10.0 (e), 10.8 (d), 10.7 (j), 11.0 (m).
WING. 3 @: 79 (2), 80 @),82"®. 1 Oe 71 © GO ee 72. (b), 72 Xe), 7% (ih), 80
(c), 84 (1), 84 (m).
STOMACH CONTENTS. Insects.
TAXONOMY. In the Snow Mountains (Rand, 1942b, p. 479), at
Telefolmin (Gilliard and LeCroy, 1961, p. 53), and in the Wahgi
246
SPECIES ACCOUNTS
Valley (Mayr and Gilliard, 1954, p. 348; Gyldenstolpe, 1955, p. 99)
this species increases in size and darkness with increasing altitude. Ot
the specimens measured in my series from the southern parts of the
Eastern Highlands, those with the longest wings (“k’, “I”, “m”) are
the three from the highest altitude. Specimens “k” and “m” are also
the darkest (on the back and flanks) and heaviest specimens (‘I was
not weighed or available for color comparison). ‘The large, dark, high-
altitude birds have been separated as lorentzit. However, this altitudinal
cline is gradual and has now been found over a stretch of 600 miles
in the Central Range. As suggested by Gilliard and LeCroy (1961,
ps 5a; 1968, p. 19), it therefore seems undesirable to recognize altitudi-
nal subspecies in this case.
DISCUSSION. R. albolimbata is the high-altitude forest fantail
of the middlestory, replacing R. hyperythra above about 4,500 ft. On
Mt. Karimui’s west peak (the one I studied in detail) the lowest sight-
ing of R. albolimbata was at 4,630 ft, and the highest of R. hyperythra
was at 4,360 ft. On Mt. Karimui’s east ridge, where moss forest
descended lower and which we crossed at 4,420 ft en route from
Soliabeda back to Karimui, it was R. albolimbata which we found at
this altitude. However, at 4,500 ft on the Sena River and at 4,250 ft
in the Okasa forest it was R. hyperythra that was present. R. albo-
limbata extends abundantly upward throughout the stunted moss
forest to above timberline, where it forages on tree ferns in alpine
grassland several hundred feet from the forest edge.
VOICE. ‘The song is a sequence of clear, detached, short, and high
notes spaced at intervals of about 1% sec, giving the impression of water
droplets falling at regular intervals. In the commonest pitch arrange-
ment the first and third notes are on the same pitch, the second note
a minor third higher, the fourth note a minor third lower. ‘This four-
note pattern is the origin of the Fore name “‘ni-ni-ke-su’”’ and is repeated
ad nauseam without pause (Fig. 27a, p. 242). Less often the song con-
sists of a five- or six-note phrase which has the same quality and high
pitch but is not repeated (Fig. 27b). One of these latter patterns, con-
sisting of five notes each one tone lower than the preceding and con-
cluding with faint twittering, is close to the song of R. rufiventris.
Rhipidura rufiventris gularis Muller
White-throated Fantail
SPECIMENS EXAMINED. Karimui: 1 ¢ (4 Aug. 1965). Bomai: 1 ¢ (7 July
1965). Soliabeda: 2 ¢ (28 and 29 July 1965).
WEIGHT. 4 @: 14.5, 15,0, 15.0, 16.0.
WING. 4 @: 85, 86, 88, 89.
STOMACH CONTENTS. Insects.
BREEDING. ‘Testes were enlarged in all four specimens and
greatly enlarged in two of them.
247
SPECIES ACCOUNTS
DISCUSSION. Whereas R. hyperythra is virtually confined to the
forest interior, R. rufiventris may be found in the forest interior but
more often occurs in natural forest clearings produced by fallen trees,
and in open parts of the forest, the forest edge, and second-growth, at
any height from the ground to the treetops. I observed or collected it
at all my collecting stations below 5,000 ft that included some second-
growth, but only up to 4,000 ft in primary forest. As one proceeds
from 4,000 ft downward, R. rufiventris becomes more common, R.
hy perythra less common. For instance, R. rufiventris reached its maxi-
mum abundance at my lowest station, Soliabeda, where R. hyperythra
was quite uncommon. R. rufiventris still occurs regularly in the flat
floodplains of the lowlands, where R. hyperythra is quite uncommon
or absent. ‘There are also differences in behavior: larger R. rufiventris
does not indulge in the pronounced tail-fanning of R. hyperythra and
R. albolimbata, and is somewhat less nervous.
VOICE. ‘The song consists of five descending notes in syncopated
rhythm, the whole song lasting slightly less than 2 sec (Fig. 27, p. 242).
The pitch is fairly high, and the successive pitch intervals are somewhat
less than a whole note. ‘The quality is a thin whistle similar to the
voice of Crateroscelis murina but less substantial and clear, and more
substantial than the songs of R. hyperythra or R. albolimbata. ‘The
pattern is similar to that of some R. lewcophrys songs and to a rare
R. albolimbata song (Fig. 27b), but the very different quality precludes
confusion. A repeated faint “whik” was once heard as a call note.
Rhipidura leucophrys melaleuca (Quoy and Gaimard)
Willie Wagtail
NATIVE NAMES. Fore: kétori. Gimi: bidii. Daribi: gédigédi.
SPECIMENS EXAMINED. Awande: 1 ¢, 2 9, 2 ? (17-20 June 1965). Karimui:
1 Q (15 July 1965).
WEIGH, Leg 2b. 3 9592310) 2E0) 20h. 2 er 22.0) 25.0:
WING? 0 air mole) 231592 95,-90,098, 27s 97 10s.
STOMACH CONTENTS. Insects.
BREEDING. ‘The Awande male had enlarged testes.
DISCUSSION. ‘The Willie Wagtail is one of the principal avian
beneficiaries of man-made ecological changes in the Eastern Highlands.
Whereas it is a lowland species elsewhere in New Guinea, in the
Eastern Highlands it is common and ubiquitous up to 6,500 ft in open
country offering perches (but not in pure grassland). Strangely, it is
only from the Eastern Highlands to Telefolmin that R. leucophrys
has succeeded in colonizing the midmontane grassland. ‘To judge from
published records, it is absent from the midmontane grassland of the
Huon Peninsula, southeastern New Guinea, and the Baliem Valley,
Ilaga Valley, and Wissel Lakes region of western New Guinea. Its
perches range from the tops of casuarina groves and of trees overlook-
248
SPECIES ACCOUNTS
ing roads and gardens to the ground. Perhaps it is the only widespread
breeding bird in the Eastern Highlands, apart from the more locally
distributed Anthus novaeseelandiac, which spends a high proportion of
its time on open bare ground (i.e., roads and closely cut lawns). Much of
its food is obtained by flycatching in midair. Willie Wagtails fly up to
and repeatedly attack perched large birds, like Accipiter fasciatus and
Corvus orru, and continue to chase them when they take off, partly
in order to pluck feathers to line the nest. ‘They also chase dogs. In
some areas Willie Wagtails leave during the dry season and return to
breed during the rains.
VOICE. Quite unlike the quality of other New Guinea species of
Rhipidura. There are two kinds of songs, which may involve notes of
two sharply different qualities: harsh or grating, low-pitched, louder
notes, somewhat like Monarcha axillaris in quality, and higher-pitched,
weaker, whistled notes. Each note has a sharp attack, is staccato, and
is at constant pitch. One of the two songs is a five- or six-note phrase
in uneven rhythm and with progressively dropping pitch (Fig. 27a, p.
242); the phrase may immediately be repeated. The other song (Fig.
27b) consists either of a three-note phrase in dotted rhythm, in the
harsh quality, and at nearly the same pitch, the phrase being imme-
diately repeated; or else (Fig. 27c) consists of this same phrase alternat-
ing with a phrase in the weaker, higher quality and consisting of four
rapid-fire notes on the same pitch. ‘These songs are occasionally heard
at night.
Monarcha axillaris fallax (Ramsay)
Black Monarch
NATIVE NAME. Gimi. luserepiyaba.
SPECIMENS EXAMINED. Awande: 1 imm. ¢, 1 ? (19 June 1964; 19 June 1965).
Okasa: 2 ¢,1 9, 2 imm. ? (20 Aug. 1964; 22-26 June 1965). Karimui: 2 imm. ¢,
I imm. 9, 1 mm. ? (10 Aug. 1964; 2-15 July 1965). Mt. Karimui Zone 1: 4 4,
3 9, Zone 2; 2g; Zone 3: 1 ¢; Zone 5: 1 @ (9 Aug.-2 Sept. 1965).
WEIGHT. 10 @ Jo.0-17.0 5.7 220.6). 4 9; 14:3, 14:7, 160,160. 3 mmm 4:
ioe, 140, 15.5. I imm,. 9: 17.0. 1 amm, ?: 12.7.
WING. NO) 63 74-82 (7 S22), 0 95.73, 745 70. Syimm. 23°73, 74, 77. 1 am.
O27.) o mime ?: 74, 79; 1D:
STOMACH CONTENTS. Insects.
TAXONOMY. ‘The nominate race of the Vogelkop, Weyland
Mountains, and North Coastal Range has much larger white pectoral
tufts. Immatures of both sexes are uniform dull slate gray, lacking the
blue-black sheen of adults. ‘They also have browner primaries, and one
specimen has a few brown feathers on the back. Adult females have
the blue-black sheen slightly less pronounced than in adult males but
are still readily distinguished from immatures.
BREEDING. ‘The gonads of the whole series were small, usually
249
SPECIES ACCOUNTS
very small, except for being slightly enlarged in two of the Mt. Karimui
males.
DISCUSSION. Monarcha axillaris is a forest species with an
altitudinal range from about 3,000 to 6,000 ft. In my study areas it
reached its maximum abundance in Zone 1 of Mt. Karimui (4,000-
4,200 ft), where it accounted for 7%, of the local avifauna, and in the
Okasa forest (3,550-4,250 ft). The contrast between its abundance on
the lower slopes of Mt. Karimui and its rareness (0.2% of the local
avifauna) on the flat basin floor at Karimui Patrol Post (3,650 ft) was
striking and in agreement with the rareness of many other hill forest
species in the Basin. Stragglers of M. axillaris turn up irregularly and
in low numbers at 6,000 ft (e.g., Awande), occasionally higher. ‘This
flycatcher provides an excellent example of the tendency of immature
birds to be found at the extremes of a species’ altitudinal range: no
immature was collected or seen in the species’ “heartland” (Zones 1
and 2) on Mt. Karimui, whereas all birds collected and seen at Karimui
Patrol Post were the gray immatures, as was one of the two Awande
specimens.
Of the New Guinea representatives of genus Monarcha, M. axillaris
is the closest to the Rhipidura fantails in its habits. It flits in the lower-
or middlestory up to ca. 20 ft above the ground, holding its body
horizontal and fanning its tail, sometimes in groups of up to four
birds. Superficially Monarcha axillaris resembles male Rhipidura atra,
so that the Fore called the former by the latters mame ( tretme );
Distinguishing features are the white pectoral tufts of the former and
white superciliary spot of the latter (not always obvious); the lower
mandible, blue-black in the former and orange in the latter; the voice;
and behavioral differences.
VOICE. ‘Three or four identical notes in rapid succession on the
same pitch, all three or four being completed within 14 sec. ‘The
quality is distinctive: dry, buzzy, rasping, fairly loud, totally un-
musical, and like an insect or the scolding of a wren (Troglodytidae).
Monarcha frater periophthalmicus Sharpe
Black-winged Monarch
SPECIMENS EXAMINED. Okasa: 4 ¢, 2 9 (22-25 June 1965). Karimui: 2 ¢,
2 @ (1 July-5 Aug. 1965). Mt. Karimui Zone 1: 1 9; Zone 2: 1 g (10 and 15
Aug. 1965).
WEIGHT, 7-42 210); 21.0) 223,225,230; 25.0, 240; 45°03 2018, 2107 2155, 215.
WENG. 6 42 65, 85, 87,188, 90; 91) b QO: 18d; S484, Sb; “tb.
TAXONOMY. ‘The black of the face extends behind the eye,
whereas it does not do so in the nominate race. In color my series is
comparable to southeastern New Guinea birds, whereas ‘Telefolmin
birds average darker gray than either on the breast and back.
250
SPECIES ACCOUNTS
BREEDING. Gonads were large in one Karimui male, somewhat
enlarged in the other, and small in all other specimens.
DISCUSSION. This is a solitary species of the middle- and upper-
stories (20-80 ft above the ground) of hill forest, between ca. 1,800 and
5,100 ft in my study areas, varying locally from common to absent.
There is no straggling to higher altitudes, and there seem to be no
records above 5,100 ft anywhere in New Guinea. Bulmer collected M.
frater at 4,000-4,500 ft in the Baiyer Valley. M. frater does not descend
to the understory and was not netted, differing in this respect from
its congener M. axillaris, much of whose altitudinal range it shares.
VOICE. The song (Fig. 28, p. 255) is a fairly loud, mellow, cheerful,
whistled note repeated several times at intervals of about | sec. Each
note is initially upslurred and then downslurred and may be preceded
by a much shorter note. The quality is reminiscent of Pitohui dichrous
or Meliphaga flaviventer. The call is a dry, buzzy “chuck-chuck-chuck.”
Monarcha guttula (Garnot)
Spot-winged Monarch
SPECIMENS EXAMINED. Karimui: 4 ¢, 3 9 (7-13 Aug. 1964; 3-16 July 1965).
Bomai: 3 ¢ (6-9 July 1965). Soliabeda: 6 ¢, 2 9 (22-28 July 1965).
WEIGHT. 10 ¢@: 14.0-18.3 (166+ 1.3). 4 9: 14.5, 16.0, 16.5, 17.0.
WING. 10 ¢: 77-83 (80+ 2). 3 9: 78, 80, 80.
STOMACH CONTENTS. Insects.
TAXONOMY. ‘The series shows no change in wing length with
altitude.
BREEDING. In August 1964 one male and one female at Karimui
were sexed, and both were found to have enlarged gonads. Out of
seven males and five females sexed in July 1965 at Karimui, Bomai,
and Soliabeda, two males and one female at Soliabeda had somewhat
enlarged gonads, and the other specimens had small gonads. This sug-
gests that the dry weather of July-August 1965 suppressed breeding,
as also noted for some other species at Karimui (p. 00).
DISCUSSION. Other localities where this tropical monarch has
been found in the Eastern Highlands are Lake Kutubu (2,450 ft) and
Baiyer River (N.G.B:S. Newsletter, No. 54, p. 2; June 1970). It did
not even get into Zone | of Mt. Karimui, and was commonest at my
lowest station, Soliabeda, where it accounted for 2°% of the local
avifauna. It was an active and solitary forest understory species, forag-
ing up to 30 ft above the ground.
VOICE. ‘The call is a dry squawk on constant pitch or slightly up-
slurred; or else a dry rasped scolding difficult to distinguish from that
of Monarcha telescophthalmus or Toxorhamphus poliopterus.
251
SPECIES ACCOUNTS
Monarcha chrysomela aruensis Salvadori
Black and Yellow Monarch
SPECIMENS EXAMINED. Karimui: 2 9 (14 July and 4 Aug. 1965). Soliabeda:
] g (24 July 1965).
WHIGTIO. 1 42 150, 2 o% 130) 1s,
—
WING. Ia: 67.) 2. 0'-266,.68.
STOMACH CONTENTS. Insects.
TAXONOMY. There are clinal west-to-east color variations in
both males and females as one proceeds from the Aru Islands and the
Fly River (race aruensis) to the Karimui area to southeastern New
Guinea and the D’Entrecasteaux Archipelago (race praerepta). In the
males, D’Entrecasteaux birds differ from the others by being generally
more yellow and less orange. ‘The crown of males is most orange in
Aru Islands and Fly River birds, least so in D’Entrecasteaux birds;
Sohabeda and southeastern New Guinea birds are intermediate, with
the Soliabeda male closer to Aru Islands-Fly River birds and south-
eastern New Guinea specimens closer to D’Entrecasteaux birds. In the
females the breast is more olive on the Fly River than in the D’Entre-
casteaux Archipelago, with southeastern New Guinea birds inter-
mediate and variable. Karimui females are somewhat nearer D’Entre-
casteaux than Fly River females in this respect but are deeper yellow
below than these or other praerepta or aruensis. Fly River females are
more olive, darker, and less yellow dorsally than D’Entrecasteaux or
southeastern New Guinea females, and in this respect the Karimui
females agree much better with Fly River females, from which they
differ in being somewhat more yellow. ‘Thus, the three specimens from
the Karimui area are on the whole closer to the aruensis than to the
praerepta end of the cline.
BREEDING. Gonads were not enlarged in the females and only
slightly enlarged in the male.
DISCUSSION. At Karimui and Soliabeda this tropical species was
very uncommon (ca. 0.2% of the local avifauna). My highest record
was a single sighting at 4,120 ft on Mt. Karimui. Except for a male
and female seen chirping at each other, the species was solitary. Its
habitat is the forest and the forest edge, particularly trees with some
open space around them. It is even more nervous and active than other
species of Monarcha and works around the tops and outside of the
crowns of trees, generally at least 30 ft above the ground, so that it was
never netted.
VOICE. The song is a loud musical jumble of mellow notes, spitted
notes, “cheep’s”, and squawks, accompanied by frequent snapping of
the bill:
252
SPECIES ACCOUNTS
Monarcha telescophthalmus harterti van Oort = M. t. henkei Meyer!
Frilled Monarch
SPECIMENS EXAMINED. Okasa: | = (23 June 1965). Karimui: 5 g, 1 ao
é (1 July-5 Aug. 1965). Bomai: 3 ¢,1 @ (7-9 July 1965). Soliabeda: 1 6, 1 Q,
Ll imm. ¢ (25-29 July 1965). Met. itr Zone 2: 1 9; Zone 3: 1 9 (14 ae 16
Aug. 1965).
WEIGHT. 8 @: 16.0-19.0 (17.1+1.0). 3 9: 15.7, 16.0, 18.5. 1 imom, @: 18.5.
WING. 10 @: 78-87 (82+ 2). 3 9: 74, 78, 79. 21mm. ¢: 79, 81.
STOMACH CONTENTS. Insects.
TAXONOMY. ‘The taxonomic analysis of this species is of in-
terest in that my series is geographically intermediate between harteri
of south-central New Guinea and henkei of southeastern New Guinea,
and in that my females are close to the former race and my males to
the latter. In the male henkei is best distinguished from harterti by the
(on the average) smaller extent of black on the chin, and in this
respect my males are closest to henkei. In the female henkei often has
a paler and more rufous back and faint rufous wash on the abdomen,
while harterti has a darker, browner, nonrufous back and a pure
white abdomen; my females agree in both respects with harterti. There
is also a slight size difference, henkei being larger than hartert:. Mayr
and Rand (1937, p. 154) give as average wing length ¢ 80.9, 9 78.7
for henkei from southeastern New Guinea, ¢ 78.5, ? 75.4 for harterti
from the Oriomo River, while Rand (1942a, p. 338) gives 4 79.5,
@ 77.0 for harterti from the Fly River. The average values for my
series (¢ 3108) a 77.0) ane mearer henkerim the male, harterti in the
female, as are the color characteristics.
BREEDING. Gonads were very small in all specimens except for
one Karimui male, in which they were slightly enlarged.
DISCUSSION. I agree with Ripley (1964, p. 61) that separation of
this monarch and its Australian relatives in a genus Arses is un-
desirable.
Monarcha telescophthalmus was seen fairly often at Soliadeda and
Karimui, where it accounted for about 1% of the local avifauna. It
was much less common on Mt. Karimui, and the highest record was
at 4,900 ft on Mt. Karimui and approximately the same altitude below
Mengino. Its habitat is the forest, edge of the forest, and second-
growth, where it is usually found 5-40 ft above the ground. A distine-
tive habit of M. telescophthalmus not shared with its New Guinea
congeners is one of standing on or coming down the sides of large
vertical tree trunks, either right-side-up or upside-down. Whet her
pursuing these nuthatch-like habits or chasing flying insects, it moves
actively.
VOICE. ‘The call is a thin, buzzy, upwards-inflected squawk. ‘The
1 Listed as Arses telescophthalmus in Rand and Gilliard (1967).
253
SPECIES ACCOUNTS
song is a dry, slow, rattled, ringing trill which progressively drops in
pitch.
Myvagra alecto subsp!
Shining Flycatcher
STOMACH CONTENTS. Insects.
DISCUSSION. Bulmer collected a specimen beside a stream at
4,200 ft in Kyaka territory, Schodde and Hitchcock report the species
in riverside shrubbery near Lake Kutubu (2,450 ft), and Kikkawa
found it in the Baiyer Valley. This lowland species of the forest edge
and second-growth may spread upward to similar habitats elsewhere
in ecologically disturbed areas of the hill slopes in the Eastern High-
lands. I follow Schodde and Hitchcock (1968) in treating the species
as a Myiagra, not as a Monarcha.
VOICE. ‘The call is a harsh rasp that crescendoes greatly. ‘The song
is a rapid trill of clear whistled notes, crescendoing somewhat, and
either all on the same pitch or else slightly rising.
Myiagra cyanoleuca (Vieillot)
Satin Flycatcher
Bulmer collected one specimen of this winter visitor from Australia
at 4,400 ft in the Baiyer Valley on 12 July 1955. Apparently the only
other records from northern New Guinea are a male collected at
Hollandia on 4 July 1938, by the Third Archbold Expedition, a male
taken by Gilliard at Aiome at the foot of the Schrader Range in May
or late April 1964, and records from Karkar (March 1914; June 1969)
and Manam (December 1913; January 1914) islands.
Machaerirhynchus flaviventer xanthogenys Gray
Yellow-breasted Flatbill Flycatcher
SPECIMENS EXAMINED. Karimui: 2 ¢,1 9 (2-16 July 1965).
WEIGHT. 2 4; 10,115, i o- 1200.
WING. 2 4: 61, 62. 1 9: 60.
TAXONOMY. ‘The forehead and eye stripes of the males are yel-
low rather than white, a character unique to the southern New Guinea
race xanthogenys. ‘The upper backs of the two males are black with
but little olive; the amount of olive in larger series of xanthogenys is
quite variable.
BREEDING. The gonads were greatly enlarged.
DISCUSSION. ‘This tropical flycatcher was seen and heard regu-
1 Listed as Monarcha alecto in Rand and Gilliard (1967).
254
SPECIES ACCOUNTS
larly at Soliabeda (2% of the local avifauna) and infrequently at
Karimui, Bomai, and up to 4,100 ft on Mt. Karimui. It was solitary,
active, and usually seen 10-50 ft above the ground, but one of the
specimens was netted in the understory. It seems commoner at the
forest edge and in open parts of the forest than in the shaded forest
interior.
VOICE. ‘The quality of the song is distinctive, a weak, high, buzzy,
unmusical and dry run-together trill. The pattern is usually that
depicted in Figure 28: two connected notes, the first dropping in
Monarcha frater:
NAAA ate. Gis ee Ee aie
“ANANANH
Machaerirhynchus flaviventer:
—
Machaerirhynchus nigripectus:
Microeca flavovirescens:
el pad load
Fic. 28. Voices of four New Guinea flycatchers.
255
SPECIES ACCOUNTS
pitch, the second steady at a higher pitch. Sometimes the trill rises
straight up the scale instead.
Machaerirhynchus nigripectus saturatus Rothschild and Hartert
Black-breasted Flatbill Flycatcher
NATIVE NAME. Fore: yaragiyéta.
SPECIMENS EXAMINED. Awande: 1 ¢ (17 June 1965). Agotu (Gimi terri-
tory): 1 ¢@ (22 July 1964). Maiba (Gimi territory): 1 9 (23 July 1964). Mt
Karimui Zone 3: 1 9 (18 Aug. 1965; Zone 4: 1 ? (23 Aug. 1965).
WEIGHT. 1 4: 120.
WING. 1 4: 61. 2 9: 60, 60.
STOMACH CONTENTS. Insects.
TAXONOMY. ‘The blackish backs of the females separate these
from harterti of southeastern New Guinea and the Huon Peninsula,
in which females are dark gray-brown above.
DISCUSSION. Machaerirhynchus nigripectus is the high-altitude
equivalent of M. flaviventer, foraging rapidly from the understory to
the treetops while calling loudly. On Mt. Karimui I found the former
from 5,210 to 6,100 ft, and the latter up to 4,100 ft. Other collectors
in the Eastern Highlands have encountered M. nigripectus regularly
between 5,000 and 8,000 ft, in forest, second-growth, or at the forest
edge.
VOICE. ‘The call consists of a loud, rapid series of musical but
buzzy notes, similar to Jfrita kowaldi or to Mohoua ochrocephala of
New Zealand. ‘The song has a similar quality (Fig. 28).
Eugerygone rubra saturatior Mayr
Red-backed Flycatcher
SPECIMENS EXAMINED. Mt. Karimui Zone 6: 1 ? (3 Sept. 1965).
WEIGHT. 9.
WING. 58.
STOMACH CONTENTS. Insects.
TAXONOMY. The specimen is presumed a female because of its
plumage. The female of the nominate rubra of the Vogelkop differs
in its browner, less olive, back.
DISCUSSION. I found this rare flycatcher only between 6,100 and
7,170 ft on Mt. Karimui. The few other Eastern Highlands records
are also mainly from forest at higher elevations (7,000-8,000 ft in the
Kubor Mountains, Wahgi Mountains, Schrader Range, and Mt.
Hagen). Songs came from solitary individuals 6-10 ft above the ground
in the stunted trees of the moss forest. ‘(Their behavior was nervous;
the tail was flicked from side to side, and the wings were continuously
fluttered. The white eyelids and white in the undertail are good field
marks. The other species with which confusion may arise in the field
256
SPECIES ACCOUNTS
because of appearance, behavior, and habitat are Gerygone cinerea,
Gerygone ruficollis, and Melanocharis verstert. ae.
Mayr and Gilliard (1954, p. 349) have argued that this bird is a
flycatcher, rather than a warbler, as previously held.
VOICE. A very high-pitched, faint, tinkling song fluctuating alter-
nately above and below the same pitch.
Niche Differences in the Genera Microeca and Tregellasta
New Guinea has four widespread species of Microeca (M. flavigaster,
M. flavovirescens, M. griseoceps, and M. papuana) and one species of
Tregellasia (T. leucops). Tregellasia resembles New Guinea Microeca
species in the color of the plumage and in the orange legs, has often
been considered congeneric with Microeca, and the two genera ob-
viously share a common derivation. Of these five species, all but M.
flavigaster have been found in the Eastern Highlands. The following
discussion of their ecological differences is based partly on my field
observations in the Eastern Highlands and North Coastal Range and
partly on the basis of published locality records from other parts of
New Guinea.
While the other four species are birds of the forest, the habitat of
M. flavigaster is savanna and open country with scattered trees, from
the lowlands up to about 2,500 ft, with one record at 4,800 ft (Tele-
folmin).
Of the four forest species, M. flavovirescens is in the lowlands and
lower hill slopes, dropping out at an altitude which varies locally
between 2,000 and 4,000 ft; M. griseoceps and T. leucops are on the
middle hill slopes from about 3,000 to 5,000 ft; and M. papuwana is in
the mountains above 5,000 or 6,000 ft. Examining local variations of
the limits in more detail, one finds that the range of M. flavovirescens
and the range shared by M. griseoceps and T. lewcops are mutually
exclusive, and that the latter range and that of M. papuana are not
only mutually exclusive but probably separated by a gap of 700-800 ft.
Local variations in the altitudinal limits of the four species are there-
fore intercorrelated. For instance, the intrusion of tropical conditions
at Karimui gives M. flavovirescens a higher upper limit, and T. leucops
a higher lower limit (somewhere between 3,650 and 4,000 ft), than
usual. Similarly, the descent of moss forest to low elevations in the
North Coastal Range gives T. lewcops a lower upper limit, and M.
papuana a lower lower limit (somewhere between 4,200 and 5,000 ft),
than usual. ‘The occasional records of M. flavovirescens above 2,000 ft
are from localities where M. griseoceps is absent. While this triple
altitudinal sequence seems to hold for virtually all the explored high
mountains of New Guinea, it does not apply to the zoogeographically
distinct Fly River delta region of southern New Guinea or to the
Cape York Peninsula of Australia, where 21 species which are con-
257
SPECIES ACCOUNTS
fined to the mountains elsewhere in New Guinea, including M.
griseoceps and T. leucops, descend to sea level.
The differences between M. griseoceps and T. leucops, the only two
of the five species sharing the same altitudinal range and habitat, are
behavioral. The four Microeca species all behave in essentially the
same manner: they are social birds, usually seen in groups of three or
four, and perch upright and motionless, sallying out from their perches
to catch insects. T. lewcops sometimes adopts the same behavior, but
it seems to spend proportionately more time in the understory; it is
often solitary, whereas I have yet to see any of the other four species as
solitary individuals; and it often perches sideways on the bark of trees,
a behavior I have never seen in the four New Guinea Microeca but
which is characteristic of T. leucops’s Australian relatives and of the
New Guinea thicket flycatchers (p. 262). In their shared altitudinal
range T. leucops is a common and ubiquitous species, whereas M.
griseoceps is uncommon and local. The uncommoness of M. griseoceps,
the middle species in the three-fold sequence M. flavovirescens-M.
griseoceps-M. papuana, with habits similar to its higher and lower
relatives, is typical of the fate of the middle species in many three-
species altitudinal sequences (p. 34). The numerous T. leucops has
evidently succeeded in avoiding this “curse of the middle species” by
adding some different habits to its behavioral repertoire. However,
the fact that 7. lewcops has not been able to invade the altitudinal
ranges of the successful M. papuana and M. flavovirescens, although
it has completely overrun the range of M. griseoceps, may imply that
many of the habits of T. leuwcops are still those of the flavovirescens-
griseoceps-papuana group.
Microeca flavovirescens cuicui (De Vis)
Olive Microeca Flycatcher
SPECIMENS EXAMINED. Karimui: 6 ¢,1 9, 3 ? (8 and 9 Aug. 1964; 12 July-
5 Aug. 1965). Bomai: 2 ¢, 2 ? (6 July 1965). Soliabeda: 3 ¢ (27 and 28 July
1965).
WHIGHT. 10 4: 140070056 = Ll). 19s 120, Serr 1S0;aS0F Ts.
WING. 9 4: 76-83 (80+2). 1 Q: 73. 2 ?: 69, 72.
STOMACH CONTENTS. Insects.
TAXONOMY. Specimens from southeastern New Guinea and the
southern slope of the Snow Mountains, and my specimens, belong to
the barely distinguishable race cuicut of most of New Guinea, and
have the yellow of the underparts just detectably less dull than in
nominate flavovirescens of the Fly River and Aru Islands. ‘The dorsal
coloration appears to be subject to postmortem changes.
BREEDING. ‘Testes were much enlarged in all three Soliabeda
males, in both Bomai males, and in three of the six Karimui males.
The three Karimui males with small testes had relatively shorter wings
(76, 78, 81 mm) and may have been subadult.
258
SPECIES ACCOUNTS
DISCUSSION. At Karimui, Bomai, and Soliabeda this lowland
forest species accounted for about 0.5-1.0% of the local avifauna. Its
upper limit at Karimui (3,650 ft) is somewhat higher than usual else-
where in New Guinea, in agreement with the general Karimui trend
and probably related to the absence of M. griscoceps. Groups of two or
three birds sat motionless on perches within about 15 ft of each other.
Periodically a bird sallied out and darted around after insects, return-
ing to the same or a nearby perch. Eventually the whole group moved
on to a new set of perches. On three occasions my collectors brought in
two birds shot together, and on one occasion three birds. ‘Phe compost-
Om G& these eroups was: |. 4,.27: | 4, 17; 2875 2¢. The specimens
of uncertain sex were probably females or young birds because of their
short wings (69 mm) or low weights (13.0 and 13.5 g).
VOICE. A faint song based on a repeated phrase (Pig. 28, p. 20d),
Microeca griseoceps subsp.
Yellow-footed Microeca Flycatcher
SPECIMENS EXAMINED. Okasa: 1 9.
WEIGHT. 12.
WING. 70.
TAXONOMY. Reassessment of the races of this species, to be re-
ported in connection with my North Coastal Range collections, indi-
cates that poliocephala is synonymous with occidentalis and that the
Cape York race kempi (type examined), described by Mathews as a
new species and genus, is not separable even subspecifically from
nominate griscoceps. Analysis of color differences in this species 1s
complicated by marked postmortem changes, as also noted by Rand
(1942b, p. 483). My Okasa specimen has the wing longer than in nomi-
nate griseoceps and the back brighter and more yellow than in com-
parative material. More material would be necessary to place the
Eastern Highlands population racially. ‘he legs are bright orange, the
upper mandible black, and the lower mandible orange.
BREEDING. ‘The gonads were small.
DISCUSSION. ‘The specimen was collected at 3,550 ft and is the
sole one reported to date in the Eastern Highlands.
Microeca papuana Meyer
Yellow Microeca Flycatcher
NATIVE NAME. Fore: kenantagure.
SPECIMENS EXAMINED. Mt. Michael: 1 g,1 9,1 imm. @ (4-12 July 1964).
Mt. Karimui Zone 5: 2 4,3 9,1; Zone 6: 1 $, 1 9; Zone 7:1 4,1 93: Zone
S39 ¢, 2 9, (2/ Aug.-8 Sept. 1965),
WEIGHT. 7 6: 13.5-15.0 (14.3 + 0.6). 7 9: 13.0-16.0 (13.9 + 1.1).
WING, 97 (Go 1242 (ise 2). bes 70, 71, 72; 74 (2). Le: 69,
STOMACH CONTENTS. Insects.
259
SPECIES ACCOUNTS
TAXONOMY. The immature bird from Mt. Michael has the
legs brownish orange (only the toes bright orange), the lower mandible
orange, and the upper mandible grayish orange, whereas the whole
leg is bright orange and the bill black in adults. ‘The uppertail of the
immature is brownish, and the underparts a dirty lemon-green instead
of the bright yellow of adults.
BREEDING. Gonads were small in all Mt. Karimui specimens.
DISCUSSION. Microeca papuana is widespread in montane forest
of the Eastern Highlands up to about 8,500 ft. In the Okapa area
M. papuana descended to 6,600 ft and T. lewcops occurred up to
5,800 ft, while the corresponding altitudes for Mt. Karimui were
6,100 and 5,400 ft. Thus, there is a gap of 700-800 ft between the two
species. ‘he transition takes place 400-500 ft higher in the Okapa area
than on Mt. Karimui, in agreement with the general trend for these
areas. M. papuana ranges from the understory to about 60 ft above the
ground in groups of three or four birds which may be in several dif-
ferent trees spread over a distance of 50 ft and which fly to a new
perch at intervals of 5-15 sec. On four occasions two birds out of a
group were collected. ‘Three of these pairs proved to be a male and a
female, while the fourth was a female plus a bird of uncertain sex,
probably another female or an immature because of its short wing
(69 mm).
VOICE. ‘The song is a high, brief, tinkling, formless warble remi-
niscent of Malurus alboscapulatus and descending in pitch. ‘The calls
are a high-pitched “‘tsee”; a very faint, sibilant upslur; and five to
eight repetitions of a dry, scolding note on the same pitch.
Tregellasia leucops wahgiensis Mayr and Gilliard
White-faced Flycatcher
NATIVE NAME. Gimi: sabarotoiya.
SPECIMENS EXAMINED. Miarosa: 1 imm. 9 (13 June 1964). Okasa: 2 ¢, 2
9, limm. ¢ (21 Aug. 1964; 22-26 June 1965). Karimui: 1 ¢ (15 July 1965). Mt.
Karinui Zone I: 2 45 Zone 2: 5 6, 2 9; Zone 3: 3 $729 (W021 Aug, 1965).
WEIGHT. 10 4: 15.3-194 (174410). 7 9: 140-180 \(1b4s214), 2 imm. ¢:
15.3, 18.0.
WING, 10 @: 75-79 (721), 5 92 69 @), 70, 71 (2). 2imms 6. 73; 7:
TAXONOMY. The Okasa series compares well with a topotypical
wahgiensis female from the Kubor Mountains and with wahgiensis
from Mafulu and the Mambare River, in the white chin and yellow
throat, the white forehead, and the color of the bill, which is pale
straw-orange except for a black tip to the upper mandible. ‘The Mt.
Karimui specimens agree in these respects but have the crown blacker,
less washed with olive. There is no size difference between the Okasa
and Mt. Karimui series.
BREEDING. ‘The gonads were greatly enlarged in all Mt. Karimui
260
SPECIES ACCOUNTS
males, in the sole Karimui male, and in two males and one female
from Okasa.
DISCUSSION. Tregellasia leucops is common and widely distrib-
uted in hill forest between about 3,000 and 5,500 ft. Its maximum
abundance was reached in the Okasa forest (3,550-4,250 ft) and in Zone
2 of Mt. Karimui (4,400-4,750 ft), where it accounted for about 2%
of the local avifauna. On Mt. Karimui it dropped out at 5,400 ft, and
the sole Miarosa specimen (5,800 ft) was an immature. It was virtually
absent at Karimui (3,650 ft) despite the favorable altitude but in line
with the general tropical pattern there. The sole Karimui specimen
was not the expected immature at the limit of the species’ range but
an adult male with large testes. ‘The habits of this flycatcher were
mentioned on p. 258.
VOICE. Whereas the four widespread New Guinea species of
Microeca have weak but musical warbled songs, I heard no song or
musical notes from Tregellasia leucops despite most of the males
being in breeding condition. ‘Vhe call is a dry, nasal “chew” or a thin,
nasal series of about nine scolding notes dropping somewhat in pitch
toward the end of the series.
Monachella muelleriana muelleriana (Schlegel)
River Flycatcher
NATIVE NAME. Fore: nintere.
SPECIMENS EXAMINED. Okasa: 1 ¢,1 9 (22 and 26 June 1965). Sena River:
1 g, 1 ? (26 July 1964). Soliabeda: 1 6,1 9 (@3 and 26 July 1965).
WEIGHT. 2 62.22.0275. 209230) and’ 235,
WING. (2 242796,.98;,, (2 Oi295;. 90:
BREEDING. Gonads were small in all specimens.
DISCUSSION. Monachella muelleriana is one of the two strictly
riparian species of passerines in New Guinea, the other being Grallina
bruijni. Its habitat is streams which (a) are rushing, (b) have many
protruding boulders, and (c) are sufficiently broad to have an air space
free of vegetation over them. In practice, I encountered Monachella
muelleriana on all Eastern Highlands streams I visited between 1,350
and 4,500 ft with beds at least 15 ft broad, but not on brooks com-
pletely covered by forest. It disappears from lowland rivers below the
fall line, where the streams start to meander and the banks and bed
are no longer strewn with boulders.
In this habitat Monachella is uncommon but widespread, perching
motionless on boulders, fallen logs, and driftwood in the torrent or
else on an overhanging branch, occasionally flicking its tail or sallying
out after insects. It is a social species, usually seen in groups of three or
four, sometimes up to eight, infrequently singly. After each member of
the group has made a few sallies, the group flies a short distance
along the river with a characteristic undulating flight, just a few feet
261
SPECIES ACCOUNTS
above the water. In both its social habits and its pattern of alternating
periods of sallying with periods of perching motionlessly, Monachella
muelleriana resembles the species of the genus Microeca.
VOICE. A high-pitched, sweet, piping note “pink” given singly or
rapidly repeated five to eight times.
Petroica bivittata bivittata De Vis
Forest Robin
STOMACH CONTENTS. Insects.
DISCUSSION. Five specimens ot this uncommon high altitude
robin have been taken in the Eastern Highlands: one collected by
Bulmer at 11,500 ft on Mt. Hagen in bushes of the alpine grassland;
and two at 11,000 ft in the Lamende Range, plus two more at 10,000-
11,000 ft on Mt. Giluwe, obtained by Shaw-Mayer. I did not find it on
Mt. Michael, nor did Gilliard on Mt. Wilhelm.
Niche Differences Among ‘Thicket Flycatchers
For convenience the name thicket flycatcher may be applied to
muscicapids of the genera Poecilodryas, Peneothello, Heteromytas, and
Pachycephalopsis, which link Petroica to Pachycephala. Most thicket
flycatchers have similar habits, size (20-40g), and body form; the generic
lines among them are not entirely clear (Mayr, 1941la). Eight species
have been found to date in the Eastern Highlands: Poecilodryas
hypoleuca, Poecilodryas placens, Poecilodryas albonotata, Peneothello
sigillatus, Peneothello cyanus, Peneothello bimaculatus, Heteromyias
albispecularis, and Pachycephalopsis poliosoma. Poecilodryas albo-
notata lives in the middle- and upperstories, but the other species live
mainly in the understory and are rarely seen more than 6 ft above the
ground, so that mistnets have proved a boon to studying their distribu-
tions. At least six (Poecilodryas hypoleuca, P. placens, P. albonotata,
Peneothello sigillatus, P. cyanus, Heteromyias albispecularis), and prob-
ably all, have the distinctive habit of perching sideways on the bark
or stems of vertical saplings or tree trunks with the body held hori-
zontal, a habit shared by some other related muscicapids of the genera
Tregellasia, Eopsaltria, Quoyornis, and Petroica. Like flycatchers of
genus Petroica, the thicket flycatchers move in a characteristic fashion:
they remain perched motionlessly for long periods, sometimes holding
apparently awkward postures and appearing tense and ready to pounce,
then abruptly fly several yards further to a new perch. Flycatchers of
genus Pachycephala, to which the thicket flycatchers show some re-
semblance and in which a few of the species were formerly placed, may
also perch motionless for short times but have less abrupt transitions
between motion and rest.
The three commonest species sort out on the basis of altitude, ap-
262
SPECIES ACCOUNTS
parently with sharp transitions and no overlap: Pachycephalopsis
poliosoma at low altitudes (ca. 3,000-5,000 ft), Peneothello cyanus at
middle altitudes (ca. 5,000-8,000 ft), and Peneothello sigillatus at high
altitudes (ca. 8,000-11,000 ft). The Pachycephalopsis poliosoma-Penco-
thello cyanus transition provides one of the few cases in New Guinea
of strict altitudinal exclusion between two species which are not 1m-
mediately derived from a common ancestor. Poecilodryas hypoleuca
is widely distributed in the lowlands and could be considered a fourth,
low-altitude equivalent of the triple sequence Pachycephalopsis polio-
soma-Peneothello cyanus-Peneothello sigillatus, but its altitudinal ceil-
ing is well below the lower limit of Pachycephalopsis poliosoma.
Poecilodryas albonotata shares most of the altitudinal range of
Peneothello cyanus and the lower part of the range of P. sigillatus,
but differs from both in its preference for the middle- and upperstories.
Heteromyias albispecularis has an altitudinal range similar to that of
Peneothello cyanus but is less common and spends more of its time
actually on the ground. Peneothello bimaculatus is local in its distribu-
tion, is largely (but not entirely) confined to the southern watershed,
and is found in hill forest in the lower part of the altitudinal range
of the heavier and commoner Pachycephalopsis poliosoma. Poecilo-
dryas placens has an extremely local distribution (only six localities in
New Guinea) below the altitudinal ranges of the other species, except
for Poecilodryas hypoleuca.
Poecilodryas hypoleuca hypoleuca (Gray)
Black and White Thicket Flycatcher
STOMACH CONTENTS. Insects.
DISCUSSION. This lowland species of the forest understory,
second-growth, and thickets has been recorded in the Eastern High-
lands only at Lake Kutubu (2,450 ft) and the Baiyer River (3,500 ft:
N.G.B.S. Newsletter, No. 54, p. 2, June 1970).
VOICE. A loud, disyllabic slur ‘“whi-chew,” the first note staccato
and at a higher pitch, the second note more explosive.
Poecilodryas placens (Ramsay)
Yellow Thicket Flycatcher
SPECIMENS EXAMINED. Karimui: 1 ¢, 1 imm. ? (10 Aug. 1964; 4 July 1965).
Bomai: 2 9, 2 ? (6-9 July 1965). Soliabeda: 3 ¢,1 9 (25-30 July 1965).
WEIGHT. 2 4: 26, 28. 3 9: 23, 24, 25. 1 ?: 28,
WING. 4 @: 90; 92, 93, 95. 3 9: 83, 84, 84. 2 ?: 84, 91. 1 imm., ?: 81.
TAXONOMY. The legs are orange or pale orange. The immature
has brown on the upperwing coverts. Comparative measurements are
summarized in ‘Table 6, from which it appears that some minor
geographical size variation exists.
263
SPECIES ACCOUNTS
TABLE 6
WING OF Poecilodryas placens
a ee eee eee eee
Locality 3 2
Batanta Island: 424, 3 Q 92-98 mm (96)1 87-91 (89)
Weyland Mountains: 84, 29 90-100 (97) 86, 88 (87)
Lake Kutubu: 1 42 96,5
Karimui: 4¢, 39 90-95 (92.5) 83-84 (83.7)
Southeastern New Guinea:
atone 92-94 (93) 85, 85 (85)
Astrolabe Bay: 1¢, 19 95 oz
1 Average values are given in parentheses.
2 From Schodde and Hitchcock (1968).
BREEDING. Gonads were greatly enlarged in all males and in
one female.
DISCUSSION. Of the passerines with a wide geographical range
on the lower hill slopes of New Guinea, few have a patchier distribu-
tion than Poecilodryas placens. It is known from six far-flung localities:
the summit of Mt. Besar on the island of Batanta off the western tip
of New Guinea, at 2,800-2,900 ft; the Menoo Valley in the Weyland
Mountains of western New Guinea, at 1,000-3,300 ft; Keku on Astro-
labe Bay in northeastern New Guinea; southeastern New Guinea
(Kubuna, Brown River, Goldie River, Mt. Cameron, and mountains
of the Kotoi District); Lake Kutubu (2,450 ft); and the Karimui area,
at 2,000-3,650 ft. At some of these localities P. placens is apparently
not uncommon, although quite unobtrusive. For example, it accounted
for 0.9% of the local avifauna at Soliabeda; Stein took 18 specimens
in the Weyland Mountains; and Gilliard collected eight on Batanta.
It is highly improbable that these six localities uniquely possess some
habitat feature required by P. placens: its habits are similar to those
of several other thicket flycatchers which are numerous, widespread in
their altitudinal ranges, and among the dominant lowerstory fly-
catchers. The only guess I can hazard is that P. placens may be too
similar ecologically to the more successful P. hypoleuca to coexist with
it and is in the process of becoming extinct. ‘his is suggested by the
fact that at four of the six localities where P. placens has been found,
P. hypoleuca is missing (Batanta, Karimui, most of southeastern New
Guinea) or else present but much rarer than P. placens (Menoo Valley).
In the Karimui area P. placens was found only in the most shaded
parts of the forest, where the understory was relatively open. Here it
remained strictly within 6 ft of the ground. Even more frequently than
other thicket flycatchers it perched on the vertical stems of small trees,
one leg an inch above the other, gripping the stem, holding the body
horizontal and leaning outward on the extended legs, and giving an
impression of intense concentration as it peered ahead. It held these
uncomfortable-looking perches without moving for long times, but
when it finally moved it did so abruptly and flew on to another stem
264
SPECIES ACCOUNTS
T
a few yards away or occasionally alit on the ground. I saw only solitary
individuals, though a male and a female were once collected together.
Apparently the only other published field observations of P. placens
are those, of Stem (1936, p. 39).
VOICE. The call is a buzzy squawk, but the song is one or the
loveliest in New Guinea. The pitch is moderately low, the volume
subdued, and the quality mellow and flute-like. Notes begin with a
clear and sharp attack, like a bell or like the pipes of a good Baroque
organ. The pattern consists of one to three connected pairs of notes
with a brief pause between each pair, delivered at a rate of about
three pairs in 2 sec (Fig. 29). The first note of each pair is higher than
the second by an interval that may be up to a major third, particularly
for the second and third pairs of a song, while the pitch interval in the
first pair is often so small as to give the impression of one longer note
on the same pitch. Usually the lower members of all two or three pairs
are on the same pitch, but in one case the third pair was entirely
below the pitch of the second pair. Once I heard a song consisting of
five single notes on progressively lower pitches but with the usual
quality.
Poecilodryas albonotata griseiventris (Rothschild and Hartert)
Black-throated ‘Thicket Flycatcher
SPECIMENS EXAMINED. Awande: 1 ¢ (a) (26 June 1964). Mt. Michael: 1 ¢@
(b) (5 July 1964). Mt. Karimui Zone 5: 3 4,2 9; Zone 8: 1 Q (28 Aug.-4 Sept.
1965).
WEIGEMIC 3° 0: 98:0. 40:7, 45.0) 5° 02 34.7... 30.5, 9957.
WENG. 5 3298 (), LOL (by), 106, 110; 12. 3 Os 102, 104, 109,
STOMACH CONTENTS. Insects.
TAXONOMY. The race correcta of southeastern New Guinea and
the Huon Peninsula differs from this series in being paler above, hav-
ing the white of the underparts more extensive and buffy, and the
black of the throat paler. The Awande specimen, from my farthest
east station, approaches correcta in having the greatest extent of
white on the underparts of my series, more than any griseiventris
specimen available for comparison and more than some correcta.
However, it lacks the buffy tinged underparts and paler upperparts
of correcta. ‘The Mt. Michael and Mt. Karimui specimens agree in
color with griseiventris except in having the throat blacker and less
brown, possibly because of foxing in the older material. P. a. correcta
averages slightly smaller than griseiventris from western New Guinea,
but the Awande and Mt. Michael males (listed as “a” and “‘b” above)
plus one male taken at Nondugl by Gyldenstolpe (1955, p. 107: wing
given as 97 mm) are smaller than the Mt. Karimui birds or any other
specimen of grisetventris and smaller than most correcta.
BREEDING. Gonads were small in all specimens.
265
SPECIES ACCOUNTS
Poecilodryas placens:
ran Gea le le or ~ Lx or mee als o ee
a
2 sec ,
Peneothello cyanus:
------ or eee ree
whistled
unmusical
Pachycephalopsis poliosoma:
a
Bs -~
ee a OF A 7
, 3 sec, pa SctSeO a 4
Fic. 29. Voices of three species of thicket flycatchers.
DISCUSSION. Poecilodryas albonotata is widely distributed but
uncommon in forest at 6,000-9,000 ft. It perches upright and motion-
less at 20-70 ft above the ground, sometimes in the understory, holding
the perch for 10-60 sec before flying with a loud fluttering of wings
to the next perch 20 ft or more away. Occasionally it slowly raises and
lowers its tail. Its insect food is either captured in midair or plucked
off the vegetation.
VOICE. ‘The song is at the highest pitch of any New Guinea bird
song: two to four identical, thin, drawn-out downslurs, repeated at
lf sec intervals (Fig. 29).
Peneothello sigillatus sigillatus (De Vis) and
P. s. hagenensis Mayr and Gilliard
White-winged ‘Thicket Flycatcher
SPECIMENS EXAMINED. Mt. Michael: 1 ¢,1 9, 1 imm. ? (8-11 July 1964).
Mi, Karim Zone 7: 1 4, 1 eo 1 pi4=7 Sept, 1965),
WEIGHT. 1 4, 2402 1 OF 213. 1 rs 2
WING, 2 2:°90,95. 2°95 87,85. Ly 9a. 1 imi. rs 57.
TAM. Us 02.9, OisG2. Le omine yor.
STOMACH CONTENTS. Insects.
TAXONOMY. ‘The race quadrimaculatus of western New Guinea
differs in having white patches on the sides of the breast; hagenensis
from Mt. Hagen differs in that the white patch in the secondaries is
266
SPECIES ACCOUNTS
much more extensive and in that the innermost secondary lacks a
black tip. The Mt. Michael and Mt. Karimui series show no dilfer-
ences in the wing patch, with white being confined to three secondaries,
which have extensive black tips. ‘The wing patch is of the same size
in nominate sigillatus from southeastern New Guinea, Mts. Wilhelm
and Kubor, and the Schrader Range, but the one available specimen
of saruwagedi from the Huon Peninsula has somewhat more extensive
black tips. Measurements of wing length gave: saruwagedi, 1 9? 88;
P. s. sigillatus, Mts. Wilhelm and Kubor, 4 ¢, 92, 93, 94 and 96, 5 ¢9@,
89, 92, 94, 95, and 96 mm. Smaller size was cited as a characteristic of
saruwagedi compared to sigillatus in the original diagnosis (Mayr,
1931, p. 680), but use of this as a racial characteristic is complicated
by an increase of size with altitude at the same location (Mayr and
Rand, 1937, p. 159; Rand, 19425, p. 485). My semes may, therefore,
be assigned to sigillatus and is quite distinct from hagenensis. It is
surprising that these two subspecies occur 30 miles apart (the distance
between Mt. Hagen and Mt. Kubor), since most other high altitude
species are represented by the same subspecies on different peaks of
the Eastern Highlands.
The immature has the uppertail largely black, the upperparts largely
black but mixed with increasing numbers of rufous feathers towards
the head, and the underparts mottled rufous and dark gray, with
rufous predominating.
DISCUSSION. Peneothello sigillatus is the thicket flycatcher of the
moss forest and seems to be the high-altitude equivalent of P. cyanus.
On Mt. Michael, where moss forest begins at 8,700 ft, P. cyanus went
up at least to 8,100 ft, and P. sigillatus extended from 11,000 ft, near
timberline, down to at least 9,500 ft. On Mt. Karimui, where moss
forest descends to 6,500 ft, the transition between P. cyanus and P.
sigillatus took place around 7,000-7,300 ft. P. sigillatus usually remains
within a few yards of the ground or on the ground itself, changes
perches at intervals of 5-20 sec, and often perches sideways on saplings.
Once I saw it in a tree fern in alpine grassland 40 ft from the forest
edge.
Peneothello cyanus subcyaneus (De Vis)
Slaty ‘Thicket Flycatcher
NATIVE NAMES. Fore: apari. Gimi: Idélori. Daribi: karodbo.
SPECIMENS EXAMINED. Awande: 3 6,1 9,1 imm. ¢ (19-27 June 1964; 15-
19 June 1965). Mt. Karimui Zone 3: 1 ¢; Zone 4: 2 6,3 9; Zone 5: 4 4,2 Q;
Zone 6: 2 6,2 9 (17 Aug.-7 Sept. 1965).
WEIGHT. 10 6: 24.0-30.0 (269+1.5). 10 9: 20.7-26.0 (23.2+1.7). 1 imm.
Gi 29:0:
WING. 10 6: 93-99 (95.5+ 1.8). 10 9: 80-90 (86.2424). 1 imm.
ae by
STOMACH CONTENTS. Insects.
TAXONOMY. ‘The color of the crown is close to that of south-
267
SPECIES ACCOUNTS
eastern New Guinea birds, and paler than in specimens from ‘Tele-
folmin, the Cyclops Mountains, or the Snow Mountains. Mt. Karimui
specimens are darker in general coloration than those from Awande.
These and other Eastern Highlands series of P. cyanus are slightly
larger than suwbcyaneus from southeastern New Guinea. There is no
size difference between Awande and Mt. Karimui birds, nor is there
a change of size with altitude on Mt. Karimui.
BREEDING. Gonads were enlarged, usually greatly, in almost all
Mt. Karimui adult males and in two of the three adult males taken
at Awande in 1965. No immature birds were taken on Mt. Karimui.
DISCUSSION. Peneothello cyanus is the thicket flycatcher of forest
from about 5,000 ft up to the level at which heavy mossing appears.
Its abundance shows puzzling variability. On Mt. Karimui (west peak),
where it occurred from 5,200 to about 7,000 ft, it was the commonest
flycatcher and the fourth most abundant bird in this zone, accounting
for about 6% of the local avifauna. At some other localities it is
uncommon or even absent.
Its behavior is similar to that of Poecilodryas placens, i.e., it gen-
erally remains within 5 ft of the ground (though I have also seen it as
much as 15 ft up) and remains motionless at a perch, appearing to peer
intently, before abruptly flying on to another perch.
VOICE. ‘The song is similar to that of Myzagra alecto: a series of
a dozen clear, musical, whistled notes at medium pitch, the first few
of which are on the same pitch, but they then progressively rise in
pitch and accelerate (Fig. 29, left, p. 266). ‘The volume remains un-
changed or else progressively increases only slightly. A less common
song with a very different quality is a loud, unmusical, three-note
phrase repeated several times and preceded by one or two other notes
(Bie. 29, mene).
Peneothello bimaculatus bimaculatus (Salvadori)
White-rumped Thicket Flycatcher
SPECIMENS EXAMINED. Karimui: 5 ¢, 4 ?, 1 imm. 4, 1 imm. ? (9-12 Aug.
1964; 1 July-6 Aug. 1965). Bomai. 3 g, 2 9 (6-8 July 1965).
WEIGHT. 7 4: % @), 26 (2), 27 (2), 28. 2 9: 21, 22. 1 imm. ¢: 24
WING. 7 42 84, 86,87 (3), 88) 89. 2 O05 977, 79. 1 amms 22 786-. mm ee.
TAXONOMY. Material available for comparison consisted of
vicarius from the Huon Peninsula, Adelbert Mountains, and the
northern slopes of southeastern New Guinea, and nominate bimacu-
latus from the Vogelkop, the Weyland Mountains, the southern slopes
of the Snow Mountains, and the southern slope of southeastern New
Guinea. The race vicarius is distinguished by having almost no white
on the belly. Snow Mountains birds assigned to bimaculatus have the
next least amount of white on the belly. ‘The other series of bimacu-
268
SPECIES ACCOUNTS
Jatus (including mine) show individual variation in the amount of
white but no average differences between series. “he extent of white
in the breast patch also shows individual variation but not average
variation between series. The depth of blackness of most of the body
plumage varies; this color has a brownish tinge in all comparative
material and in my females and immatures, but my males stand out
at once as being fully black. Foxing may be part or most of the ex-
planation in this case, since one male collected by Gilliard in the
Tamrau Mountains of the Vogelkop in 1964 is as black as my males
and blacker than prewar Vogelkop males. ‘The two immatures have
brown tips to the upperwing coverts. ‘Table 7 shows that there is little
size variation, except that Snow Mountains birds are distinctly smaller.
The Snow Mountains population may eventually require naming on
the basis of small size and reduced extent of white on the belly, but
a larger series is required to confirm these differences.
TABLE 7
WING oF Peneothello bimaculatus bimaculatus
Locality 3 Q
Vogelkop 8,895,100, OL, 91, 92) (ay., 88) 79, 80, 81, 84, 85
(av., 82)
Weyland Mountains 89
Snow Mountains 78, 80, 80, 85 (av., 81)
Karimui 84, 86, 87, 87, 87, 88, 89 77,79
(av., 87)
South slopes of south-
eastern New Guinea 84, 84, 85, 85, 86, 88, 89, 90 81, 81
(av., 86)
BREEDING. Of the eight adult males, the testes were somewhat
or much enlarged in five, small in three.
DISCUSSION. Peneothello bimaculatus resembles Poecilodryas
placens in being an “unsuccessful” thicket flycatcher with a patchy
distribution, though it is not so extreme a case. It has turned up at
about 15 scattered localities in southern New Guinea extending around
to the Adelbert Mountains in the northeast and the Weyland Moun-
tains in the northwest, but it is apparently absent in most of northern
New Guinea (i.e., no records from the northern slopes of the Central
Range, the Cyclops Mountains, or the North Coastal Range). In the
Eastern Highlands I met P. bimaculatus only at Karimui and Bomai.
When seen, it was perched a few feet above the ground in under-
brush in forest or at the edge of forest, sitting motionless except for
occasionally flicking its wings.
269
SPECIES ACCOUNTS
FHeteromyias albispecularis centralis Rand
Ground ‘Thicket Flycatcher
NATIVE NAME. Fore: yobago.
SPECIMENS EXAMINED. Awande: 2 @, 1 9 (15-18 June 1965). Miarosa: I
3,2 ? (17-24 June 1964). Mt. Michael: 2 ? (4 July 1964).
WEIGHT. 2 22°52 36. oF Sir
WINGS “3 362°96, 100, N0I, W O-Ole e038:
STOMACH CONTENTS. Insects.
TAXONOMY. My specimens differ from examples of centralis
from the Wahgi region only in having the breast whiter, less gray.
Despite their intermediate geographical position they show no ap-
proach to the brownish black crown of armiti of the Herzog Moun-
tains and southeastern New Guinea (the crown is black instead), but
the Awande and Miarosa specimens do approach slightly the brighter
and more olive back of armiti.
BREEDING. Gonads were large in the female and one male from
Awande, small in the other Awande male.
DISCUSSION. Heteromyias albispecularis is present, though un-
common, at most forest localities in the Eastern Highlands between
5,500 and 8,500 ft, but was absent on Mt. Karimui. It frequently
perches on the ground.
VOICE. A ringing, bell-like note repeated 4-8 times per second
for up to 15 sec without change of pitch or volume. A fragment of
this song two to five notes long is often given as a call. The call of
Climacteris leucophaea, a midmontane species which is absent from
the Eastern Highlands, is quite similar but crescendoes slightly. An-
other song of H. albispecularis consists of three or four clear, high-
pitched, whistled notes going up the scale and is the origin of the Fore
name “‘yo-ba-go”’.
Pachycephalopsis poliosoma albigularis Rothschild
White-throated ‘Thicket Flycatcher
NATIVE NAMES. Gimi: soy. Daribi: karoabo.
SPECIMENS EXAMINED. Okasa: 2 ¢, 1 92 (20 Aug. 1964; 22 and 26 June
1965). Sena River: 1 ¢,1 9 (26 and 29 July 1964). Karimui: 2 ¢,2 9 (7 Aug.
1964; 11 July and 6 Aug. 1965). Mt. Karimui Zone 1: 4 6,5 9; Zone 2: 1 ¢, 2
oO; Zone a: 1 @52 Oo (9-12 Aue. 196),
WEIGHT. 10 @: 35-42 (405+ 2.6). 10 9: 32-40 (35.6 + 2.7).
WING. 10 @: 100-110 (105 + 3). 10 9: 93-103 (98 + 2).
TAXONOMY. ‘The Eastern Highlands series was compared with
albigularis (type and two other topotypical specimens from the Wey-
land Mountains, plus two from ‘Telefolmin), approximans (southern
slope of the Snow Mountains, four specimens), balim (the type from
the northern slope of the Snow Mountains at higher altitudes), iden-
burg: (northern slope of the Snow Mountains, lower altitudes, four
270
SPECIES ACCOUNTS
specimens), /rypopolia (Huon Peninsula, one specimen), and nominate
poliosoma (southeastern New Guinea and Herzog Mountains, many
specimens). Eastern Highlands birds are close to topotypical albigu-
laris, particularly in the dorsal coloration, and differ mainly in the
bluer, less gray underparts and the darker uppertails, in both of which
deviations they resemble ‘Teleformin birds. It was initially surprising
when Telefolmin birds, separated from the Weyland Mountains by
300 miles and the populations of idenburgi and balim, had to be as-
signed to albigularis (Gilliard and LeCroy, 1961, p. 59). It is even more
surprising that birds from the southern slope of the Eastern High-
lands, 500 miles to the east of the Weyland Mountains and on the
opposite watershed, are also albigularis. Of the two other southern
watershed races, approximans differs in the more eray-brown and less
blue upperparts, the grayer and less blue underparts, and the much
lighter and browner uppertail, while nominate poliosoma is duller,
more gray-brown and less blue above (especially on the crown), lighter
on the exposed parts of the upperwing and tail, more gray-brown and
less blue below (especially on the breast), and has the white of the
throat more restricted in extent and more washed with gray-brown
posteriorly. Compared to Mt. Karimui specimens, Okasa specimens
show a slight tendency towards nominate poliosoma in the bufher and
less extensive throat patch, but they are still much nearer albigularis.
The race balim differs in having the back, crown, and breast more
gray-brown and the undertail coverts buffy, not gray; tdenburgi 1s
smaller, grayer below, has the extent of white on the throat more
reduced, and the margins of the remiges browner; and hypopolia is
smaller, paler, and grayer on the back and crown, more gray-brown
below, and the extent of white on the throat is more restricted. No
specimens of the race hunsteini (mountains on the upper Sepik River)
were available, but from the description it must differ in the reduced
extent of white on the throat and in other respects.
BREEDING. Gonads were small in Okasa specimens except one
male, which had greatly enlarged testes. At Karimui gonads were
greatly enlarged in males and in one female. On Mt. Karimui testes
were moderately enlarged in one male, slightly in two, not at all in
three; and ovaries were enlarged in only two females.
DISCUSSION. P. poliosoma is the common thicket flycatcher of
the hill forest (3,000-5,000 ft). However, it is so shy and hard to see
that it may be overlooked completely unless one learns its song or sets
up mistnets. Song censuses and netting returns show that it reached
its maximum density in Zone | of Mt. Karimui (4,000-4,200 ft), where
it was the most abundant bird and accounted for 12% of the local
avifauna. Between 5,100 and 5,200 ft in Zone 3 of Mt. Karimui it was
abruptly replaced by Peneothello cyanus, altitudinal exclusion between
the two species being strict. (In the Daribi language spoken at Karimui
Peneothello cyanus and Pachycephalopsis poliosoma go by the same
271
SPECIES ACCOUNTS
name, “karoabo”). At Karimui, on the floor of the Karimui Basin,
Pachycephalopsis poliosoma was much less common (0.7% of the local
avifauna), and the lowest record of it was at 3,230 ft en route to
Soliabeda. When seen, it was always solitary, in the forest interior, and
perched stationary for relatively long times within a few feet of the
forest floor.
VOICE. ‘The call is a single whistled note, often slightly upslurred,
rather loud, and with considerable carrying power and a clear, piercing
quality, like the sound produced by blowing across the open end of an
expended shotgun cartridge. The song (Fig. 29, p. 266) consists of four
to seven, usually four or five, notes with a very buzzy quality. Each note
is upwards inflected and about half a tone higher than the previous
note. Ihe time intervals are slightly less than 1 sec between the first
few notes and distinctly shorter between the last two or the last three
notes. Both the song and the call are heard often, and particularly the
song is easily recognized.
PACHYCEPHALINAE: WHISTLERS
Pachycare flavogrisea subaurantia Rothschild and Hartert
and P. f. swbpallida Hartert
Dwarf Whistler
NATIVE NAMES. Fore: yagigusaréa. Gimi: labiliyago. Daribi: bobidau.
SPECIMENS EXAMINED. Okasa: 5 ¢, 29 (23 Aug. 1964; 22-26 June 1965).
Karimui: 2 8,4 9 (13 Aug. 1964; 15 July-6 Aug. 1965). Mt. Karimui Zone 1: ]
os fone 22966. 1 2; Zone 322 6 (0-17 Auoy 1965):
WiRIGEUE 10 Ga 0-19 02a 3). Felt U2 bo Sislba7 see):
WING. 10 4: 62-68 (65+ 2). 4 9: 62, 63, 63, 64.
TAXONOMY. Pachycare flavogrisea shows considerable geographi-
cal variation in the depth of its yellow ventral color. The race randi
of the northern slope of the Snow Mountains has the deepest and most
orange color, followed by swbaurantia of ‘Yelefolmin and the southern
slope of the Snow Mountains, followed by nominate flavogrisea of the
Vogelkop, followed by swbpallida of southeastern New Guinea, the
palest and most yellow race. Where swbaurantia grades into subpallida
in the Sepik Mountains, birds resembling nominate flavogrisea are
found. Unfortunately, the color fades markedly within the first ten
years postmortem, so that subspecific assignment of fresh specimens
involves an educated guess as to what their appearance will be in ten
years. For instance, Weyland Mountains birds collected by Stein in
1931 “agreed completely” with topotypical swbaurantia in 1935 (Har-
tert, Paludan, Rothschild, and Stresemann, 1936, p. 202) but “agreed
perfectly” with topotypical P. f. flavogrisea when reexamined in 1940
(Rand, 1942b, p. 487). When I examined my specimens (collected in
1964 and 1965) in March of 1966, I found that the Karimui series was
272
SPECIES ACCOUNTS
approximately as orange as subaurantia from ‘Telefolmin (collected
in 1954, so that fading was probably completed) and was considerably
more orange than the Okasa series which agreed with nominate flavo-
grisea. In February 1967 I reexamined the specimens and found that
Karimui birds were still swhaurantia but that Okasa birds were inter-
mediate between flavogrisea and subpallida. It seems a safe guess that
the Okasa series will finally fade into suwbpallida. 1 am provisionally
listing the Karimui birds as swbaurantia, but it remains to be seen
whether they will reach the level of flavogrisea by the time they finish
fading. In any case, the general increase in orange color from east to
west (exclusive of the Vogelkop) is reflected in the difference between
Okasa and Karimui, 100 miles west of Okasa.
BREEDING. Gonads were small in all Okasa specimens except
one male; large in both Karimui males; and small on Mt. Karimui
except for being quite large in one male and slightly enlarged in
another.
DISCUSSION. Pachycare flavogrisea is a characteristic, vocal, and
common species of hill forest from about 2,800 to 5,400 ft, reaching its
maximum abundance in Zones 1, 2, and 3 of Mt. Karimui (4,000-5,400
ft, 6% of the local avifauna). There seems to be no tendency to stray
above its local altitudinal ceiling, although the level of the ceiling
shows minor local differences. While P. flavogrisea is sometimes found
in low trees in open second-growth if there is forest nearby, it is
commonest in forest. It can be seen at almost any height in the vegeta-
tional column from a few feet above the ground to the crowns of trees
but is found most often in the middlestory. It is usually solitary,
fairly active, and much more often heard than seen.
VOICE. ‘The loud, melodious, whistled, and explosive calls and
songs are among the characteristic sounds of hill forest and support
the suspected relationship of Pachycare with Pachycephala and Col-
luricincla. ‘These calls involve two kinds of notes: an upslurred
“wheep!” and a nonslurred “chew!” ‘The patterns permute a rapid
series of 2-4 “wheep’s” on the same pitch with rapid series of 2-4
“chew’s” on some other pitch, usually several notes lower. Each series
of “wheep’s” or “chew’s” progressively increases in volume. One of the
commonest patterns is: “wheep-wheep! chew-chew-chew! wheep-wheep!
chew-chew-chew! wheep-wheep! chew-chew-chew!” Figure 30 (p. 278)
gives other patterns. The rapid tempo and the repeated crescendoes are
distinctive, but confusion with the even louder and more explosive call
of Eupetes castanonotus is possible.
Niche Differences in the Genera Pachycephala, Colluricincla,
and Pitohwi
The species of these three closely related genera may conveniently
be discussed together as regards niche differences.
7 °
“49
SPECIES ACCOUNTS
Nine species of Pachycephala have been found in the Eastern High-
lands. Five of these are ecologically similar, similar-sized (weight ca.
20-25 g), rather common, forest-dwelling, insectivorous species which
form a broadly overlapping altitudinal sequence: P. griseiceps (0-4,500
{t), P. hyperythra (1,500-4,200 ft), P. soror (3,500-6,000 ft), P. schlegelit
(5,000-10,000 ft), and P. modesta (6,000-11,000 ft). As discussed on p. 00,
each species partly overlaps the altitudinal range of the species above
and below it, and the altitudinal overlap is apparently made possible
by differences in preferred vertical foraging level within the vegeta-
tional column. P. leucostigma, an uncommon forest species, has an
altitudinal range that brings it into contact with all eight other
Pachycephala species, and is similar in size to the preceding five, but
differs in that it takes considerably more fruit than insects, while the
other species take mainly insects and rarely fruit. P. rufinucha lives in
the forest and overlaps the altitudinal ranges of six of its congeners
(all those found in the Eastern Highlands except P. griseiceps and P.
hyperythra), but differs in that it is 70°% heavier (weight ca. 40 g), has
a more powerful bill and legs, lives mainly on the forest floor, and takes
larger prey than its congeners as well as fruit. P. tenebrosa, a large
(weight ca. 45 g) midmontane bird of the forest floor, is rare and known
in the Eastern Highlands from only one locality. P. rufiventris overlaps
all its congeners altitudinally but lives in very open groves, e.g., gardens
or casuarinas, a habitat shared by no other member of the genus in
New Guinea.
Colluricincla megarhyncha is common, similar in proportions to the
first five Pachycephala species discussed, overlaps most of them altitudi-
nally (ca. 0-6,000 ft), and forages for insects in the lower- and middle-
stories of the forest and second-growth, but is larger (weight ca. 35-40 g)
and more sluggish. C. harmonica is twice as large as C. megarhyncha,
comparable in size to the species of Pitohui, and lives in open groves,
a habitat in which the only other whistler is the much smaller Pachy-
cephala rufwentris.
The species of the endemic New Guinea genus Pitohwi are about
twice as large (70-100 g) as Colluricincla megarhyncha and take both
insects and fruit. All five widespread members of this genus have been
found in the Eastern Highlands. One member sorts out on the basis of
altitude: P. nigrescens is found only above 5,500 ft, where its congeners
are all absent. All four of the other species were able to coexist at
Karimui (3,650 ft). P. cristatus 1s a rare species of hill forest apparently
living mainly on the ground, whereas the other four species seem
similar to each other in their habits and occupy both the understory
and middlestory. P. kirhocephalus, P. dichrous, and P. nigrescens ap-
parently form a three-species altitudinal sequence with much overlap
between the first two and a sharp transition between the second two.
The altitudinal ranges of P. ferrugineus and P. kirhocephalus coin-
cided in the Karimui area, though P. ferrugineus has a slightly lower
274
SPECIES ACCOUNTS
ceiling elsewhere in New Guinea, and the ecological differences be-
tween these two species are not clear to me. The species “Pitohui
tenebrosus of Palau in the Caroline Islands, initially described in the
genus Recles (= Pitohui), subsequently placed in a monotypic genus
Malacolestes, sometimes placed in Collwricincla, and transferred back
to Pitohui by Mayr (1967) in Peters’ Check-list, Vol. 12, appears to me
to belong to Colluricincla and not to Pitohur.
Pachycephala leucostigma obscura (Rand)!
Mottled Whistler
SPECIMENS EXAMINED. Awande: 1 @ (28 June 1964). Okasa: 2 9, 1 ?, 1
imm. ¢ (22-26 June 1965). Mt. Michael: 1 imm. ¢ (5 July 1964). Mengino: 1 ¢
(16 July 1964). Karimui: 1 Q (17 July 1965). Mt. Karimui Zone 2: 1 9; Zone 5:
L 4, limm. 6: Zone 6: 1 @,.1 2 Ge Aug-4 Sept. 1965);
WEIGHT. 2 @: 25.0, 28.3. 5 9: 23.0, 24.5, 28.0, 29.0, 29.5. 2 imm. ¢: 25.0,
28.0. I? (9 or mmm.:), 24.0.
WING. 4 @: 85, 88, 89, 90. 5 9: 84, 84, 85, 87, 87. 3 imm. @: 85, 85, 86.
Il eke Nowe
STOMACH CONTENTS. Fruit only, ranging in diameter from 1 to 9 mm (3
d..0 ©); fruit and imsects (6 4); insects only (2).
TAXONOMY. In the western races nova and nominate leucostigma
the adult male has a mottled pattern approaching that of the female.
Compared to five adult males of obscura (one from Mt. Goliath and
four from southeastern New Guinea), my males are darker above and
darker and more olive below. The Mt. Goliath male is the next darkest
and also has obscure barring on the underparts, which is present but
less marked in my specimens and virtually absent in southeastern New
Guinea males. ‘The iris was olive in the one adult male whose eye color
I noted, brown in the females and immature males.
BREEDING. ‘Testes were greatly enlarged in the Mengino adult
and slightly enlarged in the Mt. Karimui Zone 5 adult and the Okasa
immature. In all other specimens gonads were small.
DISCUSSION. ‘The series clearly illustrates a tendency towards
altitudinal specialization according to age and sex. All four adult males
were taken above 6,000 ft (at 6,500 and 6,770 ft on Mt. Karimui, and
at 6,100 ft at Mengino and near Awande). At Okasa (3,550-4,250 ft),
Karimui (3,650 ft), and Mt. Karimui Zone 2 (4,700 ft) only females
and immature males were taken. The highest specimen was the im-
mature male from 8,000 ft on Mt. Michael. Gilliard obtained one
specimen on Mt. Hagen and another at 8,300 ft in the Schrader
Range.
I have no field observations of this whistler, whose stomach contents
consist of both fruit and insects but mainly fruit. The statement by
Rand and Gilliard (1967, p. 422) and by Stein (1936, p. 36) that its
1 Listed as Rhagologus lewcostigma in Rand and Gilliard (1967).
275
SPECIES ACCOUNTS
diet is exclusively fruit is incorrect, as already noted by Gilliard and
LeCroy (1968, p. 21). Presumably its diet is what enables it to coexist
at the same altitudes and in the same habitats as six other whistlers
(P. soror, P. schlegelii, P. griseiceps, P. hyperythra, P. modesta, and
P. rufinucha), which take fruit infrequently. All of my specimens were
obtained in forest, and 11 of the 13 were netted, suggesting lowerstory
habits.
P. leucostigma was originally described in the genus Pachycephala,
and its close affinity to the other species of this genus has never been
questioned. Although it has a distinctive color pattern and diet, I see
no reason to conceal its relationships by recognizing the monotypic
genus Rhagologus for it. Even my Fore assistants referred to P. lewco-
stigma simply as “pitna’, the Fore name for all species of genus
Pachycephala (but not of the genera Colluricincla or Pitohut).
Pachycephala soror klosst Ogilvie-Grant
Sclater’s Whistler
NATIVE NAMES. Fore: pitna. Gimi. dbuhai.
SPECIMENS EXAMINED. Awande: 2 4, 2 9 (28 June 19645 15> june 1965):
Okasa: 1 $,1 Q (2 Aug. 1964; 22 June 1965). Mt. Michael: 1 ¢,1 9 (5 and 13
July 1964). Karimui: 1 ?, 1 imm. ¢, 2 imm. ? (13 Aug. 1964; 1 July-3 Aug. 1965).
Mit anion Zone i273, Zone 2. 3563 Zone 3: lf Os Zone aie Ol
Aug.-1 Sept. 1965).
WEIGHT. 10 4: 22.5-27
290, 20. 2 imam, Pp: 20:55, 2
WING. 10 @: 88-93 (90
85.
(24,872 123) 8 Os 23.057 7,0 (24.9 420.5). 2am oie
2). i 92586-8988 20), Sammy g89), mmm
TAXONOMY. Adult males have the tail blackish (as in the western
race klossi) rather than strongly washed with olive (as in bartoni of
southeastern New Guinea and the Herzog Mountains). ‘he more east-
ward-lying Awande and Okasa birds are indistinguishable from Kari-
mui birds in this respect and thus show no approach to bartont.
Immatures have brown on the head and wings.
BREEDING. At Awande in 1965 most specimens had the gonads
much enlarged, and one female contained a nearly full-sized egg. On
Mt. Karimui one adult male had the testes somewhat enlarged, another
slightly, and the others not at all.
DISCUSSION. P. soror has a lower altitudinal limit around 3,500
ft and a ceiling at an altitude varying locally between 5,800 ft (Mt.
Karimui) and 8,000 ft (Mt. Michael). It shares the lower part of its
range with P. hyperythra, the upper part with P. schlegeli and P.
modesta. It is similar in habits to these other species except that it
spends less time in the understory than P. schlegeli and more time in
the understory than P. hyperythra or P. modesta. Three of the four
specimens at Karimui (3,650 ft), the lower limit of its altitudinal range
in this area, were immatures, in agreement with the usual pattern of
276
SPECIES ACCOUNTS
immature birds setting the lower limit of a species. Within its altitudi-
nal range P. soror is fairly common (1-2% of the local avifauna).
VOICE. I never identified the song with certainty. However, be-
tween 4,750 and 5,680 ft on Mt. Karimui, a zone in which P. soror, P.
schlegelii, and P. rufinucha were the only species of Pachycephala, I
frequently heard a distinctive song which could only have been a
Pachycephala and which was attributed by the Fore to a piina, their
name for all species of Pachycephala. Since the song of P. schlegelit
is distinctly different and I am familiar with it, and since P. rufinucha
was considerably less common, I tentatively assume that this unidenti-
fied song was P. soror and describe it here. ‘he commonest pattern
(Fig. 30) consists of a fairly rapid series (ca. three notes per second) of
8-10 very clear, whistled, high-pitched notes, all on the same pitch
and progressively increasing in volume, and usually concluding in a
loud slur, either upslurred or downslurred, which is occasionally
omitted. Variants are that the individual notes in the series of 8-10
notes may all be explosively upslurred, or else explosively downslurred,
or else disyllabic. ‘The crescendo and the bell-like clarity of the whole
song are distinctive.
Pachycephala schlegelii obscurior Hartert
Schlegel’s Whistler
NATIVE NAMES. Fore: pitna. Gimi: ébuhai.
SPECIMENS EXAMINED. Awande: 1 ¢ (17 July 1965). Mt. Michael: 1 ¢@ (2
July 1964). "Mt Karina Zone 21 imm. 2, 1 imm,. 2; Zone 3° 7s, 1 OF 1 eo or
ie. )5. Zone 4s le 40 Zone 5-5 S53, lima, S3-Zone6: 4 45 IO. 1 amm:
onconed: 1 6; Zone 8: 5° &,2 9, 1 imm. 4 (13. Aue-10 Sept. 1905);
WHIGTIE. 10) 62 200245 (224 a2 1.2). 10 0% 194-248 (224e 14), ers23.5.
4 spoon, Ge PAM, PAO), GO, PBN, lh siaokan, 38 2 e)
WING. 10 @: 81-89 (8642). 10 9: 81-86 (8441). 1 ?: 8. 5 imm. @: 84,
84, 85, 86, 87. 1 imm. 9: 86. 2 imm. ?: 83, 85.
STOMACH CONTENTS. Most stomachs contained only insects; one adult fe-
male, only seeds; two juveniles, seeds and insects.
TAXONOMY. Analysis of wing length shows no change with alti-
tude between 4,400 and 8,165 ft, although birds from 9,000 ft upward
have been shown to be larger (Mayr and Rand, 1937, p. 173; Rand,
1942... 403),
The subspecific color differences cited by Hartert in defining the
races obscurior, viridipectus, and cyclopum are very slight. In adult
males the abdomen is somewhat more ochraceous in obscurior of south-
eastern New Guinea than in other races. My males are intermediate
but nearer obscurior.
BREEDING. ‘Testes were greatly enlarged in the one male from
Awande and in seven from Mt. Karimui; and moderately enlarged in
six, slightly in four, and not at all in five adult males from Mt.
Karimui.
277
Pachycare flavogrisea:
Ne 22D DDS
--- or ad et,
wheep! wheep! wheep!
chew!
OBEN NNN NTNENI SN or
SISIISJISI eh WASNT AS TNS
Pachycephala schlegelii:
JAMANAVIIAH __
ELGG 2S
Oe a= a ee |,
a ag
Pachycephala rufiventris:
Or ------ Or ------ = Or aS
Pachycephala hyperythra:
—fo oo nr a Adee cs dite! teh
Seer ee — , LS
< whik! whik!
ee ae
_— ‘ ; : or i
aa — whik!whik!whik!
Of, pices = So ao or = or
= chew!chew! chew! ae =
Fic. 30. Voices of five New Guinea whistlers.
278
4/
|
< whik!
f/
SPECIES ACCOUNTS
DISCUSSION. The change in population composition with alti-
tude as we went up Mt. Karimui was striking. In Zone 2 (4,400-4,750 It)
my only records were of the two immatures collected, a male in female-
like plumage and with tiny testes, and an unsexed immature with
many scattered rufous feathers (a carryover from the nestling plumage).
In Zone 3 an unsexed bird in female plumage was collected at 5,080 ft,
an adult male with small testes at 5,180 ft, and an adult female at 5,210
ft. In Zone 4 (5,390-5,960 ft) one singing male was seen and heard
around 5,900 ft, adult females were collected at 5,700, 5,865, 5,870,
and 5,960 ft, and the first adult male with enlarged testes was collected
at 5,960 ft on the boundary between Zones 4 and 5. In Zone 5 an adult
female was taken at 6,100 ft and two at 6,450 ft, an adult male with
enlarged testes at 6,100 ft and another at 6,400 ft, three adult males
with small testes at 6,100 ft, 6,430 ft, and 6,530 ft, a male in nearly
adult plumage with tiny testes at 6,500 ft; and singing males began to
be encountered regularly above 6,250 ft. In Zone 6 through 8 (6,500 ft
up to the summit at 8,165 ft) males with enlarged testes accounted
for more than half the population, and singing males were common.
The contribution of P. schlegelii to the total avifauna was below 1%
up to 5,780 ft but thereafter rose progressively from 4% in the upper
half of Zone 4 (5,780-5,960) ft) to 11%, in the upper half of Zone 8
(7,610-8,165 ft), where it was the second commonest bird (after Ptzlo-
prora guiset).
P. schlegelii is abundant and widespread in the Eastern Highlands
in forest between about 4,400 and 9,700 ft, the exact limits varying
locally. Whereas it descended to 4,400 ft on Mt. Karimui, in the Okapa
area it was still very uncommon at 6,000 ft. It ranges from the under-
story to the crowns, shifts perches every few seconds while foraging,
and obtains its food by plucking it from surfaces rather than by
sallying.
VOICE. Varied, but readily recognized as a Pachycephala by the
crescendoes, and distinguished from other species of Pachycephala by
the frequent use of a characteristic slurred note and of unmusical
sounds resembling the sound of smacking one’s lips, kissing, or a cat
meowing. ‘The characteristic slur is an explosive whistled note which
suddenly rises in pitch and then immediately returns to the original
pitch, increasing in volume as it does so (Fig. 30, p. 278). ‘This resembles
somewhat a call of Monarcha frater. A typical song consists of 10 such
slurs repeated identically at intervals of slightly less than 1 sec. Another
song is a prolonged cat-like meow which increases in volume, rises
slightly in pitch, and is terminated by the explosive slur. Sometimes
this song consisting of the meow and the slur is prefixed by four
detached, identical, bell-like notes. Countersinging males snap their
bills and flare and erect the yellow nape patch.
219
SPECIES ACCOUNTS
Pachycephala griseiceps perneglecta Hartert
Gray-headed Whistler
4 9, 1 imm. Q (7-11 Aug. 1964; 1-17 July 1965). Bomai: 1 9 (7 July 1965).
Soliabeda: 5 @, 2 Q (22-28 July 196 sp
WEIGHT. 10 ¢@: 20.0-24.0 (21.7 + 1.2). 10 9: 20.5-24.3 (220+1.2). 1 imm.
OS ANE:
WING. 10 ¢@: 82-86 (84+1). 10 9: 79-85 (82+ 2). 1 imm. 9: 76
STOMACH CONTENTS. Insects.
TAXONOMY. ‘The series is close to perneglecta from the Fly River
and from southeastern New Guinea west of Port Moresby, differing
slightly in that the olive back is brighter and greener and in that the
gray of the head contrasts more with the olive back. Okasa and Karimui
specimens are virtually the same. The race dubia of southeastern New
Guinea east of Port Moresby is very distinct in its brown back and lack
of yellow on the underparts. ‘The Lake Kutubu population also belongs
to perneglecta (Schodde and Hitchcock, 1968).
BREEDING. Gonads were small in all females and in the Okasa
male. At Soliabeda the testes were slightly enlarged in three males,
small in two; at Karimui, large in two males, small in two others.
DISCUSSION. Pachycephala griseiceps will probably be found in
most areas of the Eastern Highlands from about 4,500 ft down to sea
level. It occurs in forest but is more numerous at the forest edge and
comes out into partly cut forest and dense second-growth. Its altitudi-
nal range includes the lowest part of P. soror’s range, the entirety of
P. hyperythra’s range, and the lowlands, where no other member of the
genus is present. Whereas P. hyperythra, P. soror, and P. schlegelit
spend much of their time in the understory, P. griseiceps is usually
found at least 15 ft above the ground in the middlestory or in the
crowns, descends less frequently to the understory, and is rarely taken
in mistnets.
VOICE. <A melodious chirping which is unmistakable once learned.
The pattern consists of a strict alternation between a monosyllabic,
slightly upslurred chirp and a disyllabic chirp. Each kind of chirp is
given alternately a few to a half dozen times and returns at intervals
of between 1% sec and | sec. ‘The most distinctive and charming feature
is the flexibility both of tempo and volume, reminiscent to an exag-
gerated degree of a Chopin waltz played rubato. ‘The singer unpredict-
ably and irregularly doubles his rate of delivery while increasing the
volume. At the same time the disyllabic chirp, which is at a lower
pitch than the monosyllabic chirp at a slow rate of delivery, moves to
a pitch higher than that of the monosyllabic chirp. ‘The song is ventrilo-
quial, and the singer often hard to identify even when in open view.
Only when I succeeded in observing the throat of a P. griseiceps mOov-
ing in synchrony with the song could I finally prove that it was the
author. Among other species of Pachycephala the only foreshadowing
280
SPECIMENS EXAMINED. Okasa: 1 4, 3 2 (22-26 June 1965). Karimui: 4 2,
SPECIES ACCOUNTS
I have found of this unusual delivery is an occasional slight and
temporary hesitation in the middle of the song of P. hyperythra.
Pachycephala hyperythra salvadoru Rothschild
Rufous-breasted Whistler
SPECIMENS EXAMINED. Karimui: 1 9, 4 ? (2-10 Aug. 1964; 12 July and 5
Aug. 1965). Bomai: 1 9 (6 July 1965). Soliabeda: 4 ¢, 1 @ (27-30 July 1965).
Mt. Karimui Zone 1: 1 ¢,1 9 (9 and 10 Aug. 1965).
WEIGHT. 5 @: 25.5, 28:0; 28.5, 29.0, 29.0. 3 9% 25.5, 27:0; 2910.
WING. 4 @: 89, 90 (2), 92. 4 9: 83, 85, 88,89. 1 ?: 87.
TAXONOMY. Compared with salvadori: of southeastern New
Guinea, these specimens agree in the pale, dull ochraceous underparts,
the gray throat, and the dark and slightly brown-tinged olive back,
and differ only in the slightly brighter color of the belly. ‘The races
sepikiana and nominate hyperythra differ in the brighter and more
orange underparts; retchenowi in the brighter underparts, browner
back, and whiter, less gray, throat. The Lake Kutubu population also
belongs to salvadorii (Schodde and Hitchcock, 1968). A single speci-
men, in poor condition, obtained by the Second Archbold Expedition
on Mt. Mabiom at the headwaters of the Fly River, is brighter olive
above and somewhat brighter ochraceous below than my series. ‘Thus,
there appears to be a cline on the southern watershed, with increasing
brightness from southeastern New Guinea to the Karimui area to the
Fly River to nominate hyperythra of the Vogelkop.
BREEDING. Gonads were enlarged in one Soliabeda male but
small in all other specimens.
DISCUSSION. P. hyperythra is the whistler corresponding to P.
soror and P. schlegelii in the lower hill forest, with an altitudinal
range of about 1,500-4,200 ft. From Soliabeda it extended up on Mt.
Karimui only to 4,200 ft and failed to overlap P. schlegelii (found
above 4,400 ft) but did overlap P. soror (found above 3,650 ft). Inthe
Eastern Highlands it remained at 10-30 ft above the ground, and I
never netted it. In the North Coastal Range, where P. soror was
missing, P. hyperythra was frequently seen in the understory and was
netted.
While P. hyperythra’s geographical range covers most of New
Guinea, its local distribution has the curious patchiness which one
comes to associate in New Guinea with genera containing many eco-
logically similar species (p. 25). I failed to find it at Okasa (3,550-4,250
ft), although the altitudinal range would appear suitable. In south-
eastern New Guinea it is known from few localities; it is not included
in the exhaustive collections of the First Archbold Expedition, and
neither Mayr nor Stevens encountered it in the Herzog Mountains.
There are no records at all from the southern slope of the Snow
Mountains, although the hill forest zone there has been explored at
281
SPECIES ACCOUNTS
many points by six large expeditions. At several localities where P.
hyperythra is absent, P. soror, whose lower limit in the presence of
P. hyperythra is around 3,500 ft, ranges down to 2,000 ft or lower and
appears to take over P. hyperythra’s altitudinal range.
VOICE, A typical Pachycephala song, beginning with 1-8 clear,
faint, bell-like or whistled notes that crescendo and terminate in loud,
usually repeated slurs ““whik! whik!” (Fig. 30, p. 278).
Pachycephala modesta hypoleuca Reichenow
Brown-backed Whistler
SPECIMENS EXAMINED. Awande: 1 ? (19 June 1964). Mt. Michael: 1 ¢,1 9
(30 June and 8 July 1964). Mt. Karimui: Zone 5: 1 @ (2 Sept. 1965).
WEICTIN.. So 97.
WINGS 2 Or t82, 808 lars 85%
STOMACH CONTENTS. Solely insects in most stomachs, insects and a fruit pit
in one.
TAXONOMY. Specimens from my three localities are very similar
to each other and to birds collected on Mt. Kubor by Gilliard. ‘The
nominate race is more olive and generally lighter on the back, while
telefolminensis is grayer below and darker and grayer on the back.
BREEDING. ‘The Mt. Karimui female contained a nearly full-
sized ege.
DISCUSSION. In the areas where I collected, P. modesta was the
least common Pachycephala and the one with the highest altitudinal
range, extending from 6,000 ft to timberline at 11,000 ft. While oc-
casionally seen in the understory, it usually remained in the crowns
40 or more feet above the ground, where it foraged both by plucking
and sallying and moved by frequent short hops of a few feet alternating
with short flights. At at least two localities in southeastern New Guinea
P. modesta descends to considerably lower altitudes: to ca. 3,500 ft in
the Herzog Mountains (Mayr, 1931, p. 716) and to 4,200 ft in the
Owen Stanley Range inland from Port Moresby (Gilliard, 1950, p. 30).
The only guess I can hazard for an explanation is that, since this is an
area where P. hyperythra is missing, the remaining Pachycephala
species may have expanded their niches and altitudinal ranges.
P. modesta is one of only five species of montane birds confined to
eastern New Guinea and lacking a western New Guinea representative
in the same superspecies. However, there is a close relative of P.
schlegelii, P. lorentzi, which is confined to western New Guinea, oc-
cupies there a high altitudinal range (ca. 6,000-12,500 ft) similar to
that of P. modesta in eastern New Guinea, and may fill an analogous
niche. P. modesta and P. lorentzi to date have been found sympatric
only at Telefolmin.
282
SPECIES ACCOUNTS
Pachycephala rufiventris dorsalis Ogilvie-Grant!
White-bellied Whistler
SPECIMENS EXAMINED. Okasa: 2 6,2 Q (22-26 June 1965). Lufa: 2 g,1 @,
limm. ¢, 2? (29 June-14 July 1964). Karimui: 3 ¢,1 9 (13-15 July 1965). Bomai:
1 ¢@,1 9 (8 July 1965).
WEIGHT. 5 4: 245; 2,05, 26.3; 27.5, 28:0. 3 9: 240, 25.5, 27.0,
WING. 5 4: 88, 89, 90, 92, 93. 4 9: 83, 88, 89, 89.
TAXONOMY. Although Rand and Gilliard (1967, p. 429) keep
as distinct species P. rufiventris of Australia, southeastern New Guinea,
and the Louisiade Archipelago and P. monacha of the rest of New
Guinea and the Aru Islands, the evidence summarized by Mayr and
Gilliard (1954, p. 352) for considering them conspecific seems con-
vincing. The two “‘species” intergrade in the vicinity of Port Moresby.
DISCUSSION. Prior to the ornithological exploration of the
Eastern Highlands P. r. dorsalis had been considered a rare bird, known
only from seven scattered localities. In the Eastern Highlands it is
common, loud, and conspicuous in tall casuarina trees growing on
lawns in European settlements. It may also be found, but more
erratically, in trees at the edge of the forest or trees in native gardens
and villages. Possibly it is a very recent arrival in the Eastern High-
lands and is spreading rapidly; Paran told me that it had settled at
Awande for the first time in 1963. It has also colonized (or is coloniz-
ing) settled areas in the Baliem Valley, at Teleformin, and in the
‘Torricelli Mountains.
The perches of P. rufiventris range from 15 ft above the ground to
the tops of tall casuarinas. I have seen it catching insects on the wing
and also eating a caterpillar.
VOICE. ‘The song is a typical whistled Pachycephala song but
more explosive than that of other New Guinea species. Most songs
end in an explosive, slurred (either up or down), whistled “whoop”
or ““wheep” or “chew”. This is preceded either by a single long note
on constant pitch, by a rapid series of staccato notes on the same pitch
separated by identical brief time intervals, or by a rapid series of
identical slurs. Whichever is the case, this introductory note, notes, or
slurs increase dramatically in volume to terminate in the final ex-
plosive slur. All notes are clear and whistled; there are no unmusical
notes such as occur in the songs of P. hyperythra and P. schlegelii
(Big. 30, p. 2738),
Pachycephala rufinucha niveifrons Hartert
Rufous-naped Whistler
NATIVE NAME. Gimi: kiriferiye.
SPECIMENS EXAMINED. Awande: 1 Q, 1 ? (16 and 19 June 1965). Mt.
1 Listed as P. monacha in Rand and Gilliard (1967).
283
SPECIES ACCOUNTS
Michael: 1 ¢, 1 6 (4 and 8 July 1964). Mt. Karimui Zone 3: 1 @; Zone 4: 1
6 (17 and 30 Aug. 1965).
WEHIGH YT. 2 22 380,425, Trib:
WING. 3 @: 86, 88,90. 2 9: 84 88. I P: 84.
STOMACH CONTENTS. Insects (three stomachs); insects and seeds (two); a
5 cm worm (one). Natives credit this species with the ability to take large prey.
TAXONOMY. These agree with niveifrons in the whitish tore-
head and the olive back which has no tinge of brown. Not even my
most easterly specimens (from Awande) show an approach to gamblei
of southeastern New Guinea. The males show an increase in wing
length with altitude, as also found by the First and the Third Archbold
Expeditions.
BREEDING. Gonads were small.
DISCUSSION. ‘This distinctive Pachycephala, which has a very
heavy bill and long and powerful legs, spends much time on the ground
and in the understory, and five of the six specimens were netted. Its
habitat is forest from 4,800 to at least 8,000 ft.
Pachycephala tenebrosa tenebrosa Rothschild
Sooty Whistler
The only Eastern Highlands record of this rare western New Guinea
Pachycephala is a single female collected by Btirgers on the Schrader-
berg during the German Sepik Expedition.
Colluricincla megarhyncha nea (Mayr) and
C. m. tappenbecki Reichenow!
Rufous Shrike-thrush
NATIVE NAMES. North Fore: kupési. South Fore: kokopilo. Gimi: okopido.
Daribi: hogobiya.
SPECIMENS EXAMINED. Awande: 2 ¢, 3 9 (28 June 1964; 16-20 June 1965).
Okasa: 3 &, 4 9 @l and 23 Aug, 1964 22-26 June 1905). Karimui: 2°64, 1 9 (6
Aug. 1964; 1 July and 3 Aug. 1965). Bomai: 2 ¢,1 @ (6-9 July 1965). Soliabeda:
2 4,1 (22-30 July 1965); Mt. Karmut Zone i: Yo 22> Zone 2a liga 95
Zone 2; 1) 4,0 9 Oral Aus, 1960),
WEIGHT. Awande-Okasa: 5 4, 33-41 (374+2.8); 5 9, 34-40 (36.2 + 2.0).
Karimui-Bomai-Soliabeda-Mt. Karimui: 5 @, 29-41 (35.7 +2.7); 5 9, 30-41 (34.7 +
Sab):
Hache Awande-Okasa: 5 @, 92-100 (98+ 3); 5 9, 87-95 (923 3). Karimui-
Bomai-Soliabeda-Mt. Karimui: 5 ¢, 89-99 (93 + 3); 5 9, 83-96 (90 + 3).
STOMACH CONTENTS. Insects.
TAXONOMY. ‘The series was compared with goodsoni (including
the type) from the Merauke District of southern New Guinea, wuroz
(including the type) from southern New Guinea, palmeri (including
the type) from the Fly River, despectus (including the type) from the
1 Listed as Myiolestes megarhynchus in Rand and Gilliard (1967).
284
SPECIES ACCOUNTS
south coast of southeastern New Guinea, superfluus (including the
type) from the north coast of southeastern New Guinea, nea from the
Herzog Mountains, madaraszi from the Huon Peninsula, and tap-
penbecki from northeastern New Guinea and the Kubor Range of
the Eastern Highlands. My specimens are closest to nea, with which
they agree in the dark gray bill and the gray throat, and from which
they differ only in having slightly darker upperparts and brighter
underparts. The race palmeri from the Fly River is also close but
differs in the buffier and less gray throat, the more straw-colored bill,
and the slightly more distinct breast streaks. Awande-Okasa_ birds
average very slightly brighter olive above, less heavily streaked on the
breast, and less dull or gray below than birds from the Karimui area
when large series are viewed side by side. There is no convincing
change of weight or wing length with altitude in either the Awande-
Okasa area or the Karimui-Bomai-Soliabeda-Mt. Karimui area, but
birds from the former area are slightly heavier and have longer wings
than birds from the latter area. The race goodsoni is much paler and
duller above, particularly on the crown, and much less olive above;
wuro? is paler above and less gray on the chin; despectus is paler above
and below and slightly more olive above; superfluus is paler, duller,
and less ochraceous below; madaraszi is less olive above; tappenbecki
is browner, duller, and less olive above, and the throat and upper
breast are grayer and contrast more with the belly. Birds from the
Wahgi Valley and Mt. Giluwe, collected by Gilliard, Gyldenstolpe,
and Shaw-Mayer, are tappenbechi.
BREEDING. Gonads of most specimens were large at Awande but
small at other localities.
DISCUSSION. Colluricincla megarhyncha is inconspicuous and
solitary but nevertheless common and ubiquitous from sea level up to
a ceiling which varies locally between 5,400 and 6,500 ft. On Mt.
Karimui it dropped out at 5,400 ft, but it was still common around
6,100 ft at Mengino and 6,200 ft at Awande, while Bulmer found it
up to at least 6,500 ft in Kyaka territory. Its habitat is the lower- and
middlestories (to ca. 10-15 ft above the ground) in the forest interior,
at the forest edge, and in second-growth which is sufficiently dense for
the understory to be well shaded. In these habitats it accounts for
about 3-59% of the local avifauna.
In behavior, posture, appearance, and voice C. megarhyncha is
simply a big and sluggish Pachycephala. Some specimens had to be
examined carefully to distinguish them from the dull race of Pachy-
cephala hyperythra living in the Karimui area.
VOICE. ‘The song is the origin of the South Fore and Gimi name
“kokopilo” and consists usually of four notes, the last two of which
are joined in an upslur or downslur (Fig. 31). There is a strong but not
explosive increase in volume in the last two notes. The first note is
approximately a major sixth in pitch above the second. Sometimes an
285
SPECIES ACCOUNTS
extra note is added at the beginning, creating a five-note song. The
total length is about 11% sec, the quality is mellow, and the pitch some-
what lower than that of Pachycephala rufwentris. It was amusing to
hear hidden individuals give their song after every gunshot, a useful
attribute that made it easy to determine the altitudinal range of the
species.
Colluricincla megarhyncha:
<=
,isec ko-ko-pi-lo
Colluricincla harmonica:
- - - - ---- Or - - - - ~s
PrN
r ere OW aaa oe
or Y~/47/7--~\ Of 7 Ty —<
Fic. 31. Voices of Colluricincla megarhyncha and C, harmonica.
Colluricincla harmonica tachycrypta Rothschild and Hartert
Gray Shrike-thrush
DISCUSSION. ‘This large whistler of scattered trees in open coun-
try is common in the lowlands of eastern New Guinea and has recently
been recorded in two Eastern Highlands towns, Goroka and Kainantu
(N-G.BS, Newsletter, INO. 59, p. 1) Oct, A970):
VOICE. A loud, full, mellow, Pachycephala-like song at medium
pitch, beginning with repeated notes or upslurs which are on the same
pitch but crescendo, and usually ending with a downslur (Fig. 31).
Pitohui kirhocephalus brunneiceps (D’Albertis and Salvadori)
Variable Pitohut
NATIVE NAME. Daribi: hua.
SPECIMENS EXAMINED. Karimui: 6 ¢, 1 @ (3 July-4 Aug. 1965). Bomai:
286
SPECIES ACCOUNTS
4 9,1? (6 and 7 July 1965). Soliabeda: 6 ¢,1 9 (22-29 July 1965).
WEIGHT and MEASUREMENTS. See Table 8.
TAXONOMY. Pitohwi kirhocephalus shows more striking geo-
eraphical variation than any other New Guinea bird and encompasses
among its subspecies most of the color patterns occurring in the other
members of the genus. Some races (brunneivertex, jobiensis, and
meyeri) are fairly uniform brown and resemble P. ferrugineus or I,
cristatus. Males of other races (meridionalis, nigripectus) have a black
and maroon pattern like P. dichrous. In addition, most populations,
including mine, show much individual variation. On the average, my
specimens are much closer to a topotypical and uniform series of
brunneiceps collected on the Fly River by the Second Archbold Ex-
pedition than to the 14 other races with which they were compared.
The variation among my specimens concerns six features: (1) ‘The
darkness of the fulvous, somewhat orange color of the breast and belly.
All Soliabeda birds are darker than all except one of the Karimui and
Bomai birds. (2) The darkness of the head, which is dark brown in
most specimens but is nearly black in the Soliabeda specimen labelled
“b” in ‘Table 8. (3) The distinctness of the hood, which depends upon
how marked is the contrast between the throat and breast. In the
Bomai specimen “1”, probably an immature, the throat and breast are
the same color, and there is no hood. The contrast is slight in four
Karimui specimens and one from Soliabeda. In three Soliabeda speci-
mens the throat is nearly as dark as the crown, so that the throat: breast
contrast forms a very distinct hood. (4) ‘The color of the upper tail,
which is a rich brown to olive-brown in some specimens but nearly
black in specimen “b”, while the tail feathers of some other specimens
are blackish around the shaft. (5) The color of the wings, varying from
brown to nearly black. (6) ‘he color of the back, which is a slightly
deeper and richer brown in Soliabeda and Bomai specimens than in
Karimui specimens, which agree in this respect with Fly River brun-
neiceps. ‘Vable 8 scores all my specimens and three related populations
or individuals on the first four of these points. In my specimens there
is no consistent difference between the sexes in these characters. All
my specimens have the bill black.
As summarized in ‘Table 8, most Karimui and Bomai birds are
somewhat paler below and have less distinct hoods than Fly River
brunneiceps, while some Soliabeda birds have darker underparts and
most have darker heads than brunneiceps. In addition, the wings of
Fly River birds average 3-4 mm shorter. ‘These differences are minor
in view of the considerable individual and local variation and the
much more marked differences compared to other races. Males of
meridionalis, the next race to the east (in southeastern New Guinea),
are distinct in the really black hood, wings, and tail, while females
have the head paler than in my series. As seen in Table 8, two males
from Deva Deva collected by Hamlin, which come the closest of the
287
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SPECIES ACCOUNTS
available meridionalis specimens to brunneiceps, are approached by
two of the darkest Soliabeda specimens but still differ in having the
throat tipped with black. Most specimens of nigripectus (south-west
ern New Guinea) have the head and tail even blacker than my spect-
men “b’, and in addition nigripectus males usually have a black
breast. One aberrant nigripectus male collected by Meek on the
southern slope of the Snow Mountains (A.M.N.H. No. 656,061), re
sembles my series and is scored separately in ‘Vable 8.
BREEDING. Gonads were slightly enlarged in seven males but
were small in the remaining males and all females.
DISCUSSION. I found this tropical species common (ca. 2% of
the local avifauna) at Soliabeda, Bomai, and Karimui but nowhere
else, not even in Zone | of Mt. Karimui. While it might be found in
the forest and in dense second-growth at any level from the understory
to the treetops, sometimes in fruiting trees, it preferred dense thickets
near the forest edge, where it kept out of sight. A jumble of calls would
emerge from a thicket, leaves would shake and shapes dart about,
and finally three or four P. kirhocephalus would emerge and chase
each other into a nearby thicket. In flight it had a loud, fluttering
wingbeat, as do P. dichrous and P. nigrescens. Its habits and sometimes
its voice and appearance resemble P. dichrous, of which it may be
considered the low-altitude representative, though there is a broad
overlap.
VOICE. Loud, liquid, and melodious whistles at medium pitch,
somewhat harsher than those of P. dichrous. A common element is
a connected pair of notes, the first note a fourth in pitch below the
second note. A typical song begins with one such pair, followed a
second later by two more such pairs in immediate succession, after
which the song degenerates into an irregular but melodious jumble.
Often two singers will deliver their jumbles simultaneously from
adjacent thickets, one starting after the other. his may involve rival
males singing competitively (countersong). Other instances of one
bird calling in response to another bird’s call occur frequently. A
common call note is an upslurred whistle. Elsewhere in New Guinea
this species sings duets and trios (Diamond and ‘Terborgh, 1968, py 73).
Pitohwi dichrous (Bonaparte)
Black-headed Pitohui
NATIVE NAMES. Fore: obo. Gimi: okofo.
SPECIMENS EXAMINED. Awande: 1 ¢, 1 9 (28 June 1964; 17 June 1965).
Okasa: | Q (26 June 1965). Karimui: 4 ¢,2 9 (12 Aug. 1964; 3 July-11 Sept. 1965).
Soliabeda: 2 Q (28 and 30 July 1965). Mt. Karimui Zone 1: 1 @,1 9; Zone 2: 3 2,
® 9. 1 7: Zone 3; I ¢ G-16 Aug. 1965),
WEIGHT. 10 ¢: 67-76 (72+ 2). 10 9: 67-79 (72 + 4).
WING. 10 ¢: 106-116 (110+ 3). 10 Q: 102-113 (107 + 3).
STOMACH CONTENTS. Fruit.
289
SPECIES ACCOUNTS
BREEDING. ‘The only specimens with enlarged gonads were the
1964 Karimui male, collected 12 Aug., and the last male collected at
Karimui in 1965, on 11 Sept. This may suggest that the breeding sea-
son is toward the end of the calendar year or is suppressed in dry
periods.
DISCUSSION. P. dichrous is the middle bird in an altitudinal
sequence in genus Pilohwi, occurring between the ranges of P. kirho-
cephalus and P. nigrescens. It descends regularly to 1,500 ft and reaches
sea level in a few parts of New Guinea, so that it overlaps the upper
half of P. kirhocephalus’s range. Its ceiling varies between about 5,000
and 6,000 ft, and at this ceiling it excludes P. nigrescens, probably
strictly in undisturbed forest. In the ecologically disturbed forest edge
at Awande both species were netted at approximately the same alti-
tude (6,200 ft). The following figures for the contribution of P. dich-
rous to the avifauna show that it is most numerous in hill forest above
the range of P. kirhocephalus: Soliabeda, 1,350-2,000 ft, 0.3%; Kari-
mui, 3,650 ft, 0.7% (P. kirhocephalus present at both Soliabeda and
Karimui); Mt. Karimui Zone 1, 4,000-4,200 ft, 2% (P. kirhocephalus
absent at this and higher altitudes); Zone 2, 4,400-4,750 ft, 5.69%; Zone
3a, 4,750-5,080 ft, 2.6%; Zone 3b, 5,080-5,150 ft, 0.7%; mot observed
above 5,150 ft.
Like P. kirhocephalus, P. dichrous may be seen at any height in the
vegetational column in the forest or in dense second-growth, from the
understory up to the crowns of tall trees, as solitary individuals or in
groups of three or four. ‘Terborgh observed it in trees with small fruits.
VOICE. Loud and mellow whistles at medium pitch, similar to
those of P. kirhocephalus but with no harsh quality. “wo common
types of vocalizations are: a rapid series of a half dozen identical up-
slurs delivered at the rate of 4 per second; and a song consisting of
whistled notes, upslurs, and downslurs in an interestingly irregular
and hesitating rhythm, and usually beginning with two notes on the
same pitch followed by an upslur (Fig. 32).
Pitohui dichrous:
sid haat ett
| 3 sec }
Fic. 32. Voice of Pitohui dichrous.
290
SPECIES ACCOUNTS
Pitohui nigrescens subsp.
Black Pitohui
NATIVE NAME. Fore: koroli.
SPECIMENS EXAMINED. Awande: 4 4, 1 ? (28 June 1964; 17-20 June 1965).
Mt. Michael: 1 ? (5 July 1964). Mt. Karimui Zone 4: 1 ¢; Zone 5: 3 g@ (27 Aug.-
5 Sept, 1965).
WEIGHT. Awande: 2 @, 73, 78; 1 ?, 78. Mt. Karimui: 4 ¢, 74, 76, 82, 86.
WING. Awande: 4 4, 131, 135, 135, 137; 1 ?, 138. Mt. Karimui: 4 ¢@, 130, 130,
[ody Toe.
STOMACH CONTENTS. Insects (four stomachs), insects and seeds (two), 4 mm
fruit (one).
TAXONOMY. | Both unsexed birds are black and probably males,
though black females are known (Gilliard and LeCroy, 1970, p. 17).
As subspecific characters are based mainly on the female, I cannot
make a racial assignment of this series, which was compared with
males of nominate nigrescens (Vogelkop), wandamensis (Wandammen
Peninsula), meeki (Weyland and Snow Mountains), and schistaceus
(southeastern New Guinea). Males were found to differ in wing length,
hue (blacker or browner), and depth of color (darker or lighter). As
shown in Table 9, males from Awande, lying to the east of Mt. Karti-
mui, have the longest wings, whereas birds from southeastern New
Guinea, still farther to the east, have the shortest wings. Both Awande
and Mt. Karimui birds are more black, less brown, than any other
available for comparison. The possibility of foxing cannot be ex-
cluded, as all other material was at least 28 years old. The races
schistaceus and nominate nigrescens are lighter than meeki or wanda-
mensis. Mt. Karimui birds are comparable to meeki in this respect
and are darker than four of the five Awande specimens, which are in
turn darker than schistaceus. On the basis of color Mayr and Gilliard
assigned their only specimen, a long-winged female from Mt. Hagen,
TABLE 9
WING LENGTH OF Pitohui nigrescens MALES
schistaceus,
southeastern New Guinea 124 (2), 125 (4), 126 (4), 127, 128, 129 mm (av., 125.8)
subsp. ?, Awande lol, 155 (2), 17, 138 (avs, 135.2)
subsp. ?, Mt. Karimui [30)(2), 16, 13d(av., 151.0)
meeki,
Snow Mountains 133
Weyland Mountains 129, 130 (3), 131, 136, 138 (av., 132.0)
wandamensis,
Wandammen Peninsula 132
nigrescens,
Vogelkop 123, 126 (2), 128, 130, 131 (2), 182, 133 (av., 128.9)
291
SPECIES ACCOUNTS
to buergersi, which was described from the Schraderberg but was un-
available for comparison and is doubtfully distinct from meeki. More
females from the Eastern Highlands, including topotypical females
from the Schraderberg, are necessary to assess the validity of buergersi
and the status of the Eastern Highlands population, but it is definitely
closer to meeki than to schistaceus.
BREEDING. Gonads were small in all specimens.
DISCUSSION. PP. nigrescens is an uncommon forest species be-
tween about 5,600 and 8,200 ft, ranging from the understory to about
35 ft above the ground. The few that I saw were solitary and flew
with a loud wing beat like P. kirkocephalus and P. dichrous.
VOICE. An unmusical, buzzy “cher-wee’” repeated two or three
times in succession. The syllable “cher” is at a lower pitch than “wee”.
Also, a musical chirp “kyew’’, repeated eight times at 14 sec intervals.
Pitohui ferrugineus ferrugineus (Bonaparte)
Rusty Pitohui
NATIVE NAME. Daribi: hia.
SPECIMENS EXAMINED. Karimui: 5 ¢, 2 9 (11 and 14 Aug. 1964; 3 July-4
Aug. 1965). Bomai: 3 ¢, 2 9 (6-7 July 1965). Soliabeda: 4 ¢, 4 9 (22-29 July
1965).
etary LO Se SODIO (GS se 8) 97 Oe 77-9990 a5 7).
WINGE LO as 1308S (Al Se DS) 8 Ons 2413 8 (sorte):
STOMACH CONTENTS. Insects and fruit.
TAXONOMY. Specimens from the Karimui area are closest in
depth of color to nominate ferrugineus from the Vogelkop and upper
Fly River, and are also close to birds from the Idenburg River and
southern slope of the Snow Mountains. ‘The race lewcorhynchus is
much darker and has a whitish bill; clarws of southeastern New Guinea
and the lower Fly River is paler; holerythrus of Japen and the Wey-
land Mountains is darker; and brevipennis of the Aru Islands has the
black slightly darker. P. f. ferrugineus thus has a checkerboard distribu-
tion. The bill was black and the iris straw-colored in my specimens.
BREEDING. Gonads were moderately enlarged in one Soliabeda
male, greatly enlarged in another, slightly enlarged in one Karimui
male, and small in other specimens.
DISCUSSION. ‘To date, this low altitude species has been found
in the Eastern Highlands only in the Karimui area and at Lake
Kutubu. Although the number of specimens indicate that it was fairly
common (1.7% of the local avifauna at Soliabeda and Bomai, ORDO 75 al
Karimui), I observed it only infrequently, from the understory to 40
ft above the ground in the forest interior. ‘The altitudinal ranges of
P. ferrugineus and P. kirhocephalus are coincident in the Karimui
area (0-3,650 ft). P. ferrugineus is slightly larger, more sluggish in its
movements, quieter, and more limited to the forest interior, whereas
ye) 4
SPECIES ACCOUNTS
P. kirhocephalus is noisy and often seen at the forest edge and in dense
second-growth as well as in the forest interior.
Pitohui cristatus arthuri Hartert and P. c. kodonophonos Mayr
rested Pitohui
NATIVE NAME. Daribi: sigo (?).
SPECIMENS EXAMINED. Karimui: 1 ? (6 Aug. 1964).
WING. 108.
TAIL. 85.
EXPOSED CULMEN. 24.
TAXONOMY. The short wing, tiny gonads, and rufous edges to
the upperwing coverts suggest that this specimen was an immature.
‘The bill was dark gray.
The Karimui specimen is closest to the type and another topotypical
specimen of arthuri from the Cyclops Mountains and is equally olive
on the back but differs in being generally slightly darker, distinctly
darker on the crown, and more rufous on the uppertail. The race
kodonophonos of southeastern New Guinea, to which Schodde and
Hitchcock referred a Lake Kutubu specimen, approaches the Karimui
specimen in the darkness of the crown but is considerably paler below.
‘The nominate race from the Vogelkop and southern slope of the
Snow Mountains is darker below, more brown and less olive above.
When the crest lies flat, P. cristatus is superficially similar to P. fer-
rugineus but may be distinguished by the rufous uppertail, the dark-
ened breast band, and the brown instead of straw-colored or whitish
iris.
DISCUSSION. Pitohui cristatus is a rare hill forest species known
in the Eastern Highlands only in the Karimui area and at Lake
Kutubu. I heard its suspected song a dozen times in the understory at
Karimui and in hill forest between 2,600 and 4,300 ft.
VOICE. If the statements of several natives on this matter are
correct, then P. cristatus is the author of the most extraordinary bird
song in New Guinea.
‘The song consists of a long series of identical notes which are
initially all on the same pitch and at equal time intervals. The pitch
lies approximately an octave above middle C. One of the two remark-
able features of the song is its length. One song which I timed and
which seemed to be of average length lasted 175 sec without interrup-
tion. As the song proceeds, the notes gradually accelerate and also
gradually drop in pitch. Thus, the rate of delivery was timed at five
notes per second at the start of the song, 13 notes per second at the
end; and the final pitch may be about half an octave below the initial
pitch. ‘he song ends more or less abruptly in the middle of the series,
and may then be given again after a pause.
The other and more remarkable feaure is the unusual, throbbing
293
SPECIES ACCOUNTS
quality, suggestive of either the ‘ ‘singing’ sound produced by high-
voltage transmission lines or else of the sound produced by blowing
across the narrow mouth of a large open vessel. As a result of this
quality, the song blends so well into the surroundings and is so
unlike any other bird song that it does not catch one’s attention,
least of all if one is listening for bird calls. Although the song is muted
and not loud, it carries for long distances of up to half a mile that
are impossible to estimate unless the singer happens to be very close.
Eulacestoma nigropectus De Vis
Wattled Shrike-tit
SPECIMENS EXAMINED. Mt. Karimui Zone 5: 2 ¢; Zone 6: 2 ¢,1 9; Zone
7: 1 @ (28 Aug.-7 Sept. 1965).
WEIGH ET. 5 8 1007, 205,. 2052217, 22:0. 1 2 2L3.
WING. 5 4: 69; 70, 70, 71, 71. 1 9: 72.
STOMACH CONTENTS. Insects.
‘TAXONOMY. ‘The males are blacker and less brown on the
breast than nominate “nigropectus’ from the Mambare River and
Wharton Range or “clara” from Mt. Goliath and the Kubor Moun-
tains, with the exception of one male from Mt. Tafa (Wharton Range),
which is equally black. The female has the underparts yellow-olive
with obscure barring on the throat and breast. A juvenile has the
back and upperwing coverts rufous. With the description of clara
(Stresemann and Paludan, 1935, p. 44), topotypical specimens of clara
from the Weyland Mountains, and southeastern New Guinea birds
before me, I was unable to detect the differences cited in the descrip-
tion of clara, which had already been suspected as being synonymous
with the nominate race by Mayr and Gilliard (1954, p. 354) and Rand
and Gilliard (1967, p. 438).
BREEDING. ‘he gonads of all specimens were small.
DISCUSSION. Eulacestoma nigropectus belongs to a monotypic,
distinctive, poorly-known, endemic New Guinea genus. ‘Though com-
mon in southeastern New Guinea, it is uncommon and local in the
Eastern Highlands at 6,400-8,000 ft, and I met it only on Mt. Karimul.
It obtains its food from limbs and branches, particularly from the
broken-off tips of dry dead branches, between the understory and 30 ft
above the ground. In searching, every few seconds it hops a few inches
(in dense vegetation) up to four feet (on leafless branches) along a
limb, frequently leaning over the limb head downwards and nuthatch-
like to inspect the underside but never hanging upside-down from
the underside. The ends of twigs bend down under its weight, so that
it must be weight limited in its choice of feeding station. Occasionally
it stops to pry, pound, or dig vigorously at a branch, and chips of
wood and pieces of moss fly as it works. I judge its heavy bill to be
extremely powerful, having been bitten by it.
294
SPECIES ACCOUNTS
LANIIDAE;: SHRIKES
Lanius schach stresemanni Mertens
Schach Shrike
NATIVE NAME. Fore: kaito.
SPECIMENS EXAMINED. Awande: 1 ¢,1 9,1 ? (15-20 June 1965).
WEIGHT. I 4746. 1-0: 48. I Py 46.
WING. 1 9: 96.
BREEDING. ‘The male, taken in June, had small gonads, but on
18 Aug. 1964, two nestlings were shown to me at Awande. I was told
that the kato disappeared from the Okapa area during the wet months
of Janury, February, and March. Perhaps it breeds in the dry season,
like many grassland birds, and then undertakes a local migration in
the wet season.
DISCUSSION. ‘The Schach Shrike has a peculiar geographical
range, namely, midmontane grassland of eastern New Guinea as far
west as Telofolmin but not farther west. There is no obvious reason
why it should be missing from the midmontane grassland of the Baliem
Valley of western New Guinea. It is fairly widespread in the Eastern
Highlands (but is absent at Karimui) and perches on trees and fences
in grassland, gardens, and villages, quickly flicking its tail as it waits
to pounce.
VOICE. According to the Fore, this shrike mimics the voices of
other birds.
ARTAMIDAE: WOOD SWALLOWS
Artamus maximus Meyer
Greater Wood Swallow
NATIVE NAMES. Fore: éseyabi. Daribi: tamani.
SPECIMENS EXAMINED. Mengino: 1 6,2 9 (14 July 1964). Karimui: 4 ¢,
2 Q (10-15 Aug. 1964; 16 July-4 Aug. 1965). Bomai: 2 ¢, 1 9 (6-7 July 1965).
Soliabeda: 3 ¢,4 Q (23-29 July 1965).
WEIGHT. 7 4: 54-64 (59+ 4). 5 Q: 52-59 (57 + 3).
WING. 7 @: 152-163 (158+ 4). 7 9: 154-164 (158 + 4).
STOMACH CONTENTS. Medium-sized insects.
TAXONOMY. ‘The wing averages shorter and the weight averages
lower than in a series from the Wahgi Valley and other parts of New
Guinea, perhaps because of the low altitude. The race wahgiensis de-
scribed by Gyldenstolpe is invalid, as pointed out by Sims (U95G,, Po
422) and by Gilliard and LeCroy (1961, p. 63).
BREEDING. Gonads were small in all specimens except for the
latest one collected, a male taken at Karimui on 15 Aug. 1964, which
had greatly enlarged testes.
DISCUSSION. A. maximus was present at all my Eastern High-
295
SPECIES ACCOUNTS
lands stations that offered some cleared land, up to 8,500 ft. Its original
habitat was hill forest and midmontane forest, and one still sees it
occasionally at an exposed perch on a protruding branch or at the top
of a dead tree in primary forest. It is far more common, however, in
settled areas and must have enjoyed a great increase in population
since the advent of agriculture. It is usually seen soaring without
flapping, sometimes quite high, or else perched on a branch in tightly
packed rows of two to six birds, each apparently touching its neighbors.
VOICE. The call of A. maximus is a pleasant chirp which I can-
not distinguish from the call of its low-altitude equivalent, A. leuco-
rhynchus. ‘The rarely heard song, delivered perched from the crown of
a tall tree, is a prolonged soft jumble including imitations of other
birds and very similar to the songs of New World mockingbirds
(Mimidae). Elements include squawks, chirps, and short trills, each
repeated 2-5 times.
SL URNIDAE? STAREINGS
Aplonis cantoroides (Gray)
Singing Starling
STOMACH CONTENTS. Fruit.
DISCUSSION. Gilliard and Gyldenstolpe were surprised inde-
pendently to find large flocks of this lowland species at Nondugl
(5,200 ft), evidently in the same grove of trees but in different years.
Specimens taken in April and May were not in breeding condition,
while those taken in late August and September were entering breeding
condition and beginning courtship activities. I observed groups of
four at Goroka on 14 Sept. 1966 and 31 July 1969. According to Bell
(pers. comm.), this starling breeds in the vicinity of Port Moresby in
the rainy season (January-May). Perhaps A. cantoroides is undertaking
postbreeding migrations from the lowlands to the Eastern Highlands
in increasing numbers.
VOICE. A clear, sweet downslur.
Aplonis metallica metallica (lemminck)
Metallic Starling
STOMACH CONTENTS. Fruit up to 12 mm in diameter; rarely, insects.
DISCUSSION. Shaw-Mayer collected a female of this lowland
starling in May 1951, at 7,000 ft on Mt. Giluwe and found it plentiful
there at 5,300 ft. No other Eastern Highlands records are known to me.
VOICE. A harsh downslur readily distinguished from the call of
A. cantoroides.
296
SPECIES ACCOUNTS
Mino dwmnontit dumonti Lesson
Yellow-faced Myna
NATIVE NAMES. Fore: yatta. Gimi: kaho.
SPECIMENS EXAMINED. Karimui: 5 g,8 9? (3 July-5 Aug. 1965). Soliabeda:
1 4,2 2 (25-27 July 1965).
WEIGHT. 4 4: 197, 198, 209; 235. 10 9: 140-198 (176 = 21),
WING. 6 @: 146-157 (152 +4). 10 9: 136-153 (145 + 5).
STOMACH CONTENTS. Fruit, 6-19 mm in diameter.
TAXONOMY. The iridescent color is green in eight females, pur-
ple in two, and green in two males, blue-green in two, and purple in
two. Differences in the iridescent color were among the characters used
to separate a north coast race violaceus, but Mayr and Rand (1937,
p. 187) and Junge (1939, p. 72) showed that the various colors can
all be found in a single population (as also true at Karimui) and prob-
ably change with wear.
BREEDING. ‘The gonads were small in all specimens.
DISCUSSION. ‘This lowland species has been found in the Eastern
Highlands at Lake Kutubu and in the Karimui area (Soliabeda,
Bomai, and Karimui). At Karimui (3,650 ft) it was fairly common
(1.2-1.5% of the local avifauna), and its presence there constitutes an
altitudinal record. It was usually seen in small groups of several (up
to four) birds, and often in pairs, on conspicuous perches in the tops
of tall trees (either leafy or dead) at the edge of forest and in second-
growth. ‘Terborgh found up to 10 M. dwmontii eating small fruits in
a strangling fig.
VOICE. A disyllabic, low-pitched, penetrating, totally unmusical
croak, with the second syllable on the lower pitch. ‘There are also
several other equally weird and unmusical notes. If one hears what
sounds like a bullfrog at the top of a tall tree, either M. dumontii or
Eurystomus orientalis is in the vicinity.
ORIOLIDAE: ORIOLES
Oriolus szalayi (Madarasz)
New Guinea Oriole
NATIVE NAMES. Gimi: itakuréfo. Daribi: unandburo.
SPECIMENS EXAMINED. Okasa: 3 @ (red bill); 1 ¢, 1 9 (black bill) (22-26
June 1965). Karimui: 4 ¢, 2 Q (red bill); 4 ¢, 1 @ (black bill) (12 and 15 Aug.
1964; 1-18 July 1965). Bomai: 1 g (red bill); 1 Q (black bill) (22 July 1965).
Soliabeda: 1 @ (red bill); 1 @ (black bill) (22 July 1965).
WEIGHT. 8 ¢: 91, 92, 93, 95, 98, 98, 101, 106; 2 9: 101, 115 (red bill). 6 @:
79, 84, 90, 95, 99, 100; 2 9: 88, 91 (black bill).
WING. 10 ¢: 134 (2), 136, 137, 138 (2), 140 (2), 142, 144; 2 9: 138 (2) (red bill).
a7: 123, 129; 132, Uso, las, 1355.5 Oe 129; 129; 132 (black bill),
STOMACH CONTENTS. Fruit.
297
SPECIES ACCOUNTS
TAXONOMY. The bill is either black or else very dull blood-red.
Because the wing is longer in all females with red bills than in any with
a black bill, and is also longer in all but two of the red-billed ma_es
than in any back-billed male, I presume that a black bill is a sign of im-
maturity. Three of the red-billed males have the underparts buffer
than the rest.
BREEDING. The only specimen with enlarged gonads was a red-
billed male from Okasa.
DISCUSSION. Oriolus szalayi is present in many ecologically dis-
turbed areas at the edge of the Eastern Highlands up to about 4,000
or 5,000 ft and is particularly common at Karimui, but occurs in
primary forest only at much lower altitudes. Its habitat is the upper-
story and middlestory of tall trees at the edge of the forest, in second-
growth, gardens, and in partly cut forest, i.e., wherever it finds perches
in tall trees adjacent to open spaces. Usually several individuals were
seen together.
Plumage convergence of Philemon novaeguineae and O. szalayi is
discussed on pp. 382-383.
VOICE. ‘There are three kinds of vocalizations. The commonest is
the song, a liquid, medium-pitched, fairly loud, run-together series of
pairs of notes or slurs. ‘he second note is the lower note in the first
few pairs but is usually the higher note in the last pair (Fig. 33). ‘The
second vocalization is a call with the same liquid quality, consisting of
a single note which is initially slurred downward and then upward,
Oriolus szalayi:
E | or L
Or
—
Fic. 33. Voice of Oriotus szalayt.
298
SPECIES ACCOUNTS
and at the same time undergoes a crescendo and decrescendo, with the
maximum volume on the lowest pitch (Fig. 33). The third call is a
single harsh, hoarse, downslurred “chew”.
GRALLINIDAE: MUDNEST BUILDERS
Grallina bruijnt Salvadori!
Torrent Lark
NATIVE NAMES. Fore: nintere. Gimi: neterewiya. Daribi: sénili.
SPECIMENS EXAMINED. Awande: 1 Q, 1 ? (18 and 19 Aug. 1964). Sena River:
1 4,1 ? (26 and 28 July 1964). Mt. Karimui Zone 3: 1 ¢ (17 Aug. 1965).
WEIGHT. 1 4: 38.4.
WING. 2°42 102; 107. 1 oe 200:
TAXONOMY. I follow Amadon (1950) in considering the ‘Torrent
Lark congeneric with the Magpie Lark, Grallina cyanoleuca, of Aus-
tralia, rather than meriting recognition in the monotypic genus
Pomareopsis.
Both unsexed birds were in female plumage.
DISCUSSION. ‘This is one of the two strictly riparian passerines
in New Guinea, the other being the less uncommon Monachella muel-
leriana. Both species require rushing, rocky, mountain torrents, may
often be found on the same stretch of the same stream, and are
absent from the flat lowlands. I found G. bruijni in these montane
streamside habitats from 2,700 to 5,150 ft, and Shaw-Mayer and Bulmer
observed it at 6,500 ft or higher. A male with small testes was shot
by one of my assistants at 5,150 ft in forest on the crest of a ridge of
Mt. Karimui and many hundred feet above the streams on either side
of the ridge, a habitat in which I never saw G. bruijni myself and
where my assistant was equally surprised to find it. The bird was
probably crossing the ridge from one stream to the other.
G. bruijni is a striking and active bird which alights on boulders in
torrents and walks (not hops) up the sides of or along the boulders,
wagging its seemingly loosely-hinged tail from side to side. It spins
about, turns, and displays like a fantail (Rhipidura). Once while I
was watching a group of three that had alighted on a large boulder,
one bird faced the other two, reared up on its legs, raised its head
and body towards the vertical position, and opened its wings wide to
show to its partners the striking black and white pattern of the under-
parts. The sexes differ markedly with respect to this pattern.
VOICE. ‘The call is a penetrating, buzzy, harsh upslur or upwards-
inflected squawk, quite different from the sweet, high-pitched note of
Monachella muelleriana.
1 Listed as Pomareopsis bruijni in Rand and Gilliard (1967).
299
SPECIES ACCOUNTS
CRACTICIDAE: BUTCHERBIRDS and
BELL-MAGPIES
Cracticus cassicus cassicus (Boddaert)
Black-headed Butcherbird
NATIVE NAMES. Gimi: koiki. Daribi: hdérabe.
SPECIMENS EXAMINED. Karimui: 1 ¢, 2 9, 1 imm. 4, 3 juv. ? (12-16 Aug.
1964; 12 July 1965). Bomai: 1 ¢,1 9 (6 and 9 July 1965). Soliabeda: 3 4, 3 9
(23-30 July 1965).
WEIGHT. 38 4: 140, 145, 155. 4 9: 130, 136, 141, 147.
WING. 5 @: 159, 162, 164 (2), 166, 3 02 156, 158) 159°
STOMACH CONTENTS. Fruit and insects.
TAXONOMY. ‘The immature male from Karimui has some brown
striations on the black of the head and back.
BREEDING. In 1964 at Karimui two of the three sexed adults
had greatly enlarged gonads. On 13 Aug. 1964 I found a nest contain-
ing one nestling, located on a horizontal tree branch 40 ft above the
ground in second-growth forest and constructed of sticks. On the
same day a native brought in two nestlings from another nest. ‘Testes
were greatly enlarged in the Bomai male and in one of the three
Soliabeda males, small in the two others. Evidently breeding is con-
centrated in the dry season.
DISCUSSION. ‘The preferred habitat is the crowns of trees bor-
dering open spaces (never the interior of primary forest), so that this
butcherbird is much less common in undisturbed forested regions
than in areas occupied by man. In the areas of primary forest at
Bomai and Soliabeda small groups of C. cassicus concentrated in river
gorges at the forest edge. The black and white pattern of the upper-
parts as it flew across the gorges, and the loud calls echoing back and
forth across the gorge, made it conspicuous. Other Eastern Highlands
records are from the Schrader Range, up to 4,500 ft in cultivated
areas of Kyaka territory (Bulmer), at 3,500 ft in Baityer Valley (N.G.
B.S. Newsletter, No. 54, p. 2, June 1970), and Lake Kutubu (2,700 ft).
All these locality records in the Eastern Highlands are rather high for
C. cassicus, which seldom ranges above 2,000 ft elsewhere in New
Guinea.
VOICE. A loud, jumbled bugling and yodeling, which combines
bell-like notes and liquid and rollicking phrases with hoarse notes,
eurgles, and musical croaks. The song of the Australian cracticid
Gymnorhina tibicen is somewhat similar, and the only New Guinea
song with a comparable volume and combination of different quali-
ties is that of Cracticus quoyi, which differs in having a more consis-
tent and organized pattern. Once I heard a particularly long (several
minutes in duration) and varied jumble, including imitations of other
birds, in typical C. cassicus habitat (edge of a gorge) and with the
quality of the C. cassicus song, so that I think it probably came from C.
300
SPECIES ACCOUNTS
cassicus even though I did not see the singer. Among the species
imitated were Cracticus quoyi, Paradisaea vaggiana, and Meliphaga
spp. The closely related Cracticus nigrogularis of Australia is known
to be a mimic.
Cracticus quoyt quoyi (Lesson)
Black Butcherbird
NATIVE NAME. Daribi: gauwadi.
SPECIMENS EXAMINED. Karimui: 4 ¢, 3 @, 2 ? (6-14 Aug. 1964; 1 July-6
Aug. 1965). Bomai: 1 @ (8 July 1965). Soliabeda: 3 Q (26-29 July 1965).
WEIGHT. 3°22: 158, 104, 166; 5.20%, 131, 145, dol, 16k 170.
WING, 5 65 lop, 170) (2), 172, 078, (6°92 158, 166, los, FAO), AZ/AL, Whesiil
TAXONOMY. ‘These specimens agree with the widespread New
Guinea race C. g. guoy? in the short wing and in the breadth of the
bill at the base as viewed from above. ‘The Australian race spaldingi
reaches the Aru Islands, and possibly the Digul River and Fly River
bulge of southern New Guinea (Mees, 1964, p. 32). Other Eastern High-
lands records are from the Jimmi Valley of the northern watershed
(Gyldenstolpe) and from Lake Kutubu (Schodde and Hitchcock).
BREEDING. ‘The gonads were very small in all specimens of C.
quoyt, whereas C. cassicus was breeding at this time (August 1964,
July-August 1965).
DISCUSSION. C. quoyi differs ecologically from C. cassicus in
being (1) found in the forest interior, (2) solitary, (3) much less com-
mon in most parts of New Guinea, and in (4) spending much of its
time in the lower story. ‘he latter difference is illustrated by netting
results: C. cassicus was never netted, whereas four of the 13 specimens
of C. quoyi were collected with nets.
C. quoyi was more common and conspicuous at Karimui than at
any other locality in New Guinea I have visited. Occasionally an in-
dividual would come out of the forest, perch up to 20 ft above the
ground in a roadside tree, and give its distinctive call, to be answered
by a couple of other individuals in the distance. More often it skulked
and was glimpsed only briefly as it flew through the undergrowth a
few feet above the ground. On Mt. Karimui it was heard occasionally
in Zones | and 2 between 4,030 and 4,370 ft; this probably represents
its altitudinal ceiling.
VOICE. The call and the song are equally loud, ringing, and
distinctive and carry considerable distances. ‘The call consists of four
notes within a time span of | sec (Fig. 34). The first of these four
notes is a hoarse upslur, while the second, third, and fourth notes are
ringing. Ihe time interval between the first and second notes is twice
as long as the other intervals. ‘The second and fourth notes are on the
same pitch, and near the pitch of the first note, while the third is a
fourth lower in pitch. ‘he bold and ringing song has a striking rhythm
301
SPECIES ACCOUNTS
and interval pattern (Fig. 34). It begins with three notes in a dotted
rhythm, followed by a series of three to six three-note phrases. The
second note of all the phrases is on the same pitch, and the third note
is always on another pitch a third below the second note. The first note
of the phrase is alternately a third and a fourth higher than the second
note. ‘he time interval between the first and second note equals that
between the second and third note when the pitch interval is a third,
and is twice as long when the pitch interval is a fourth. The whole
song lasts 5 sec. While these descriptions may sound complicated, both
the call and the song are in practice “catchy”, unmistakable, and
consistent, and may be made clearer by Figure 34. An additional call
consists of pairs or longer series of hoarse, harsh, and more or less
musical upslurred croaks.
Cracticus quoyi:
cal: 7 -~
song:
| 5 sec
Fic. 34. Voice of Cracticus quoyi.
DICRURIDAE: DRONGOS
Dicrurus hottentottus carbonarius Bonaparte
Spangled Drongo
NATIVE NAME. Daribi: sigiyarasare.
SPECIMENS EXAMINED. Karimui: 4 ¢ (7 Aug. 1964; 18 July-5 Aug. 1965).
Bomai: 1 ¢,3 2 (5-9 July 1965). Soliabeda: 2 9 (23 and 24 July 1965).
WEIGHT. 3 4: 76, 80, 80. 4 9: 64, 65, 68, 73.
WING. 5 4: 145, 145, 153, 154, 155. 4 Q: 138, 142, 143, 143.
EXPOSED CULMEN TO EDGE OF FEATHERING ON DORSAL MIDLINE.
5 f: 18, 21, 22, 23, 27. 4 9: 20, 22, 22, 24,
302
SPECIES ACCOUNTS
CULMEN FROM BASE. 4 4: 30, 32, 33, 35. 4 9: 31, 31, 32, 33.
STOMACH CONTENTS. Insects.
TAXONOMY. The specimens all belong to the resident New
Guinea race carbonarius, not to the Australian race (possibly a separate
species) bracteatus, which winters and possibly may breed (Rand,
1938c) in southern New Guinea. D. h. bracteatus has a green instead
of purple gloss and has less of the bill covered by feathering at the
base than does carbonarius. My measurements of the exposed culmen
give: carbonarius (Fly River and Idenburg River, collected by the
Second and the Third Archbold Expeditions), $ 22-25, 9, 20-22;
bracteatus (Fly River, collected by the Second Archbold Expedition,
and Cape York Peninsula of Australia), 4 96-33: @ 27-30. For the
whole culmen from the base: carbonarius (Fly River), g 31-33, 9 3()-
32; bracteatus (Fly River and Cape York), $ 33-38, 9 32-34 mm.
BREEDING. One Bomai female had enlarged ovaries, but gonads
in all other specimens were small.
DISCUSSION. In the Eastern Highlands the Spangled Drongo has
been recorded from the Karimui area (to 3,650 ft), and from 3,500 ft
in the Batyer Valley (N.G.B.S. Newsletter, No. 54, p. 2, June 1970);
where it 1s above its normal altitudinal range, and also at Lake Kutubu
(2,450 ft). In habits it was solitary and to be found in the middlestory
of trees at the edge of the forest and in second-growth but not (or
rarely) in the forest interior. It held its perches for relatively long
times, frequently flaring its tail and flicking the tail with each call.
VOICE. ‘The call consists typically of three ringing disyllabic notes
‘“‘k-sing”’, the second and third notes on the same pitch and the first on
a higher pitch. ‘The first syllable is very brief, virtually a grace note to
the second syllable, and has a sharp and explosive attack. The longer
second syllable is more nasal. Each note lasts not quite 1% sec. This
call is surprisingly similar to the usual call of the parrot Geoffroyus
geoffroyi and sometimes not safely distinguishable from it. Other calls
are a rapid series of nasal notes on the same pitch, and notes with a
strange quality as a siren or creaking door.
Chaetorhynchus papuensis Meyer
Mountain Drongo
SPECIMENS EXAMINED. Okasa: 2 ¢,2 92 (22-26 June 1965). Sena River: 1 9,
(6 July 1964). Karimui: 3 ¢,1 9, 1 imm. ? (1 July-5 Aug. 1965). Bomai: 1 4,
3 2 (6-8 July 1965). Soliabeda: 1 9, 1 imm. ? (22 July 1965). Mt. Karimui Zone 1:
ea, 9; Lone 27) ¢ 5 Zone 3: 1.4 (9-16 Ame, 1965).
WEIGH. 10 @% 30-45: (42 2-2), 10 GO: 27-39 (86 54). Lamm. Py 84.
WING. 10 4: 109-122 (115 + 3), 10 9: 102-110 (106 + 2).
TAXONOMY. The iris is brown, whereas it is orange in Dicrurus
hottentottus, the other drongo in New Guinea. Most but not all speci-
mens have a well-concealed white patch at the base of the scapulars.
303
SPECIES ACCOUNTS
BREEDING. Gonads of all specimens were small, often very small.
DISCUSSION. Chaetorhynchus papuensis, a monotypic genus en-
demic to New Guinea, is a solitary and inconspicuous but ubiquitous
species confined to the interior of hill forest. It reached its maximum
abundance in Zone | of Mt. Karimui (4,000-4,200 ft, 3.59% of the local
avifauna). At Soliabeda (2,000 ft), which is probably at the lower limit
of its altitudinal range, it was quite uncommon, and its ceiling on
Mt. Karimui was around 4,800 ft. There is no tendency for individuals
to stray above the ceiling in this species. As usual, only or mainly males
are present near the ceiling, while the lowest stations (Soliabeda and
Bomai) had mainly females and immatures. I generally found C.
papuensis perched stationary and upright, like a large flycatcher, on a
horizontal branch in the understory or up to 40 ft above the ground.
On one occasion I saw the usually-concealed white scapular patch on
a bird in the field, suggesting that it may be used for display or
recognition purposes.
VOICE. Whe calls are a strong, explosive “pik” or “peep, given
either once or five to seven times in rapid-fire succession; and a down-
slurred, whistled “‘pi-yew’’. ‘The song is a loud, prolonged jumble.
CORVIDAE GROWS
Corvus tristts Lesson and Garnot!
Bare-faced Crow
NATIVE NAMES. Fore: kai. Gimi: lata. Daribi: tadi.
SPECIMENS EXAMINED. Okasa: 1 ¢ (a) (22 June 1965). Karimui: 3 ¢ (b, ¢,
d), 2 @ (e, f) (15 July-5 Aug. 1965). Bomai: 2 9 (g, h) (7 and 9 July 1965).
WING. 4 2: 320(b), 327 (a), 338 (c), 352 (d). 3 Q: 325 (e), 334 (£), 334 (2).
TAXONOMY. Whe amales “a, ~e>, and dl znecolored) dark
brown, glossy above, and had large or greatly enlarged testes. ‘he
female ‘‘g” is slightly less dark and glossy. The male “b”, which had
small testes, and the females “e’ and “f’, are paler brown with little
or no gloss. In each sex the dark birds have longer wings than the pale
birds. Taken alone, this might suggest that darkness is correlated with
age, but other larger series taken by other collectors have suggested
that it is a matter of individual variation.
DISCUSSION. Up to 5,000 ft, rarely higher, this crow is present
in low numbers in most forested areas. I met it in flocks of four to six
individuals, perched in trees in the forest or else flying slowly near the
forest edge. Several sightings were in river gorges. ‘Lerborgh found it
feeding on small fruits in a strangling fig at Okasa.
VOICE. A hollow, short, and nasal “ka”, rather weak and high-
pitched for a crow.
l Listed as Gymnocorvus tristis in Rand and Gilliard (1967).
504
SPECIES ACCOUNTS
PARADISAFIDAE: BIRDS OF PARADISE
Generic Revision of the Paradisaeidae
The family Paradisaeidae consists of 25 species or superspecies. Vir-
tually all recent classifications, e.g., the ones by Mayr (1962) in Peters’
Check-list, Vol. 15, and in the book by Gilliard (1969), distribute these
25 species and superspecies among 20 (sometimes 21) genera, of which
11 are monotypic in the strictest sense, six are monotypic in the sense
of consisting of a single superspecies, and only three (15% of the
genera) contain more than a single species or superspecies. While this
degree of generic splitting and this low species-to-genus ratio (1.25) is
unparalleled among bird families of comparable size, there are even
more finely split alternative classifications of the Paradisaeidae, such
as that of Iredale (1948), who lists many subspecies and hybrids as
separate species and genera and thereby arrives at 40 genera.
These classifications deprive the genus taxon of meaning. While the
species taxon corresponds to a biological reality, namely, a group of
interbreeding populations, the genus taxon and other higher categories
are arbitrary groupings erected for taxonomic convenience in indicat-
ing relationships. ‘The higher categories are based on a_ biological
reality insofar as the distances between species of a phylogenetic tree
are not equal. However, these interspecies distances vary continuously,
while the number of levels of higher categories is finite and small.
Hence delineation of higher categories always involves some degree of
arbitrariness and is dictated by practical considerations (see Mayr,
1942, pp. 275-291, especially his Fig. 29, for further discussion). A
monotypic genus tells nothing about the relationship of the species in
it, and the adding of the genus name to the species name conveys no
new information. A practical justification for a monotypic genus
nevertheless arises in two cases: if the affinities of a species are un-
known; or if a species differs much more markedly in several inde-
pendent characters from its relatives than its relatives differ from each
other.
On neither of these two grounds is the recognition of many mono-
typic genera among the Paradisaeidae justified. ‘There are only three
species in the family about whose affinities not enough is known to
place them near some other species. Except in respect to one class of
characters (the ornamental plumes of adult males), most paradisaeid
species fall into compact groups. Females and immature males of dif-
ferent “genera” are often so similar that even experts such as Strese-
mann have misidentified specimens as to “genus” (cf. Lophorina
superba pseudoparotia, p. 312, and the nest attributed to Loria loriae,
p. 000). Intergeneric hybrids are known from 10 of the 20 ‘general’.
The historical reason for the high degree of generic splitting in the
Paradisaeidae has been the ornamental feathers of adult males, which
are more bizarre and diverse than in any other family. ‘hese orna-
305
SPECIES ACCOUNTS
ments include wire-like feathers growing out of the tail or head,
central rectrices several times as long as the rest of the bird, scalloped
occipital plumes twice as long as the rest of the bird, wattles, dorsal
capes, breast shields, and pectoral fans. However, these ornaments are
poor generic characters, because species whose adult males appear at
first glance so drastically dissimilar as to suggest they belong to dif-
ferent families may prove to be closely related in most other respects
(e.g., Crcinnurus regius and Diphyllodes magnificus; Lophorina superba
and the Parotia superspecies). Marked variation in these ornaments
occurs even within the same superspecies and was used by Iredale to
place members of the same superspecies in different genera. For in-
stance, Diphyllodes respublica has a bare crown of naked blue skin,
D. magnificus does not; Parotia wahnesi has a tail twice as long as the
other three members of its superspecies; and Paradigalla carunculata
has a tail three times as long as that of Paradigalla brevicauda. Adult
male displays can be equally divergent. For instance, Paradisaea
rudolphi and P. guilielmi display upside-down, which the other Para-
disaea forms do infrequently or never; Parotia constructs dance areas
on the ground, while the closely related Lophorina displays in trees;
and Diphyllodes similarly constructs ground courts, unlike the arbo-
really displaying but related Cicinnurus.
A generic classification of the Paradisaeidae that indicated relation-
ships more effectively than the oversplit one in current use would be
desirable for several reasons. No New Guinea bird family offers such
clear and numerous examples of intermediate stages in speciation. As
noted by Bock (1963, p. 100), the Paradisaeidae “‘could provide an
excellent example of adaptive radiation in feeding habits’, but this
has been “one of the least discussed features of the birds of para-
dise. .. .” They could also illustrate clearly the types of niche differ-
ences which make coexistence among closely related species possible.
The parallel geographic variation in females of Lophorina superba
and the Parotia superspecies is remarkable but has never been dis-
cussed. ‘The acceptance of predominantly monotypic genera has been
an important reason why the illuminating examples provided by birds
of paradise with respect to these ecological and evolutionary questions
of general interest have not received more attention. A prerequisite
for exploitation of the Paradisaeidae in this sense is a better under-
standing of relationships among paradisaeid species, as expressed in
generic grouping.
An adequate generic classification should be broadly based rather
than relying on single characters. In the following tentative revision
I have taken into account the following types of characters: female
plumage; similarities rather than differences in adult male plumage;
the anatomical studies of Bock (1963) and Stonor (1938); altitudinal
range, voice, and observations of behavior and diet derived from my
field experience of 17 of the 20 currently recognized genera; geo-
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SPECIES ACCOUNTS
graphical distribution; hybridization; and displays. Preliminary efforts
also to draw conclusions from eggs were not productive because the
variation within the whole family seemed not much greater than the
variation within one genus or superspecies. The point of considering
altitudinal range is that, in other New. Guinea families, species with
mutually exclusive altitudinal ranges and similar habits almost always
prove to be closely related and congeneric (p. 27). Jt is explicitly
assumed that no differences in male ornamental plumage, no matter
how exaggerated, should disqualify two birds of paradise from being
considered congeneric if this is the conclusion indicated by other kinds
of evidence. In this family in which morphological divergence has
spectacularly outpaced the development of reproductive isolation, it
is important to realize that one must accept genera exhibiting a much
greater range of morphological variation among adult males than in
morphologically more conservative families.
The starting point for this revision is Bock’s (1963) discovery that
the Paradisaeidae can be divided into two or three groups by anatom1-
cal criteria: the subfamily Cnemophilinae, consisting of the “genera”
Cnemophilus, Loria, and Loboparadisea; the subfamily Paradisaeinae,
with 16 of the 17 remaining ‘genera’; and the monotypic genus Mac-
gregoria, which appears in several respects intermediate. I consider in
turn the following problems: (1) The relation between Manucodia
and Phonygammus. (2) ‘The relation between Epimachus and Drepa-
nornis. (3) The relation between Cicinnurus and Diphyllodes. (4) ‘The
relations between Lophorina, Parotia, and Ptiloris. (5) ‘The affinities
of groups 3 and 4 to each other, and of Astrapia, Pteridophora, and
Seleucidis. (6) ‘The affinities of the remaining groups of Paradisaeinae
(Paradisaea, Paradigalla, Semioptera, Lycocorax, and Macgregoria).
(7) Evaluation of genera of Paradisaeinae. (8) ‘The relations among the
three genera of the Cnemophilinae.
1. Relation between Manucodia and Phonygammus.
The birds of paradise, popularly known as manucodes, comprise four
closely related species or superspecies, currently called M. chalybatus
and M. comri (probably constituting a superspecies), Manucodia ater,
M. jobiensis, and Phonygammus keraudrenii. Apparently all are
monogamous and form stable pair bonds, unlike most other birds of
paradise. Correlated with this, no hybrids involving these species are
known. All are medium-sized (wing 160-240 mm), nearly uniform
glossy black, and sexually monomorphic, and have heavy, crow-like
bills. All live in the middle- and upperstories of the forest. Probably
all have a peculiarity called a convoluted trachea, namely, a loop of
windpipe in adults extending onto the breast between the skin and
muscles, although the erroneous assumption is sometimes made that
this is peculiar to keraudrenit. Definitive information about the
presence of a convoluted trachea seems to be unavailable for the semi-
307
SPECIES ACCOUNTS
species comrii, but I have dissected specimens of chalybatus, jobiensis,
and keraudreni with convoluted tracheas, and van Oort (1909, p.
105) published a drawing of the convoluted trachea of ater. ‘The
description of the call of jobiensis by Ogilvie-Grant (1915, p. 9) is
similar to the calls I have heard from keraudrenii, and Rand (1938d,
p- 1) suggested similarity of the call of jobiensis to that of ater.
In the areas where I have collected, there has been a_ three-fold
altitudinal sequence, with keraudreni at the highest elevations,
chalybatus at medium elevations, and ater and jobiensis sharing the
lowest elevations. Among the many normally montane species of New
Guinea birds that descend to sea level at the mouth of the Fly River
are chalybatus and keraudrenii. ‘The five forms differ in the adult
mainly in some details of the ornamentation, which is little developed
in manucodes compared to most other paradisaeids. Perhaps the most
distinct form is comrii, which has twisted central tail feathers, a raised
mat of feathers on the crown, and very curled and crinkled feathering
of the neck and underparts. Crinkling is present but less marked in
chalybatus and jobiensis, which are difficult to distinguish even as.
adults. In keraudreni there are elongated head and neck feathers and
a tuft of feathers on the crown whose length varies subspecifically,
being longest in the races hunsteint and in jamesiz of the Eastern
Highlands and shortest in the race neumanni and the nominate race.
‘The breast feathers of ater are slightly curled and pointed. Immatures
and subadults of all species are very similar and have occasionally
been misidentified. “The most similar species are chalybatus and its
lowiand sibiing jobiensis, which appears to be the product of a recent
speciation and is still largely confined to the north coast of New
Guinea but has had its whole geographical range reinvaded by chaly-
batus, the altitudinal ranges of the two species being mutually ex-
clusive.
The monotypic genus Phonygammus has long been accepted for the
species keraudreni. No other currently used paradisaeid genus seems
to me so devoid of justification. ‘he manucodes form a compact group
very similar to each other and distinct from other birds of paradise,
and constitute an ideal genus. The head tufts of Manucodia keraudreni
are greatly reduced in some subspecies, could not be considered a
character of generic value even if long and constant, and are much less
distinctive than the ornamental feathering of the semispecies M. comyrit.
Even such keen observers of birds of paradise as my Daribi native
assistants used the same name, “pagonabo”, for M. keraudreni and
the other species of manucode in their native valley, M. chalybatus.
2, Epimachus and Drepanornis.
Eleven species or superspecies of the Paradisaeinae (10 currently
recognized genera) constitute a related group eight of whose members
308
SPECIES ACCOUNTS
hybridize with at least one and sometimes with as many as four other
members of this group. The females are all brown dorsally, with bars
or else markings derived from bars ventrally. The males share to vary-
ing degrees the features of a glossy black coloration, a soft velvet
texture to the dorsal feathers due to a distinctive feather structure, and
a green iridescent crown. This group consists of the “genera” Epi-
machus, Drepanornis, Lophorina, Parotia, Ptiloris, Diphyllodes, Cicin-
nurus, Astrapia, Pteridophora, and Seleucidis. Within this association
the most specialized and strikingly distinct forms as regards a func-
tionally significant character are the related genera Epimachus and
Drepanornis.
The genus Epimachus consists of two similar species, £. fastosus of
western New Guinea and E. meyeri of eastern New Guinea. ‘They are
sympatric in most of central New Guinea with mutually exclusive
altitudinal ranges (E. fastosus, ca. 4,500-7,500 ft; E. meyert, ca. 7,500-
9,500 ft) and are probably products of a recent speciation and rein-
vasion according to the mechanism discussed on p. 34. ‘The genus
Drepanornis consists of two similar species whose geographical ranges
are still allopatric and which may therefore be considered a super-
species: D. bruijnii of the western New Guinea lowlands north of the
Central Range between long. 136 and 141°E, and D. albertisii, which
lives in much of New Guinea except for the area inhabited by D.
bruijnit and ranges from 2,000 ft up to a ceiling varying geographically
between ca. 5,000 and 7,000 ft. ‘Che most distinctive features of all
four forms is the long and slender sickle-shaped bill (culmen from base
65-80 mm), a feature possessed by no other paradisaeid. Epimachus
and Drepanornis are superficially similar and have always been placed
next to each other in classifications.
The similarity appears to be more than superficial. Bock (1963,
p. 101) stated that the skulls of the two genera were virtually identical
except for the lacrymal being larger in Epimachus. Stonor (1938, pp.
468-472) noted that details of the feather tracts were similar, e.g., the
extreme narrowing of the terminal portion of the ventral tract. The
four forms are of similar sizes (wing 150-210 mm) and share the follow-
ing features of plumage, several of which distinguish them from such
related genera as Lophorina, Parotia, and Astrapia; a solid gray-brown
(E. meyeri and both Drepanornis forms) or black (E. fastosus) breast
in adult males; similar lateral tufts and pectoral tufts in adult males
(cf. Stonor, 1938), but not the distinct breast shields of Lophorina or
Parotia or the different shield of Astrapia; a solid velvet-black chin in
adult males; a brown unstriped head in Epimachus males and in both
sexes of Drepanornis, unlike the black striped head of Lophorina and
Parotia; and barred underparts in the females. The most marked
plumage differences between Drepanornis and Epimachus concern the
much longer tail of Epimachus, and the upperparts of adult males,
309
SPECIES ACCOUNTS
which are brown in Drepanornis (as in females of all four forms) but
velvet-black on the back and iridescent blue-green on the crown in
Epimachus, as in Lophorina, Parotia, and Astrapia.
Little is known about the habits of D. bruijnii, but the other three
forms live in the forest crown and use their long bills to probe for
insects and pick berries. The stomach contents given on labels of
Shaw-Mayer’s specimens of these three forms in the American Museum
of Natural History include grasshoppers, other insects, berries, and
fruit; the stomach contents of E. meyeri which I dissected included
insects, grubs, and fruit; and van Heurn found fruit and seeds in the
stomach contents of D. bruijnii (Hartert, 1932). ‘The two genera to-
gether may provide a threefold altitudinal sequence, D. albertisii-E.
fastosus-E. meyeri. On Mt. Karimui I found that the range of D.
albertistt was wholly below that of E. fastosus, and this is also sug-
gested by Stein’s (1936) records for the Weyland Mountains, with the
transition altitude around 4,200-5,000 ft. In southeastern New Guinea,
where the middle-altitude E. fastosus is absent and only the high-
altitude E. meyert remains, and on the Huon Peninsula, where both
species of Epimachus are absent, labels of specimens taken by Anthony
and Shaw-Mayer show D. albertisii as high as 7,000 ft, suggesting that
it may range higher in the absence of competition from £. fastosus.
Since the four sicklebills are closer to each other than to other
paradisaeids and share the most specialized bills of the family, they
form a natural genus, with the name Epimachus having priority and
Drepanornis available as a subgeneric name. Presumably the common
ancestor speciated to form the two lines or subgenera, with each line
speciating again to yield the present four forms.
3. Relation between Cicinnurus and Diphyllodes.
The genus Diphyllodes consists of a superspecies, D. magnificus
on the New Guinea mainland and D. respublica on the islands of
Waigeu and Batanta. Cicinnurus is monotypic (C. regius). At first
glance one is struck by dissimilarities in the adult males, Cicinnurus
being scarlet and white, Diphyllodes green, velvet, red, and yellow. ‘The
displays are also different, Diphyllodes clearing courts on the ground,
Cicinnurus displaying in the trees. However, the two genera are similar
in most other respects and have been placed next to each other in
recent classifications. ‘The affinity is attested by the fact that more
hybrid specimens of the D. magnificus x C. regius cross are known
than for any other paradisaeid hybridization.
The morphological resemblances include the following. Adult males
of both genera have two slender, greatly elongated central rectrices,
which cross at the base (as in no other paradisaeid), are curved at the
tip, and have barbs restricted to one side of the shat. Whe feather
tracts are similar (Stonor, 1938). The skulls are very similar in most
respects (Bock, 1963). In both genera the back of the adult male has
310
SPECIES ACCOUNTS
scarlet areas, and lacks black areas, whereas the other related para-
disaeids in which the female also has barred underparts (Lophorina,
Parotia, Astrapia, etc.) have the back largely velvet-black in the adult
male and lack scarlet areas. The females are similar generally, and
particularly in having brown unstriped heads, unlike the blacker
striped heads of Lophorina, Parotia, and Ptiloris. Cicinnurus regius
is closer to D. respublica than to D. magnificus, e.g., in size (wing Ca.
110 mm in D. magnificus, 95 mm in the other two species), in the ven-
tral coloration of the female (the barring is closer and the ground color
less ochraceous in D. magnificus than in the other two species), and in
the red dorsal coloration of the male (least extensive and dullest in
D. magnificus).
Both Cicinnurus and Diphyllodes eat mainly fruit and have rela-
tively short bills. Both spend much time in the lowerstory and are
regularly trapped in mistnets but also range up to the forest crown,
I have not heard the call of Cicinnurus regius adult males, but the
description of it by Gilliard (1969) is similar in pattern to the calls I
heard from D. magnificus adult males, a series of loud, harsh notes on
the same pitch. Most of the altitudinal range of D. magnificus (5,000 ft
down to the lower hill slopes, adult males generally uncommon below
2,000 ft) lies above that of Cicinnurus regius (the flat lowlands up to
about 2,000 ft in most areas). ‘The ecological differences between these
two similar species in the zone of altitudinal overlap are probably
related in large part to size, since D. magnificus is 60°% heavier than
G. neous:
Consideration of the taxonomic status of Diphyllodes and Cicinnurus
is postponed until the remaining species of Paradisaeidae have been
discussed.
4. Relations between Lophorina, Parotia, and Ptiloris.
The genus Lophorina is monotypic (L. superba), while Parotia con-
sists of a superspecies (P. sefilata, P. carolae, P. lawesi, and P. wahnest)
and Ptiloris also consists of a superspecies (P. magnificus, P. victoriae,
P. paradiseus). Parotia is distinct in that adult males clear display
courts on the ground and have six wire-like plumes on the head. ‘Chere
are also some differences in the bills and hence in the feeding habits
and skulls of these three groups. Parotia has a bill similar to that of
Cicinnurus or Diphyllodes and feeds largely on fruit. Lophorina has
a more slender bill and takes not only fruit but also beetles, grass-
hoppers, and other insects. The bill is somewhat longer in Ptiloris
(more so in P. magnificus than in the other two semispecies) and is
used to obtain insects from bark and crevices as well as to take fruit.
In plumage, however, both in the female and in some ornaments of
the adult males, these three groups are extremely similar. So detailed
is the resemblance that Stresemann (1923, p. 31) initially listed the
Lophorina females collected by Burgers on the Hunsteinspitze as
oll
SPECIES ACCOUNTS
Parotia carolae, and only 10 years later did he recognize them as a
(thinly differentiated) new race of Lophorina superba, which he ac-
cordingly named L. s. pseudoparotia. Hybrids are known between
Lophorina and Parotia, and between Lophorina and Ptiloris.
The plumages of the adult males are basically similar except for
the wire-like plumes of Parotia. The coloration is mainly black, with
glosses of various shades. The dorsal feathers have a soft, velvet-like
texture and, in Lophorina, are laterally prolonged into a cape. The
crown is iridescent blue-green (in Parotia this is true only distally).
All three species or superspecies possess a distinctive ventral ornament
shared by no other paradisaeid, an iridescent blue-green breast shield of
flattened, stiff feathers which have black central dots in some popula-
tions of each group (e.g., the dots are present in Parotia, Ptiloris
magnificus, and the Lophorina superba races from the Central Range,
but they are absent in Ptiloris victoriae, Ptiloris paradiseus, and the
races Lophorina s. superba and L. s. niedda of the Vogelkop and
Wandammen Peninsula). The shield is markedly prolonged laterally
in Lophorina and slightly in two of the three Ptiloris semispecies. ‘The
iridescence of the shield varies from green to purplish among the
Parotia semispecies. The feather tracts in the three “genera” are
similar (Stonor, 1938).
The females share a basic color pattern (brown back, blackish head
and moustache with whitish superciliary and gape stripes, close black
and white ventral barring on an ochraceous background), and similar
types of marked geographic variation are encountered in the three
groups. ‘The upperparts vary from rufous (Ptiloris magnificus, Parotia
wahnest and P. lawest) to olive-brown (Ptiloris paradiseus and P.
victoriae, Parotia carolae, Lophorina superba latipennis). Rufous
edges are present on the remiges of Ptiloris magnificus, Parotia wah-
nest and P. carolae, and the Lophorina races of the Central Range;
reduced in Ptiloris paradiseus, Ptiloris victoriae and Parotia lawes?;
and nearly absent in Parotia sefilata and the Lophorina races superba
and niedda. The underparts have a pronounced ochraceous wash in
Ptiloris victoriae and Parotia lawesi and P. wahnesi, less of a wash in
Ptiloris paradiseus, Parotia carolae, and Lophorina s. feminina, still
less of a wash in Parotia sefilata and Lophorina s. minor, and the
ground color is virtually white in Ptiloris magnificus. ‘The light super-
ciliary stripe is conspicuous in Ptiloris, Parotia carolae, and the
Lophorina races from the Snow Mountains (feminina) eastwards; re-
duced to a narrow streak posterior to the eye in Parotia wahnesi; and
completely or nearly absent in Parotia sefilata and P. lawesi and the
Lophorina races L. s. niedda and L. s. superba. The crown is black in
the Parotia semispecies sefilata, lawesi, and wahnesi and in the
Lophorina races L. s. superba and L. s. niedda, brown in Ptiloris
magnificus, Parotia carolae, and Lophorina s. feminina. In the area
immediately below the eye the light gape stripe is absent in the
312
SPECIES ACCOUNTS
Lophorina races L. s. superba and L. s. niedda, while the dark
moustache is conspicuous in the Parotia semispecies sefilata, lawesi,
and carolae, reduced in Ptiloris magnificus and Parotia wahnesi, vir-
tually absent in Ptiloris paradiseus, and entirely absent in the
Lophorina races of the Central Range. The markings of the under-
parts are bars in Parotia, Lophorina, and Ptiloris magnificus, crescent-
shaped in Ptiloris paradiseus, and reduced to sparse dots in P. victoriae.
One of the most remarkable and overlooked features of this geo-
eraphic variation of the females is that it runs parallel in some popula-
tions of Parotia and Lophorina. Thus, Parotia sefilata is confined to
the Vogelkop and Wandammen Peninsula. The Lophorina races in
this region (nominate superba on the Vogelkop, niedda on the Wan-
dammen Peninsula) resemble a small version of this Parotia semi-
species in the black head, dark back, reduction of the rufous edges
of the remiges, and obliteration of the superciliary stripe, and differ
only in the absence of the gape stripe. In the Snow Mountains the
crown is brown and the superciliary conspicuous both in the Parotta
semispecies (carolae, the one for which Stresemann and van Heurn
independently mistook Lophorina) and the Lophorina race (feminina),
and the olive back, the rufous edges of the remiges, and the color of
the underparts match well. In eastern New Guinea the species deviate,
since the underparts are much less ochraceous, and the superciliary is
much more conspicuous, in Lophorina than in Parotia (P. lawesi and
P. wahnes).
‘The altitudinal ranges of Lophorina and Parotia largely coincide
with each other (usually ca. 3,500-6,500 ft), and largely exclude that
of Ptiloris magnificus (sea level to ca. 3,000 or 4,000 ft). Ecological
segregation between Parotia and Lophorina probably depends on
Parotia’s greater size (nearly twice as heavy as Lophorina) and
Lophorina’s longer bill and greater consumption of insects.
5, Affinities of groups 3 and 4, and of Astrapia, Pteridophora, and
Seleucidis.
While Cicinnurus and Diphyllodes are particularly close, and Ptiloris,
Lophorina, and Parotia are very close, these two groups are also re-
lated to each other. ‘This is illustrated by the basically similar pattern
of the females (brown dorsally, barred underparts; remiges edged
rufous in Diphyllodes respublica and in Cicinnurus but not in D.
magnificus) and by the resemblances between the skull and bill of
Parotia on the one hand and of Diphyllodes and Cicinnurus on the
other (Bock, 1963). Field observations suggest that Diphyllodes magnifi-
cus and Lophorina are close. Adult males of these two common species
have mutually exclusive altitudinal ranges: on Mt. Karimui I found
that the lowest adult males of Lophorina appeared about 35 ft above
the highest males of Diphyllodes. While Lophorina females and im-
mature males penetrated 1,000 ft into the range of Diphyllodes, the
313
SPECIES ACCOUNTS
abundances of the two species varied complementarily in the overlap
zone so that the sum of their abundances remained relatively constant.
The calls of adult males are quite similar. The species differ only by
13% in weight, and both spend much time in the understory but also
range into the forest crown. Hybrids are known.
There are three other birds of paradise whose relationship to the
Lophorina and Cicinnurus groups is indicated by female plumage,
and whose relationship to the Lophorina group is further indicated
by male plumage. These are the Astrapia superspecies, Pteridophora
alberti, and Seleucidis me’anoleuca:
Astrapia consists of five midmontane and high-altitude semispecies
(A. nigra, A. splendissima, A. mayeri, A. stephaniae, and A. rothschildt).
The adult male plumage conforms closely to the Lophorina-Parotia-
Ptiloris type (black with various glosses, the crown iridescent blue-
green, the back with a soft velvet-like texture). The central rectrices
of the adult male are greatly elongated as in Epimachus and (less
markedly) in Parotia wahnesi, and the feathers of the nape are
lengthened to varying degrees in different subspecies. However, Astra-
pia lacks specially modified ornamental feathers. ‘The females are
brown dorsally, barred ventrally, and brownish-black on the head and
upper breast. Most of the altitudinal range of Astrapia is shared by
the smaller Pteridophora, which weighs only half as much, and by
the sicklebilled Epimachus meyert. ‘The lower part of Astrapia’s range
is shared with Lophorina and Parotia. Hybrids with Epimachus
fastosus are known. Although Astrapia would be regarded as wildly
aberrant in any other family, by the standards of the Paradisaeidae
it must be considered as having few distinctive features of significance.
The long tail of the adult male cannot be regarded as a good generic
trait within this family, because of the marked variation in tail length
among the semispecies of Parotia or Paradigalla and between the re-
lated Epimachus and Drepanornis. Astrapia has generally been placed
next to Epimachus in classifications because of the long tail, but
lengthened tails have apparently been acquired independently in four
groups (Parotia, Paradigalla, Astrapia, Epimachus) among the 11 spe-
cies or superspecies of the brown-female complex (listed on p. 309).
There is no other reason to suspect that Astrapia is closer to Epimachus
than to other members of the complex.
The monotypic genus Pteridophora (P. alberti) has particularly
bizarre ornamental feathers in the adult male, a pair of long and
scalloped occipital plumes. As in Lophorina, the upperparts are black
with a soft velvet texture, and the feathers of the upper back are
lengthened into a Lophorina-like cape, but the lower breast and belly
differ from Lophorina in being yellowish. Females are brown dorsally,
lack stripes on the head (cf. Cicinnurus), and are whitish ventrally
with dark curved bars similar to those of Ptiloris paradiseus females
but less deeply curved. The weight of Pieridophora is approximately
314
SPECIES ACCOUNTS
the same as that of Lophorina. The altitudinal range is almost entirely
above that of Lophorina, both in my experience and in that of Gilliard
(1969). The only competing species of Paradisaeinae at these altitudes
are the heavier Astrapia and the sicklebilled Epimachus. As gauged
from stomach contents, the diet is largely fruit, with seeds reported
once. Pieridophora may be thought of as the high altitude member of
the four-species altitudinal sequence Pteridophora-Lophorina-Diphyl-
lodes-Cicinnurus. ‘The bill and skull are much as in Crcimnurus or
Parotia (Bock, 1963). Pteridophora may be less close to the Lophorina
group, particularly in the colored underparts of the male, than is
Astrapia, but it is still a relatively undistinctive form (apart from the
occipital plumes).
The monotypic genus Selewcidis (S$. melanoleuca) lives in lowland
swamp forest, has an elongated bill, is medium-sized (160-200 g), and
feeds on seeds, fruit, insects, and nectar. ‘The female plumage conforms
extremely closely to the Lophorina-Parotia-Ptiloris pattern: the back
and wings are rufous (cf. Ptiloris magnificus, Parotia wahnesi), the
head solid black (cf. Lophorina s. superba and L. s. niedda), the under-
parts pale ochraceous with close barring (as close as in Ptiloris magni-
ficus and Diphyllodes, somewhat closer than in Lophorina or Parotia).
The male shares with Lophcrina and its allies the glossy black upper-
parts with a soft velvet texture. The breast shields of Seleucidis,
Ptiloris, Lophorina, and Parotia males form a series: all feathers are
stiff and flattened in Lophorina and Parotia, fewer are stiff and flat-
tened in Ptiloris magnificus, stull fewer in Ptiloris paradiseus, fewer
again in Ptiloris victoriae, and none in Seleucidis, in which the whole
shield is composed of soft feathers. The male of Selewcidis differs
from Lophorina in its yellow belly which fades to white postmortem
and in its lengthened white flank plumes with 12 wire-like feathers,
but these features are paralleled by the yellowish belly of Pteridophora
and the lengthened purple flank plumes of Ptiloris magnificus. Hybrids
with Ptiloris and Paradisaea are known. The bill is somewhat more
specialized than in other forms of groups 3, 4 and 5 but is not nearly
so specialized as in Epimachus and is quite like that of Ptiloris,
especially Ptiloris magnificus. The skull is undistinctive and typical of
the Paradisaeinae (Bock, 1963). Particularly the female plumage, but
also the above-mentioned features of the male plumage and the bill,
place Seleucidis near its allies Ptiloris and Lophorina.
The taxonomic status of these three “genera” is considered later.
6. Remaining species of Paradisaeinae.
Five genera of Paradisaeinae remain to be considered. The first
three of these are “good” monotypic genera whose affinities are obscure.
Macgregoria pulchra is a high-altitude, monogamous, sexually mono-
morphic species of particularly obscure affinities, in that it is not even
$15
SPECIES ACCOUNTS
clear whether it belongs to the subfamily Cnemophilinae or Para-
disaeinae (Bock, 1963).
Lycocorax pyrrhopterus is confined to the Molucca Islands, where
the only other paradisaeid is Semioptera. Like the manucodes, which
it suggests superficially in its bill, size, and uniform glossy dark brown
color, Lycocorax is monogamous and sexually monomorphic. There is
no reason to suspect a particular affinity of Lycocorax to any other
member of the Paradisaeinae, unless Manucodia. Whether its super-
ficial similarity to Manucodia represents true relationship or conver-
gence is unclear (Bock, 1963).
Semioptera wallacei is the other paradisaeid of the Moluccas. The
female is uniform brown. The male differs only in having a pair of
long white plumes arising from each shoulder, and a green, laterally-
expanded breast shield superficially reminiscent of Diphyllodes magni-
ficus but basically quite dissimilar (Stonor, 1938). The form of the
ventral feather tract is shared with no other paradisaeid (Stonor,
1938). ‘The bill is elongated and heavy. Bock (1963) states that the
skull is typical of the Paradisaeinae and especially resembles that of
Seleucidis. Clear indications of a particular affinity of Semioptera to
some other paradisaeine genus are lacking, and the female plumage
is distinct.
The Paradigalla superspecies (P. carunculata and P. brevicauda) 1s
sexually monomorphic, uniformly black, and adorned only by facial
wattles. This plumage is distinctive for a paradisaeid female (super-
ficially suggested only by Macgregoria). However, the crown has stif-
fened blue-green feathers as in Lophorina, Ptiloris, Epimachus, and
Astrapia males. Hybrids with Lophorina, Parotia and Epimachus fur-
ther imply affinities to this large group of paradisaeines. ‘The skull is
apparently typical of the Paradisaeinae (Bock, 1963).
The last genus, Paradisaea, consists of a superspecies (P. apoda, P.
rubra, P. raggiana, P. minor, P. decora) plus P. rudolphi of eastern
New Guinea and P. guilielmi of the Huon Peninsula. ‘The latter two
are sympatric with the superspecies and inhabit altitudinal ranges that
lie largely or entirely above that of the superspecies. Some affinities to
the large group of Paradisaeinae with barred females are suggested by
the distinctly barred underparts of P. rudolphi females and the ob-
scurely barred underparts of P. decora females, as well as by hybrids
with Ptiloris, Diphyllodes, and Seleucidis. Diphyllodes and Cicinnurus
may be the members of this group to which Paradisaea is nearest, as
suggested by the following shared features: elongated wire-like central
rectrices with barbs reduced or absent, the presence of red or yellow
and absence of black on the back of males (except in P. rudolphi), and
the upside-down displays of males of P. guilielmi, P. rudolphi, and
occasionally Cicinnurus regius and P. raggiana.
316
SPECIES ACCOUNTS
7. Evaluation of genera of Paradisaeinae.
Having surveyed the 22 species or superspecies of Paradisaeinae, we
may now consider how best to draw generic lines.
Macgregoria pulchra, Lycocorax pyrrhopterus, and Semioptera wal
lacei are each isolated forms without clear affinities to another species.
These three, and only these three, members of the Paradisaeinae fulfill
the criteria of good monotypic genera. Manucodia is an isolated and
very homogeneous group of four species (one a superspecies). In the
case of Paradisaea (three species, one a superspecies) there seem to be
clear affinities with “Diphyllodes” and “Cicinnurus”’, but Paradisaea
is still sufficiently distinct and compact that its right to stand as a
genus is unlikely to be disputed. Similarly, the Paradigalla super-
species is related to the Lophorina complex but constitutes a distinctive
and acceptable genus.
The remaining 11 species or superspecies form a related group, as
indicated by the female plumage of all members, the male plumage of
most, and hybridization. From this group Epimachus (including
Drepanornis; three species, one a superspecies) deserves to be sub-
tracted as a separate genus, because its species share the most strikingly
distinct bill in the family and form a compact group which has
radiated in three easily reconstructed speciations.
The assignment of the other eight species or superspecies of the
group provides the most recalcitrant obstacle to a satisfactory generic
revision of the Paradisaeidae. It may be recalled that within this
group “Diphyllodes’ and “Cicinnurus’ form one closely-knit sub-
group, “Lophorina”, “Parotia’, and “Ptiloris” form another subgroup,
and “Astrapia’, “Seleucidis”, and “Pteridophora” are each separate.
One solution would therefore be to recognize five genera, two polytypic
and three monotypic. Against this solution one may object that it
fails to express taxonomically the interrelationships within the group,
that it exaggerates the distinctness of the members by comparison with
the other much more distinct paradisaeid genera, and that neither
Astrapia, Pieridophora, nor Seleucidis has characters warranting erec-
tion of a monotypic genus, if one rejects ornamental plumes as a
proper basis for monotypic genera. Alternatively, the eight species
could be placed in a single genus (Lophorina) of five subgenera
(Cicinnurus, Lophorina, Astrapia, Seleucidis, and Pteriodophora), as
illustrated in Table 10 (p. 321). This has the disadvantage that eight of
the 22 members of the subfamily Paradisaeinae would be placed in one
rather large genus, and that the differences between some of the
species in this genus would be greater than the differences among
congeneric species in the other polytypic genera (Manucodia, Para-
disaea, Epimachus). ‘The former point is less a disadvantage than
simply a recognition of a biological reality, that the subfamily does
consist of a group of eight species plus several more isolated groups or
317
SPECIES ACCOUNTS
species. ‘Vhe latter difficulty is inevitable in translating a phylogenetic
tree into hierarchical nomenclature with a finite number of levels: the
possible distances between species of the tree vary continuously, and
it is therefore impossible to attain a nomenclature in which the dis-
tances between all units on the same level are equal. On the whole
the arrangement of Table 10 appears to me the least unsatisfactory
one I can devise at present, in expressing relationships and in minimiz-
ing variation in differences between taxonomic units on a given level.
8. Subfamily Cnemophilinae.
As presently construed, this subfamily consists of three monotypic
genera, Cnemophilus (macgregori), Loria (loriae), and Loboparadisea
(sericea). Bock (1963) showed that these three species resemble each
other, and differ from the species of Paradisaeinae, in several features
of skull construction sufficiently distinctive to warrant placing these
three birds in a separate subfamily.
Among these three species Loria loriae, which De Vis initially de-
scribed in the genus Cnemophilus, and Cnemophilus macgregorii
resemble each other, and differ from Loboparadisea, in several morpho-
logical respects. Loboparadisea is a smaller bird (weight 62-77 g vs.
80-100 g for the other two species). ‘he underparts of the adult male
are yellow in Loboparadisea, velvet-black (Loria) or brownish velvet-
black (Cnemophilus) in the other two. ‘The females of Cnemophilus
and Loria are unpatterned and rather uniform-colored birds: Cnemo-
philus, olive-brown, slightly paler below; Lora, olive, paler and more
yellow below, olive-brown on the wings and tail. Both Cnemophilus
and Loria females present a slightly scaled appearance due to obscure
dark feather margins. Loboparadisea females have a distinct pattern
dorsally, in that the head and upper back are olive-brown, the lower
back and rump yellow, and the tail rufous-brown; the underparts are
yellow. Loboparadisea has a distinct immature plumage which is
streaked ventrally and which is otherwise more like adult females of
Cnemophilus than of Loboparadisea, whereas in the cases of Loria
and Cnemophilus immatures resemble the adult female. Cnemophilus
and Loria females are sufficiently similar that when Shaw-Mayer and
Loke Wan Tho discovered the nest of Cnemophilus and obtained a
high quality close-up photograph of the female at the nest in sharp
focus, the bird was misidentified as Loria, Sims (1956, p. 426) described
the nest under Loria, Gyldenstolpe (1955, Pl. XI) published a 7” > 5”
reproduction of the photo labeled as Loria in his report on his collec-
tions, and the misidentification was still not recognized until years
later. In dorsal coloration adult males of Loria and Cnemophilus differ,
since Loria is velvet black dorsally as ventrally, while Cnemophilus
is yellow-orange anteriorly becoming browner posteriorly and on the
wings. In addition, Cnemophilus has a short, slender crest of six
feathers. Loboparadisea adult males are distinctively patterned as the
318
SPECIES ACCOUNTS
adult females but with brighter colors, and have pale blue-green wat-
tles at the base of the bill.
Ecological and distributional evidence alone would suffice to indicate
a close relation between Cnemophilus and Loria. Both species live
mainly in the understory (unlike the more arboreal Loboparadisea)
and feed on fruit. The local altitudinal ranges of the two species
appear to be mutually exclusive, with the transition altitude around
7,600 ft on Mt. Karimui (Cnemophilus at higher altitudes, Lorva
lower). The geographical range of Cnemophilus is confined to south-
eastern New Guinea and the Eastern Highlands, a distribution shared
by only five other montane species, all products of a recent speciation.
One of these, the semispecies Parotia lawesi, is still allopatric to its
relatives; Astrapia stephaniae is largely allopatric to its relatives but
overlaps A. mayeri at the western extremity of its range, the two forms
living in mutually exclusive altitudinal bands; while the geographical
ranges of the other three, Ptiloprora guisei, Amblyornis subalaris, and
Paradisaea rudolphi, are shared completely with their more wide-
spread congeners Ptiloprora perstriata, Amblyornis macgregoriae, and
Paradisaea raggiana, with the two forms segregating altitudinally in
each case. These five cases represent stages in a shared speciation pat-
tern, in which the range of a southeastern New Guinea isolate of a
superspecies is reinvaded and completely overrun by the next isolate
to the west, the southeastern isolate also gradually extends westward,
and sympatry is made possible by altitudinal segregation. By analogy
Cnemophilus probably evolved as a southeastern New Guinea isolate
of a superspecies including Loria. ‘The course of speciation seems to
have proceeded equally far in the cases of Cnemophilus macgregorit,
Ptiloprora guisei, and Paradisaea rudolphi, all three of which now
share their entire geographical ranges with the sister species and have
extended westward to somewhere between long. 143°30’ E and 142°E.
For these reasons the species loriae and macgregorit should again be
considered congeneric, as De Vis did initially. Cnemophilus has priority
as the genus name, and Loria becomes a synonym. Loboparadisea
remains as an allied but monotypic genus.
Conclusions.
The classification tentatively arrived at is summarized in Table 10.
The 25 species or superspecies are now distributed among 10 rather
than 20 genera, yielding a species-to-genus ratio of 2.5. Of the 10 genera,
five are monotypic, one contains two species, two contain three species,
one contains four species, and one contains eight species. All of the
10 discarded genera are monotypic, five in the strictest sense, five in
the sense of consisting of a single superspecies.
Nomenclatural problems which would result from this classification
are that the species name of “Ptiloris” magnificus (described as Fal-
cinellus magnificus by Vieillot in 1819) would become preoccupied by
319
SPECIES ACCOUNTS
that of “Diphyllodes” magnificus (described as Paradisaca magnifica
by Pennant in 1781) as a result of both being transferred to the genus
Lophorina; and that the species name of “Pteridophora’ alberti (de-
scribed under the same name by Meyer in 1894) would become pre-
occupied by the race “Ptiloris” magnificus alberti (described as Ptiloris
alberti by Elliot in 1871) as a result of both being transferred to
Lophorina. However, coining of new names seems unwise, and likely
to lead only to confusion, until these and perhaps other proposals
for generic reclassification of birds of paradise have been more widely
discussed and until more evidence has accumulated. For the present,
in referring to paradisaeid species whose familiar generic names I
proposed to discard, I shall continue to cite this discarded name in
parentheses: e.g., Lophorina (Ptiloris) magnifica, Lophorina (Diphyl-
lodes) magnifica, Epimachus (Drepanornis) albertisii, and Cnemophilus
(Loria) loriae.
Cnemophilus macgregorii sanguineus Iredale
Multi-crested Bird of Paradise
SPECIMENS EXAMINED. Mt. Karimui Zone 8: 1 ¢, 1 subadult ¢ (2 and 6
Sept. 1965).
WEIGHT. 1 6:94 isubadult 4: 36-
WING. 1 ¢@: 114. I subadult @: 112;
STOMACH CONTENTS. Fruit, 3-12 mm in diameter.
TAXONOMY. ‘The subadult male has the deep velvet underparts
of the adult, with only two small olive patches, and has a full-sized
crest, but the upperparts are olive-brown as in the female, with areas
of orange on the crown and upper back.
The Eastern Highlands race sanguineus is quite distinct from nomi-
nate macgregoru of southeastern New Guinea in the orange rather
than yellow back. ‘The race kuboriensis described from the Kubor
Mountains was characterized as having paler upperparts and blacker
underparts than sanguineus (Mayr and Gilliard, 1954, p. 361). Since
examination of the type of kwboriensis showed that it was extremely
similar to sanguineus, and since only one other species (Peneothello
sigillatus) exhibits subspecific differences between Mt. Hagen and the
Kubor Range (50 miles from Mt. Hagen), detailed comparison of the
following material was undertaken: two adult males (including the
type of kuboriensis) from the Kubor Mountains, the two Mt. Karimui
specimens, and seven topotypical adult males of sanguineus from Mt.
Hagen. Average wing lengths from the three localities are the same
within 1 mm. Two Mt. Hagen specimens and the Mt. Karimui adult
are either as dark and black ventrally or slightly more so than the two
Kubor birds, five Mt. Hagen birds are slightly paler, and the Mt.
Karimui subadult is the blackest. ‘he back and uppertail coverts are
slightly paler and less orange in the Kubor specimens, the Mt. Karimui
320
SPECIES ACCOUNTS
TABLE 10
GENERIC CLASSIFICATION OF THE PARADISAEIDAE
i i000
Proposed Classification
Usual Classification
Subfamily Cnemophilinae
Cnemophilus macgregorit
Cnemophilus loriae
Loboparadisea sericea
Cnemophilus macgregorit
Loria loriae
Loboparadisea sericea
Subfamily Paradisaeinae
Macgregoria pulchra
Lycocorax pyrrhopterus
Manucodia ater
Manucodia chalybatus1-comrii
Manucodia jobiensis
Manucodia keraudrenii
Semioptera wallacei
Paradigalla carunculata-brevicauda
Epimachus fastosus
Epimachus meyeri
Epimachus [Drepanornis|? albertisii-
bruijnii
Lophorina superba
Lophorina sefilata-carolae-lawesi-
wahnest
Lophorina magnifica-paradisea-
victoriae
Lophorina [Cicinnurus| magnifica-
respublica
Lophorina [Cicinnurus] regia
Lophorina [Astrapia] nigra-splendissima-
mayeri-stephaniae-rothschildi
Lophorina [Pteridophora] alberti
Lophorina |Seleucidis| melanoleuca
Paradisaea rubra-apoda-raggiana-minor-
decora
Paradisaea guilielmi
Paradisaea rudolphi
Macgregoria pulchra
Lycocorax pyrrhopterus
Manucodia ater
Manucodia chalybatus-comrit
Manucodia jobiensis
Phonygammus keraudrenii
Semioptera wallacei
Paradigalla carunculata-brevicauda
Epimachus fastosus
Epimachus meyert
Drepanornis albertisti-bruijnii
Lophorina superba
Parotia sefilata-carolae-lawesi-wahnesi
Ptiloris magnificus-paradiseus-
victoriae
Diphyllodes magnificus-respublica
Cicinnurus regius
Astrapia nigra-splendissima-mayeri-
stephaniae-rothschildi
Pteridophora alberti
Seleucidis melanoleuca
Paradisaea rubra-apoda-raggiana-
minor-decora
Paradisaea guilielmi
Paradisaea rudolphi
1 Names of members of a superspecies are connected by hyphens for purposes of brevity.
2 Subgeneric names are given in brackets.
adult, and one Mt. Hagen bird than in the other six Mt. Hagen
birds, and the Mt. Karimui adult is slightly deeper orange on the head
than the Mt. Hagen birds, but these differences are also trivial. Hence,
kuboriensis must be synonymized with sanguineus.
BREEDING. ‘The testes of the subadult male were moderately
large.
DISCUSSION. Cnemophilus macgregorii occurred from 7,620 ft
to the summit (8,165 ft) on Mt. Karimui, and has also been collected
by Gilliard, Shaw-Mayer, Bulmer, and Hitchcock between 8,000 and
11,000 ft on Mt. Giluwe, Mt. Hagen, and the Kubor Range in moss
forest. The western limit of its range must lie somewhere between Mt.
321
SPECIES ACCOUNTS
Giluwe and Telefolmin but remains to be determined. It is usually
seen in fruit trees, sometimes up to 60 ft above the ground but more
often in the understory. Its movements are characteristically abrupt, its
wingbeat loud. Once I watched a group of two adult males and two
birds in female plumage, moving singly or together 3-40 ft above the
ground, often with all four birds within 3 ft of each other, periodically
stopping to eat berries. They occasionally gave low harsh calls, and
the wings whirred loudly in flight, but the two males appeared not to
react to each other, and there was no suggestion of a display. South-
eastern New Guinea natives on Mt. Albert-Edward told me that C.
macgregortt does not carry out a display (“he no sing-sing, he walk-
about nothing that’s-all’’), and Gilliard was given the same information
in the Eastern Highlands.
VOICE. Quiet harsh and rasping sounds.
Cnemophilus (Loria) loriae amethystinus (Stresemann)}
Loria’s Bird of Paradise
NATIVE NAME. Gimi: arudimi.
SPECIMENS EXAMINED. Awande: 1 ¢ (19 June 1965). Mt. Karimui Zone 3:
Das Zone ali Ge Zone, OZ Omeone 7 Ay Zoomers ale ee Uh (17 Aug.-8 Sept.
1965).
WEIGHT. 5 @: 80-101 (90+8). 3 9: 91, 94, 95.
VWEING 2 or os 99-107 (N02 93) 3 O96, 995 00:
TAXONOMY. Subspecies are founded on the iridescent color of
the inner secondaries of adult males viewed in good light: solid violet
in amethystinus, blue-green in inexpectatus, between violet and blue-
green in nominate loriae. Eastern Highlands specimens are unmis-
takably amethystinus. Gyldenstolpe’s (1955, p. 142) suggestion, ad-
vanced without comparative material being available, that Eastern
Highlands birds might be C. /. loriae is untenable. ‘The gape is white.
BREEDING. ‘The testes were moderately enlarged in four of the
males, small in the other three.
DISCUSSION. Although Cnemophilus loriae was apparently not
uncommon between 5,300 and 7,600 ft on Mt. Karimui, I never saw
it in the field and obtained no evidence of its presence on Mt. Michael.
The frequency of netted individuals indicates lowerstory habits. Gil-
liard, Gyldenstolpe, Shaw-Mayer, Bulmer, and Hitchcock all collected
Cnemophilus loriae at various altitudes in forest between 6,000 and
9,000 ft. My highest record of it on Mt. Karimui came 20 ft below the
lowest record of Cnemophilus macgregoru, and the altitudinal distri-
butions of these congeners also appear to complement each other at
other localities, with little overlap.
1 Listed as Loria loriae in Rand and Gilliard (1967).
oan
SPECIES ACCOUNTS
Loboparadisea sericea sericea Rothschild
Yellow-breasted Bird of Paradise
SPECIMENS EXAMINED. Soliabeda: 1 imm. ? (22 July 1965). Mt. Karimui
Zone 1: 1 imm. 3; Zone 2: 3 4; Zone 3:1 ¢, 1 imm. 6; Zone 4: 1 g,1 @ (10-
31 Aug. 1965).
WEIGHT. 6 4: 70, 72, 72,78, 75. 1 oO: 77, Timm, 765. limm: g* 7,
i imm. ?; 66.
WING. 5 @: 94 (3), 95, 96. I°9: 93. 1 imm. 4:96, 1 imm. 9: 93. 1 imm.
>: 98.
TAIL. 2 hs 61163.
TAXONOMY. Material available for comparison was as follows.
L. s. aurora: Herzog Mountains, 1 ¢. L. s. sericea: type, 1 ¢ (prob-
ably from the Weyland Mountains); Weyland Mountains, 7 6, 1 @,
limm. ¢, 3 imm. @; Mt. Goliath, 2 ¢; Utakwa River, 1 imm. 9;
Telefolmin, 1 imm. ¢; unknown locality in Dutch New Guinea, | ¢.
In color the adult males from Mt. Karimui are indistinguishable from
nominate sericea. They differ from aurora in that the top of the head
is olive-brown darker than the back, rather than olive scarcely darker
than the back; in that the back is less pale and less yellow; and in that
the primaries are darker. In the initial diagnosis of aurora, Mayr
(1930, p. 147) also cited paler yellower underparts with a green in-
stead of orange tone. The underparts of the one aurora specimen
available to me are paler than the Mt. Karimui males or two Weyland
Mountains males but can be matched exactly by other Weyland
Mountains males, five of which have a green tone.
Measurements of wing length are given in Table 11. There is ob-
vious variation, but it is too irregular to use as a subspecific character.
In the Weyland Mountains females are larger than males, but this is
not the case on Mt. Karimui. A single adult male which was received
by Gyldenstolpe from the Jimmi River and which was described as
closer to nominate sericea in color has a wing length equal to the
average value for Mt. Karimui males (Table 11).
What Rand and Gilliard (1967, p. 505) describe as the adult female
is actually the immature plumage. Adult females have the same pat-
tern as adult males but duller colors. ‘The wattles of the male are very
pale blue-green in life.
BREEDING. Gonads were enlarged in three of the five adult
males, small in all other specimens.
DISCUSSION. Loboparadisea sericea exemplifies to an extreme
degree the patchiness of distribution characteristic of numerous New
Guinea birds, especially some birds of paradise. It was common on
the west ridge of Mt. Karimui, but Gyldenstolpe’s Jimmi River speci-
men is the only other Eastern Highlands record. It appears from
published records to be common on Mt. Kunupi of the Weyland
Mountains, on the Utakwa River of southwestern New Guinea, and at
5235
SPECIES ACCOUNTS
TABLE 11
COMPARATIVE WING LENGTH OF Loboparadisea sericea
ot ee eee
g Q imm. @ imm. Q
aurora
Herzog Mountains! 96
, 95 (2), 96 (2)2
sericea
Mt. Karimui 94 (3), 95, 96 93 96 93
Jimmi River 953
Telefolmin Oo
Mt. Goliath 97598
Weyland Mountains 89, 90, 91 (type), 98 96 94,96, 98
G52, 939)
unknown locality 100
1 Localities listed successively from east to west.
2 From Mayr (1930, p. 147).
® From Gyldenstolpe (1955, p. 145).
Dawong in the Herzog Mountains, but to be absent at some nearby
localities and in many large and well-explored areas. In all, it is known
from only about nine localities (Wau and Dawong in the Herzog
Mountains, Mt. Karimui and Jimmi Valley in the Eastern Highlands,
Telefolmin, Utakwa River and Mt. Goliath in the Snow Mountains,
and Mt. Kunupi and the Gebroeders Range in the Weyland Moun-
tains).
I never observed Loboparadisea in the field and can offer only the
statement of my native assistants that they found it in the treetops,
which is consistent with no specimens having been netted. As with
some other hill forest species like Charmosyna pulchella and Rhipidura
atra, the intrusion of the flat Karimui Basin has given it a discon-
tinuous altitudinal range: present at Soliabeda (2,000 ft), absent at
Bomai (3,250 ft) and Karimui (3,650 ft), present between 4,000 and
5,890 ft on Mt. Karimui. ‘The population structure follows the usual
pattern, with two of the three immatures coming from lower altitudes
than any of the adults, and the Soliabeda immature (2,000 ft) a full
2.400 vertical feet below the lowest adult (4,400 ft).
Manucodia chalybatus (Pennant)
Crinkle-collared Manucode
NATIVE NAMES. Daribi: pagonabo (adults), tagari (immatures).
SPECIMENS EXAMINED. Karimui: 4 ¢,5 @ (10-13 Aug. 1964; 2 July-5 Aug.
1965). Soliabeda: 1 imm. ¢ (28 July 1965). Mt. Karimui Zone 1: 1 ¢ (9 Aug. 1965),
WEIGHT. 5 47 158-> 210) 5 °9% 167-181 (175 225).
WING. 5 @: 154-174 (16247). 5 9: 150-164 (155 + 4).
TAIL, 5 &: 121-168 (161 3-5). 5 Qs W19-163 (12> 5),
EXPOSED GULMEN, 5 42 34-39 (672-1). 5 Qs 30-50"G8=2 1).
TAXONOMY. Apparently iris color changes from brown to red-
524
SPECIES ACCOUNTS
orange with age: the iris was orange to red in adult males (with
crinkled breast and throat feathers) and in most females examined,
two of which were in breeding condition, but was orange-brown or
brown in subadult and immature specimens. A convoluted trachea
was present in adult males.
Subadults and immatures of M. chalybatus can be very difficult to
distinguish from subadults and immatures of M. ater, because the
distinctive glosses and feather shapes of the adults are lacking. Hence
measurements were made of both species in southern New Guinea
(Fly River) and southeastern New Guinea, with the results tabulated
in Table 12. The most consistent difference is the longer tail of M.
ater. In southeastern New Guinea M. ater has a longer wing than
M. chalybatus, but on the Fly River there is much overlap. On the
Fly River the dorsal sheen is a useful character except in immatures,
since the sheen is consistently more green in M. ater there and more
purple in M. chalybatus. ‘This character is of less use in southeastern
New Guinea, where M. ater is more purple. M. ater is mainly a low-
lands species, M. chalybatus a hill forest species.
BREEDING. ‘The two females examined at Karimui in 1964 had
greatly enlarged ovaries with a nearly full-sized egg, and one crinkled
Karimui male in 1965 had the testes slightly enlarged, but the gonads
were small in other specimens.
DISCUSSION. Single individuals or pairs of M. chalybatus were
TABLE 12
COMPARATIVE MEASUREMENTS OF Manucodia chalybatus AND Manucodia ater
IN SOUTHERN AND SOUTHEASTERN NEW GUINEA
Exposed
Culmen Wing Tail
Manucodia chalybatus
southeastern New
Guinea, 3 4
southeastern New
39.5-41.0 (40.0)1 169-177 (173) 138-142 (140)
Guinea, 5 9
Fly River, 2 3
Fly River, 39
Karimui, 5 2
Karimui, 5 9
34.5-37.0 (35.5)
37.0-38.0 (37.5)
34.0-35.0 (34.5)
34.0-39.0 (36.5)
33.0-36.0 (34.5)
154-167 (158)
166-177 (172)
160-162 (161)
154-174 (164)
150-164 (158)
120-137 (130)
138-139 (139)
123-128 (126)
121-138 (130)
119-133 (124)
Manucodia ater subalter
southeastern New
Guinea, 5 2
southeastern New
38.0-40.0 (39.0) 192-205 (198) 162-174 (167)
Guinea, 59
Manucodia ater ater
Fly River, 42
Fly River, 69
36.0-38.0 (37.5)
35.5-36.0 (36.0)
35.0-36.0 (35.5)
182-197 (189)
169-182 (176)
168-178 (174)
158-164 (160)
144-153 (149)
144-155 (149)
SSS ——————————— ee
1 Average values are given in parentheses.
325
SPECIES ACCOUNTS
not uncommon at Karimui and in Zone 1 of Mt. Karimui (1% of the
local avifauna) but were inconspicuous, quiet, and remained in the
middle- and upperstories. At Soliabeda, which was probably near the
bottom of the altitudinal range, the sole record was an immature
brought in by natives, and the ceiling on Mt. Karimui was around
4,750 ft. Bulmer also found the species present but very inconspicuous
in Kyaka territory up to at least 5,500 ft, and collected a pair of breed-
ing birds at a nest at 5,000 ft. Schodde and Hitchcock obtained it at
Lake Kutubu (2,450 ft). The silhouette in flight is similar to that of
Paradisaea raggiana, and the flight movements are also similar: several
quick, convulsive flaps while jerking the tail, then a short glide.
VOICE. ‘The only call I heard was a staccato “tuk” like a raindrop
falling in a shallow pan of water.
Manucodia keraudrenit jamesii (Sharpe), M. k. purpureoviolaceus
(Meyer), and M. k. neumanni (Reichenow)!
‘Trumpetbird
NATIVE NAMES. North Fore: kaukdbara. South Fore: kautyaba. Daribi:
pagonabo (adults), tagari (immatures).
SPECIMENS EXAMINED. Awande: 1 ¢, 1 9 (15 and 17 June 1965). Okasa:
265) 2 4 Aue 1964; 23 and 24 June 1965). Kani 1 aimm, 4 (4 Aug. 1965);
Mt Karmur Zone Il @, 0 Oo, 1 imm.: 635 Zone 2) 1-9 (G-16 Aus: W965):
WEIGHT. Awande-Okasa: 2 ¢, 168, 174; 2 9, 131, 171. Karimui-Mt. Karimui:
tg. 19032 OF ea7, Wi2; 2 imine (og lias 76:
WING. Awande-Okasa: 2 ¢, 164, 167; 2 9, 151, 161. Karimui-Mt. Karimui:
La 64 2 Ooo, 1b6562 mm Ss blo:
EXPOSED CULMEN. Awande-Okasa: 3 @, 29, 32, 33; 2 9, 30, 32. Karimui-Mt.
Kamm. hy 2, sls O29; 3052 iim: 4, 30; 20:
TAXONOMY. ‘The series was compared with purpureoviolaceus
of southeastern New Guinea, jamesi of the Fly River, Aru Islands,
and Baroka (southeastern New Guinea lowlands west of Hall Sound),
neumanni from the Wahgi and Sepik Mountains, and nominate
keraudrenit from the Weyland Mountains and Vogelkop. Awande-
Okasa birds have rather green underparts and blue-violet wings, tail,
and back and are closest to Fly River jamesit, from which they differ
in having the underparts, wings, and tail slightly, and the back con-
siderably, more purple and less green. However, in the color of the
back they are only slightly more purple than Aru Islands jamesii.
Of the Karimui-Mt. Karimui birds, two are quite green above, as are
most from the Fly River; one is more blue-violet above and can still
be matched exactly by some Fly River birds; and the remaining two
are still more blue-violet, both on the back and on the wings, like
Awande-Okasa birds. ‘These differences indicate that my series is inter-
mediate morphologically, as it is geographically, between the Fly River
1 Listed as Phonygammus keraudrenii in Rand and Gilliard (1967).
326
SPECIES ACCOUNTS
(jamesii) and southeastern New Guinea (purpureoviolaceus); and that
the more westerly-lying Karimui series is nearer jamesi, as one would
expect. The differences are more marked compared to purpureo-
violaceus, which is much more purple on the underparts, back, wings,
and tail. One specimen collected by the Sixth Archbold Expedition in
the Kassam Pass belongs to purpureoviolaceus. ‘The head plumes in
my series are unusually long. M. k. newmanni is a very deep dull
violet on the lower back, wings, and tail and is the race to which
belongs one Nondugl aviary specimen (given to Gilliard) of unknown
provenance but presumably from the northern watershed of the
Eastern Highlands. Thus, birds in the southern, northern, and eastern
parts of the Eastern Highlands are distinct and referrable to three
races: jamestt, neumanni, and purpureoviolaceus.
The immature has less gloss and a smaller crest and beard.
BREEDING. ‘The 1964 Okasa male had greatly enlarged testes,
but gonads were small in all other specimens examined.
DISCUSSION. Although in some other parts of New Guinea
(notably the mouth of the Fly River) Manucodia keraudrenii regularly
reaches sea level, it is a hill forest species in the Eastern Highlands,
ranging from 3,500 ft up to a variable ceiling at 6,200 ft (Okapa area)
or less (4,600 ft on Mt. Karimui). The only record at Karimui (3,650 ft),
near the bottom of its range, was of the one immature male. Most
of the altitudinal range of M. keraudrenii lies above that of M. chaly-
batus. Sightings were of solitary individuals in the middle- or upper-
story of the forest, except for a pair at Miarosa in a casuarina grove
a few hundred yards from the forest edge. One was seen in a fruiting
tree at Miarosa.
VOICE. A prolonged, hoarse, harsh note with a peculiar quality,
similar to the sound humans produce in response to a doctor’s request,
“Open your mouth and say a-a-ah.” Also, a musical, downslurred,
slightly nasal “kyew”, similar in quality to the call of Philemon nova-
equineae.
Paradigalla brevicauda Rothschild and Hartert
Short-tailed Paradigalla
NATIVE NAME. Gimi: ibinimi.
SPECIMENS EXAMINED. Mt. Karimui Zone 3: 1 9 (18 Aug. 1965).
WING. 151.
TAIL... 89,
DISCUSSION. ‘This is one of the birds of paradise with a very
local distribution. ‘The specimen, collected at 5,160 ft in the forest
middlestory, was the only representative of this species I saw. Gilliard
collected three in the Hagen and Bismarck Mountains and reported
it as ‘common in the forest crown between 6,500 and 8,000 ft but diffi-
cult to collect” (Mayr and Gilliard, 1954, p. 355). In Kyaka territory
327
SPECIES ACCOUNTS
Bulmer (pers. comm.) found it “common at forest edge and in forest
from below 6,000 to at least 7,000 ft’.
Epimachus fastosus stresemanni Hartert
Black Sicklebill
NATIVE NAMES. Fore: kumpi. Gimi: kurikuri. Daribi: padukwa.
DISCUSSION. Like several other birds of paradise, Epimachus
fastosus is at some localities common and at other similar localities
absent. On Mt. Karimui, I heard E. fastosus males calling most days
from about 5,200 to 7,300 ft and saw a female or immature male at
the top of a tree at 5,400 ft. Neither Shaw-Mayer nor Gyldenstolpe
collected it; Gilliard encountered only one male at 7,500 ft on Mt.
Hagen; in Kyaka territory Bulmer (pers. comm.) found it “fairly
common at forest edge and in forest 6,000-7,000 ft’; and Burgers
found it common on the Schraderberg.
VOICE. The male’s call consists of two identical, sharp, liquid
notes in immediate succession, with considerable carrying power:
“buk-buk”, like the call of a foraging domestic chicken but much
sharper and louder. On Mt. Karimui males never called on the ridge
crest but only from the steep flanks of the ridge, which I also found to
be the case with the isolated population of Epimachus fastosus ultimus
on Mt. Menawa in the North Coastal Range (Diamond, 1969) and with
Epimachus meyer.
Epimachus meyert bloodi Mayr and Gilliard
Brown. Sicklebill
STOMACH CONTENTS. ‘Two mm fruits (two stomachs); insects and 9 mm fruit
(one); 7 mm fruit pits and two 40 mm grubs (one).
DISCUSSION. ‘This sicklebill occurs at 8,000-9,500 ft on Mts.
Hagen, Kubor, and Wilhelm, the Wahgi Mountains, and the Schrader
Range. Its altitudinal range and that of its lower-altitude congener
E. fastosus are mutually exclusive. In southeastern New Guinea, where
E. fastosus is absent, I have found E. meyerz down to 5,000 ft. It
ranges from 10 ft above the ground to the crowns, and congregates in
fruiting trees.
VOICE. Adult males call from the steep flanks of ridges, two or
three individuals within earshot of each other. ‘he reverberating,
totally unmusical call lasts 2 sec, is repeated at intervals of about 2
min, and consists of three double notes. ‘he quality may be com-
pared to the sound of a pneumatic drill, a burst from a machine gun,
or a trash can lid being slammed on the pavement and its sound being
quickly damped out.
328
SPECIES ACCOUNTS
Epimachus (Drepanornis) albertisii cervinicaudus (Sclater)!
Black-billed Sicklebill
NATIVE NAME. Daribi: sodadi.
SPECIMENS EXAMINED. Karimui: 2 imm. ¢ (15 Aug. 1964; 5 Aug. 1965).
Mt. Karimui Zone 1: 1 ¢ (10 Aug. 1965).
WEIGHT, i ¢: 9.
WING, I 4: 154 2 imm, ¢: 147, 148.
TAM 2 ss 13052 3mm. Ss 22, 130:
CULMEN FROM BASE: 1 ¢: 75. 1 imm.°¢: 77.
TAXONOMY. Reexamination of available material indicates that
only two thinly differentiated races are worth recognizing: nominate
albertisii (including geisleri) from the Vogelkop, Wandammen Penin-
sula, and Huon Peninsula; and cervinicaudus (including inversus)
from the Central Range. In the adult male, adult female, and imma-
ture male cervinicaudus has a slightly paler upper tail and slightly
lighter, duller, more olive back than albertisii. I cannot detect the
alleged slight color differences between the populations of the Huon
Peninsula (‘“geisleri’”) and the Vogelkop (nominate albertisi) after
comparing adult males, adult females, and immature males from both
areas. The Karimui population is slightly darker on the back than
cervinicaudus of southeastern New Guinea but otherwise matches.
DISCUSSION. Epimachus albertisii is an uncommon and local
species, especially in the main body of New Guinea, where the Karimui
series provides the first record for the southern watershed. ‘There ap-
pear to be three locality records for the northern watershed (Mt.
Kunupi in the Weyland Mountains at long. 135°30’E, the Lordberg
in the Sepik Mountains at long. 143°E, and the Herzog Mountains at
long. 147°E), in addition to records for four peripheral areas (the
Vogelkop, Wandammen Peninsula, Huon Peninsula, and southeastern
New Guinea). Rand and Gilliard (1967, p. 470) and Mayr (1941b, p.
170, and in Mayr and Greenway, 1962, p. 189) state that the range
includes the northern slopes of the Central Range from the Weyland
Mountains to the Lordberg, but this is an incorrect assumption: there
are no records at all between the Weyland Mountains and the Lord-
berg. ‘This area includes the entire range of the closely related and
equally uncommon Epimachus (Drepanornis) bruijnii (ca. long. 136-
141°E in the lowlands of the northern watershed). The hill slopes in
this area have been intensively surveyed by the Third Archbold Ex-
pedition around long. 139°E (northern slope of the Snow Mountains),
by Mayr around long. 140°30’E (Cyclops Mountains) and by me be-
tween long. 141°30’E and 143°30’E in the North Coastal Range, and
(less intensively) by van Heurn on the Doormanpad (long. 138°30’E)
and by Gilliard at ‘Velefolmin (long. 141°30/E), so that the apparent
1 Listed as Drepanornis albertisii in Rand and Gilliard (1967).
329
SPECIES ACCOUNTS
absence of E. albertisii in this area is probably real, and it and E.
bruijnit are probably still allopatric and constitute a superspecies. In
addition, their altitudinal ranges are different, since E. bruijnii is
known only near sea level and E. albertisii only in the hills and moun-
tains (3,650-4,200 ft in the Karimui area). On Mt. Karimui the highest
record of E. albertisii was 1,000 ft below the lowest record of E. fasto-
sus, so that the altitudinal ranges of these two similar species are mu-
tually exclusive.
VOICE. Karimui natives pointed out to me a call from the tree-
tops which they said belonged to this species. It consisted of a rapid
and loud series of over a dozen musical notes which progressively
increased in volume and dropped slightly in pitch. This call is similar
to the description of the call of this species by Stein (1936, p. 26) and
of E. bruijnii by Ripley (1964, p. 48).
Lophorina superba latipennis Rothschild
Superb Bird of Paradise
NATIVE NAMES. Fore: néni (male), piyé (female and immature male). Gimi:
nine-6ba (male), araro (female and immature male).
SPECIMENS EXAMINED. Awande: 3 ¢, 2 imm. ¢ (14-20 June 1965). Mt.
Karimui Zone 1: 1 imm. ¢; Zone 3: 2 9; Zone 4: 2 9; Zone 5: 1 @ (10 Aug.-l Sept.
1965).
WY ERG, iG 0508 (BOSE) 4 Ol ONR75 (Oyo) 8 Cm Oneida ols
WING, =4° 45 los2s8(ISGs-2).9 40S IS 23 (Sie 3). Saimin oe es leas
eA
STOMACH CONTENTS. Fruit, 6-8 mm in diameter.
TAXONOMY. Iredale (1948) founded a race addenda on a single
adult male from the Hagen district, citing apparently striking differ-
ences in the cape, the frontal shield, and the breast feathers. Mayr and
Gilliard (1954, p. 358) and Gyldenstolpe (1955, p. 133) did not recog-
nize addenda, because adult males from the Hagen district and other
parts of the Eastern Highlands proved indistinguishable from males
from other parts of the Central Range. However, there has been no
agreement as to the correct assignment of the Eastern Highlands popu-
lation, which has been variously placed with feminina (Mayr and Gil-
liard, 1954; Mayr, in Mayr and Greenway, 1962, p. 193; Rand and Gil-
liard, 1967), with pseudoparotia (Gilliard and LeCroy, 1961), or with
a resuscitated addenda (Gilliard, 1969) on the basis of female char-
acters. In fact, comparison of Eastern Highlands females with those
of other races shows that they are clearly closest to latipennis of the
Huon Peninsula, from which they differ only in being slightly darker
above. From feminina they differ obviously in the crown being blacker
and less brown, and in having much less white speckling, the under-
parts having much less ochraceous wash, and the back being darker
330
SPECIES ACCOUNTS
and more olive. The race minor of southeastern New Guinea differs
in the crown being slightly more blackish and the back much warmer
brown; nominate superba (Vogelkop) and niedda (Wandammen Penin-
sula) in the crown being an unspeckled black and the superciliary
stripe greatly reduced. ‘Topotypical specimens of pseudoparotia (Sepik
Mountains), which were not available to me for comparison, were
noted by Stresemann (in Gyldenstolpe, 1955, p. 132) and by Mayr and
Gilliard (1954, p. 358) to be browner and less olive dorsally than
Eastern Highlands females. The race connectens of the Herzog Moun-
tains 1s eintually indistinguishable from latipennis and must be con-
sidered a synonym. Eastern Highlands males have distinct black
centers to the feathers of the breast shield, as do males of all other
populations except nominate superba and niedda.
BREEDING. ‘The gonads were much enlarged in one female,
small in the others; small in immature males, and enlarged in most
adult males.
DISCUSSION. In my study areas Lophorina superba was the com-
monest bird of paradise within its altitudinal range (ca. 3,500-7,000 ft),
and accounted for 3°% of the local avifauna at 5,400-6,000 ft on Mt.
Karimui. There was marked change in population composition with
altitude on Mt. Karimui. Zone 1 yielded only an immature male, and
Zone 2 two females or immature males. Zone 3 yielded females, and
the first adult male was heard calling at 5,115 ft. Adult males were
commonest in Zone 4, where females remained until 5,700 ft. Above
5,700 ft few Lophorina superba were encountered, all adult males. At
the bottom of the forest on Mt. Michael (ca. 7,000 ft) there were a few
Lophorina superba, all adult males. ‘This change in sex ratio with alti-
tude, though true of many other species, is particularly marked in birds
of paradise, as initially reported by Stein (1936).
There are also some differences in the habits of adult males as op-
posed to immatures and females. Adult males call and display high in
the crowns, though not at the very tops, of leafy trees. In ecologically
disturbed areas these display trees are often at the forest edge or in
partly cut areas, to a greater extent than with the other midmontane
birds of paradise. Females and immatures spend proportionately twice
as much time’in the understory as adult males. Females and adult
males were never seen together, and Paran said that nestlings were fed
solely by the female. ‘Verborgh found Lophorina superba feeding in
fruit trees at Okasa and Miarosa. It is generally solitary and moves
slowly.
VOICE. The call of the adult male is a series of seven to ten very
harsh, raucous, utterly unmusical notes within a span of 5 sec. Suc-
cessive notes in the series become lengthened and louder and decelerate
greatly. A given individual repeats the series from the same perch at
intervals of about 10 min.
Sal
SPECIES ACCOUNTS
Lophorina (Parotia) carolae subsp.1
Queen Carola’s Six-wired Bird of Paradise
To date there are three Eastern Highlands records, all from the
northern watershed: a single female at the Nondugl aviary, possible
from the Jimmi River region, which Mayr and Gilliard (1954, p. 358)
assigned to the race chrysenia despite some differences; a male and
a female from the Jimmi Valley, which Gyldenstolpe was unable to
assign to an existing subspecies; and one native-prepared skin Bulmer
obtained in the Schrader Range, where he also obtained Lophorina
(Parotia) lawest.
The four semispecies of “Parotia” form a superspecies, with two
members (L. sefilata of the Vogelkop and L. wahnesi of the Huon
Peninsula) confined to peripheral mountain “islands”. In the main
body of New Guinea, L. carolae is the western semispecies and L.
lawesi the eastern semispecies. From present distributional information
it appears that the ranges of the two forms meet around long. 144-
145°E. The details of the transition may prove as interesting as in
the case of the Lophorina (Astrapia) mayeri-L. (Astrapia) stephaniae
transition.
Lophorina (Parotia) lawesi lawesi (Ramsay)?
Lawes’s Six-wired Bird of Paradise
NATIVE NAMES. Fore: kiara. Gimi: kiara. Daribi: papa.
SPECIMENS EXAMINED. Awande: 1 ¢,1 9 (15 and 19 June 1965). Okasa: 1
6 (22 June 1965), Mt. Karimui Zone 1: I) imm. 4; Zone 2: 2-9, 1 imm: 4; Zone
oe. 1 Os Zone 4. lO (e2 7 Anres 1905).
WEIGHT. Dos Vibe -bigk 139-664 (bl =29). 2 imme ose 7s:
WING. 2 @: 157, 158. 5 Q: 142-149 (144+1). 2 imm. ¢: 149 (2).
TAXONOMY. Iredale (1948, p. 162) founded the race exhibita on
a single female from the Mt. Hagen district, remarking that a single
male was indistinguishable from nominate lawesi. His diagnosis reads:
“But the female shows valid colour differences in the marking of the
under-surface, the whole being rather deep rufous, closely crossbanded
throughout from chin to under tail-coverts, while the upper surface is
also a deeper red brown.” Since subsequent collectors obtained no
females, the validity of exhibita has remained unchecked. ‘The above
series from the Eastern Highlands was carefully compared with
nominate lawesi from southeastern New Guinea. Adult males were
found to be indistinguishable except, apparently, for a slightly longer
wing in Eastern Highlands birds. ‘The females showed no difference
in the closeness of the barring of the underparts nor in the color of
the upperparts. Of seven Eastern Highlands females examined, three
1 Listed as Parotia carolae in Rand and Gilliard (1967).
2 Listed as Parotia lawesi in Rand and Gilliard (1967).
332
SPECIES ACCOUNTS
agreed in the color of the underparts with most southeastern New
Guinea females, while the other four were slightly duller and less
rufous below. Hence exhibita cannot be recognized. An adult male
collected by the Sixth Archbold Expedition at Purosa near Okapa
has a wing of 158 mm, and Gyldenstolpe (1955, p. 131) gives 150 mm
and 154 mm for the wings of two males from the Jimmi Valley (north-
ern watershed). My measurements of southeastern New Guinea lawest
give average wing lengths of 151.6 mm for adult males and 142.5 mm
for females, so that birds from the southern watershed of the Eastern
Highlands average slightly larger.
I examined the color of the iris in eight individuals from the Eastern
Highlands (1 ad. ¢, 2imm. ¢,5 ¢), and in all cases it was blue. Eye
color is indicated on the labels of 40 specimens from southeast New
Guinea in the American Museum of Natural History (21 ad. ¢, 12
mm: 450 Os) ?), Im 27. cases a ade 4,78 imi: 655) O01 7) cits
listed as blue; in 10 cases (7 ad. 6, 3 imm. ¢ ) there is a blue inner ring
and yellow outer ring, or the (el simply states yellow and blue; in
two cases (1 imm. ¢?, 1 ¢) the label states yellow; and in one case
(limm. ¢), brown. In Lophorina (Parotia) carolae (3 ad. ¢, 8 imm.
6,6 ¢@) the iris color is given as yellow-green, yellow, or whitish. ‘The
statement by Rand and Gilliard (1967, p. 483) that the iris is yellow in
the adult male of L. lawesi and pale blue in the female of L. carolae is
therefore incorrect and reversed. The gape in the specimens I ex-
amined was yellow.
BREEDING. Gonads of all specimens were small.
DISCUSSION. On Mt. Karimui Lophorina lawesi occurred be-
tween 4,000 and 5,700 ft, while it occurred up to at least 6,000
ft in the Okapa area. There are at least three locality records from
the northern watershed: (1) Bulmer collected it in Kyaka territory in
forest and at the forest edge, at 5,300 and 6,000 ft; (2) Bulmer collected
two females or immatures in the Schrader Range; (3) Gyldenstolpe
was given two males from somewhere in the Jimmi Valley. As dis-
cussed under L. carolae, these records are of interest in making it
likely that the ranges of L. carolae and L. lawesi meet.
Lophorina (Ptiloris) magnifica subsp. and L. m. magnifica (Vieillot)!
Magnificent Riflebird
NATIVE NAME. Daribi: to-owa.
SPECIMENS EXAMINED. Karimui: 1 9 (a), 1 imm. ¢ (14 July 1965).
beda: 1 92 (b) (29 July 1965).
WEIGHT. 2 9: 120 (a), 147 (b). limm. @: 168.
WING, 2 9; 142 (a), 15k Go). Lum, 4s 473;
EXPOSED CULMEN. 2 9: 46 (a), 50 (b). limm. ¢: 57
Solia-
1 Listed as Craspedophora magnifica in Rand and Gilliard (1967) and as Ptiloris
magnificus by Mayr (1962) in Peters’ Check-list, Vol. 15.
333
SPECIES ACCOUNTS
TAXONOMY. My measurements of females give: the eastern race
intercedens, 9 ¢ from southeastern New Guinea, exposed culmen 37-
41 (39.9), wing 141-155 (148.8); the western race magnifica, 8 9 (Fly
River, southern slope of Snow Mountains, Weyland Mountains, Hum-
boldt Bay, Idenburg River), exposed culmen 50-53 (51.6), wing 148-157
(151.8). ‘The Soliabeda adult female is nearer magnifica, but a firm
racial assignment will have to await examination of adult males.
Schodde and Hitchcock assigned the Lake Kutubu population to nomi-
nate magnifica.
BREEDING. The Soliabeda female examined was an adult with
enlarged ovaries, whereas the smaller Karimui female had very small
ovaries.
DISCUSSION. The Magnificent Riflebird was uncommon in the
Karimui area (0.2-0.3% of the local avifauna) up to about 4,000 ft on
the lowest slopes of Mt. Karimui. Females or immature males were
observed singly a few times in the middlestory of the forest. Adult
males were never seen, but their powerful calls showed that they had
a regular but very sparse distribution at intervals of about one-half
mile to a mile.
On two occasions at Karimui I observed a display between two birds
in female-like plumage (probably one a female and the other an
immature male. ‘The immature male and the female with small ovaries
were collected together at Karimui with bow and arrow by a native
and may also have been displaying). In the first case (7 Aug. 1964) the
two birds alighted next to each other on a branch 15 ft above the
ground at the top of a bush at the edge of the forest. One bird faced
the other, reared up on its legs, spread out its wings and bent them
backwards, threw back its head, and remained in this uncomfortable
posture for some time. ‘The second bird then assumed this posture, and
finally both assumed it simultaneously before flying off. No calls had
been given. On the second occasion (12 Sept. 1965) two birds perched
facing each other on a branch 30 ft above the ground in forest. One
tilted its body up until it was vertical, held the tail back at right
angles to the body so that it was horizontal, opened the wings and
bent them at the shoulder so that they nearly met behind the back
and the breast was arched towards the partner, and then rose up and
down on its legs like a horseman posting on a trotting horse. ‘The
pair flew off to a branch of another tree and repeated the display. A
less powerful version of the usual adult male call was periodically
given. The display of a captive adult male is apparently similar (Rand
and Gilliard, 1967, p. 487).
VOICE. The colossal whistles of the adult male are among the
most powerful and beautiful bird songs in New Guinea. ‘These calls
consist of either two or three upslurred whistles, each lasting nearly
1 sec and followed by the next at an interval of less than 1 sec. Each
note starts at approximately the same pitch (an octave or less above
334
SPECIES ACCOUNTS
middle C). The first slur sweeps a full octave upscale, the second slur
has a smaller sweep (of a sixth or a seventh), and the third note (if
there is one) a still smaller sweep. When the singer is close, one can
hear that the beginning of each slur is slightly hoarse, but the quality
is otherwise full and mellow. The singer remains hidden in the crown
of a tall tree and calls every few minutes. In Zone 2 of Mt. Karimui,
700 feet above the last riflebird, these whistles were still readily audible
coming up from below, and each singer probably hears his next rival
a half mile or more distant.
Lophorina (Diphyllodes) magnifica hunsteint (Meyer) and
L. m. chrysoptera (Elliot)
Magnificent Bird of Paradise
NATIVE NAMES. Fore: tdrotaro (male). Gimi: yaro. Daribi: mébudali.
SPECIMENS EXAMINED. Karimui: 4 9,4 imm. ¢ (1 July-6 Aug. 1965). Mt.
Karimui Zone 1: 1 ¢; Zone 2:1 ¢ (11 and 14 Aug. 1965).
WHICH r. 2 6: 88,94. 4 °0:60-93 (7/7 7). 4 1mm. ¢@: 77-108 (O02 7),
WING, 2 3: 115, Ile. 4 oO: 105-113 (108 =- 2). 4 imm. Gy 107-114 (UT a2 2),
STOMACH CONTENTS. Fruit.
TAXONOMY. ‘The adult males are closest to hunsteini of south-
eastern New Guinea, from which they differ only in that the back is
brighter and lighter red than most (but matched by some) hunsteznz.
They differ from chrysoptera in the brighter brick-red midback, and
the brighter, more orange, less brown crown; and from nominate
magnifica and intermedia in the brighter orange scapulars and inner
secondaries. Gyldenstolpe found that birds from south of the Wahgi-
Sepik Divide were hunsteini and those north of the Divide were
chrysoptera, which seems reasonable, since the passes all lie above the
altitudinal ceiling of the Magnificent Bird of Paradise and it has no
tendency to wander above this ceiling. ‘here are few or no racial dif-
ferences in the females.
BREEDING. ‘The two adult males had large testes. Gonads were
small in most females and in immature males in female-like plumage.
DISCUSSION. ‘The Magnificent Bird of Paradise is the commonest
bird of paradise and one of the most abundant birds in hill forest,
even in ecologically disturbed areas, dropping out towards sea level
and with a ceiling around 4,000-5,000 ft. In the Karimui area the
population composition and abundance were as follows: Soliabeda
(2,000 ft), 1.79% of the local avifauna, immature males twice as numer-
ous as females, few adult males; Karimui (3,650 ft), 2.7%, females
slightly more common than immature males, few adult males; Mt.
Karimui Zone | (4,000-4,200 ft), 2.4%, and Zone 2 (4,400-4,750 ft),
1 Listed as Diphyllodes magnificus in Rand and Gilliard (1967).
335
SPECIES ACCOUNTS
2.9%, adult males increasingly common; lower half of Zone 3 (4,750-
5,080" it), 123°, mainly adult males; no higher records.
In my experience the variations in the altitudinal limits of Lopho-
rina (Diphyllodes) magnifica are correlated with those of Lophorina
superba, and the two species seem to represent each other altitudi-
nally, a conclusion consistent with records of expeditions in other
parts of New Guinea. On Mt. Karimui the lowest adult males of L.
superba appeared (5,115 ft) just above the altitude where the highest
males of L. (Diphyllodes) magnifica dropped out (5,080 ft). Some fe-
males and immatures of L. superba extended 1,000 feet downward
into the range of L. (Diphyllodes) magnifica. From 2,000 to 6,000 ft
in the Karimui area the sum of the contributions of these two species
to the avifauna was approximately constant (1.6-3.7%), with only L.
(Diphyllodes) magnifica below 4,000 ft, only L. superba above 5,000 ft,
and their abundances in the overlap zone changing in complementary
fashion. The absence of L. (Diphyllodes) magnifica at Okasa was cor-
related with the downwards extension of L. superba’s range at Okasa
to well below 4,000 ft. Although the plumage and mating display of
the adult males are quite different, the calls of the adult males, the
plumage of the females, and the behavior of these two species are
very similar and reinforce the impression of close relationship drawn
from the altitudinal distribution.
Females and immature males are solitary and move about sluggishly
in the middle- and lowerstory. Adult males were much less often
seen than heard, but also seemed to spend a good deal of their feeding
time in the understory. As with L. superba, females and adult males
were rarely or never seen together and may meet only to mate. An
adult female was once seen feeding a juvenile male. Individuals some-
times perch sidewise on the trunks of saplings in the manner of thicket
flycatchers, and often hang upsidedown to feed.
Natives showed me two display areas maintained by adult males,
one at the Sena River (4,500 ft) and one at 4,350 ft on Mt. Karimui.
In each case an area of ground 6-8 ft in diameter had been cleared
completely of leaves and twigs. In this area stood two or three dead
saplings with their leaves removed. According to natives, the resident
male displays on these saplings.
VOICE. The call of the adult male, given from high concealed
perches im trees, 1s similar to that of L. superba and consists of a half-
dozen loud, harsh, buzzy, downslurred, low-pitched notes on the same
pitch, progressively increasing in volume. The L. superba call is
harsher and decelerates much more markedly. Several males may be
heard calling in the same general vicinity. I heard one calling near
one of the two display courts I visited. ‘Ihe females and immature
males call one to three harsh, downslurred, and quiet “chew’s’”, i.e. a
shortened and soft version of the male’s call.
336
SPECIES ACCOUNTS
Lophorina (Cicinnurus) regia rex (Scopoli) and
L. r. similis (Stresemann)!
King Bird of Paradise
NATIVE NAME. Daribi: naburo.
SPECIMENS EXAMINED. Karimui: 2 ¢, 2 9? (9 and 11 Aug. 1964; 1 July and
11 Sept. 1965). Soliabeda: 1 @ (22 July 1965).
WEIGHT. “3 @7 48, 5b, 57. I OG? 515.
WING. S 2°95, 96, 07. 2° 9%: 94. 96,
STOMACH CONTENTS. Fruit, 6-12 mm in diameter.
TAXONOMY. These belong to the widespread race rex (possibly
synonymous with regius; Mees, 1964). One immature male with en-
larged testes was largely in female-like plumage but had the head
largely red and some red on the primaries and uppertail coverts.
Gyldenstolpe received a specimen of L. r. similis from the Jimmi Valley
of the northern watershed, Schodde and Hitchcock took two rex at
Lake Kutubu (2,450 ft), and Bulmer collected one specimen at 2,000-
3,000 ft in the Baiyer Valley of the northern watershed.
BREEDING. All adult males had enlarged testes.
DISCUSSION. ‘The King Bird of Paradise was fairly common at
Soliabeda (ca. 1.39% of the local avifauna) but very uncommon at
Karimui (ca. 0.19%), where it was above its usual altitudinal ceiling.
The population structure shows the usual change with altitude: im-
mature males made up about half of the Soliabeda population, with
adult males and females also present, whereas only adults were found
at Karimui. My limited observations were of single individuals, male-
female pairs, or pairs in female-like plumage (females and/or imma-
ture male) 10-60 ft above the ground in the forest and at forest edge.
Schodde and Hitchcock (1968, p. 61) remark that at Lake Kutubu
L. (Cicinnurus) regia frequented second-growth and the forest edge,
while L. (Diphyllodes) magnifica lived in primary forest.
Lophorina (Astrapia) stephaniae ducalis (Mayr) and
L. s. feminina (Neumann)!
Princess Stephanie’s Astrapia
NATIVE NAMES. Fore: tawanta (male), 6kai (female and immature male).
Gimi: mélo. Daribi: kwekwe.
SPECIMENS EXAMINED. Mt. Michael: 1 9 (9 July 1964). Mt. Karimui Zone
oo 1 65 Zone 5: 1 & (16-27 Aug, 1965).
WEIGHT. 1 ¢@: 153. 1 9: 145.
WING. 1 62 162. 293 148, 161.
TVAUL.. I @: 560: 2 9% 294529,
STOMACH CONTENTS. Fruit 3-14 mm in diameter (five stomachs); fruit and
a large, hard insect (two); a 2 cm insect (one).
1 Listed as Cicinnurus regius in Rand and Gilliard (1967).
1 Listed as Astrapia stephaniae in Rand and Gilliard (1967).
337
SPECIES ACCOUNTS
TAXONOMY. Lophorina (Astrapia) stephaniae occurs from Mt.
Giluwe (long. 144°E) eastwards, with birds from Mts. Giluwe, Hagen,
Kubor, Karimui, and Michael referrable to the race ducalis and those
from the Wahgi Mountains, Mt. Wilhelm, and the Schraderberg to
feminina. ‘Vhe validity of ducalis as distinct from nominate stephaniae
of southeastern New Guinea is doubtful and has also been questioned
by Gillard and LeCroy (1968, p. 25). The supposed difference in
blackness may be due to foxing, and the difference in wing length is
slight.
DISCUSSION. L. stephaniae has been encountered by all collectors
at virtually all forested localities in its altitudinal (5,000-8,000 ft) and
geographical range, but its abundance is subject to marked local varia-
tion. I found it uncommon on Mt. Karimui (5,100-8,100 ft) and Mt.
Michael (7,000-8,000 ft) and never saw it in the Okapa area, whereas
Bulmer found it very common on Mt. Hagen and the Schraderberg.
I observed individuals or groups of two or three 10-70 ft above the
ground, prying in thick moss on trunks, or abruptly hopping up
branches and vertical trunks, remaining stationary between hops. ‘The
long tail is flicked to the side as the bird turns, and looks as if it is
about to fall off. The convulsive flight consists of rapid flaps alter-
nating with a glide.
VOICE. I heard only a single quiet, harsh, frog-like note. In Gil-
liard’s experience as well it seems to have been one of the quieter
birds of paradise.
Lophorina (Astrapia) mayeri (Stonor)}
Ribbon-tailed Astrapia
The range of L. mayer extends eastward to Mt. Hagen and Giluwe,
where Gilliard, Shaw-Mayer, and Bulmer found it plentiful to abun-
dant at 8,000-10,000 ft. As discussed by Mayr and Gilliard (1952a), its
geographical range overlaps that of L. stephaniae on Mts. Hagen and
Giluwe, where hybridization occurs to a limited extent (Shaw-Mayer,
fio) Sis, ISG, py 420).
A point of particular interest is that on both Mts. Hagen and Gil-
uwe, astrapias at lower altitudes are apparently pure L. stephaniae or
hybrids closest to it, while those at higher altitudes are pure L. mayeri
or hybrids closest to it. The transition altitude is at 8,000-8,500 ft,
as shown by Shaw-Mayer’s observations on Mt. Giluwe and Bulmer’s
and Gilliard’s on Mt. Hagen. The geographical ranges of L. mayeri
and L. stephaniae are still largely allopatric and overlap only in this
limited area. These two species thus illustrate the earliest stage in the
evolution of two allopatric members of a superspecies group into
sympatric species with mutually exclusive altitudinal ranges (p. 22,
1 Listed as Astrapia mayeri in Rand and Gilliard (1967).
338
SPECIES ACCOUNTS
p. 34). The western limit of L. mayeri, where it is replaced by L.
splendissima, must lie somewhere between Mt. Giluwe and ‘Telefolmin
and provides one of the major unsolved distributional problems in the
western part of the Eastern Highlands. As at the eastern limit, the
details of the transition may prove of interest.
Lophorina (Pteridophora) alberti (Meyer)!
King of Saxony Bird of Paradise
NATIVE NAME. Fore: warale.
SPECiMENS EXAMINED. Miarosa: 1 imm. ¢ (24 June 1964).
WING. “19:
TAXONOMY. The specimen is largely in female-like plumage but
has scattered velvet-black patches on the crown and orange on the
breast.
BREEDING. ‘The testes were enlarged.
DISCUSSION. Although two races of this species (buergersi and
hallstromt) have been named on the basis of differences in female
plumage, reexamination of available material, including the type
and three paratypes of hallstromi (Mt. Hagen) and two nearly topo-
typical specimens of buergersi (Schrader Range, 50 miles from Mt.
Hagen) shows that the variation is irregular and probably too slight
to warrant taxonomic recognition. ‘The only differences worth men-
tioning are that the ventral barring is slightly heavier in the Snow
Mountains, and slightly more widely spaced in the Eastern Highlands,
than in the Weyland Mountains (albertt), and that the measurements
are slightly larger for the Wahgi population (average wing length of
females: Wahgi region, 116.0 mm (range, 8 ¢@, 107-122.5); Schrader
Range, 111.3 (4 ¢, 109-113); Snow Mountains, 113.2 (9 ¢, 110-115);
Weyland Mountains, 112.2 (6 ¢@, 110-114)). Gilliard and LeCroy (1968,
p. 26) already noted that the female which Gilliard collected in the
Schrader Range (the type locality of Rothschild’s race buwergerst) is
indistinguishable from some Wahgi specimens, which had been sep-
arated from buergersi as hallstromi. The trivial differences on which
collectors have often founded subspecies of birds of paradise have
obscured a biologically important and still unexplained fact, namely,
that geographical variation within many species of birds of paradise
is remarkably slight compared to the striking differences between dif-
ferent semispecies or species.
The King of Saxony Bird of Paradise is widely but unevenly dis-
tributed in forest between about 6,000 and 9,000 ft. On Mt. Michael
an adult male was observed “‘singing’”’ from a perch 30 ft above the
ground at 7,800 ft. On Mt. Karimui two adult males “sang” regularly
1 Listed as Pteridophora alberti in Rand and Gilliard (1967).
339
SPECIES ACCOUNTS
within hearing distance of each other, one at 6,500 ft. and the other at
6,200" ft.
VOICE. “The “song” lasts about 3 sec, gradually increases in vol-
ume, and has such a weird quality that one is unlikely to suspect a
bird as the author. It is a very dry rattling, a spitted jumble of
insect-like notes poured out at a machinegun pace and suggestive of
bad static on the radio, which briefly turns into a twittering at the
climax of the crescendo.
Paradisaea raggiana salvadorii Mayr and Rand
Ragegiana Bird of Paradise
NATIVE NAMES. Fore: to. Gimi: é6romo. Daribi: puri.
SPECIMENS EXAMINED. Karimui: 1 ¢,4 9,4 imm. ¢ (3 July-5 Aug. 1965).
WEIGHT. 4 @: 164-203 (182 + 14).
WING, 1 @: 177. 2 QO: 147-169 (b8i=- 8), 4 umm. f= 159-179 (7126):
TAXONOMY. ‘The male specimen is not yet fully adult and has
tail wires 410 mm long but no flank plumes. Birds from the Karimui-
Okapa area as well as those from the Wahgi Valley are pure P. rag-
ginan salvadoru and show no approach to P. apoda novaeguineae of
the middle Fly River (whitish flank plumes, nape maroon rather than
yellow) nor to P. r. raggiana of southeastern New Guinea (upper back
yellow).
BREEDING. Gonads were small in almost all birds examined.
DISCUSSION. Paradisaea raggiana is common in the Eastern
Highlands up to a ceiling varying locally between about 4,500 and
5,600 ft. At the so-called Hybrid Gap at the western end of the Wahgi
Valley, where the pass between the Baiyer River drainage of the
northern watershed and the Wahgi drainage of the southern watershed
has been deforested, Paradisaea raggiana has been able to cross into the
Baiyer Valley and hybridize with P. minor, according to Blood (cited
by Mayr and Gilliard, 1954, p. 321) and information given to Bulmer
by native informants. It is considerably more numerous in ecologically
disturbed areas (casuarina groves, second-growth, and the forest edge)
than in the forest interior, and its unusually high ceiling in the Eastern
Highlands and its invasion of the northern watershed have probably
resulted from native agriculture.
Fully plumed adult males were very rare in my study areas, un-
doubtedly due to native hunting pressure. Even subadults with tail
wires but no flank plumes were encountered only twice. That the
species nevertheless remains common in settled areas is probably due
to subadult males with none of the prized ornamental plumage regu-
larly mating with females in the absence of fully adult males. The
birds remain in the middle- and upperstory (none was netted), and
were often associated in noisy groups of a half-dozen. ‘Verborgh re-
corded P. raggiana both in fruiting trees and flowering trees in the
340
SPECIES ACCOUNTS
Karimui area. The characteristic flight consists of several convulsive
flaps followed by a brief soar.
VOICE. The common call is a series of a dozen or more loud
“caw’s” which progressively rise in pitch and decelerate. Often this
series concludes with two more “caw’s’” at lower pitches and longer
intervals (Fig. 35).
Paradisaea raggiana:
-- Or a aS
4 sec
Fic. 35. Voice of Paradisaea raggiana.
Paradisaea minor finschi Meyer
Lesser Bird of Paradise
STOMACH CONTENTS. Fruit.
DISCUSSION. ‘This is the northern watershed equivalent of P.
raggiana, hybridizing with it in the upper Baiyer Valley. Bulmer found
P. minor common in settled areas of Kyaka territory up to 5,500 ft,
though fully adult males were as seldom seen as in the case of P. rag-
giana and presumably for the same reason.
VOICE. A series of loud “caw’s” very similar to the call of Para-
disaea raggiana.
Paradisaea rudolphi margaritae Mayr and Gilliard,
and P. r. rudolphi (Finsch)
Blue Bird of Paradise
NATIVE NAMES. Fore: kongonamu. Daribi: barimoi.
SPECIMENS EXAMINED. Awande: 1 ¢, 2 imm. ¢, 1 ? (16-20 June 1965).
Karimui: 1 ? (4 Aug. 1965).
WERIGIED, Legs 1/8.- 2mm Gs lo6; 1725 2 Ps 138. 175,
WING, 1 4: 156, 2 imm. ¢: 149, 152. 2 Py 138, 150.
TAXONOMY. In females and immature males of the race marga-
ritae, originally described from the Wahgi Valley, the underparts, in-
cluding the central belly and flanks, have black barring, whereas
rudolphi of southeastern New Guinea has only the lower belly dis-
tinctly barred and the chest only faintly barred. ‘The Karimui specimen
is as barred as margaritae and is assigned to it. One of the Awande
immature males is barred but more obscurely, and the other Awande
341
SPECIES ACCOUNTS
immature and the unsexed bird have as little barring as rudolphi, so
that the more easterly Awande population is nearer rudolphi. ‘The
adult male from Awande is indistinguishable from rudolphi.
DISCUSSION. ‘The Blue Bird of Paradise lives in forest between
about 3,600 and 6,500 ft and is generally uncommon. ‘The population
structure apparently follows the usual pattern, with adult males seen
and calls heard only in the upper parts of the altitudinal range.
Unlike P. raggiana and P. minor, it has not been able to colonize
second-growth extensively and hence has been driven from much of
its range by native agriculture (some of Bulmer’s records, however,
came from the forest edge and nearby garden areas). In undisturbed
forest the altitudinal range of P. rudolphi lies largely or wholly above
that of P. raggiana, as also true for P. guilielmi vis-a-vis P. minor on
the Huon Peninsula. The western limit of the geographical range of
P. rudolphi in the western part of the Eastern Highlands remains to
be determined.
VOICE. The call pointed out to me by natives consisted of a
series of notes somewhat similar to the call of Paradisaea raggiana, but
with longer intervals between the notes, successively lower pitches,
and a bell-like quality. The resemblance of each note to the syllable
“kong” is the origin of the Fore name “kongonamu”.
PTILONORHYNCHIDAE: BOWERBIRDS
Archboldia papuensis sanford: Mayr and Gilliard
Archbold’s Bowerbird
This extremely local bowerbird is known only from a small area
between 8,500 and 9,000 ft on the southwestern slope of Mt. Hagen,
where Gilliard and Shaw-Mayer collected it, and between 8,500 and
9,000 ft on Mt. Giluwe, where Shaw-Mayer collected it. Neither Bul-
mer nor Gilliard found it on the northern slopes of Mt. Hagen, and
Bulmer’s native informants from the southeastern slopes said that
it occurred only farther west.
Amblyornis macgregoriae macgregoriae De Vis
MacGregor’s Bowerbird
NATIVE NAMES. Fore: 6nkena (male), antau (female). Gimi: dse.
SPECIMENS EXAMINED. Awande: 1 ¢,1 9 (15 and 19 June 1965). Okasa: 2
imm. ¢ (22 and 25 June 1965). Mt. Karimui Zone 4: 1 @, 1 imm. ¢; Zone 5: 1
imm. ¢ (19 Aug.-2 Sept. 1965).
WHIGHT. 2 &: 110,123. 1 0: ITS. Samm. Gi. Ulo-1o2 (128 = 7),
WING. 2 #: 187,139. 1 °Q: 126. 2 umm, 6% 128; 129.
TAM, 2948s) 90, 1.O7o. “2m, ee Se - 88,
STOMACH CONTENTS. Fruit, 4-7 mm in diameter.
TAXONOMY. Gilliard and LeCroy (1961, p. 74) showed that the
342
SPECIES ACCOUNTS
eastern race macgregoriae differs from the western race mayri in pos-
sessing a slightly shorter crest and tail. ‘They measured the crest length
from the posterior base, while ‘Table 13 summarizes measurements of
crest lengths I obtained from the anterior margin of orange feathering
laid flat. The conclusion is still that western birds have longer crests,
though there is some overlap.
TABLE 13
Crest LENGTH OF ADULT MALES oF Amblyornis macgregoriae
mayrt
Weyland Mountains, 5 2 90-103 mm (av., 96.2)1
Lake Habbema, 2 ¢ 90, 102 (av., 96)
Mt. Goliath, 1 g 100
macgregoriae
Mt. Hagen, 5 ¢ 74-94 (av., 88.8)
Kubor Mountains, 3 2 80, 81 (av., 80.7)
Nondugl, 1 2 76
Mt. Karimui, | 2 83
Awande, | 2 2
Southeastern New
Guinea, 7 2 71-93 (av., 82.4)
1 Measurements are from the anterior margin of orange feathering.
‘The crests of both my adult male specimens are a deep orange com-
parable to that of the much shorter crest of A. swbalaris, and darker
than in any of the available museum specimens of A. macgregoriae,
which have a paler and yellow-orange crest.
DISCUSSION. Amblyornis macgregoriae is widespread, though
uncommon, in forest from 9,000 ft down to a lower limit varying
locally between 3,500 ft (Okasa) and 5,200 ft (Mt. Karimui). One bower
was found at about 6,800 ft in forest in the Okapa area, while there
were four bowers on the west ridge of Mt. Karimui, all on the flat
crest of the ridge: two within a few feet of each other at 5,390 ft, one
at 5,820 ft, and one at 6,060 ft. An immature male in female plumage,
with tiny testes and no trace of a crest, was collected while working on
a bower.
I spent two mornings observing from a blind at one of these bowers
(Fig. 36). It was located on a broad, flat, ridge crest, and was con-
structed around a sapling 16 ft tall and 3/4 inches in diameter. Sticks
were criss-crossed about the trunk of the sapling to 8 ft above the
ground, subtending for the most part 1 ft, occasionally up to 2 ft.
Above 8 ft the sapling had branches with leaves, but below this height,
where the sticks were criss-crossed, there was only one leafless branch
(the bird probably having stripped the leaves) and a vine from an
adjacent tree with a few leaves. About the base of the sapling the bird
had constructed a circular moss platform 54 inches in diameter, with
343
SPECIES ACCOUNTS
a raised rim around the periphery surrounding a moss basin 5 in deep
and 30 inches in diameter. In the center of the basin a moss base 10
inches in diameter rose immediately about the sapling. Outside the
platform was a small pile of chips of black, burnt wood. ‘The surround-
ing forest was tall and open, with some particularly large Araucaria
cunninghamii. Most of the bower was in shade, though some sunlight
reached the bower floor.
On the first day both a crested adult male and a bird in female
plumage were in the vicinity when I arrived at 10:20, and the birds
moved about within 30 ft of the bower. The female never actually
came to the bower nor within 10 ft of the male and perched between
12 and 35 ft above the ground. The male appeared squat and short-
tailed with his center of gravity and much of his body apparently well
forward of his legs. His orange crest remained inconspicuous, the base
concealed, only the distal part visible. While perched, he seemed to
peer intently, and his movements as he suddenly turned on his perch
or suddenly flew off with a loud wing beat were ludicrously abrupt.
Until 10:40 he assumed various perches 4-15 ft above the ground but
not at the bower and was silent except for some kissing and scolding
notes. Eventually he approached the bower, giving soft, downslurred,
whining notes and rapidly repeated scolding notes. From 10:40 to
10:46 he stood around the base of the bower and continued to make
whining and scolding sounds. Until 11:01 he periodically appeared
and reappeared 20 ft from the bower, perched 4-15 ft above the ground,
and once hopped vigorously up the trunk of a 2 inch sapling to a
height of 15 ft. Between 11:01 and 11:06 A.M., when observations
were terminated, no birds were visible.
On the next day I arrived at 09:28, placed 20 dead leaves on the
moss basin to see what the bird would do with them, and went into
the blind. At 09:30 the male appeared with a rustle of wings, and re-
mained in the vicinity until 10:58 without the female. At first he
took up various perches 15 ft or more from the bower, once perching
sideways on a vertical sapling like a thicket flycatcher. At 09:47 he
gave nasal downslurred calls like a Melidectes honeyeater; at 09:49,
neighed like a horse; and at 09:51 gave a call like the crack of a
bushknife striking a tree. Finally, at 09:57 he alighted on the rim of
the bower with white moss or lichens in his bill, occasionally turning
his head. After a minute he picked up one of the dead leaves I had
left in the basin and dropped it on the rim, carried another leaf 3 ft
off, and threw two over the rim with a toss of his head. He then spent
a minute on the ground just outside the rim, during which he briefly
raised, flared, and exposed his whole orange crest, which was otherwise
held flat, furled, and basally concealed. At 10:01 he flew off and re-
turned to the bower at 10:06. This time he threw out the leaf he had
previously dropped on the rim, then threw out some smaller leaves
by carrying them to the rim and tossing them several inches outside,
344
SPECIES ACCOUNTS
sticks
moss
platform
@ sapling
base
basin
Fic. 36. Bower of Amblyornis macgregoriae (dimensions in inches), Above: cross-
section in vertical plane through the central sapling. Below: cross-section in hori-
zontal plane through the moss platform. See text, p. 343, for details.
345
SPECIES ACCOUNTS
and at 10:07 picked up the largest leaf and flew off with it. In 26 min
he appeared again briefly, and at 10:40 reappeared with a whirring of
the wings and perched 2 ft up on a vertical sapling behind the bower.
He rapidly shifted perches to different saplings several times, quickly
opening and closing his wings while perched. At 10:44 he flew off 15 ft
to a perch 7 ft up and remained stationary for 3 min, during which he
once again raised and flared his crest. After another shift of perch he
began quiet, broken, nasal downslurs, coughing sounds, and chirps
for 5 min. The sounds stopped at 10:58, and he remained away until
at least 11:30, when I left. There were still 12 dead leaves remaining
in the bower.
Paran found a nest with one nestling near the bower, 7 ft up in a
pandanus. It was a bowl 5 inches deep and 51% inches in diameter,
lined with small twigs, and woven of strips of dry blades 3/4 inch broad
and 1-2 ft long brought from the adjacent grassland, plus a few dry
pandanus leaves and Castanopsis leaves. In Paran’s experience, the
nests of Amblyornis macgregoriae generally contain one nestling, which
is fed by the female alone.
[Sericulus aureus ardens (D’Albertis and Salvadori)"]
Golden Bowerbird
Iredale (1948) lists a skin from an unspecified locality in the Mt.
Hagen district. This probably originated from somewhere in the Fly-
Strickland-Kikori drainages and reached the Mt. Hagen district by
native trade routes. There are no authentic records from the Eastern
Highlands.
Chlamydera lauterbachi uniformis Rothschild
Lauterbach’s Bowerbird
NATIVE NAME. Fore: kairo.
TAXONOMY. ‘The ranges of the subspecies have been reevaluated
by Gilliard (1969).
BREEDING. A female brought in at Okasa by a native had en-
larged ovaries.
DISCUSSION. ‘This grassland bowerbird was present at Okasa
and Miarosa but absent at Karimui. Gilliard, Gyldenstolpe, Shaw-
Mayer, Bell, Bulmer, and Hitchcock all found it common in grassland
at various localities in the Wahgi Valley and neighboring areas.
1 Listed as Xanthomelus aureus in Rand and Gilliard (1967).
346
SPECIES ACCOUNTS
Ailuroedus crassirostris melanocephalus Ramsay and
A. c. guttaticollis Stresemann
Green Catbird
NATIVE NAMES. Gimi: waraléa. Daribi: buziba.
SPECIMENS EXAMINED. Mt. Karimui Zone 2: 1 ¢ (14 Aug. 1965).
WEIGHT. 205.
WING. 152.
TAXONOMY. Specimens of melanocephalus from southeastern
New Guinea vary somewhat in the darkness of the underparts, and
the Mt. Karimui bird agrees with the darker specimens. ‘he race
melanotis of southern New Guinea is paler and greener below, while
guttaticollis of the Sepik Mountains, to which Gyldenstolpe assigned
with reservations a single specimen from the Jimmi River (northern
watershed), has buffer spotting on the crown and nape.
BREEDING. ‘The testes of the specimen were very small.
DISCUSSION. ‘The altitudinal ranges of A. crasstrostris and A.
buccoides are mutually exclusive. In most areas of New Guinea the
transition takes place around 2,500 ft, but in the Karimui area it is
around 4,000 ft because of tropical conditions in the Karimui Basin.
I found A. crassirostris between 4,300 and 5,450 ft on Mt. Karimui and
also at Okasa. My observations were of solitary individuals in the
middlestory of the forest (at least 20 ft above the ground), where their
movements were easily traced by the calls. However, the single speci-
men was netted, probably a victim of its habit of eating birds and
bats trapped in nets.
VOICE. A peculiar, not loud, mewing sound “wa-a-a-a-a-a” which
lasts 114 to 2 sec and is repeated at 10- to 15-sec intervals. Each note
fluctuates up and down in pitch a few times within a span of about
two whole tones.
Ailuroedus buccoides stonit Sharpe, A. b. geislerorum Meyer,
and A. b. cinnamomeus Mees
White-eared Catbird
NATIVE NAME. Daribi: buziba.
SPECIMENS EXAMINED. Karimui: 2 ¢,3 9 (8-12 Aug. 1964; 1-15 July 1965).
Bomai: 2 g,1 Q (6-8 July 1965). Soliabeda: 2 Q (23 and 29 July 1965).
WEIGHT. 3 ¢@: 134, 142, 146. 4 9: 126, 136, 140, 150.
WING. 4 @: 135, 135, 138, 142. 6 9: 126, 127, 130, 130, 133, 134.
STOMACH CONTENTS. Fruit.
TAXONOMY. ‘The series agrees most closely though not perfectly
with stoni of southeastern New Guinea. The crown averages darker
in my specimens, though the darkest stonzz and darkest Karimui birds
are similar. ‘he underparts also average darker in the Karimui area,
347
SPECIES ACCOUNTS
though the darkest stonii are comparable. There is good agreement
in the small size of the spots on the underparts and their sparseness on
the abdomen. The races oorti, geislerorum, and buccoides all have
much paler crowns, geislerorum and oorti have much paler under-
parts, and the spots on the underparts are larger in oorti and buccoides.
Gyldenstolpe assigned a single specimen from the northern watershed
to geislerorum, while Schodde and Hitchcock placed the Lake Kutubu
population with cinnamomeus.
BREEDING. Gonads were small in all individuals examined in
1965, and were enlarged in one male and one female and small in the
other male in 1964.
DISCUSSION. In the Karimui region Ailwroedus buccoides was
rarely seen but frequently netted because of its habit, shared with
A. crassivostris, of eating birds trapped in mistnets. The head of the
victim was always eaten, and sometimes other parts as well.
NEOSITTIDAE: AUSTRALIAN NUTHATCHES
Daphoenositta miranda kuboriensis Mayr and Gilliard
Pink-faced Nuthatch
DISCUSSION. ‘This nuthatch of high-altitude moss forest has
been recorded four times in the Eastern Highlands at 8,000-10,000 ft:
in the Kubor Mountains by Gilliard, in the Lamende Range by Shaw-
Mayer, and in the Schrader Range by Gilliard and by Bulmer. Its
habits, and those of Neositta chrysoptera, seem to me more like those of
a warbler than of Sitta. Its progress is neither systematic nor slow; it
spends only a few minutes in a tree before flying off, usually out of
sight. It hops rapidly along a branch of either a dead or a leafy tree
at about 1 ft per sec, leaning over and peering first to one side then
another, and flits off to another branch. While it occasionally clings
to the underside of a horizontal branch or goes up a vertical one, much
more often it remains on the upper side of horizontal branches. I
watched one digging and pounding repeatedly in bark, another shak-
ing and eating a 3 cm grub. It was always in flocks, of three to five
birds.
VOICE. Faint, sweet, sucked-in, slightly squeaky contact calls
which are louder in flight than when perched; and a faint chatter
when two individuals came together.
Neositta chrysoptera subsp.+
Papuan Sittella
NATIVE NAME. Fore: yaloto.
SPECIMENS EXAMINED. Miarosa: 1 ? (24 June 1964). Awande: 1 ¢,1 9 (4
July 1967).
1 Listed as N. papuensis in Rand and Gilliard (1967), see Mayr (1950).
348
SPECIES ACCOUNTS
WING, 1 45°77, 1 @78,
STOMACH CONTENTS. Insects.
TAXONOMY. My three specimens have the whole head and
upper breast pure white except for a few brown streaks on the chin
and crown of the male, and the white markings on the inner edge of
the primaries are more prominent than in any other New Guinea
examples of the species available for comparison. Four birds which I
observed in the Karimui area also had the whole head white.
The three geographically nearest races are alba of the Idenburg
slopes (male with the whole head white, female unknown), toxopeusi
of the Snow Mountains (male with the head and neck streaked, female
apparently with the head and neck white), and albifrons of south-
eastern New Guinea (male with more dark areas and streaking on the
head, female with a whitish head with brown shaft streaks). White
spotting as on the primaries of the Miarosa specimen is present but
reduced in albifrons, toxopeusi, and papuensis and is nearly absent in
alba. Descriptions of three other Eastern Highlands adults have been
published: a single white headed adult male collected by Shaw-Mayer
at 8,000 ft on Mt. Giluwe, which Sims (1956, p. 430) found to match
alba; a single adult male collected by Gilliard at 8,300 ft on Mt. Hagen,
which is in general intermediate between albifrons and toxopeust
except for lacking the primary spots, and which Mayr and Gilliard
refrained from identifying subspecifically; and a single adult female
collected by Gyldenstolpe at Nondugl, which was nearest toxopeusi
but had more dark areas on the head and which he described as a
new race, wahgiensis. Descriptions of one specimen collected by
Bulmer at ‘Tari, one collected by Blood at Wabag, and five collected
by Hitchcock in the Kubor Range had not been published at the time
of writing. In view of the individual variation among the Eastern
Highlands specimens collected to date, it is not possible to assign the
material or to evaluate the validity of wahgiensis until an adequate
series can be compared with adequate series of the other races (alba
and toxopeusi are each known from only four specimens).
DISCUSSION. ‘This species may be the middle-altitude equivalent
of Daphoenositta miranda. 1 observed small flocks at 8,000 ft on Mt.
Michael, at 6,000 ft in Gimi territory, and at logoramalu (3,630 ft) in
the Karimui Basin, in addition to collecting the Miarosa (6,800 ft)
and Awande (6,000 ft) specimens. These flocks were in tall trees with
little foliage in forest clearings and gardens. Each flock spent a few
minutes at one tree and then moved on as a group with undulating
flight to a neighboring tree or else out of sight. As they worked actively
over a tree, they frequently crawled on the undersides of branches,
hung upside-down from branches, or else assumed upside-down. posi-
tions on vertical trunks. Paran said that Neositta chrysoptera disappears
from the Okapa area during the rainy season.
349
SPECIES ACCOUNTS
NECTARINIIDAE: SUNBIRDS
Nectarinia sericea sericea (Lesson)
Black Sunbird
SPECIMENS EXAMINED. Soliabeda: 1 ¢, 1 9 (26 and 29 July 1965).
WiRIGIH. <l w-9o7, “) or Oi:
\WENG, a6 ror 57.
STOMACH CONTENTS. Principally insects, occasionally fruit.
DISCUSSION. ‘The Black Sunbird was common at Soliabeda, and
rare at Karimui, at the forest edge and in gardens and second-growth.
‘The Karimui records are much higher than the species normally occurs
elsewhere in New Guinea. Schodde and Hitchcock also found the
Black Sunbird at Lake Kutubu (2750 tt). It perched) ims treetops;
hovered in hummingbird fashion at flowers, probed while hanging
upsidedown, or alighted on fallen logs and fences in gardens, moving
rapidly and seldom remaining in one place for long. It fed on the
surface of vegetation, never inside the forest.
VOICE. ‘There are a variety of common calls, all very high-pitched,
thin, sibilant, and rather faint. Common patterns are a rapid series of
identical upslurs; one or several upslurs followed by a slow trill at a
lower pitch; a breezy, slower upslur like North American goldfinches
(Carduelis sp.); and a rapid series of notes alternating between two
different pitches (Fig. 37). Some of these calls are similar to those of
the honeyeater Conopophila albogularis, and others to the calls of the
flowerpecker Dicaeum geelvinkianum, another tiny black bird of the
treetops.
Nectarinia sericea:
4AIS)/ Or 4As or aa
=— be porr
Fic. 37. Voice of Nectarinia sericea.
MELIPHAGIDAE: HONEYEATERS
Timeliopsis fulvigula meyer (Salvadori)
Mountain Straight-billed Honeyeater
SPECIMENS EXAMINED. Awande: 1 9 (14 June 1965). Mt. Michael: 1 ¢, 1
? (6 and 7 July 1964). Mt. Karimui Zone 2 6; Zone 5:1 9 28 Aucel Sept
1965).
Which, 2 22 20;.20. 2 Oo: 16; 19.
WING. 9 7 7% cl, 82. 2 O73, 77.
STOMACH CONTENTS. Insects.
350
SPECIES ACCOUNTS
TAXONOMY. The four races of this species were separated on
the basis of trivial differences in color shades. The Vogelkop popula-
tion (nominate fulvigula) has the olive of the back slightly duller and
darker, and the throat and breast browner, than the population of the
Central Range (meyeri). The population of the Snow Mountains is
much too similar to the southeastern New Guinea population to
warrant the name montana (type and two paratypes from Mt. Goliath
examined). The fourth race, fuscicapilla, is also probably a synonym
of meyeri.
‘The iris was orange in all specimens.
BREEDING. ‘Testes were enlarged in the Mt. Karimui males.
DISCUSSION. .The Awande specimen came from 6,200 ft, the
Mt. Michael ones from 8,000 ft, and the Mt. Karimui ones from 5,780
to 6,500 ft. Four of the six specimens were netted, suggesting under-
story habits, in agreement with the observations of Stein (1936, p. 30).
This species does not use flowering trees. Other Eastern Highlands
records are one specimen each from the Wahgi Region (Gilliard)
and the Schrader Range (Bulmer).
Melilestes megarhynchus megarhynchus (Gray)
Long-billed Honeyeater
NATIVE NAMES. Fore: éro. Gimi: koyage. Daribi: yoborusudbe.
SPECIMENS EXAMINED. Awande: 1 ¢ (28 June 1964). Karimui: 4 ¢, 2 9
(30 July-14 Aug. 1964; 2 July-5 Aug. 1965). Bomai: 1 ¢, 3 Q@ (7-9 July 1965).
Soliabeda: 1 3, 4 Q (22-29 July 1965). Mt. Karimui Zone 1: 1 @; Zone 2: 1 4;
fonerd. 1g, 1 Oo 9-17 Aue: 1965),
WEIGHT. 10 g: 43.0-49.5 (466+1.7). 10 Q: 36.3-47.5 (41.1 + 3.3).
WING, 10) 23 92-100 (M00== 3), 1093 387-047 (9 222),
STOMACH CONTENTS. Insects.
TAXONOMY. ‘These specimens belong to the widespread nomi-
nate race, and differ obviously from stresemanni of northern New
Guinea in the more olive, less gray underparts. ‘The iris was orange.
There may be a very slight increase in size with altitude: average
wing lengths on Mt. Karimui and at Awande (4,000-6,200 ft) are
é 101.8, 9 93.0; at Karimui (3,650 ft), ¢ 101.2, ¢ 90.6; at Soliabeda
(2,000 ft), 2 LOO. o O10;
A yellow eye ring, a sign of immaturity, is present in several speci-
mens. Iwo of these specimens also have streaked throats, short wings
(96 and 97 mm), and tiny testes and must be younger birds. Of the
males with eye rings but without streaked throats, two had the testes
somewhat, one considerably, enlarged.
BREEDING. One Karimui female in 1964 had greatly enlarged
ovaries. Testis size ranged from small to large, with a higher proportion
of enlarged testes on Mt. Karimui.
DISCUSSION. Melilestes megarhynchus is a common but. un-
ool
SPECIES ACCOUNTS
obtrusive tropical species ranging up to 5,000, rarely 6,000 ft. In the
Karimui area, where it accounted for 1-2% of the local avifauna, the
usual change in sex ratio with altitude is apparent: females predomi-
nate at the two lower stations (Solhabeda and Bomai), males at the
two higher stations (Karimui, Mt. Karimui). In habits this is a solitary,
shy, slow-moving, and silent bird of the lower- and middlestory (up
to about 30 ft above the ground) in forest and second-growth. Propor-
tionately more of the specimens were taken in nets at Soliabeda and
Bomai than at Karimui, and still fewer were netted on Mt. Karimui.
Since the Mt. Karimui specimens were mostly males in breeding con-
dition, they may spend less time in the lowerstory. M. megarhynchus
tended to feed by working its way up a tree, gleaning (presumably for
insects) close to the trunk and branches. It also came to flowering trees.
‘Terborgh observed a few individuals in a tree with small fruits at
Mengino and in the tree at Miarosa with hard-shelled fruits chipped
open by parrots (see under Trichoglossus haematodus, p. 143).
VOICE. A harsh but quiet downslur “chur-r-r’’ or mewing sound
is occasionally given.
Niche Differences in the Genera Toxorhamphus and Oedistoma
The genus Toxorhamphus has usually been considered to consist
of three very similar species, T. poliopterus, T. novaeguineae, and T.
tliolophus. However, I concur with the decision of Salomonsen (1967,
p- 342) to remove zliolophus to the genus Oedistoma, formerly con-
sidered monotypic (O. pygmaeum).
The niche differences among these ecologically similar honey-
eaters are as follows. T. poliopterus and the very similar T. novae-
guineae live mainly in the understory and do not visit flowering
trees. Their altitudinal ranges are mutually exclusive (7. polio-
pterus at higher altitudes than T. novaeguineae) where both occur,
and the former descends much lower in the absence of the latter. T.
novaeguineae has not been recorded from the Eastern Highlands and
is probably absent on the southern watershed east of the Fly River
but should turn up below 4,000 ft on the northern watershed, where
Gilliard collected it at the foot of the Schrader Range. The similar-
sized O. iliolophum, in addition to foraging in the understory, also
ranges into the treetops and visits flowering trees, which T. poliopterus
and T. novaeguineae never do in my experience. O. pygmacum is
similar in habits to O. iliolophum but spends even more time in
flowering trees and less time in the understory, weighs slightly less
than half as much, and has a relatively shorter bill. O. pygmaeum and
O. iliolophum range infrequently down to sea level, T. poliopterus
never. O. pygmaeum has the lowest altitudinal ceiling, O. iliolophum
higher, and T. poliopterus the highest.
352
SPECIES ACCOUNTS
Toxorhamphus poliopterus poliopterus (Sharpe)
Slaty-chinned Longbill
NATIVE NAMES. Fore: antabo. Daribi: tuniabe.
SPECIMENS EXAMINED. Awande: 2 ¢, 2 9 (20-21 June 1964; 14-18 June
1965). Karimui: 2 ¢, 3 2 (31 July-15 Aug. 1964; 1 July-3 Aug. 1965). Bomai: 1
& (6 July 1965). Soliabeda: 2 @, 1 9 (24-30 July 1965). Mt. Karimui Zone 1k
6: Zones: Je, 2 9: Zone 3: 1 4,2 0 (1-16 Aug. 1965),
WEIGHT. 10 @: 10.5-14.3 (124+1.0). 10 9: 9.3-11.5 (10.4 + 0.7).
WING. 10 @: 63-73(69 = 3). 10 9: 61-68 (64 - 3),
CULMEN FROM BASE. 10 @: 29-35 (32.00+1.5). 10 9: 27-35 (29.9 + Moe
STOMACH CONTENTS. Insects.
TAXONOMY. ‘Table 14 summarizes measurements for populations
from the eastern half of New Guinea. The first four entries, pertaining
to my Eastern Highlands specimens, or else the second through fourth,
pertaining only to the Karimui area, show a significant increase in bill
length and wing length with altitude. ‘There also appears to be some
local variation in the bill-to-wing ratio (compare ratios for Nondugl,
Mt. Karimui, and Awande, within 80 miles of each other and at similar
altitudes). Greenway (1935, p. 98) pointed out that there is a wide
range of variation in bill length and crown color (more gray or more
green) in specimens from a given locality. Huon Peninsula birds were
separated from southeastern New Guinea birds (nominate poliopterus)
as septentrionalis on the basis of a shorter bill, slightly longer wing,
lower bill-to-wing ratio, and more greenish, less pure gray crown
(Mayr and Rand, 1935, p. 14). ‘This race may be insufficiently distinct
to merit recognition. ‘he average difference in the color of the crown
is slight, though detectable; the average difference in the culmen shown
by ‘Table 14 is slight; and the difference in wing length shown in ‘Table
14 is difficult to evaluate without better knowledge of the altitudes
at which the specimens were collected, since both the Huon Peninsula
and southeastern New Guinea series fall within the range of averages
for Eastern Highlands populations at various altitudes. On the basis
of the average color of the crown (virtually the same as in southeastern
New Guinea) and the bill-to-wing ratio, Eastern Highlands birds would
have to be considered nominate poliopterus rather than septentrionalis,
if septentrionalis were recognized. ‘The race maximus of western New
Guinea has a darker crown, and perhaps larger dimensions.
BREEDING. All ovaries were small, but testes at most localities
were enlarged. ‘The main exception was the small testes of Soliabeda
males, suggesting that, as usual, immatures or nonbreeding birds are
concentrated at the bottom of the species’ altitudinal range.
DISCUSSION. The altitudinal range of T. poliopterus is roughly
1,500-6,000 ft, but both the lower limit and the ceiling show marked
variation. ‘The variations in the lower limit are correlated with the
presence or absence of the related T. novaeguineae, which occurs from
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SPECIES ACCOUNTS
sea level to about 4,000 ft in western New Guinea, extending east
through the Sepik Basin on the northern watershed but only as far as
the Fly River on the southern watershed. Where the two species are
sympatric, T. poliopterus does not descend below 4,000 or 5,000 ft
(Sepik Mountains, Snow Mountains, Weyland Mountains, and pos-
sibly the northern watershed of the Eastern Highlands). Where 1.
novacguineae is absent, T. poliopterus descends to about 1,500 ft
(southeastern New Guinea, southern watershed of the Eastern High-
lands at Soliabeda, and possibly Herzog Mountains and Huon Penin-
sula). The variations in the ceiling depend upon the local elevation
of the moss level, so that the ceiling was lower on Mt. Karimui (5,800
ft) than in the Okapa area (up to at least 6,800 ft) and still higher on
Mt. Michael (8,100 ft).
Within its altitudinal range, particularly in the upper part of its
range, T. poliopterus is among the most abundant forest birds, some-
times the most abundant species (e.g., 99% of the local population at
4,000-5,600 ft on Mt. Karimui). One would never guess this, however,
without nets or knowledge of the song, since the bird is infrequently
seen. Its habitat is the forest interior, forest edge, or second-growth.
It may be seen up to 30 ft above the ground but concentrates in the
lowerstory, as evidenced by the numbers in which it saturated our
mistnets (see under Oedistoma pygmaeum for further discussion of
its habits). It darts about rapidly in the undergrowth, pecking at
branches and sometimes hanging upside-down.
VOICE. Vocalizations are varied, frequent, weak, similar in quality
to the calls of the sunbirds Nectarinia sericea and N. jugularis, and
fall into two categories. Some calls have a dry quality, such as a dry
squawk, a dry scolding, and a disyllabic sneeze-like note; but most
have a sweet quality. The commonest call is a sweet “‘tsip’; another
is “tsee-tsee-tsee-tsee-tsee’. ‘The songs with sweet quality consist of
slurs, disyllabic notes, and repeated notes (Fig. 38). One of these songs
resembles a weak version of the song of Meliphaga auga and may be
confused with it.
Toxorhamphus poliopterus:
— or I
nae ato 5 alls Gel Gea
i Se
—— SSS a SS OF yy
Fic. 38. Voice of Toxorhamphus poliopterus.
500
SPECIES ACCOUNTS
Oedistoma iliolophum flavum (Mayr and Rand)!
Gray-bellied Honeyeater
NATIVE NAME. Daribi: piye.
SPECIMENS EXAMINED. Okasa: 1 ¢ (22 June 1965). Karimui: 5 ¢, 2 9
(30 July-13 Aug. 1964; 1 July-5 Aug. 1965). Bomai: 1 9 (6 July 1965). Soliabeda:
1 g (23 July 1965). Mt. Karimui Zone 1: 1 9; Zone 2: 3 4, 3 9; Zone 3: 1 @
(10-16 Aug. 1965).
WEIGHT. “10 @= 13.0-15.7 (42 == 10)9), 7 O% 1107120 (11 2 033).
WING. 10 4: 65-72 (68+ 2). 6 Q: 61-65 (63 + 2).
TAXONOMY. The flanks are more brightly lemon-colored than
in nominate liolophus. The wing lengths of both the Okasa specimens
(2 6, 68 and 70) and of the remainder from the Karimui area agree
with southeastern New Guinea birds (3 66-70, 9 62-64) and are larger
than on the Fly River (4 63-67, 9 58-62).
BREEDING. Gonad condition varied from small to moderately
enlarged.
DISCUSSION. O. iliolophum was present at all my collecting
stations up to 5,125 ft, and at none above this altitude. It was com-
monest at Karimui (3,650 ft), much less common at Soliabeda (2,000 ft),
and is either absent or quite uncommon in flat country at sea level
(as with many hill forest species, this last statement does not apply
to the flat lowlands of the Fly River basin).
O. iliolophum and Toxorhamphus poliopterus have nearly the
same wing length; O. tlolophum weighs slightly more; and T.
poliopterus has a considerably longer bill (culmen from base ca.
30 vs. ca. 20 for O. iliolophum). ‘T. poliopterus has brighter colors,
but the coloration of O. iliolophum is sufficiently similar that field
identification is usually difficult. There is broad altitudinal overlap
from 1,500 to 5,000 ft, with 7. poliopterus extending above and O.
tliolophum below these limits. However, there are two striking niche
differences. First, in flowering trees we never saw T. poliopterus but
regularly saw O. tliopholum, which accounted for 1.39% of the bird-
usage in flowering trees at Karimui. The other difference involves
netting returns: nets accounted for 57°% of our specimens of O. zliolo-
phum but 88% of our specimens of 7. poliopterus. Evidently both
species spend much time in the understory, but the tendency is de-
veloped to an extreme degree in T. poliopterus. In flowering trees
O. iliolophum was seen as often in the upper half of the tree (includ-
ing the canopy) as in the lower half. It was also frequently seen
within 10 ft of the ground (as one would suspect from the netting
returns) and had a particular affinity for the clusters of small wild
bananas at Karimui. Its behavior was often nuthatch-like, feeding
close to branches and perching upside down on tree trunks and banana
1 Listed as Toxorhamphus iliolophus in Rand and Gilliard (1967); see p. 352.
356
SPECIES ACCOUNTS
clusters. However, it also hovered at the banana clusters, like a sun-
bird (to which it bears a striking superficial resemblance) or a hum-
mingbird.
Oedistoma pygmaeum pygmaeum (Salvadori)
Pygmy Honeyeater
SPECIMENS EXAMINED. Karimui: 1 ¢, 1 @ (1 and 17 July 1965).
WEIGHT: Tf gPo0: 179? 5.7.
WING. 1 4: 48. 1 9: 47.
TAXONOMY. Ina paper preliminarily naming a total of 28 races
of meliphagids, based on reexamination of older material, Salomonsen
(1966) described three new races of O. pygmaeum. ‘The birds collected
by Stevens on the Watut River in 1932 and already analysed by
Greenway were described as olivascens: “Very similar to nominate
pygmaeum but under parts with a conspicuous grayish olivaceous
tinge and upper parts slightly darker’ (Salomonsen, 1966, p. 2). Ex-
amination of the type series shows that the type (a male) stands out
obviously among the eight males collected by Stevens as having the
least yellow belly and lower flanks. Apart from this individual variant,
Stevens’ series is not distinguishable from other New Guinea popula-
tions. The name flavipectus was proposed for southern New Guinea
birds, with the type a bird collected by Rand in 1934 on the First
Archbold Expedition and already analysed by Mayr and Rand.
Southern New Guinea birds do average a shade paler above and
yellower below than the Vogelkop and northwestern New Guinea
population, but many specimens can be matched, and the average
difference is too slight for recognition.
DISCUSSION. ‘The Pygmy Honeyeater occurred from about 2,000
to 4,500 ft. It was commoner than the two specimens indicate, and
gathered in flowering trees, where it often accounted for 10 to 30%
of bird-usage and was twice as common in the upper half as in the
lower half of the crowns. In flocks of up to 10 it called incessantly and
was in constant motion. One was seen to hover. ‘The tailless silhouette
(the tail extends no farther than the wings) is confusable only with
Gerygone warblers, from which the hypernervous and social behavior
and the voice serve to identify it.
VOICE. A constant, weak, rapid train of spitted notes on the same
pitch.
Niche Differences in the Genus Myzomela
The five members of this genus contribute heavily to the bird-usage
of flowering trees in New Guinea (Terborgh and Diamond, 1970).
Apart from the erratically distributed second-growth species M.
357
SPECIES ACCOUNTS
adolphinae, the only form above 5,000 ft is M. rosenbergu. ‘Three
species (M. cruentata, M. nigrita, and M. eques) occur at lower allti-
tudes, and at Karimui these three plus M. rosenbergii were present
and often seen in the same tree. At Soliabeda M. nigrita, M. cruentata,
and M. eques fed in the same tree, while M. nigrita, M. cruentata, and
M. rosenbergii fed together at the Sena River. M. nigrita, M. cruentata,
and M. rosenbergii are very similar in size and habits, confining them-
selves to the top of the canopy to an even more marked degree than
does Oedistoma pygmaeum. Among the three species of myzomelids
at lower altitudes, M. eques sorts out ecologically on the basis of its
larger size (twice as heavy as M. nigrita and M. cruentata), presence
at sea level (where M. nigrita and M. cruentata are rare or absent), and
less extreme tendency to stay in the treetops. M. nigrita and M.
cruentata are very similar ecologically, and the only difference I have
noticed is that the altitudinal range of M. cruentata lies on the
average higher.
Myzomela eques karimuiensis Diamond
Red-spotted Myzomela
SPECIMENS EXAMINED. Karimui: 1 ¢ (3 July 1965).
WECHT. 18.
WING. 75.
EAL] 162:
CULMEN FROM BASE. 21.
STOMACH CONTENTS. Insects.
TAXONOMY. ‘This is another of the dark races from Karimui
(Diamond, 1967a, p. 9).
BREEDING. ‘The gonads of the unique type were somewhat
enlarged.
DISCUSSION. Of the five myzomelids in the Eastern Highlands.
M. eques is the largest (in the male) and is the only one to extend
regularly down to sea level. It was present at Soliabeda (2,000 ft) as
well as Karimui (3,650 ft) but not at any higher station. ‘Verborgh’s
observations at Karimui show that it accounted for 1-7% of the bird-
usage in flowering trees and was consistently several times less numer-
ous than M. nigrita but more numerous than M. cruentata or M.
rosenbergit. A possible indication of niche specialization among
myzomelids is provided by ‘Terborgh’s observation that M. eques,
although mainly a canopy species, spent about one-third of its time
in the lower half of the crown, whereas M. cruentata and M. nigrita
were virtually confined to the upper half of the crown. In addition, the
larger M. eques is somewhat slower moving than M. cruentata, M.
nigrita, or M. rosenbergit.
358
SPECIES ACCOUNTS
Myzomela adolphinae Salvadori
Mountain Red-headed Myzomela
DISCUSSION. ‘This myzomelid of second-growth, open country,
and gardens between 4,000 and 6,400 ft varies radically and unpre-
dictably in its abundance from place to place and from time to time,
perhaps due to local movements associated with flowering of trees.
Eastern Highlands records are from Kyaka territory and the Baiyer
Valley (Bulmer), Nondugl (Gyldenstolpe, Shaw-Mayer), and the Kubor
Mountains (Gilliard, Hitchcock). This species is present in the Vogel-
kop and Eastern Highlands but absent in the entire western half of
New Guinea outside the Vogelkop, a distribution shared only by
Zosterops novaeguineae. I have seen a singing male M. adolphinae
and a male M. rosenbergii in the same tree.
VOICE. ‘The song, repeated at 4 sec intervals, is suggestive of an
Empidonax flycatcher (Tyrannidae, North America). It consists of two
high-pitched notes, the first higher than the second, the first often
upslurred, the second often trilled. The song of Myzomela sclateri
is very similar.
Myzomela cruentata cruentata Meyer
Red Myzomela
SPECIMENS EXAMINED. Karimui: 4 ¢, 2 9 (2-12 July 1965).
WEIGHT. 3 ¢: 9.0 (3). 2 9: 7.0, 7.3.
WING. 3 4: 56, 57, 59.
BREEDING. ‘The gonads were greatly enlarged in three of the
males, very small in the fourth.
DISCUSSION. ‘The Red Myzomela was observed at Soliabeda
(2,000 ft), Karimui (3,650 ft), and the Sena River (4,500 ft). In flowering
trees at Karimui and Soliabeda it accounted for 1-4% of the bird-
usage and was much less numerous than M. nigrita or M. eques. At the
higher altitude of the Sena River M. eques was absent, M. nigrita
very uncommon, and M. cruentata and M. rosenbergit equally common
(16% of the bird-usage each). Like M. nigrita, M. cruentata is vir-
tually confined to the top of the canopy. The only ecological difference
I can observe between these two species is that the altitudinal range
of M. cruentata is centered slightly higher (M. nigrita commonest at
1,500-3,500 ft, M. cruentata commonest at 2,500-4,500 i)
Myzomela nigrita meyeri Salvadori
Black Honeyeater
SPECIMENS EXAMINED. Karimui: 4 ¢,1 9, 2 imm. 4, 2? (3 July-5 Aug.
1965). Soliabeda: 2 ¢ (23-27 July 1965).
coy
SPECIES ACCOUNTS
WEHRIGHT. 6 @: 9.8, 100 45105, 1192833, ~2 mmm. 6:8:3, 1008 2729012),
WING. © 4: 58 (4),.59) Gl. iO; 54, 2)imm, 62.57 (2):
TAXONOMY. In size these are near the borderline between the
widespread race meyeri (wing, ¢, 58-64) and the smaller Aru Islands
and southern New Guinea race nigrita (wing, ¢, 55-58), but are
within the range of meyeri. ‘The female and one of the unsexed birds
are Olive-brown to gray-brown, with a rose chin and forehead, while
the adult males are all black. The two immature males are black except
for a small rose area on the chin.
BREEDING. ‘Testes were greatly enlarged in one adult male,
somewhat in two, at Karimui (3,650 ft); not enlarged in the two from
Soliabeda (2,000 ft); and somewhat enlarged in one of the two im-
mature males, not in the other, at Karimui.
DISCUSSION. In flowering trees at Karimui and Soliabeda this
was the commonest myzomelid, nearly three times as common as the
other three species combined. On the average, it accounted for 10%
of the bird-usage. At the Sena River it was much less common than
M. cruentata and M. rosenbergiu and accounted for only 1% of the
bird-usage. Like M. cruentata, M. nigrita is a canopy species: out of 77
individuals whose position in the upper or lower half of the tree was
noted, 73 were in the upper half.
Myzomela rosenbergi rosenbergit Schlegel
Red-collared Myzomela
NATIVE NAME. Fore: pani.
SPECIMENS EXAMINED. Mt. Michael: 1 ¢ (3 July 1964). Karimui: 2 9 (3
July 1965). Mt. Karimui Zone 2: 6 ¢,1 9,2 imm. ¢; Zone 3: 1 imm. ¢; Zone
4; 1 43 Zone 5: > ¢, 1 mmm. ¢; Zone 7: 1 O53 Zones’: 2 Us Aug Sept a86p):
WEIGHT. 15 4: 9.3-12.7 (11.0+ 2.5). 4 2: 9.0-10.7 (9.7 + 0.6).
WING. 15 ¢@: 61-67 G41). 4 92 58:61 69-1).
STOMACH CONTENTS. Insects.
TAXONOMY. Gyldenstolpe (1955, p. 155) separated Eastern High-
lands birds (wahgiensis) from nominate rosenbergii of the Vogelkop.
However, comparison of my adult males with adult males collected
in the Vogelkop (topotypical rosenbergit), Wahgi Valley (topotypical
wahgiensis), southeastern New Guinea, and at Telefolmin shows that
all series are identical except for the more recently collected specimens
being slightly more pure black, less blackish-brown below, due _pre-
sumably to foxing. The race wahgiensis accordingly must be considered
synonymous with rosenbergii, as concluded by Gilliard and LeCroy
(JOGU pe 75);
BREEDING. In the Karimui area gonads were greatly enlarged in
all the many adult males examined, but not in the females.
DISCUSSION. ‘The sex ratio on Mt. Karimui is very unbalanced
(males outnumber females 7:1), as is frequently true in museum col-
360
SPECIES ACCOUNTS
lections of myzomelids. Two factors may contribute, in addition to
the possibility that the ratio really is unbalanced in nature. (1) Rand
and Gilliard (1967, p. 533) suggest that “males and females have
different habitat preferences; the females instead of visiting flowering
trees more often feed singly among the foliage, and perhaps lower 1n
the forest.” Most myzomelids are, in fact, collected in flowering trees
where they congregate in large numbers and are easily secured, so
that a preference of males for such trees might explain the unbalanced
ratio in museums. I found that on Mt. Karimui males outnumbered
females 12:1 in flowering trees but only 5:3 away from flowering trees.
Corresponding figures on Mt. Albert Edward were 3:2 and 1:1, as on
Mt. Karimui and in agreement with the suggestion of Rand and Gil-
liard. (2) Males and females may have different altitudinal preferences.
Bulmer found that the proportion of adult males was much lower at
4,000-5,000 ft than at the higher altitudes up to 10,000 ft. At Karimui
(3,650 ft) both specimens and most birds seen were females (or im-
matures).
Myzomela rosenbergii is fairly common from about 4,000 to at least
10,000 ft in forest and second-growth. Its local abundance appears to
vary enormously in time and space according to the presence of flower-
ing trees. On Mt. Karimui’s west ridge during the month we were
there, there were four such trees: at 4,250 ft, just above Zone 1 and
beyond collecting and censusing limits; at 4,450 ft in Zone 2; at 6,490
ft in the upper half of Zone 5; and a fourth in Zone 3 on the side of
the ridge, where we collected only very briefly. This is reflected in the
relative abundance figures: Zone 1, 0.0% of the local avifauna; Zone 2,
In, Zones, [97 Zone 4.27, lower halt ot Zones), 0.09.5 awpper
halt of Zone, 97,; Zone.6,,0.0%,. Zone 7, 29,; Zones, 129. IM. resem
bergu reached the bottom of its altitudinal range at Karimui and was
the rarest of the four myzomelids there.
M. rosenbergii was netted frequently in high-altitude moss forest on
Mts. Karimui and Michael, but not at lower altitudes, indicating that
it stays mainly in the treetops until the forest becomes stunted.
VOICE. The call is a high-pitched upslurred “tswi” or a high
“ts-ts”. ‘he song is an energetic, breathless, high-pitched sibilant trill
or rapid alternation between two notes on different pitches.
Meliphaga analoga group
Anyone who has had to work with the nine exceedingly similar
honeyeaters of the Meliphaga analoga group on mainland New Guinea
will agree that they pose the most difficult problems of taxonomy, field
identification, and niche definition in the New Guinea avifauna. All
can be briefly described as small honeyeaters with olive upper parts,
gray underparts, and a yellowish or whitish auricular spot. Six of these
nine species occurred sympatrically and abundantly in the Karimui
361
SPECIES ACCOUNTS
area. My general observations on taxonomy, voice, ecology, and field
identification are summarized here and are followed by the individual
species accounts.
Taxonomy.—One’s first impression on starting to work with this
group is that the forms of New Guinea Meliphaga are a hoax per-
petrated by previous workers and constitute just a single form showing
continuous and minor individual and geographical variation. With
practice, however, it becomes possible to identify many specimens in
the hand without measurements and to identify most (but not all!)
of the rest by measurements (cf., Table 15). The grouping of popula-
tions in different areas into species was long a subject of controversy
and confusion until clarified in a fine review by Rand (1936), the
validity of most of whose conclusions has survived subsequent dis-
coveries of new populations. Rand’s classification appears to me at
present to require modification in only two or three respects: the
southern New Guinea populations auwga and setekwa, which Rand
grouped under M. montana, constitute a distinct species, M. auga
(p. 368); the population citreola of northern New Guinea belongs to
M. orientalis, not to M. analoga (Diamond, 1969, pp. 38-46); and
M. auga may be a synonym of the form known as M. albonotata. ‘The
characters by which I distinguished the forms of the six species oc-
curring in my study areas are listed below. It is necessary to realize
that the minor differences between different species at one place are
often less marked than the minor differences between different popula-
tions of the same species, so that this list will not necessarily be
adequate in other parts of New Guinea.
I. M. auga auga. Readily identified by the whitish auricular spot
of the adult, contrasting with the yellow spot of other species. ‘The
only possible sources of confusion are that the spot is pale lemon to
yellowish in immatures, and that it is pale lemon to whitish in the
uncommon M. flavirictus. ‘Vhe underparts are a dull, uniform, rather
dark olive, very similar to the other species except M. flavirictus. ‘The
underwing coverts are olive-ochraceous, slightly more yellow and paler
than in M. mimtkae, averaging more olive than in M. orventalis, and
distinct from the yellower coverts of the other three species. ‘The inner
edges of the primaries are pale olive, sometimes olive-ochraceous. ‘The
size (wing, tail, and weight) is similar to M. mimikae and M. aruensis,
and larger than the other three species (Table 15).
2. M. flavirictus flavirictus. The rarest of the six species. Best dis-
tinguished by the auricular spot, which is pale lemon to nearly white
(more yellow anteriorly); the marked and bright yellow rictal streak
(a streak extending from the base of the bill to below the eye), better
developed than in the other species; the distinctly yellow chin, more
so than in M. aurensis or M. analoga; the lower mandible, which is not
black, as in adults of the other species, but horn-colored; and the legs,
which are less dark than in the other species, orange on the rear sur-
362
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SPECIES ACCOUNTS
face and dull olive on the front surface. None of these characters is
infallible taken alone (e.g., immatures of other species may duplicate
the bill and leg color), but the combination is distinctive. The upper-
parts are pale, dull, and gray and fairly distinct from the more olive
and brighter color of the other species. The underwing coverts are
whitish to yellow, not at all olive or ochraceous. The inner edges of
the primaries are pale yellow. The underparts are pale, clear, and
rather lemon as in M. analoga and M. aruensis and unlike the other
three species. The size (wing, tail, bill, weight) is nearly as small as
M. orientalis, somewhat smaller than M. analoga, and significantly
smaller than the other three species.
3. M. mimikae bastille. The underparts are darker and more un-
even (more spotted) than in the other species but are approached by
M. orientalis. The underwing coverts are olive-ochraceous (occasionally
more olive-yellow), and the inner edges of the primaries are near
ochraceous (sometimes olive-buff to ochraceous). ‘The auricular spot is
yellow and of medium size. The bill is stout. The size is large, as in
M. auga and M. aruensis. No single mark of this species is infallible,
but the spotted underparts are the best mark, and the combination
of the above traits is generally diagnostic. The most similar species is
M. orientalis, from which M. mimikae may be recognized by the
significantly larger size and stout bill.
4. M. arvensis aruensis. The auricular spot is yellow, and averages
larger than in the other species. ‘The feathers of the rump form a
blackish tuft which stands up somewhat. ‘This offers the best, though
not an infallible, mark. ‘The underparts are pale and evenly colored
with a yellowish chin, and resemble M. analoga and M. flavirictus.
‘The underwing coverts are yellow or yellow-orange, not ochraceous or
olive. ‘Ihe inner edges of the primaries are yellow, sometimes yellow-
buff. The size is large, as in M. auga and M. mimikae, and the length
of the wing and tail averages greatest in M. aruensis. However, the
stout bill is shorter than in the other five species, including the three
smaller ones (M. analoga, M. orientalis, M. flavirictus). ‘The face is
rather blackish between the eye and the auricular spot. ‘The most
similar species is M. analoga, from which the combination of the rump
tuft, stout bill, longer wing and tail, and larger auricular spot separate
i,
5. M. orientalis facialis. ‘The combination of small size, spotted
underparts, long and slender bill, and somewhat ochraceous under-
wing are useful characters. ‘Ihe wing and tail are shorter, and the
weight less, than in other species except M. flavirictus (those attempt-
ing to use this key in other areas should be warned that M. gracilis,
which did not occur in my study areas, is equally small, and that some
populations of M. orientalis in other parts of New Guinea are not so
small). The slender bill is as long as in the large M. auga and M.
mimikae, and more slender than in these or M. aruensis. The under-
564
SPECIES ACCOUNTS
parts are darker and more uneven (more spotted) than in other species
except M. mimikae, which is slightly darker and more spotted. ‘J he
auricular spot is smaller than in most other species and bright yellow.
The underwing coverts are variable, and usually rather orange, gen-
erally ochraceous olive to ochraceous yellow, occasionally more yellow.
The inner edges of the primaries are generally ochraceous olive. ‘The
most similar species are M. mimikae, which has similar underparts but
is larger and has a stouter bill; M. analoga, which is somewhat larger
and differs in the underparts and underwing; and M. gracilis, which
has considerably paler and duller upperparts and paler and even-
colored underparts.
6. M. analoga analoga. This species has no single feature that could
be called distinctive, and must be recognized by a combination of
characters, each shared individually with other species. ‘The underparts
are pale, even, and somewhat yellowish on the chin, as M. aruensis
and M. flavirictus and unlike the other three species. ‘The size is some-
what smaller than M. auga, M. mimikae, and M. aruensis and some-
what larger than M. orientalis and M. flavirictus. The underwing
coverts are yellow, similar to M. arwensis but slightly paler. ‘The inner
edges of the primaries are yellow, as M. aruensis. ‘The yellow auricular
spot is slightly smaller than in M. aruensis and larger than in M.
orientalis. The bill is slender and averages longer than in all the other
species. The most similar species are M. aruensis, M. orientalis, and
M. flavirictus, under which the distinguishing marks are listed.
V oice.—Regretably, meliphagas are as similar and undistinctive in
voice as in appearance. Meliphagas call frequently. With one exception,
all the calls I heard were a single, short, and musical note which can
be rendered as “chip”, “chup”, “tup”, or “tuck” and is either on con-
stant pitch, slightly upslurred, or slightly downslurred. In the Eastern
Highlands I identified this call positively from M. auga and M.
orientalis, with high probability from M. mimikae, and elsewhere in
New Guinea from M. montana and probably from M. aruensis and
M. analoga. ‘The one good vocal field mark is that M. auga has in
addition a distinctive song.
Ecology.—Of the six meliphagas in the Karimui area, five coexisted
at Karimui, four at Soliabeda, and three at Bomai. Niche differences
involve habitat preference, altitudinal range, vertical distribution in
the vegetational column, and probably feeding habits. (1) M. auga
was strikingly confined to second-growth and the forest edge and was
completely absent from the forest interior. (2) Much of the altitudinal
range of M. orientalis lies above that of all other meliphagas, though
it meets four other species at the lower extremity of its range (Karimui).
M. orientalis and M. analoga are ecologically similar and have mutually
exclusive altitudinal ranges. M. mimikae and M. auga are also hill
forest species which rarely reach sea level and have a higher ceiling
than the other species except for M. orientalis. Of the three species
365
SPECIES ACCOUNTS
which reach sea level, M. analoga has a slightly lower ceiling than M.
aruensis or M. flavirictus, and the ceiling of M. gracilis is so low that
it failed to reach Soliabeda, my lowest station. (3) Netting returns
suggest some differences in vertical preferences. M. auga, M. mimikae,
and M. aruensis were often netted, M. orientalis much less often, and
M. analoga and M. flavirictus never. (4) The differences in the bills
suggest differences in feeding habits. The bills of the three lowerstory
species (M. auga, M. mimikae, and M. aruensis) are stout, those of the
three middlestory species (M. orientalis, M. analoga, and M. flavirictus)
slender. Meliphagas in general glean for insects, visit flowering trees,
and visit fruit trees, but deciding just which species is doing what 1s
usually difficult. At Karimui Terborgh found meliphagas accounting
for 7-30% of bird-usage in flowering trees, 66-76% in trees with small
fruits, and 44% in trees with woody fruits opened by parrots. M.
flavirictus was shot both in a flowering tree and in a tree with small
fruits, and M. orientalis was shot in a flowering tree and observed at
Miarosa in fruiting trees. The white-eared M. auga was also recognized
in flowering trees. Far more information on stomach contents and on
what specimens were doing when shot is necessary.
Iield identification.—M. auga can usually be identified by its white
ear and by its song, though not by its call. Field identification of the
yellow-eared meliphagas is at present impossible unless one has care-
fully compared specimens of the forms known actually to be present
in the study area, and then only after much practice and under the
most favorable conditions of light and proximity imaginable. On two
such occasions I felt confident that I was watching M. mimikae be-
cause of the uneven underparts. On one occasion I felt that I was
watching M. aruensis because of the large and bright yellow auricular
spot and dark face. ‘The small and bright yellow auricular spot of
M. orientalis might prove of some use as a field mark. Identification of
yellow-eared meliphagas held in the hand, or prepared as study skins,
should not be attempted without comparative material.
Meliphaga aruensis aruensis (Sharpe)
Puff-backed Meliphaga
NATIVE NAMES. North Fore: augarié. South Fore: agerabite. Gimi: loho.
Daribi: sobadage. (The same native names are applied to all species of the
Meliphaga analoga complex).
SPECIMENS EXAMINED. Karimui: 2 ¢,. 1 @ (8 Aug. 1964; I and 11 July
1965). Soliabeda: 1 g,2 Q (22 July 1965).
WEIGHT, 2°62 27,29, 3 O20, 255 Ain
WING. 3 42 87, 90, 94. 3 O72 83, 57; 38.
TAM. 8: AxG7, 74, 771. Oe O75 Ty As
GULMEN FROM BASE. 3 @: 19.0, 200; 205. 3 9% 18.5, 1910; 195.
EXOSED CULMEN. 3 @: 145; 15.0) 165. 38 Oi 140, I5.0)\@);
TAXONOMY. The yellow auricular patch is square, rather than
elongated as in the race sharpeti.
366
SPECIES ACCOUNTS
BREEDING. The testes were small in one male, somewhat en-
larged in another.
DISCUSSION. Three of the specimens were netted. This large,
stout-billed meliphaga of the lowerstory reached its altitudinal ceiling
at Karimui (3,650 ft).
Meliphaga auga auga Rand
Southern White-eared Meliphaga
SPECIMENS EXAMINED. Karimui: 19 ¢,17 9,1 imm. 9, 1 imm. ? 1-16 Aug.
1964; 1-17 July 1965). Bomai: 3 ¢,2 9 (6-8 July 1965). Soliabeda: 1 g,4 @ (23-
28 July 1965).
WEIGHT. 14 4: 27-34.(50.7 =: 2.3), 13 9: 22-29 (25.6.ce 1.8).
WING. 7 2 62-9) (86.932 158), 18 9: 77-85 (79-5 =e ING),
TAIL. 17 4: 66-78 (71.8 + 3.0). 17 9: 63-70 (67.1 + 1.9).
CULMEN FROM BASE. 17 ¢: 2010-225 @l.1 +07), 18 Q: 1810-220 (199 4.
0.8).
EXPOSED CULMEN, 17 @: 16.5-19.0, (17.6 3: 0.7). 18 9: 150-180 (60 22 0:8).
CULMEN FROM ANTERIOR EDGE OF LATERAL FEATHERING. 13 ¢:
15.5-17.0 (16.1 +0.5). 16 9: 14.0-16.5 (14.7 + 0.8).
TAXONOMY. The range and average values of measurements for
five males (including the type) and four females of nominate auga
from southeastern New Guinea are: wing, ¢ 82-88 (86.0), @ 75-80
(77.8); tail, $ 68-75 (72.4), @ 65-67 (66.0); culmen (from base), g 20-22
(21.3), @ 19-20 (19.3). Birds from the Karimui area thus average a
trifle larger, and are very slightly duller and darker above, but other-
wise match auga well. The type and paratype of setekwa differ in the
slightly more orange axillaries, dirtier breast, and smaller size. Wahgi
birds were separated by Gyldenstolpe and Gilliard (Gyldenstolpe, 1955,
p. 166) as gretae, on the basis of having “‘a longer tail and somewhat
longer wing; general coloration above and on the chest richer green”.
Specimens of gretae were not available for comparison, but average
sizes calculated from the measurements cited in the original diagnosis
(wing ¢ 89.7, 9 81.0; tail g 72.4, 9 68.0 in gretae) are only slightly
larger than in Karimui or southeastern New Guinea birds, and I doubt
that gretae differs from auga. Gilliard and LeCroy (1967, p. 78) tenta-
tively referred 14 white-eared meliphagas from the Adelbert Moun-
tains to gretae, but examination shows that these specimens are close
to the distinct and heavily spotted form M. montana huonensis (of
which Gilliard and LeCroy lacked comparative material) and could
not belong to the unspotted gretae (= auga?).
BREEDING. At Karimui testes were moderately or greatly en-
larged in 1964, slightly or moderately enlarged in 1965. Ovaries were
generally small except for being enlarged in one 1964 Karimui female.
One nesting female was collected at Soliabeda. The nest was 15 ft
above the ground in a bamboo thicket at 1,400 ft near a small river
and held two young. In shape it was a cup 21% inches bigh and with
367
SPECIES ACCOUNTS
an inside diameter of 2 inches. The outside was constructed of green
moss; the inside, of dry strands of grass arranged concentrically, with
a few paper-thin chips of dry bark; and there was a lining of an un-
identified, fine, soft, white, cotton-like material (possibly the same as
the material Rand (1942a, p. 362) found in a nest of this species and
identified as “plant down from the silk cotton tree’).
DISCUSSION. In my study areas Meliphaga a. auga differed strik-
ingly in its ecology from other species of Meliphaga in being strictly
confined to second-growth and the forest edge and completely absent
from forest interior remote from the edge. It was common in the
Karimui area from 1,350 to 4,000 ft but absent at Okasa (3,550-4,250
ft), despite suitable habitat. Gilliard collected one specimen, and
Gyldenstolpe 14, at Nondugl (5,200 ft). Gyldenstolpe’s (1955, p. 167)
comments on habitat duplicate my experience: “..... rather common
in patches of second-growth around Nondugl. Never observed in the
forests of the Wahgi Divide.” Schodde and Hitchcock (1968, p. 66)
met M. auga in similar habitat at Lake Kutubu (‘in the lower stages
of tall secondary and marginal primary forest’). In the feeding trees
studied by Terborgh he found white-eared meliphagas (M. auga) out-
numbering yellow-eared individuals (four species) by about two-to-one.
Solitary individuals of M. auga were also common in shrubbery 3-15 ft
above the ground, where one was seen eating an insect, and were
frequently netted.
In his review of the meliphagas Rand (1936) assembled eight white-
eared forms, viz., auga, setekwa, montana, sepik, steini, germanorum,
huonensis, and aicora, to which must be added a subsequently dis-
covered form margaretae (Greenway, 1966, p. 22), into the single
species M. montana. Rand noted, however, that the resulting “species”
showed more geographic variation than any other in the Meliphaga
analoga complex. More recent evidence suggests that two species are
involved which are largely allopatric but overlap at Huon Gulf and
possibly on the Huon Peninsula and in the Vogelkop and Weyland
Mountains: a second-growth species with a more yellow-olive under-
wing in southern New Guinea, M. auga (races auga and setekwa, the
former possibly a synonym of albonotata; see next paragraph); and
a forest species with a more ochraceous underwing in northern New
Guinea, M. montana (races montana, margaretae, steini, sepik, ger-
manorum, huonensis, and aicora). ‘The evidence is as follows. (1) Of
the two forms of which I have field experience, auga (southern slopes
of Eastern Highlands) and seprk (North Coastal Range), auga is con-
fined to the forest edge and second-growth, while sepik lives in the
forest interior. The taxa auga and sepik have very distinct calls, unlike
each other’s or that of any other meliphaga. Ecologically, sepik ap-
pears closer to the yellow-eared species M. mimtkae of southern New
Guinea. Except for its yellow ear, the large, stout-billed, spotted M.
mimikae suggests (and is allopatric to) the northern New Guinea
368
SPECIES ACCOUNTS
populations grouped under M. montana and is suspiciously similar to
margaretae, as already noted by its describer, Greenway. However,
M. mimikae bastille also lacks the distinctive call of sepik, so that I
do not think that sepik and M. mimikae bastille are actually con-
specific. (2) The inner edges of the primaries are ochraceous in the
populations I group under M. montana, predominantly yellow-olive
in auga and setekwa. (3) A white-eared meliphaga collected at Wau
(Territory of Papua and New Guinea Museum. No. B 334) belongs
unmistakably to auga. Wau lies halfway between the type localities
of the quite different, ochraceous-underwinged races aicora (85 mules
to the south) and hwonensis (80 miles to the north). ‘Vhus, M. montana
and M. auga are probably sympatric at Huon Gulf. If auga is a
synonym of albonotata (see next paragraph), then the two species are
also sympatric on the Huon Peninsula and in the Vogelkop and Wey-
land Mountains. Field observations of voice and habitat preference
of the other populations besides auga and sepik will be necessary to
test this proposed grouping.
The remaining white-eared meliphaga is the form known as M.
albonotata, described in 1876 by Salvadori from D’Albertis’ collection
on the southern slope of southeastern New Guinea and subsequently
reported from the Vogelkop and Weyland Mountains by many col-
lectors, from southwestern New Guinea by Ogilvie-Grant (1915), and
from the Huon Peninsula by Mayr (1931). I have examined Ogilvie-
Grant’s specimens, which are in the British Museum of Natural History
and are close to M. auga setekwa; and I have also examined 13 speci-
mens from the Vogelkop and Weyland Mountains in the American
Museum of Natural History, which are also close to M. auga setekwa,
differing only in the slightly brighter olive back and very slightly
shorter wing and more slender bill. ‘The type of M. albonotata from
southeastern New Guinea was not available to Rand when he de-
scribed auga from southeastern New Guinea, nor was it available to
me, but Salvadori’s (1881, Vol. 2, p. 333) description of M. albonotata
fits the type of auga well. All white-eared meliphaga specimens in the
American Museum of Natural History and in the Territory of Papua
and New Guinea Museum from the southern slope of southeastern
New Guinea apparently belong to the same population as the type of
auga. It may turn out that auga is a synonym of albonotata, and that
the Vogelkop-Weyland population is conspecific with auga (= ? albo-
notata) and setekwa and requires naming.
VOICE. — Since white-eared meliphagas were identifiable by sight
in the field, the vocalizations of this species could be identified with
certainty. ‘There are two call notes: a short, cheerful, musical, and
fairly loud “chip” or “tup” with a sucked-in quality (perhaps more
sucked-in than the call of M. orientalis?); and a very similar disy]-
labic note. ‘These calls are not distinguishable from the calls of the
yellow-eared meliphagas. ‘I'he song is a series of 10-12 thin and quieter
369
SPECIES ACCOUNTS
notes at the rate of about 5 per second, dropping in pitch towards the
end and often rising slightly in pitch at the beginning. No yellow-
eared meliphaga gave this song.
Meliphaga mimikae bastille Diamond
Large Spot-breasted Meliphaga
SPECIMENS EXAMINED. Okasa: 1 4,1 9 (22 and 24 June 1965). Sena River:
4 8,1 9 (26 and 27 July 1965). Karimui: 18 ¢,17 9 (10-14 Aug. 1964; 1 July-6
Aug. 1965). Bomai: 1 ¢, 14 9 (6-9 July 1965). Soliabeda: 6 6,9 9 22-30 July
1965). Mt. Karimui Zone 1: 4 ¢,1 9 (9-12 Aug. 1965).
WEIGHT. 25 4: 24-33 (28.5 + 2.0). 41 9: 29-99 (25.4 + 1.7).
WING. 30 @: 82-91 (864+ 2.7). 42 9: 75-82 (79s = 109),
TAIL. 31 4: 67-75 (710+2.6). 43 9: 61-68 (65.2 + 1.8).
CULMEN FROM BASE. 27 4: 20.0-23.0 (ZIT =5 O01). ©4219): W8ib-2 0 ba(20 eee
0.8).
EXPOSED CULMEN. 26 @: 17:0-2010 (8 Jie (058). 39°; 15,0218-5) (1677 = 09}.
TAXONOMY. ‘The diagnosis of this race has been given previously
(Diamond, 1967a, p. 12).
BREEDING. ‘Testes were large in males taken in 1964 (Sena River
and Karimui). In 1965 testes were slightly enlarged in three out of four
males on Mt. Karimui and in two males at Soliabeda but were small
in most males from Karimui and Bomai.
DISCUSSION. M. mimikae was the commonest forest meliphaga,
and the commonest yellow-eared meliphaga, in my study areas up to
4,200-4,500 ft, above which altitude it disappeared. Netting figures
suggest that it spends much time in the lowerstory, since 48% of the
specimens were netted. The species is confined to the hills of southern
New Guinea, mainly between 500 and 4,500 ft, and consists of three
races known from the following localities. M. m. mimikae, south-
western New Guinea: Mimika, Kapare, Wataikwa, Iwaka, Setekwa,
and Utakwa Rivers, from sea level to 2,500 ft but with the great
majority of the specimens collected in the hills (Ogilvie-Grant, 1915,
p. 67); upper Setekwa River and southern slopes of the Snow Moun-
tains, 3,000 ft (collected by A. S. Meek); Alkmaar on the Noord River,
unspecified but probably low elevation, one specimen reported (Junge,
1939, p. 60); Fly River, only on the hilly upper reaches at elevations
above 260 ft (collected by Rand, Second Archbold Expedition). M. m.
bastille, southern slopes of the Eastern Highlands, 1,350-4,500 fe:
Karimui and Okapa areas (my collections). M. m. granti, southeastern
New Guinea, no records below 2,500 ft: Mafulu, 4,100 ft (collected by
Rand, First Archbold Expedition); Aroa River, 4,000-6,000 ft (collected
by A. S. Meek); Hydrographer Range, 2,500 ft.
An adult male yellow-eared meliphaga (A.M.N.H. 342,965) collected
by Rand at Bernhard Camp (elevation, 160 ft) in the northern New
Guinea lowlands during the Third Archbold Expedition has been
used by Salomonsen (1966, p. 4) as the unique type of a new race,
370
SPECIES ACCOUNTS
M. mimikae rara. Rand (1942, p. 508) identified this specimen as M.
analoga flavida and pointed out that it differed in color from the 28
other specimens he collected of this form but that it was virtually
identical to M. a. analoga from the upper Fly River. Examination of
this specimen shows that it is darker-backed than other M. a. flavida
specimens and superficially not unlike but also not closely similar to
the known races of M. mimikae. My measurements of it yield: wing,
83; tail, 69; exposed culmen, 18.5; culmen from base, 22.0. ‘These
measurements lie towards the upper limit for M. analoga and at the
extreme lower limit for M. mimikae (only three of 31 males of M.
mimikae measured have shorter wings). Against the attribution of this
specimen to M. mimikae are the additional facts that the type locality
would provide the sole northern New Guinea record; the locality is
300 miles along the hill slopes from the nearest place of known oc-
currence of M. mimikae; the elevation would be unusually low, though
not unprecedented, for M. mimikae; and only a single specimen was
taken although Rand’s collection was exhaustive and M. mimikae has
proved abundant at most localities where it occurs. Given these facts,
the difficulty of identifying even large series of New Guinea meli-
phagas, and the likelihood that this specimen is merely an individual
variant of another species, it seems inadvisable to take a single speci-
men as evidence for a new race and large range extension of M.
mimtikae.
VOICE. Since M. mimikae could rarely be distinguished from other
yellow-eared meliphagas in the field, the call specifically of this species
was not identified with certainty. However, at localities in the Karimui
region where M. mimikae and other yellow-eared meliphagas were
present, all vocalizations of yellow-eared meliphagas consisted of a
single very similar call note, described on p. 365. Since 60-95% of the
yellow-eared meliphagas were M. mimikae, I assume that it was re-
sponsible for most of these vocalizations.
Meliphaga orientalis facialis Rand
Small Spot-breasted Meliphaga
SPECIMENS EXAMINED. Awande: 3 ? (20 June 1964). Miarosa: 1 ? (28 June
1964). Okasa: 2 ¢,1 9 (24-26 June 1965). Karimui: 5 @,6 9 (8 Aug. 1964; |]
july-> Aug. 1965). Mt. Karimui: Zone 1: 1 9; Zone 2: 1 @, 2 9; Zone 3: 2 4,1
@ (11-17 Aug. 1965).
WEIGHT. 9 4: 16-20 (1861.7). Il 9: 16-20 (176 = 1.2).
WING. 10 @: 71-78 (7482 2.0). 12 9: 66-71 (68.8 + 1.5).
TAIL. 10 ¢: 56-62 (59.9+1.9). 12 9: 52-58 (56.4 + 1.9).
CULMEN FROM BASE. 10 ¢: 20.5-23.0 (21.6+ 0.8). 11 9: 19.0-21.0 (20.24
m7).
eOseD CULMEN. 10 6: 175-195 (184+ 0.8). 11 9: 16.0-18.5 (17.5 + 0.9).
CULMEN FROM ANTERIOR EDGE OF LATERAL FEATHERING. 3
15.0; 17.0, 185. 4 95 16, 17.0, L7-5, 1755.
—
ns
TAXONOMY. ‘The various populations grouped under facialis
371
SPECIES ACCOUNTS
are not identical, but the differences are not sufficiently marked or
well-established to justify naming of new races. The principal differ-
ences are that birds from the Vogelkop (3 ¢, including the type: wing,
71, 75, 77; tail, 56, 59; 61: culmen from base, 21.5, 22.0) and the
Weyland Mountains (2 6, wing, 78, 79; tail 63; 06; culmenurom: base,
22, 22) have longer wings and tails than those from the southern slope
of the Snow Mountains (wimer so 745 2 =o OoRandy legen alae cs
2 2 56 and 57; culmen from base, 1 ¢ 22.0,2 2 20.0 and 20.0) or my
series. In addition, my birds show some approach to nominate
orientalis of southeastern New Guinea in that the underparts are less
spotted, and the upperparts darker, grayer, and less bright, than in
other facialis populations, but they are still nearer facialis than
orientalis in these respects. The nearly black postocular stripe of
orientalis is greatly reduced in facialis, including my specimens. ‘The
race becki is considerably larger and still less spotted below, while
citreola (formerly considered a race of M. analoga; Diamond, 1969)
is yellower.
BREEDING. Gonads were small in all specimens.
DISCUSSION. In my study areas the altitudinal range of M.
orientalis lay for the most part above those of its congeners. In the
Okapa area it was the only meliphaga at Awande (up to 6,200 ft) and
Miarosa (5,800 ft), and shared Okasa (3,550-4,250 ft) only with M.
mimikae. In the Karimui area it descended to Karimui (3,650 ft),
where four other species were present, but not to Bomai (3,250 ft); it
shared Zone | of Mt. Karimui (4,000-4,200 ft) only with M. mimikae;
and it was the only meliphaga on Mt. Karimui above 4,400 ft until it
dropped out around 5,600 ft. Its maximum abundance (5-79% of the
local avifauna) was at 4,400-5,600 ft. The fairly abrupt disappearance
above this limit may have been correlated with the increase in abun-
dance of Ptiloprora guiset, while the decline in abundance at lower
altitudes may have been correlated with the appearance of the other
meliphagas, particularly the similar M. analoga.
Two of the specimens on Mt. Karimui were shot in the flowering
tree at 4,450 ft. The habitat appears mainly to be the forest interior,
though one was collected and another seen in a tree in a garden at
Miarosa. Only four specimens were taken in nets. Of these four netted
birds, three were obtained on a single day at Awande near the forest
edge, the fourth at Karimui, and none was netted in the undisturbed
forest of Mt. Karimui. Six specimens were in trees or the tops of trees
when shot. Probably M. orientalis is more of a middlestory and upper-
story than lowerstory species in the forest interior, and utilizes the
lowerstory mainly at the forest edge. Its altitudinal range excludes
that of the similar M. analoga, the slender-billed middlestory forest
meliphaga of lower elevations (cf., also, Diamond, 1969).
VOICE. Since this was the only member of the genus in Zones 2,
Biz
SPECIES ACCOUNTS
3, and 4 of Mt. Karimui, calls of meliphagas in these zones are as-
sumed to be of this species. ‘here are three similar call notes: a
bright, short, monosyllabic “tuck”; a bright, short, slightly disyllabic
(or else slightly upslurred) note, with the second syllable slightly higher;
and a similar disyllabic call except with a thinner quality and with the
second note lower. These calls were indistinguishable from calls of
yellow-eared meliphagas at Karimui, Bomai, and Soliabeda, assumed
to come mostly from M. mimikae.
Meliphaga analoga analoga (Reichenbach)
Mimic Meliphaga
SPECIMENS EXAMINED. Bomai: 1 ? (8 July 1965). Soliabeda: 3 6,4 Q (23-
30 July 1965).
WEIGHT, 3 6: 24,25, 26. 4-021 (6), 22), Meer 21,
WING. 3 4: 80; 93, 83. 4 0: 75, 75, 77, 78. 1 ?2 80!
TAIL. 3 @: 64, 65, 70. 4 9: 60, 61, 63, 64. 1 ?: 65.
CULMEN BROM BASE. 3. 6:°23.0; 235,240) 3 02-21,0, 20h, 22.0)" 1 ri92 ip,
EXPOSED “GUEMEN, 3 4: [80 185, 185. 3.9% 160ml io, so, Ars 180.
STOMACH CONTENTS. Fruit.
TAXONOMY. In his review of the meliphagas Rand (1936, p. 14)
pointed out, “All of the populations included in this race [analoga|]
are not identical but the differences between various groups of in-
dividuals are not sufficient to be used in separating races.” “Iwo new
races, connectens (from Beck’s Madang series) and papuae (from
Rand’s Wuroi series) have been described by Salomonsen (1966, p. 5)
from the material already discussed by Rand. Gilliard and LeCroy
(1968, p. 36) reaffirmed that connectens is not separable from flavida.
The type of flavida (type locality Japen) is the yellowest of the 15
Japen specimens I examined, and the average color differences among
flavida populations from Japen, Madang, Wewak, Hollandia, and the
Idenburg River are irregular and trivial. Several of the characters used
to separate papuae (type and paratypes examined) from analoga had
already been pointed out by Rand and are equally irregular and
trivial.
BREEDING. Gonads were small in all specimens.
DISCUSSION. None of the specimens was netted. ‘This meliphaga
appears to replace M. orientalis as the slender-billed meliphaga of the
forest middlestory at lower elevations. In the Karimui area the transi-
tion lay somewhere between 3,250 ft (highest record of M. analoga)
and 3,650 ft (lowest record of M. orientalis).
Meliphaga flavirictus flavirictus (Salvadori)
Yellow-gaped Meliphaga
SPECIMENS EXAMINED. Mengino: 2 ¢ (16 July 1964). Karimui: 1 9, 1 ?
(3 July and 3 Aug. 1965).
373
SPECIES ACCOUNTS
WEIGHT I 262-20, 1 227.
WING, 2 245 Tie, lire 7s.
TEST 92. So Gos OF) TL rho:
CULMEN FROM BASE. 2 @: 21.0, 21.5. 1 ?: 19.0.
EXPOSED GULMEN, 2 4: 1710, 17%. 1 2: 14.0,
TAXONOMY. The main difference between the three available
specimens of crockettorwm (Idenburg River, Sepik River, and southern
slope of Snow Mountains) and six specimens of nominate flavirictus
(southeastern New Guinea and Fly River) is the yellow auricular spot
of the former as opposed to the paler, lemon-whitish spot of the
latter. On the basis of this character my specimens are unequivocally
flavirictus, as expected on geographical grounds.
DISCUSSION. None of these specimens was netted. The Mengino
specimens were among many meliphagas and other honeyeaters feed-
ing in a tree with small berries at 4,600 ft.
Meliphaga flaviventer rubiensis (Meyer)!
‘Tawny-breasted Honeyeater
NATIVE NAMES. Gimi: abigému. Daribi: kesoabo.
SPECIMENS EXAMINED. Okasa: 1 ¢, 1 juv. @ (22 and 26 June 1965).
Kamm: 6 ¢,3 9 4-14 Aug. 1964; 2 July-3 Aug 1965). Boma: 1 4,1 9 (6-5
July 1965). Soliabeda: 3 ¢, 2 9 (22-30 July 1965). Mt. Karimui Zone 1: 1 9;
“one Zool fo On Zone or 1 O(a Anes 96b):
WEIGH, 12 6540-55473 s242).5 VOM = 3849 (420) Soa) Meu ae ues
WING: 12 42 96108 (OI 62] 25). 10) 0 90-99, (90.05 3.5), Wu. 0b:
STOMACH CONTENTS. Mainly fruit, some insects.
TAXONOMY. Comparison with specimens of all nine races from
the New Guinea mainland yielded the unexpected result that birds
from both the Karimui area and Okasa are closest to rubirensis, the
Weyland Mountains race, from which they differ only in having
slightly darker upperparts. One might have expected them to be
giulianettiz, the southern watershed race both to the east and to the
west, to which Mayr and Gilliard (1954, p. 369) assigned their two
specimens (from an unstated locality) without comment and to which
Schodde and Hitchcock also assigned Lake Kutubu birds. However,
specimens of giulianetti from Mafulu, Baroka, and Kubuna in south-
eastern New Guinea and from Palmer Junction on the upper Fly
River are paler above, much paler below, and duller brown on the
breast than the Karimui and Okasa series. Specimens of saturatior
from the lower Fly River and Aru Islands are also paler and duller
below and have the edges of the wing coverts less bright rufous. Birds
from the Setekwa River on the southern slope of the Snow Mountains,
1 Listed as Xanthotis chrysotis in Rand and Gilliard (1967). This and the next
three species, placed in the genera Xantholis and Oreornis by Rand and Gilliard,
are included in Meliphaga by Salomonsen (1967, pp. 386-391) in Peters’ Check-list,
Vol. 12, but do not belong to the Meliphaga analoga complex.
374
SPECIES ACCOUNTS
assigned to saturatior, are slightly darker below than Fly River and
Aru Islands specimens and show a tendency towards rubiensis 1 this
respect. The races chrysotis, madaraszi, spilogaster, kumusit, philemon,
and vis? all show marked differences.
BREEDING. Gonads were small in most specimens except 1n
1964 at Karimui, where most testes were enlarged.
DISCUSSION. Meliphaga flaviventer is one of the most abundant
bird species in the lowlands, becoming less common with increasing
altitude until it disappears completely around 5,000 ft. Its contribution
to the avifauna in the Karimui area was 7% of the local avifauna at
Soliabeda, 5% at Karimui, and 1.8% on Mt. Karimui, where the
highest record was at 4,850 ft.
Both flowering and fruiting trees were utilized. Meliphaga flavi-
venter accounted for 5-25% of the bird-usage in flowering trees at
Karimui, 10-40°% in trees with fruits smaller than 5 mm at Karimui,
Okasa, and Mengino, and 23% in the tree at Karimui with hard-
shelled fruit opened by parrots. It was absent in trees with soft fruits
larger than 10 mm. In these feeding trees it could be seen in both the
upper and lower half of the tree but was twice as numerous in the
lower half. Outside of fruit trees it was commonest from 5 to 50 ft
above the ground, so that relatively few specimens were netted. Chases
of one individual by another took place occasionally, and one pair
was observed attempting copulation in flight.
VOICE. ‘The song is a phrase of usually three notes, repeated five
to 10 times in succession and reminiscent of the songs of caprimulgids.
In the commonest version all notes are somewhat upslurred, particu-
larly the first; the first note is highest and the third lowest in pitch;
and the interval between the first and second note is twice as long as
that between the second and third note or between repetitions of the
pattern (Fig. 39). ‘The pitch is in the lower middle registers, the
quality clear, and the volume loud. When one individual starts sing-
ing, others nearby may begin after a few repetitions of the pattern and
continue more or less in synchrony. The song is suggestive of that of
the honeyeater Philemon novaeguineae in the use of a repeated pat-
tern, in the pattern itself, and in quality. Songs I heard near Port
Morebsy and Lae were very similar to those in the Eastern Highlands,
but songs in the North Coastal Range differed by being delivered
nearly twice as fast, suggesting that a comparison of songs and morpho-
logical subspecific characters at different localities might prove interest-
ing.
Meliphaga polygramma lophotis (Mayr)!
Spotted Honeyeater
NATIVE NAME. Daribi: bulugagi.
SPECIMENS EXAMINED. Karimui: 2 ¢, 2 9 (10 July-4 Aug. 1965). Bomai:
1 Listed as Xanthotis polygramma in Rand and Gilliard (1967).
375
SPECIES ACCOUNTS
Meliphaga flaviventer ( = 'Xanthotis ch rysotis"’)
aA a; a7
i y re oA
Y = ~ 4
a Sac i called ee eae eee iy
Sse
Fic. 39. Voice of Meliphaga flaviventer.
I g¢ (9 July 1965). Mt. Karimui Zone 1: 1 9; Zone 2: 2 ¢ (10-15 Aug. 1965).
WEIGHT. 3 @: 19.7, 20.0, 23.5. 3 Q: 18.0, 19.5, 19.5.
WING. 5 @: 75 (3), 76,78. 3 9: 70, 73 (2).
TAXONOMY. ‘These specimens agree with lophotis of the Huon
Peninsula and southeastern New Guinea, which has a large yellow
ear patch. ‘The patch is smaller in potkilosternos and nearly lacking in
septentrionalis, while candidior has smaller black spots on the under-
parts.
BREEDING. The latest male collected at Karimui (4 Aug.) and
one of the two males on Mt. Karimui (15 Aug.) had slightly enlarged
testes, while gonads were small in the other specimens.
DISCUSSION. ‘This hill forest species was present up to 4,500-
4,600 ft but did not occur as low as Soliabeda (2,000 ft). It was also
recorded from Lake Kutubu (2,450 ft).
In the Karimui area Meliphaga polygramma occurred regularly but
usually singly in feeding trees. It was observed in all seven flowering
trees studied at Karimui and in the one at 4,400 ft on Mt. Karimui,
where it made up 5-1%% of the bird-usage. It accounted for 1-2% of
the bird-usage in trees with small fruits at Karimui and Mengino, and
16% in the Karimui truit tree with hard-shelled fruits opened by
parrots. Its vertical distribution through the feeding trees appeared
nearly uniform, but it was rarely seen in the understory or netted. In
behavior it is silent and unobtrusive, and easily overlooked if one does
not seek it in feeding trees.
VOICE. Neither Stein (1936, p. 32) nor I ever heard a sound from
this honeyeater.
Meliphaga subfrenata salvadoriu (Hartert)!
Black-throated Honeyeater
SPECIMENS EXAMINED. Mt. Karimui Zone 2: 1 ¢@, 1 9; Zone 3: 1 ?; Zone
4s 4,1 O35 Zone: 1 6,8 Qs Zone 7: 1 @ Gt Auge Sept. 1965),
1 Listed as Oreornis subfrenatus in Rand and Gilliard (1967).
376
SPECIES ACCOUNTS
WEIGHT. 4 4: 32, 32, 34, 37. 5 Q: 26, 26, 27, 28, 28. 1 ?: 27.
WING. 4 @; 100, 101, 102, 104. 5 9: 89, 91,91, 92,98. 1 ?: 92,
STOMACH CONTENTS. Usually insects, but 3% of stomachs examined con-
tained fruit.
TAXONOMY. These are nearer to the dark southeastern New
Guinea race salvadorti than to melanolaema.
BREEDING. ‘Testes were slightly enlarged in two males, con-
siderably enlarged in the other two. One female had an enlarged
ovary. Birds in song were heard from the top to the bottom of the
altitudinal range.
DISCUSSION. At higher altitudes Meliphaga subfrenata is the
medium-sized honeyeater in flowering trees, which it shares with the
small Myzomela rosenbergii and the large Melidectes rufocrissalis (or
M. belfordi). It occurs up to timberline, where it sometimes perches
in tree ferns in alpine grassland near the forest edge. Its lower limit
is usually about 7,000 ft, but it ranged down to 4,400 ft on Mt.
Karimui, where it was fairly common (1.5% of the local avifauna).
Its vertical distribution is mainly from 10 ft above the ground
to the crowns, infrequently in the understory, once seen on the ground.
Away from flowering trees I have seen it flycatching, and also clumsily
hopping up a large vertical trunk like a would-be woodpecker, probing
the bark, sipping a couple of times, then flying to the base of another
trunk and ascending it in the same fashion. Males and females are
equally common in flowering trees and away from them and do not
differ in altitudinal distribution nor in vertical distribution through
the vegetational column.
VOICE. As described elsewhere (Diamond and Terborgh, 1968,
p. 74), the song is “a rapid, loud, bubbling, and cheerful series of notes
which first rises and then falls in pitch, progressively decelerating and
sometimes concluding in three slurs on the same pitch, after which the
pattern is immediately repeated one or two times.” The quality and
pattern of the song as well as the frequency of countersinging are
quite similar to the song of Meliphaga versicolor.
Meliphaga obscura (De Vis)!
Obscure Honeyeater
SPECIMENS EXAMINED. Karimui: 6 Q (2 and 7 Aug. 1964; 4 July-5 Aug.
1965). Soliabeda: 4 ¢,1 9 (23-27 July 1965). Mt. Karimui Zone 2: 1 g, 1 ? (13
and 14 Aug. 1965).
WEIGHT. Gu G25, )20) 20) 293. 5; Os: Ba (2)y25, 26,27. Te 27.
WING, 5 ¢= 85,92 (3),,93, 7 Or 80 (2); 88, 85: (2), 86 (2). .1) 2: 92,
TAXONOMY. Due to the lack of Vogelkop material I am unable
to assess the proposed race viridifrons (Salomonsen, 1966).
1 Listed as Oreornis obscurus in Rand and Gilliard (1967).
377
SPECIES ACCOUNTS
BREEDING. Gonads were small in all specimens.
DISCUSSION. The unsexed Mt. Karimui specimen is probably
male, from its wing length. The local sex ratios are very unbalanced
(only females at Karimui, mainly males at Soliabeda).
I found this uncommon honeyeater of the hill forest only at the
three above-cited localities where specimens were collected (2,000-4,630
ft). It was regularly seen in low numbers in trees with small fruits and
in flowering trees, which it shared with up to 11 other species of
meliphagids and where it accounted for 2-5% of the bird-usage. Meli-
phaga obscura was silent, slower in its actions than members of the
Meliphaga analoga complex, and fed strictly in the lower branches and
middlestory, never ascending into the upper half of the vegetational
column. In addition, I have a few sightings of single individuals 5-10
{t above the ground in second-growth and gardens. Males frequented
flowering trees and were never netted, while females were netted in
the understory away from flowering trees.
VOICE. No vocalization was heard.
Pycnopygius cinereus marmoratus (Sharpe)
Marbled Honeyeater
NATIVE NAME. Fore: osara.
SPECIMENS EXAMINED. Awande: 1 ¢ (20 June 1964). Okasa: 6 @ (22-26
ume 1905). Mengino: Ig, 2°? (10 July 1962). Karmomi 93" 65 a. Ole 7) imly
1965). Mt. Karimui Zone 2: 1 9 (13 Aug. 1965).
WEIGHT. 7 @: 40-58 (48:823:9). 4 9: 36, 38) 4b, 46,
WING. 10) 67 96-1127(106.6 32 533) 4 o> 96, 101) 102, 105:
STOMACH CONTENTS. Fruit, 2-7 mm in diameter (seven stomachs); insects
(seven); fruit and insects (one); flower remains (one).
TAXONOMY. ‘The races cinereus and dorsalis are less heavily
spotted below. ‘There may be an increase in size with altitude, since
the two birds with the longest wings (111 and 112) were from the
highest altitudes (Awande, 6,200 ft, and Mengino, 5,800 ft) and since
the Mt. Karimui female (4,750 ft, wing 105) was larger than the three
Karimui females (3,650 ft, wing 96, 101, 102).
BREEDING. The testes of one Karimui male were slightly en-
larged, but gonads in all other specimens were small.
DISCUSSION. Pycnopygius cinereus occurs in the forest interior
but most often at the forest edge and in second-growth between about
3,500 and 6,500 ft, and is commonest in the lower part of its altitudinal
range. The only record in the undisturbed forest of Mt. Karimui was
the single specimen at 4,750 ft. In behavior it was a silent, solitary,
sluggish, and inconspicuous bird of the upperstory, frequenting trees
with flowers or small fruits. Only one specimen was netted. At a
Miarosa fruit tree it was repeatedly driven off by another honeyeater,
378
SPECIES ACCOUNTS
Melidectes torquatus, which is nearly the same size. Besides similar
sizes, Pycnopygius cinereus and Melidectes torquatus have somewhat
similar habits (upperstory, commonest in ecologically disturbed areas,
attracted to fruit trees), but generally Melidectes torquatus is far more
abundant. At the two localities where I found Pycnopygius cinereus
common, Melidectes torquatus was either uncommon (Okasa) or absent
(Mengino), suggesting that the more aggressive Melidectes torquatus
may limit its distribution.
VOICE. As with its congener P. ixoides, I never heard any vocali-
zation from P. cinereus, nor did Stein (1936, p. 32).
Pycnopygius ixoides ixoides (Salvadori)
Brown Honeyeater
SPECIMENS EXAMINED. Mengino: 1 ? (15 July 1964). Karimui: 5 ¢, 6 9
(10 Aug. 1964; 1 July-5 Aug. 1965). Bomai: 3 @ (6-9 July 1965). Soliabeda: 2 ¢,
2 Q (23-27 July 1965).
WEIGHT: 4 47°28) 28; 29, 33." 8 OF 22722, 24, 24:26, 28, 20) 31.
WING, 4 @2 86, 86, 87,90. 3) 93°79, 79,180.
TAXONOMY. ‘These agree well with nominate ixoides from the
upper Fly River and Mimika River except in having the edges of the
primaries more olive and the ventral coloration averaging slightly
brigher and richer. ‘The underwing coverts are bright ochraceous,
and the feathers of the crown are edged gray. The race finschi from
the north coast of southeastern New Guinea is so much brighter rufous
below that at first glance one might not have placed it in the same
species. here appears to be only one record from the south coast of
southeastern New Guinea (N.G.B.S. Newsletter, No. 35, p. 2, Sept.
1968). I have no Vogelkop material (topotypical ixoides) by which to
assess the proposed southern New Guinea race cinereifrons (Salomonsen,
1966).
BREEDING. Gonads were slightly enlarged in one Karimui male,
small in other specimens.
DISCUSSION. Pycnopygius ixoides is the low-altitude representa-
tive of the somewhat larger P. cinereus, to which it is very similar in
behavior. ‘The altitudinal ranges of the two species overlap around
3,500-4,500 ft. P. ixoides was fairly common in the Karimui area up to
3,900 ft, while P. cinereus extended down to 3,650 ft. The only other
Eastern Highlands record to date is the Mengino specimen, taken at
4,600 ft within the vertical range of P. cinereus. Like P. cinereus, P.
ixoides is solitary, slow-moving, inconspicuous, and silent, and fre-
quents the middle- and upperstories. It was never netted. At Karimui
there were a few observations in flowering trees and in trees with small
fruits.
VOICE. No vocalizations were heard.
379
SPECIES ACCOUNTS
Philemon meyeri Salvadori
Meyer’s Friarbird
SPECIMEN EXAMINED. Soliabeda: 1 2 (30 July 1965).
WEIGHT. 57.
WING. 111.
STOMACH CONTENTS. Mainly fruit, some insects.
BREEDING. ‘The testes were moderately enlarged.
DISCUSSION. I met this small friarbird, which is much less con-
spicuous than its noisy congener Philemon novaeguineae, only at
Soliabeda, where a pair was seen in the treetops on two occasions.
The specimen was collected at the top of a flowering tree inside the
forest.
Philemon novaeguineae novaeguineae (Miiller)
New Guinea Friarbird
SPECIMEN EXAMINED. Soliabeda: 1 9 (23 July 1965).
WEIGHT. 133.
WING. 144.
STOMACH CONTENTS. Fruit.
‘TAXONOMY. ‘The color is darker, and the knob on the culmen
smaller, than in most but not all Fly River specimens.
While the race jobiensis from the north coast is fairly distinct, dif-
ferences among the populations of the Vogelkop and south coast are
minor and complicated by individual variation and effects of wear
on color. Birds in this area have been variously distributed among
three races, nominate novaeguineae, brevipennis, and aruensis, on the
basis of differences in wing length, bill, and depth of color, without
any agreement being reached among authors (Rothschild and Hartert
(1913, p. 513), Ogilvie-Grant (1915, p. 78), Mayr and Rand (1937, p.
Zoe), Wand (19424, p. 064), and Junge (1959) p. (eb; 1953p, 73) ne
cently Salomonsen (1966, p. 9) described the southeastern population,
which Mayr (1941b, p. 211) had refrained from naming, as fretensis, on
the basis of material already reviewed by Mayr and Rand and using
the same characters (fretensis described as slightly paler and much
larger) extensively discussed previously. I have therefore reviewed the
material in the American Museum of Natural History from the west-
ern Papuan Islands, Vogelkop, Setekwa River, Fly River and south-
eastern New Guinea.
Within this area differences in slenderness of the bill and the size of
the knob on the culmen seem as inconsistent and slight to me as they
did to Ogilvie-Grant, Mayr and Rand, and Junge. As noted previously,
average differences in color are of little help as subspecific characters
in this species due to nongenetic variation: “Wear and fading produce
such great changes in the color of the plumage of this species that it is
380
SPECIES ACCOUNTS
difficult to compare any but specimens which are actually molting”’
(Mayr and Rand, 1937, p. 234). Rand (1942a, p. 364) found that birds
from more arid areas were paler, due apparently to more pronounced
wear. Table 16 assembles measurements of wing length in 43 males
available to me (including the types of brevipennis and fretensis) and
from measurements of other authors. It appears from these figures
that birds from the Vogelkop and western Papuan islands average the
largest, with Junge’s specimens standing out as especially large; those
from Etna Bay, 35 miles from the type locality of novaeguineae (Lobo
on ‘Triton Bay), average about 4 mm smaller; those from southeastern
New Guinea (including the type of fretensis) average 1 mm shorter;
Fly River birds average 1-4 mm shorter; and those from southwestern
New Guinea exclusive of Etna Bay average another several millimeters
shorter. Thus, wing length decreases clinally eastward from the Vogel-
kop to Etna Bay to the southwestern New Guinea coastal plain and
then increases clinally further eastward to the Fly River and to south-
TABLE 16
WING LENGTH OF Philemon novaeguineae MALES
Vogelkop and western Papuan islands
De 151-164 (av. 155)
Ona 151, 155, 158
442 145-158
743 160-164
2 a4 154-156
Southwestern New Guinea
Etna Bay, 3¢5 150; 154, 1b5 (av. 1d3)
Noord River, 1443 144-151
Mimika, Wataikwa, Setekwa Rivers,
13 ¢2 139-150
Setekwa River, 32 142, 143*, 144 (av. 143)
Fly River
he 139-156 (av. 148)
10 46 146-157 (av. 151)
| Aus 153
Southeastern New Guinea
82 140-158 "(av. 1b2)**
Bi a 152, 156, 158
2468 147, 1590
nen
1 From Gyldenstolpe (1955b).
2 From Ogilvie-Grant (1915).
3 From Junge (1939).
* From Mayr and De Schauensee (1939).
5 From Junge (1953).
6 From Mayr and Rand (1937).
7 From Rand (1942a).
8 From Rothschild and Hartert (1913).
* Type of brevipennis.
** Including type of fretensis.
381
SPECIES ACCOUNTS
eastern New Guinea. In separating fretensis from brevipennis Salo-
monsen cited the longer wing of the former, but the specimen selected
as the type of fretensis is the largest specimen from southeastern New
Guinea, while the type of brevipennis is the second smallest specimen
from southwestern New Guinea. The smallest population (southwest-
ern New Guinea) overlaps somewhat with the largest population
(Vogelkop) and extensively with the intermediate populations.
‘There seem to be only two reasonable alternatives: to consider south-
western New Guinea riverplain birds as brevipennis and the rest as
nominate novaeguineae, or else to consider all the Vogelkop and south
coast populations as novaeguineae. Because the former procedure gives
novaeguineae a discontinuous range and the sizes overlap, the latter
procedure may be preferable. In either case fretensis must be consid-
ered a synonym of novaeguineae.
DISCUSSION. I met Philemon novaeguineae, one of the noisiest,
most conspicuous, and characteristic species of the New Guinea low-
lands, only at Soliabeda, where several individuals frequented the
crowns Of isolated trees. It was surprising not to find it in the Karimui
Basin, which so many other lowlands species had reached. On the
northern watershed the race jobiensis seems to have been much more
successful at colonizing midmontane cultivated areas: Bulmer found
it to be present in the upper Wahgi Valley and common in the Batyer,
Jimmi, and Kaironk valleys up to about 6,000 ft. Possibly this coloniza-
tion is actively underway, since Ripley (1964, p. 72) found P. n.
jobiensis in the Baliem Valley of western New Guinea in 1960, where
the Third Archbold Expedition had not found it in 1938 and 1939
despite exhaustive collections. Schodde and Hitchcock (1968, p. 67)
report that the population of Philemon novaeguineae at Lake Kutubu
is migratory, disappearing during the wet season there (May to
September).
The problem of the parallel geographical variation in the forms of
Philemon and of the oriole genus Orzolus through the Molucca-Timor-
New Guinea area has attracted attention ever since Alfred Russel
Wallace described this variation a century ago in his classic The
Malay Archipelago (1962) as a case of mimicry. Briefly, the Philemon
populations inhabiting seven islands in this area (Ceram, Buru, ‘Ven-
imber, Halmahera, Wetar, Timor, and New Guinea) show great inter-
island variation, and the Oriolus populations show equally great vari-
ation, but on each island the Oriolus form and the Philemon form
resemble each other, often to such a stunning degree that specimens in
the hand can be distinguished only with difficulty. In the field I have
frequently confused the New Guinea forms, Philemon novaeguineae
and Oriolus szalayi, even though the morphological resemblance is
less close than on the other six islands (both forms largely brown, the
bare black facial skin in Philemon paralleled by heavy black streaking
on the face and sides of the throat in Orrolus, Oriolus more distinctly
382
SPECIES ACCOUNTS
streaked below). The morphological similarity on New Guinea is
reinforced by similarity in habits, viz., in that both forms inhabit the
middle- and upperstory of second-growth and rarely the forest interior,
have approximately the same altitudinal range, and are social. On
the island of Ceram Oriolus forsteni and Philemon subcorniculatus
are virtually indistinguishable (large birds, mostly unstreaked brown,
with a yellow-olive wash on the breast, gray collar on the nape, dark
ear coverts). Oriolus b. bouroensis and Philemon m. moluccensis on
Buru, and Oriolus b. dicipiens and Philemon m. timorlaoensis on
Tenimber are largely brown without a yellow breast wash, with a re-
duced pale nape collar, and with a bare black facial patch of Philemon
duplicated by a similar area of black feathering in Oriolus. On Halma-
hera Philemon fuscicapillus and Oriolus phaeochromus are entirely
dark brown, the bare black facial patch of Philemon much reduced
and the black facial feathering of Oriolus absent. Philemon buceroides
pallidiceps and Oriolus viridifuscus finschi of Wetar, and Philemon Db.
buceroides and female (but not male) Oriolus v. viridifuscus of ‘Timor,
are pale below, unlike the previously mentioned forms, and a bare
black facial patch contiguous with a bare black patch on the side of
the neck in Philemon is duplicated by similar areas of black feathering
in Oriolus. Further details of plumage are given by Wallace (1962,
pp. 305-307) and by Stresemann (1914b, pp. 395-400).
Wallace (1962) attributed this remarkable parallel variation to
Batesian mimicry of Philemon by Oriolus, arguing that hawks avoid
the noisy, pugnacious Philemon with its strong claws and beak. Strese-
mann (1914b) rejected this interpretation on the grounds that selection
by hawk predation was probably insignificant, and argued instead that
the convergence was a coincidence due to common selective forces act-
ing on both forms and that the appearance of either Oriolus or
Philemon on an island would have been the same even if the other
form had been absent. However, the resemblances in pattern are so
detailed and striking that Stresemann’s interpretation appears to me
thoroughly implausible. A more promising approach to the problem
stems from recent discussions by Moynihan (1968) and by Cody (1969),
who have pointed out numerous similar cases of convergence in plum-
age between two unrelated bird species, usually ones which utilize
similar food resources. Cody suggests that the convergence facilitates
interspecific territoriality, while Moynihan refers to the convergence
as “social mimicry’, e.g., to facilitate interspecific flocking. The selec-
tive forces for convergence in the Philemon-Oriolus case remain un-
known.
VOICE. ‘The songs are the most characteristic bird sounds of the
New Guinea lowlands. ‘They consist of a somewhat nasal pattern of
three or four slurred notes, repeated several times with pauses of
less than a second between repetitions and with successive repetitions
somewhat louder (Fig. 40).
383
SPECIES ACCOUNTS
Philemon novaequineae:
Or = a a ae a etc,
Fic. 40. Voice of Philemon novaeguineae.
Ptiloprora meekiana subsp.
Meek’s Streaked Honeyeater
SPECIMENS EXAMINED. Mt. Michael: 1 9 (7 July 1964).
WING. 75.
TAIL. 67.
TAXONOMY. Im color this specimen is intermediate between
the western race occidentalis and the eastern race meekiana. It is nearer
occidentalis in the brighter and greener upperparts and in the brighter
yellow underparts and equidistant from the two races in the throat
(slightly darker gray in occidentalis). Wing measurements for nine
occidentalis females from the north slope of the Snow Mountains are
78-84 (79.9), and for two females of meekiana from the Huon Penin-
sula and southeastern New Guinea, 73 and 75. Since my specimen is
nearer occidentalis in color but matches meekiana in size, racial assign-
ment will have to await further material.
DISCUSSION. ‘The specimen, which represents the sole record for
the Eastern Highlands, was netted at 8,000 ft on Mt. Michael, where
P. guisei was common. Except on the northern slopes of the Snow
Mountains, this is a rare and local species throughout its wide range.
The two races have very different altitudinal ranges: meekiana of
southeastern New Guinea lives at 4,600-7,500 ft (mainly 5,000-7,000
ft), while occidentalis of western New Guinea lives at 7,200-9,200 ft.
Ptiloprora guisei umbrosa Mayr
Brown-backed Streaked Honeyeater
NATIVE NAMES. Gimi: habi. Daribi: kwe.
SPECIMENS EXAMINED. Awande: 1 ¢, 1 ? (14 and 15 June 1965). Mt.
Kati zone 2: I 4, 3 ©; Zone 3: |) 9G. Zone 4) 5 O. Zone sos o.oo
9; Zone 6: 4 $,5 9; Zone 7: 4 4,3 9; Zone 8: 3 6,4 @ (13 Aug.-8 Sept. 1965).
WEIGHT. 10° 4: 216-2777 (242-2177), 10 OF 170-20" (20 Aa Ut).
WING. 10 4: 85-94 (89.34 2.2). 10 9: 76-84 (79.2 + 1.6).
STOMACH CONTENTS. Usually insects, very rarely fruit seeds.
384
SPECIES ACCOUNTS
DISCUSSION. Relations in the Ptiloprora guisei-perstriata com-
plex, formerly considered a single species, were clarified by Mayr and
Gilliard (1954, p. 369; Gilliard and LeCroy, 161, p. 80). P. guiser 1s
present at medium altitudes in eastern New Guinea and reaches its
western limit somewhere between Mt. Giluwe and ‘Telefolmin. P.
perstriata occurs at medium and high altitudes in western New
Guinea but lives in eastern New Guinea only at high altitudes above
P. guisei. I take this situation to mean that P. perstriata originated in
western New Guinea as a form allopatric to P. guiset and reinvaded
the latter’s range relatively recently. P. mayri, a member of the same
complex, lives on two of the north mountain “islands” (Cyclops Moun-
tains and North Coastal Range). The very similar P. erythropleura
may constitute the Vogelkop representative (Diamond, 1969). .
On Mt. Michael the transition between P. guisei and P. perstriata
took place around 9,500-10,500 ft. Our records suggest that the two
species might overlap by several hundred feet, but this is uncertain.
Gilliard found that the transition on Mts. Hagen, Kubor, and Wilhelm
was around 8,000 ft.
P. guisei is the most abundant and characteristic bird of moss forest,
and is tied sufficiently strongly to this vegetational structure or the
associated climatic conditions that the lower limit of its altitudinal
range shows marked and predictable local variation. On Mt. Michael,
where heavy mossing began at 8,700 ft, P. guiset was common from
8,000 ft upwards but not at lower altitudes. On our walk from Mt.
Michael to Karimui we passed through a very local tongue of moss
forest which descended to 6,000 ft between Agotu and Maiba, and
immediately encountered P. guisei, after not having found it at the
same and higher elevations during a week at Mengino. In the Okapa
area, where we failed to encounter mossy conditions in our study zone
(5,800-7,000 ft), the two specimens at Awande (6,000 ft) constituted
the sole record. On Mt. Karimui, where the moss level was at 6,500 ft,
P. guise: became common a few hundred feet below that altitude,
remained common up to the summit at 8,165 ft, and trickled down to
4,400 ft in low numbers.
Census figures for P. guise? on Mt. Karimui clearly illustrate its
association with mossy conditions: Zone 2 (4,400-4,750 i); 13697, OF ihe
local avifauna; Zone 3 (4,750-5,390 ft), 0.59%; Zone 4 (5,390-5,960 ft),
as
2.3%; lower half of Zone 5 (5,960-6,250 ft), 1.4%; upper half of Zone
5 (6,250°0,500 it), 92%; Zone 6 (6,500-7,080 it), 14.4%; Zone 7 (7,080-
2,020 10), 24.47,; Lome $-(7,610-6,166: it), 27%.
The abundance of P. guisei on Mt. Karimui made it possible to
determine its population structure in detail. Zone 2 yielded three
females and a male with tiny testes; Zone 3, one female; Zone 4, five
females, one of them (netted at the border of Zones 4 and 5) with
somewhat enlarged ovaries; and the lower half of Zone 5, a short-
winged young male with tiny testes. In the upper half of Zone 5,
385
SPECIES ACCOUNTS
near the moss level, songs were heard commonly, none at all having
been heard at lower altitudes; and the sex ratio suddenly approached
equality, with the testes of all males but one enlarged. Songs remained
common, and the sex ratio remained near equality, in Zones 6 through
8. All males in Zone 6 had enlarged testes. In Zones 7 and 8, testes
were much enlarged in half of the males, small or tiny in the others.
Thus, nonbreeding birds show some accumulation at the top of the
range in addition to descending 1,500 ft below breeding birds.
P. guisei seems to take over the niche filled by the genus Meliphaga
at lower altitudes, as the common meliphagid gleaning for insects in
the lower and middle stories. It is occasionally seen in flowering trees
but does not concentrate in them. In behavior it is solitary and moves
rapidly (1-3 ft every second or every few seconds), often hanging up-
side-down to probe.
VOICE. A variety of simple and quiet two- or three-note calls, the
individual notes of which usually contrast in quality with each other.
‘The characteristic elements include: a plaintive whistled slur, which
may be upslurred, downslurred, or more complex; a staccato “chip”;
and a short buzzy note. ‘The commonest combination is for a short
staccato or buzzy note to be followed by a slur, or vice versa. A call is
given every 2-3 sec for 1 min or more (Fig. 41). The voices of P. mayrt
and probably of P. perstriata and P. erythropleura are quite similar.
Ptiloprora perstriata perstriata (De Vis)
Black-backed Streaked Honeyeater
SPECIMENS EXAMINED. Mt. Michael: 2 ¢, 1 9, 6 ?, 1 imm. ? (30 June-10
July 1964).
WING. 2 42100) 10K 1 2°91, Tammy 72789:
STOMACH CONTENTS. Insects.
TAXONOMY. As discussed elsewhere in a review of the genus
Ptiloprora (Diamond, 1969), the Snow Mountains population is so
Ptiloprora guisel:
z Or a Or ed
Bk aa :
staccato, | piercing staccato] piercing buzzy| whistle
buzzy | whistle
ae
~ = NN cat aa aa Nae ae
r= Or he or
re 2 SEC,
psec
Fic. 41. Voice of Ptiloprora guisei.
386
SPECIES ACCOUNTS
close to nominate perstriata of southeastern New Guinea that the race
lorentzi cannot be recognized. ‘The race perstriata must be considered
as encompassing the entire continuous 900-mile range of the species
on the Central Range, with the second race praedicta isolated on the
Wandammen Peninsula.
BREEDING. One of the males had enlarged testes.
DISCUSSION. Most of the specimens were netted.
P. perstriata lives at altitudes just above its sibling P. guisei on the
higher peaks of the Eastern Highlands. On Mt. Michael I found it
down to 9,500-10,000 ft in forest and all the way up to the summit at
12,300 ft, where it was common in the tongues of stunted trees and
bushes reaching onto the summit cap of alpine grassland. Gilliard
reported it above 8,000 ft (but apparently most commonly at 10,000
to 11,000 ft, judging from the altitudes given on his specimen labels)
on Mts. Wilhelm, Hagen, and Kubor. On Mt. Karimui (8,165 ft) it
was absent.
VOICE. Calls of P. perstriata that I heard on the summit of Mt.
Michael were very similar to the calls of P. guisei. Halcyon torotoro
and H. megarhyncha provide an additional example of congeners
which replace each other altitudinally and are very similar vocally as
well as morphologically.
Niche Differences in the Genus Melidectes
A review of interrelationships among the 10 species (all but M.
whitemanensis confined to New Guinea) of the genus Melidectes is
necessary as background for discussion of two questions under the
species accounts: Why is the range of Melidectes princeps restricted
to the Eastern Highlands? How did the unusual patterns of distribu-
tion and hybridization in the M. belfordi-M. rufocrissalis group arise?
On the basis of song, plumage, and bill form the species fall into
two groups. ‘The six members of the first group (M. torquatus, M.
ochromelas, M. rufocrissalis, M. leucostephes, M. foersteri, and M.
belfordi) have throat and/or gape wattles, a large bare patch of facial
skin, pale edges on the back, are large (40-105 g), have relatively stout
bills, and have similar loud nasal songs which frequently involve
duets. ‘he first four of these forms are confined to middle elevations
(ca. 3,000-8,000 ft); M. foerstert occurs both at middle elevations and
in the alpine zone; and M. belfordi occurs at middle elevations
throughout its range, and also in the alpine zone in some parts of its
range but not in others. M. rufocrissalis, M. leucostephes, and M.
foersteri constitute a superspecies. ‘The four members of the second
group (M. fuscus, M. nouhuysi, M. princeps, and M. whitemanensis)
lack throat or gape wattles, have a very small bare patch of facial skin,
lack pale edges on the back, are medium-sized (25-60 g), have relatively
slender bills, and have similar soft songs which do not involve duet-
387
SPECIES ACCOUNTS
ting. M. fuscus and the M. nouhuysi-M. princeps superspecies are re-
stricted to the New Guinea alpine zone. M. whitemanensis is confined
to the island of New Britain, and was originally described as a mono-
typic genus Vosea, but its song, behavior, and morphology suggest that
it is a Melidectes nearest M. fuscus (Diamond, 1971).
Only at alpine elevations does one find more than one species num.
erous in a given habitat. The details of distribution are such that each
alpine area on the Central Range supports two species, no more and no
less (“compound checkerboard allopatry’’). Southeastern New Guinea
has M. fuscus and an alpine race of M. belfordi: the Eastern Highlands
have M. fuscus and M. princeps but no alpine M. belfordi; the north-
ern slope of the Snow Mountains has M. nouhuysi and another alpine
population of M. belfordi (M. fuscus is known on the northern slope
only from a single specimen); and the southern slope of the Snow
Mountains has M. nouhuysi and M. fuscus, but not M. belfordi,
which apparently remains below 8,000 ft, from published reports of
Ogilvie-Grant (1915, p. 61) and Junge (1939, p. 57). The isolated moun-
tains of the Huon Peninsula, on which the area at alpine elevations is
very small, have only M. foersteri at these elevations. Niche differences
in the alpine zone are probably behavioral. M. belfordi frequents
flowering trees and remains in the crowns; M. fuscus is half as large
and gleans throughout the vegetational column, not in flowering trees;
M. nouhuyst remains in alpine shrubbery at and above timberline,
often on the ground; and the behavior of M. princeps has not been
reported.
Niche differences between the species of the first group at middle
elevations involve spatial segregation, in the sense that all the species
occupy the same kind of niche in different habitats or geographical
areas or at different altitudes. M. torquatus occurs on all the main
mountain ranges of New Guinea in open second-growth, whereas the
remaining midmontane species are mainly forest birds. ‘The geographi-
cal ranges of M. ochromelas, the M. rufocrissalis superspecies, and M.
belfordi are all peculiar when considered individually, in that each
extends widely over New Guinea but is missing from large areas of
the total range. When considered together, however, these ranges are
found to provide a peculiar case of checkerboard allopatry, in that
most areas have two species, never more and usually no less, and the
altitudinal ranges of the two species in a given area are mutually ex-
clusive. The details are as follows: southeastern New Guinea, M. bel-
fordi at higher elevations, M. ochromelas at lower elevations (my
observations on Mt. Albert-Edward); Huon Peninsula, M. foersteri
(rufocrissalis superspecies) at higher elevations, M. ochromelas at lower
elevations (Mayr, 1931; Mayr, 1941b, p. 198; labels of specimens col-
lected by Beck); Schrader Range and Star Mountains, M. belfordi at
higher elevations, M. rufocrissalis at lower elevations in ecologically
undisturbed areas (Bulmer, personal communication; Diamond, 1967,
388
SPECIES ACCOUNTS
p. 12; Gilliard and LeCroy, 1968, pp. 33-35; Mees, 1964, p. 27); Wey-
land Mountains and southern slope of the Snow Mountains, M. bel-
fordi at higher elevations, M. ochromelas at lower elevations (Ogilvie-
Grant, 1915, p. Gly Stem, 1926, pp. 30-31); Vogelkop, M. ochromelas
apparently at higher elevations, M. leucostephes (rufocrissalis super-
species) apparently at lower elevations. ‘The Herzog Mountains, most
of the Eastern Highlands, the eastern Snow Mountains (Mt. Goliath),
and the northern slope of the Snow Mountains may have initially sup-
ported M. belfordi and M. rufocrissalis, but what one now finds is a
hybrid population in the Herzog Mountains (“M. b. stresemannv’),
most of the Eastern Highlands, and Mt. Goliath (“M. b. griseirostris’”’)
(Mayr and Gilliard, 1952c), and nearly pure M. belfordi with only a
few signs of rufocrissalis genes on the northern slopes of the Snow
Mountains (Gilliard, 1959b, p. 8). Only M. belfordi was reported
from the Wissel Lakes (Junge, 1963, p. 70), and only M. ochromelas
from the Wandammen Peninsula (Hartert, 1930, p. 46), but in both
cases the ornithological survey was not intensive and an additional
species might have been present but overlooked. Wherever M. belfordi
coexists with either of the other two species, it occupies the higher
elevations. M. ochromelas lives below the representative of the rufocris-
salis superspecies on the Huon Peninsula but possibly above the
Vogelkop representative.
While there are other examples of checkerboard allopatry and of
altitudinal exclusion among New Guinea birds, the case of the mid-
montane forest Melidectes is unique in two respects. First, there is
no other instance of a species pair in which the altitudinal sequence
in one area is inverted with respect to another area, comparable to the
apparent relations of M. ochromelas and the rufocrissalis superspecies
on the Huon Peninsula and Vogelkop. Second, except among the al-
pine Melidectes I am unaware of another instance of compound
checkerboard allopatry such that each area supports two species of a
group of three, with the identity of the missing species showing ir-
regular geographical variation. This situation stems ultimately from
the facts that the forest Melidectes are too similar ecologically to co-
exist except by altitudinal exclusion; and that altitudinal series of
three species in New Guinea are unstable and often lead to the elimi-
nation of one bird (cf., p. 34). The unique complexity of the Melidectes
situation may be in part responsible for its providing the only instance
of hybridization in a New Guinea altitudinal sequence outside the
family Paradisaeidae.
Melidectes fuscus fuscus (De Vis)
Sooty Honeyeater
STOMACH CONTENTS. Insects.
TAXONOMY. Salomonsen (1966) described the Eastern Highlands
389
SPECIES ACCOUNTS
population as a new race gilliardi, based on size characters of material
collected by Gilliard. Mayr and Gilliard (1954, p. 507, Hable 7) nad
already published critical comparative measurements of this material
and concluded that it did not warrant separation from nominate
Juscus, a decision also adopted by Sims (1956, p. 432) in treating Shaw-
Mayer's material. The western population occidentalis is smaller than
the southeastern population fuscus, and the geographically intermedi-
ate Eastern Highlands population is intermediate in wing, tail, culmen,
and tarsus measurements. Since gilliardi overlaps fuscus in all four
measurements in both sexes except apparently in the culmen of males,
and since no differences other than size separate the populations, gil-
liardi should be considered synonymous with fuscus.
DISCUSSION. Gilliard, Shaw-Mayer, and Schodde collected this
honeyeater at 10,000-11,000 ft on Mts. Hagen, Wilhelm, Giluwe, and
Kubor in moss forest below timberline, but I failed to find it on Mt.
Michael. It gleans, darting rapidly and hopping along branches, from
the crowns to the understory, is frequently netted, and does not con-
gregate in flowering trees.
VOICE. ‘The long, monotonous song is based on a pair of high-
pitched notes, the second either trilled or slurred and at a lower pitch
than the first. ‘(he pair is repeated 90 times per minute for 1 min or
more. The song of Melidectes (“Vosea’) whitemanensis is similar but
involves only 3-8 repetitions.
Melidectes princeps Mayr and Gilliard
Long-bearded Honeyeater
NATIVE NAME. Labagai (the language spoken at Lufa): abayo.
SPECIMENS EXAMINED. Mt. Michael: 1 9, 2 ? (10 July 1964).
MEASUREMENTS. See Table 17.
TAXONOMY. ‘There is a naked area of skin about the eye, with
the eye at its anterior edge. In life the upper half of this area is
yellow-orange, the posterior lower quadrant yellow, and the anterior
lower quadrant light gray-green. ‘The chin and throat whiskers are
white. ‘Table 17 gives dimensions of the one Mt. Michael specimen
prepared as a skin and of Gilliard’s material. ‘There are no plumage
differences between birds from different mountains.
DISCUSSION. ‘This is one of the two semispecies endemic to the
Eastern Highlands (the other being Lophorina (Astrapia) mayer?) and
occurs in high-altitude moss forest up to timberline, between 10,000
and 12,000 ft. ‘The existing material consists of specimens collected on
Mts. Wilhelm, Kubor, and Hagen by Gilliard, on Mt. Hagen and the
Kubor Range by Shaw-Mayer, on Mt. Kinkain and the Minj-Nona
Divide of the Kubor Range by Hitchcock, and on Mt. Michael by
Terborgh and me. One of my three specimens was netted.
390
SPECIES ACCOUNTS
TABLE 17
MEASUREMENTS OF Melidectes princeps
Culmen Ex posed
Wing (from base) Culmen
Mt. Michael, 19 114 44 59
Mt. Hagen, ag 120, 120, 121 49,49, 51 44,44, 45
1Q? Vip 49 45
Mt. Wilhelm, 20 EME, We 43,45 38, 39
1 gl 104 40 BD
Mt. Orata (Kubor
Range), 36 LOS, 100, 125 41,41, 44 45, 45, 48
19 108 4] 46
1 Probably immature.
In the Snow Mountains M. princeps is represented by M. nouhwyst,
but there is no member of the superspecies in southeastern New
Guinea. Only four other species and superspecies (p. 46) are known
from elevations above 5,000 ft in the Snow Mountains and the Eastern
Highlands but not from southeastern New Guinea. Rand and Gilliard
(1967, p. 546) suggest that the absence of the M. princeps-M. nouhuyst
superspecies from southeastern New Guinea might be due to “perhaps
the annual burning of the alpine grasslands there, which has destroyed
the alpine shrubbery and has eaten into the forest, artificially lowering
the timberline”. This or other explanations involving some general
ecological condition seem to me improbable because no other wide-
spread timberline species or superspecies is missing in southeastern
New Guinea, whereas three are missing in the Eastern Highlands.
The alternative interpretation I suggest is that the alpine zone in
New Guinea provides only two Melidectes niches, as shown by the
distributional evidence (p. 388); and that the establishment of M.
fuscus and M. belfordi in southeastern New Guinea therefore left no
room for M. princeps.
Melidectes belfordi subsp. and M. b. schraderensis Gilliard and LeCroy
Belford’s Melidectes
STOMACH CONTENTS. Usually insects. Of the stomachs examined 14% also
contained fruit.
DISCUSSION. M. belfordi remains in the crowns, is rarely netted,
and congregates in flowering trees. It feeds by hopping along the
branches or up the trunk, often leaning head downwards. Its distribu-
tion is discussed on pp. 388-389 and pp. 393-396.
VOICE. On Mt. Albert-Edward in southeastern New Guinea M.
belfordi’s loud repeated calls included hoarse rasps, nasal “caw’s’, and
clear, high, piping notes. Frequently two birds facing each other on
a branch duetted by calling “caw” in rapid and perfect alternation up
391
SPECIES ACCOUNTS
to 34 times. I heard similar duets from the related M. ochromelas in
southeastern New Guinea and M. rufocrissalis in the Eastern High-
lands. The similarity of these antiphonal duets of three Melidectes
species to the antiphonal duet of the Giant Fijian Honeyeater, Gymno-
myza viridis (Diamond, 1972) supports the postulated relationship of
these two genera (Mayr, 1944).
Melidectes rufocrissalis thomasi Diamond and
M. r. rufocrissalis (Reichenow)
Reichenow’s Melidectes
NATIVE NAMES. Fore: wai. Gimi: waiyd. Daribi: kirigo.
TAXONOMY. A previous report (Diamond, 1967a, p. 9; 1969,
p. 55) lists specimens examined, weights, and measurements of this
race from the southern slopes of the Eastern Highlands. ‘The nomi-
nate race lives in the Schrader Range.
DISCUSSION. On Mt. Karimui population structure varied with
altitude in the usual fashion. Zone 2 yielded females and one male
with tiny testes, and Zone 3 yielded only females. From Zone 4 up-
ward males were common, and all without exception had enlarged
testes. One of the females in Zone 5, and both in Zone 6, had enlarged
ovaries.
In my collecting areas M. r. thomasi ranged up to 8,000 ft and down
to a lower limit varying locally between 4,000 ft (Mt. Karimui) and
6,000 ft (Mt. Michael). Bulmer found M. rufocrissalis x M. belfordi
hybrids down to about 5,000 ft in Kyaka territory, Bulmer and Gilliard
found M. rufocrissalis down to 5,000 ft in the Schrader Range, and
Mackay (N.G.B.S. Newsletter, No. 28, p. 3, Feb. 1968) found it common
at Pureni near ‘Tari.
The habitat of M. r. thomasi is the forest and the forest edge, includ-
ing partly cut forest with small gardens intruding, but not isolated
groves in open areas, a niche left to M. torquatus. Within this habitat
and altitudinal range it is not only one of the noisiest but also one of
the most abundant birds, as the following census figures on Mt. Kari-
mui show: Zone 2, 2.3%, ot the local avitauna; Zone 3, 3:99; Zone 44
S49; Zone 5, 27%; Lonel, loo%,; Zone 7; no records; Zones, Lot
It remains in the middle- and upperstory, with the result that only two
of the specimens were collected with nets. Noisy chases between sev-
eral birds in the treetops were a common occurrence. It was the
largest of the several kinds of honeyeaters attracted to the flowering
trees at 4,450, 4,700, and 6,430 ft on Mt. Karimui.
VOICE. The loud vocalizations include: a series of hoarse “caws”’
on the same pitch, similar to Lophorina superba but with no decelera-
tion; and a series of two kinds of notes strictly alternating with each
other, one note clear and on a slightly higher pitch, the note immedi-
oo2
SPECIES ACCOUNTS
ately preceding and following it more complex. This latter call is so
similar to the common call of M. torquatus as to be not safely distin-
euishable. There is also a duet of alternating caws (Diamond and
Terborgh, 1968, p. 74), similar to that of M. belfordi.
Melidectes belfordi & Melidectes rufocrissalis hybrids
The problem of hybridization between these two species in the
Eastern Highlands was recognized and greatly clarified in the studies
of Mayr and Gilliard (1952c) and of Gilliard (1959b). Additional dis-
coveries since Gilliard’s 1959 paper shed new light on the problem.
The starting point of Mayr and Gilliard’s studies was that one
finds in the Wahgi region hybrid swarms containing individuals close
to pure M. rufocrissalis, others close to M. belfordi, and a whole
spectrum of intermediates. At higher elevations (> 8,000 ft) these
hybrid populations gradually yield to typical M. belfordi. Material
collected earlier in the Herzog Mountains (“M. b. stresemannv”’) and
on Mt. Goliath (“M. b. griseirostris”) was also very variable and evi-
dently represented hybrid populations with a preponderance of
belfordi genes. Subsequently (in 1954) Gilliard obtained 12 specimens
at ‘Telefolmin (northern watershed), of which seven were nearly pure
belfordi, three nearly pure rufocrissalis, and two hybrids. Pure M.
rufocrissalis had previously been known only from the type locality
of the Schraderberg on the northern watershed.
The four newer findings are as follows:
1. Melidectes honeyeaters on Mt. Karimui, at Okapa, and on Mt.
Michael (all in the southern watershed) proved to belong to a new
face of M. rufocrissalis, M. 7. thomas (Diamond, 1967a, p. 9; 1969,
p. 55), identical to M. r. rufocrissalis except that the forehead was
black, not white. Since a black forehead is a belfordi character that
tends to predominate in the hybrid swarms, the origin of M. r. thomasi
was probably a hybrid one. However, M. r. thomas: differed from the
other hybrid swarms and hybrid races in being uniform and showing
no other belfordi traits, suggesting that the hybridization leading to
M. r. thomasit had occurred considerably earlier and that other
belfordi genes had been eliminated. Since the southern watershed of
the Eastern Highlands remains little explored ornithologically above
3,000 ft from Mt. Karimui for 300 miles westwards to the Star
Mountains, and since M. rufocrissalis has been observed near Tari
140 miles west of Mt. Karimui, the new race may have an extensive
range.
2. ‘Iwo specimens collected in the Star Mountains on the southern
watershed were discussed by Mees (1964, pp. 26-27). Of the one col-
lected at the lower elevation (ca. 4,000 ft), he reported, “This specimen
has all the characters of M. rufocrissalis, except that the supercilium is
yellow.” Of the one collected near 6,000 ft: “This bird agrees with M.
393
SPECIES ACCOUNTS
belfordi, except that the bill is much larger and grayish in color. The
feathers over the eye are white. These two characters point to hybrid-
ization with the next species [M. rufocrissalis].’ Actually, the super-
cilium is yellow in pure rufocrissalis and white in pure belfordi, so that
the rufocrissalis specimen was in fact pure and the belfordi one nearly
SO.
3. In 1966 I found that no species of genus Melidectes was present
in the North Coastal Range (Prince Alexander Mountains, Torricelli
Mountains, Bewani Mountains) although the maximum altitudes in
the North Coastal Range (up to 6,200 ft) were high enough for several
of the species. In 1959 Gilliard had found no Melidectes species in
the Adelbert Mountains, perhaps because they rise only to ca. 4,500 ft.
Mayr had also found no Melidectes in the Cyclops Mountains, which
rise to-7, LOO) it.
4. In 1963 and 1964 Bulmer (pers. comm.) and Gilliard (Gilliard
and LeCroy, 1968, pp. 33-35) discovered a population of M. belfordi
in the Schrader Range of the northern watershed, the type locality of
M. r. rufocrissalis. M. belfordi lived alone in the beech forest above
7,500 ft, M. rufocrissalis lived alone in the forest and disturbed areas
below 7,000 ft, and the two species overlapped only in a narrow band
at 7,500-8,000 ft. Most specimens were pure, but a few showed signs
of hybridization.
‘Two major problems, in addition to a host of minor ones, remain
unsolved. Why and how long has hybridization been occurring? Where
did M. rufocrissalis and M. belfordi originate?
Regarding the origin of hybridization, Gilliard (1959b) suggested that
it might stem from the extensive habitat disturbance caused by native
agriculture. ‘The evidence for altitudinal exclusion in undisturbed
habitats, and the wholesale hybridization in the extensively dis-
turbed Wahgi Region, are in general accord with this suggestion. If
this hypothesis is correct, massive hybridization may have started only
a few centuries ago. Information as to whether any hybridization at all
takes place where both species occur together in completely undis-
turbed areas is sorely lacking. I would guess that some hybridization
has been going on for a long time in undisturbed areas in the very
narrow band of altitudinal overlap, since M. +. thomasi is a uniform
and presumably not recent race even in the virgin forest of Mt. Kari-
mui; and that habitat disturbance has drastically accelerated hybridiza-
tion by creating much larger zones of contact.
Regarding the evolution of the group, Gilliard suggested “that wat-
tle birds [the M. rufocrissalis superspecies] are essentially forest-edge
birds of the northern watershed and that the black bills [M. belfordz]
are essentially pure forest birds of the central range and southern
watershed” (Gilliard, 1959b, p. 24). In particular, he suggested, the
M. rufocrissalis group might have originated on the “north moun-
tain islands” (Huon Peninsula, Adelbert Mountains, North Coastal
394
SPECIES ACCOUNTS
Range, Cyclops Mountains, and Van Rees Mountains) north of the
Mamberano-Sepik-Ramu-Markham gulch and might have reinvaded
the central range. While this hypothesis accounted for the facts avail-
able before 1959, it is untenable in the light of the more recent evi-
dence. M. rufocrissalis has at least as extensive a range on the southern
watershed (Okapa to the Star Mountains) as on the northern water-
shed (Huon Peninsula to Telefolmin); M. belfordi coexists with M.
rufocrissalis at its type locality on the northern watershed; and three
of the four “north mountain islands” explored to date (Adelbert
Mountains, North Coastal Range, and Cyclops Mountains) lack not
only M. rufocrissalis but any member of genus Melidectes. M. rufo-
crissalis’s habitat is primarily the lower altitudes and only incidentally
the forest edge; it simply happens that the forest edge habitats created
by native agriculture lie mainly in its altitudinal range because the
range of M. belfordi is mostly too high for agriculture. Finally, analy-
sis of New Guinea bird distributions suggests that colonization across
the Mamberano-Sepik-Ramu-Markham gulch by midmontane birds
has been practically a one-way affair, i.e., that the large Central Range
has provided many midmontane species to colonize the small “north
mountain islands” but that reverse movement of midmontane birds
has been almost nonexistent.
Most pairs of New Guinea montane birds with mutually exclusive
altitudinal ranges seem to have originated as eastern and western
populations of a polytypic species on the Central Range (p. 34), and
this was presumably the case in the M. ochromelas-M. belfordi-M. rufo-
crissalis group as well. ‘There are more marked differences among the
semispecies of the rufocrissalis superspecies than among the subspecies
of M. ochromelas or among the subspecies of M. belfordi, but the dif-
ferences between M. belfordi, M. rufocrissalis, and M. ochromelas are
greater than those within the rufocrissalis superspecies. Morpholog-
ically, M. rufocrissalis is no closer to M. belfordi than M. ochromelas
is to either, and one probably could not have predicted a priori in
which of the three cases (M. ochromelas-M. belfordi, M. ochromelas-
M. rufocrissalis, M. belfordi-M. rufocrissalis) isolating mechanisms
were most likely to break down. While reconstruction of the evolu-
tionary history inevitably involves some guesswork, the simplest ex-
planation of these facts is that the common ancestor first broke up
into three allopatric populations, the ancestors of M. belfordi, M. rufo-
crissalis, and M. ochromelas; and that M. rufocrissalis was the first to
expand, reinvaded the ranges of the other two species by displacing
them altitudinally, and then differentiated geographically. ‘The per-
ipheral distribution today of M. ochromelas may indicate that it was
the second to expand, reaching the mountains of the Huon Peninsula
and Vogelkop, which M. belfordi as the last species to expand may
not have had time yet to colonize. Present-day distributions would owe
their final form to the instability of three-species altitudinal sequences,
395
SPECIES ACCOUNTS
causing the extinction of M. rufocrissalis in some areas (southeastern
New Guinea, Weyland Mountains, southern slope of the Snow Moun-
tains) and of M. ochromelas in others (Eastern Highlands and northern
slope of the Snow Mountains).
Melidectes torquatus polyphonus Mayr
Cinnamon-breasted Wattlebird
NATIVE NAMES. Fore: ikitora. Gimi: kogidédo.
SPECIMENS EXAMINED. Awande: 4 ¢, 1 9 (16-20 June 1965). Okasa: 3 2
(24-26 June 1965). Mt. Karimui Zone 2: 2 @ (13 and 15 Aug. 1965).
WEIGHT. 7 6: 47, 50) 51, 58, 53, 55,58. 3 92.34 37, 44.
WING] 7 62 HO, US i Wis) OP 9h 3 Or 05. 07 Om.
TAXONOMY. In color these match a single male of M. t. poly-
phonus from Wau (Herzog Mountains), which has a wing of 116 mm.
Mayr’s (1931, p. 660) measurements of his type series of polyphonus
(2s 3.123, amd? 125, 1 or 113) are larger than my series, six males
which I measured from Gilliard’s Wahgi series (115, 116, 117, 117,
119, and 123), or Gyldenstolpe’s (1955, p. 164) Wahgi series (6 ¢, 110-
121, 8 ¢, 103-111, according to his measurements). ‘The races torqua-
tus, nuchalis, mixtus, and cahni are all paler and less ochraceous be-
low, while emilii is more ochraceous below, has larger throat wattles,
and the white throat patch reduced in size.
DISCUSSION. ‘This species and Gerygone ruficollis are probably
the midmontane birds which have profited most greatly from the
activities of man. My only records from undisturbed forest were
of the two Mt. Karimui specimens (4,400 ft) and of several seen at
the Sena River (4,500 ft). In undisturbed forest in other parts of New
Guinea as well (e.g., southern slopes of the Snow Mountains, Iden-
burg slopes) it seems rare, to judge from other expedition reports. In
settled and cleared midmontane areas, however, both in the Eastern
Highlands and in the Baliem Valley of western New Guinea it is
very common in trees of gardens, the forest edge, open second-growth,
and casuarina groves up to 5,500 or 6,000 ft. ‘he lower limit may be
somewhere around 3,500 ft, since it was uncommon at Okasa (3,550-
4,250 ft) and sighted only a few times at Karimui (3,650 ft), despite
plenty of forest edge habitat in both areas. ‘Testes were large in three
Awande males but tiny in all three Okasa males, suggesting that Okasa
may be supporting immatures at the lower fringe of the range.
Melidectes torquatus is usually seen in trees 15 ft or more above
the ground, though two were netted at the forest edge. ‘he Karimui,
Sena River, and Mt. Karimui records were all from flowering trees.
At Miarosa Terborgh found M. torquatus accounting for 90°% of
the bird-usage in a tree with a hard-shelled fruit that had been chipped
open by the parrot Trichoglossus haematodus (ct., p. 143).
VOICE. Loud, raucous, and very similar to some calls of Melidectes
396
SPECIES ACCOUNTS
rufocrissalis. Vhe pattern is difficult to describe but consists of a com-
plex disyllabic gurgle with the first syllable on the higher pitch.
Melipotes fumigatus goliathi Rothschild and Hartert
Common Melipotes
NATIVE NAMES. Fore: obéti. Gimi: érai. Daribi: yare.
SPECIMENS EXAMINED. Awande: 2 @ (14 and 20 June 1965). Okasa: 1 ¢
(22 June 1965). Karimui: 2 ? (1 and 11 July 1965). Mt. Karimui Zone 2: ae ae
@; Zone 3: 1 @, 1 9; Zone 4: 1 ¢, 1 9: Zone 5: 2 6,1 9; Zone 6: 1-3, 1 9;
Zone 7: 2 9; Zone 8: 1 4, 3 9 (13 Aug.-8 Sept. 1965).
WEIGHT. 10 @: 46-68 (56.1 45.7). 10 9: 44-57 (4919! =3-7):
WING. 10 4: 100, 102, 108-119 (113-44). 10 9: 101-108 (105 + 2).
STOMACH CONTENTS. Fruit, 2-8 mm in diameter (14 stomachs); fruit and
insects (four); insects (one),
TAXONOMY. Among the adults there is no more than a slight
increase of wing length with altitude on Mt. Karimui. ‘There are no
average differences in the darkness of the underparts or upperparts
between birds from the lowest zones of Mt. Karimui, from the highest
zones of Mt. Karimui, and from Awande. These Eastern Highlands
birds average slightly darker and blacker above than topotypical
goliathi from Mt. Goliath or from the northern slopes of the Snow
Mountains, and more so compared to nominate fumigatus of south-
eastern New Guinea, though the difference in blackness may involve
foxing (Gilliard and LeCroy, 1961, p. 76). Birds from higher altitudes
(> 8,000 ft) on Mt. Hagen and in the Snow Mountains have consider-
ably longer wings (¢ 120-128, 9 108-117) and, at least on Mt. Hagen,
have darker breasts and lighter bellies.
DISCUSSION. ‘The population structure deduced from details of
gonad condition and from weights and wing lengths is as follows. At
the bottom of the altitudinal range are immatures: a male with tiny
testes at Okasa, two specimens with very short wings (102) and undis-
cernible gonads at Karimui, and three males with short wings (100,
102), low weights (46, 49), or tiny testes in Zone 2 of Mt. Karimui.
The first adult males and females appear in Zone 2, but males with
enlarged testes do not appear until Zones 5 and 6 (ca. 6,000-7,000 ft)
of Mt. Karimui and at Awande (6,000 ft). Zones 7 and 8 (7,080-8,165
ft) contain largely or solely females.
Melipotes fumigatus is one of the commonest Eastern Highlands
forest birds from about 4,000 to 11,200 ft. Below 4,500 or 5,000 ft one
usually finds only immatures in low numbers, but in the main part
of its altitudinal range (e.g., Zones 2 through 8 of Mt. Karimui) it
accounts for about 3.59% of the local avifauna. It remained common
through the stunted moss forest both on Mt. Karimui (6,500-8,165 ft)
and on Mt. Michael (8,700-11,200 ft), and disappeared on Mt. Michael
only where the moss forest gave way to alpine grassland. It also ex-
tends to timberline in western New Guinea, but not in southeastern
397
SPECIES ACCOUNTS
New Guinea. In behavior Melipotes fumigatus is silent, slow-moving,
and solitary, and observed much less often than one might have ex-
pected from its abundance. In the forest interior it remains mainly in
the middle- and upperstories, but it often descends to the understory
at the forest edge. It is the only forest honeyeater at higher elevations
whose diet is mainly (but not solely) fruit. Several individuals were
collected in the flowering tree at 6,500 ft on Mt. Karimui, but these
were probably seeking insects attracted to the flowers rather than the
flowers themselves (Mayr “and Rand, 19377, pv 221).
VOICE. On only one occasion during four expeditions to areas
where Melipotes fumigatus was abundant have I heard a sound from
it: a series of a half dozen identical, rather slow, high-pitched, not loud
notes, upslurred at the end and with a quality as of sleighbells or
some parrots (e.g., Domicella lory).
DICAEIDAE: FLOWERPECKERS
Dicaeum geelvinkianum rubrocoronatum Sharpe
Red-capped Flowerpecker
NATIVE NAMES. Fore: isawanotaba. Gimi: férete. Daribi: dinai.
SPECIMENS EXAMINED. Okasa: 2 ¢@ (23 and 25 June 1965). Karimui: 7 ¢,
4 9 (10 Aug. 1964; 1 July-5 Aug. 1965). Bomai: 2 9 (6 July 1965).
WEIGHT. 6 ¢: 6.0, 7.0 (3), 72,73. 6 Q: 5.3, 65, 6.5, 7.0, 7.3, 7.5.
WING. 6 4: 51, 51, 54, 54, 57, 57. 3 9: 47, 49, 49.
STOMACH CONTENTS. Insects (two stomachs); fruit (two).
TAXONOMY. ‘The dorsal coloration is deep glossy violet. ‘The
races rubrigulare and albopunctatum on the Fly River have the red
throat patch of the male larger.
BREEDING. ‘Testes were enlarged in four males, not enlarged in
three others.
DISCUSSION. ‘This inconspicuous, tiny flowerpecker occurred at
all my collecting localities up to 5,700 ft in forest and up to 6,200 ft in
cultivated areas. Single individuals, occasionally pairs, ranged from
the treetops to 10 ft above the ground. I never netted the species.
VOICE. ‘The two inconspicuous and quite different calls usually
provide the only evidence for the presence of the species in forest. One
is a short, dry, buzzy, insect-like note. ‘The other is a high, sibilant
upslur confusingly similar to the call of the sunbird Nectarinia sericea.
However, the sunbird’s call is usually repeated, whereas that of
Dicaeum is given once.
Melanocharis nigra chloroptera Salvadori
Black Berrypecker
NATIVE NAME. Daribi: sisitabi.
SPECIMENS EXAMINED. Karimui: 5 ¢,3 9,5 imm. ¢ (30 July-15 Aug. 1964;
398
SPECIES ACCOUNTS
1 July-3 Aug. 1965). Bomai: 1 ¢,1 9,1 imm. ¢ (6 and 7 July 1965). Soliabeda:
2 4,1 9,1 imm. ¢ (22-30 July 1965). Mt. Karimui Zone 1: 1 ¢ (12 Aug. 1965).
WEIGHT. 9 @: 115-147 (19.22 1.1). 5 9: 140-165 (15.2+ 0.7). 6 imm. ¢:
13.5-15.3 (14.3 + 0.8).
WING. 9 @: 64-69 (666+1.6). 5 9: 63-69 (670+ 24). 6 imm. ¢: 62-69
(65.1 + 1.7).
TAIL. 9 @: 43-49 (463+2.0). 8 9: 41-47 (44.7+42.1). 6 imm. ¢: 41-47
(43.6 + 1.5).
TAXONOMY. There appears to be a slight increase in size with
altitude, as suggested by the following measurements of adult males.
In each case the average wing length is given first, then the average
tail length, then the weight: Soliabeda (1,350-2,000 ft), 65.6 mm, 45.8
mm, 12.2 g; Karimui (3,650 ft), 67.0 mm, 46.9 mm, navies
As in the case of M. versteri, females and immature males are heavier
than adult males, though the difference is less marked in the case of
M. nigra.
The race unicolor of northern and southeastern New Guinea is
entirely blue-black in the adult male and differs strikingly from the
races chloroptera and nominate nigra of southern and western New
Guinea and from my series, in which the male’s underparts are gray.
Nigra lacks the olive edges on the remiges of the male, present in my
series and in chloroptera. The only color difference between my series
and chloroptera from the Aru Islands, Fly River, and southwestern
New Guinea is that Karimui males average very slightly darker below
and that this is somewhat more pronounced at Soliabeda. As seen in
Table 18, my specimens have wings and tails as long as in southeastern
New Guinea wnicolor, slightly longer (particularly in the female) than
TABLE 18
MEASUREMENTS OF Melanocharis nigra ADULTS
Males Females
Wing Meal Wing Tail
M. n. chloroptera
Karimui area, 94,
59 64-69 (66.7)1 43-49 (46.4) 63-69 (67.0) 41-47 (44.6)
Fly River, 14,¢, 179 62-66 (63.7) 40-45 (43.1) 59-64 (61.2) 37-41 (40.8)
Aru Islands, 3g, 29 65-66 (65.7) 45 63, 65 (64) 40; 43 (41.5)
Aru Islands2 64-66 42-46 60-65 36-42
Setekwa River (south-
western New
Guinea), 5 ¢, 29 62-64 (63.0) 40-44 (41.6) 61,63 (62.0) 37, 39 (38.0)
Southwestern New
Guinea? 62-66 (63) 40-42 (41) 60-64 (62) 39-41 (40)
M. n. unicolor
Southeastern New
Guinea® 63-69 (65.7) 45-49 (47.3) 56-58 (65.9) 39-41 (40)
—_——X—a_—Xs_—-iE ?_eeee=e_eeeeeeeee nnn
1 Average values are given in parentheses.
2 From Junge (1939).
3 From Mayr and Rand (1937).
399
SPECIES ACCOUNTS
topotypical chloroptera from the Aru Islands, which are in turn
shehtly larger than the Fly River and southwest New Guinea popula-
tions. ‘he Lake Kutubu population also belongs to chloroptera and
has essentially the same measurements as the Karimui series.
The racial assignment of the Karimui population is of interest from
an evolutionary point of view. The genus Melanocharis contains a
triple altitudinal sequence: M. nigra in the lowlands to 3,000 or 4,000
ft, M. longicauda from the upper limit of M. nigra to about 5,000 ft,
and M. versteri from the upper limit of M. longicauda to near timber-
line. Subspecific variation is much less marked in M. longicauda and
M. versteri than in M. nigra, suggesting that M. nigra is the oldest
form, isolates of which have in the past reinvaded each other’s ranges
with altitudinal displacement to give rise to the other two species.
The color difference between the black-bellied males of M. nigra wnt-
coior and the gray-bellied males of the other M. nigra races is more
striking than the differences between these gray-bellied races and the
other two species. ‘The songs of the black-bellied and gray-bellied races
are detectably different to my ear. ‘The black-bellied and gray-bellied
populations must meet somewhere on the eastern shore of Geelvink
Bay on the north coast and somewhere on the Gulf of Papua on the
south coast.
On the north coast the gray-bellied race M. n. nigra has been taken
as far east as the Wanggar River at the head of Geelvink Bay, while
the black-bellied race wnicolor has been taken as far west as Pionier-
bivak on the Mamberano River, leaving an uncollected gap of 180
miles between these two localities on the New Guinea mainland. Japen
Island, which lies about 15 miles offshore in this gap, is the type
locality of wnicolor. On the south coast unicolor is known as far west
as Hall Sound, and chloroptera was previously known as far east as
Sturt Island on the Fly River. ‘The finding of chloroptera at Karimui
extends its range 200 miles farther east to within 100 miles of wnicolor.
When further ornithological exploration closes these two gaps, it will
be interesting to examine whether the black-bellied and gray-bellied
races finally intergrade in the short remaining unexplored distance
or whether they behave as good species, furnishing a model for the
evolution of M. longicauda and M. verstert.
BREEDING. Gonads were small in almost all specimens.
DISCUSSION. M. nigra is a common but inconspicuous species
(about 2% of the local avifauna in the Karimui area) which is heard
or netted much more often than it is seen. Individuals, pairs, or
eroups of up to six flit rapidly through the forest or second growth
2-20 ft above the ground. Adult males, adult females, and immature
males spend the same proportion of the time in the understory, as
judged by netting results. On Mt. Karimui, where M. longicauda was
400
SPECIES ACCOUNTS
absent, M. nigra occurred up to 4,300 ft, above which elevation it was
replaced by M. versteri. At Okasa, where M. longicauda was present,
collections down to 3,550 ft failed to yield M. nigra.
VOICE. The distinctive song is a very rapid series of thin twitter-
ing notes whose pitch describes a perfect sine wave (Fig. 42). ‘There
are up to two waves per second and more than a half dozen waves
per song. The call is a weak, brief, formless, slightly buzzy twittering
somewhat reminiscent of Peltops montanus in quality.
Melanocharis nigra:
ne < ss ors,
Fic. 42. Voice of Melanocharis nigra.
Melanocharis longicauda captata Mayr
Midmountain Berrypecker
SPECIMENS EXAMINED. Awande: 2 ¢,1 9 (15 and 16 June 1965). Okasa:
2 6,2 9 (23 Aug. 1964; 23-26 June 1965). Mengino: 1 ¢, 2 ? (15 July 1964).
Bomai: 1 ¢,1imm. ¢ (6 July 1965).
WEIGHE. oi h5 140) 15:05 lois 17.0 Sr Os 1510; 1525, 1b el mime 455 100:
WING, £42 65, (65, 66, '66.. 3.9: (64, 66, 67: 1 ami. yr 6D,
OAMIGL 4 59, DO, oy, 08. 2 Ooo, 04> al immo) ol.
TAXONOMY. My specimens as well as three collected by Gilliard
in the Wahgi Valley clearly belong to the race captata of the Huon
Peninsula, with which they agree in the pattern of white on the tail;
the outer edge of the outermost tail feathers is white, broadened dis-
tally into a bar, as illustrated by Salomonsen (1960). The race ori-
entalis of southeastern New Guinea, to which previous collectors
tentatively assigned their more limited material, differs in having a
smaller white spot distally.
BREEDING. Gonads were small in all specimens.
DISCUSSION. ‘This species is best distinguished from its congeners
by the yellowish wash of the underparts, the lemon pectoral tufts, and
the tail length (longer than M. nigra, shorter than M. versteri). It
forms the middle member of a three-species altitudinal series with
M. nigra and M. verstert, both of whose altitudinal ranges it overlaps
slightly, and is local and uncommon, as are the middle members of
401
SPECIES ACCOUNTS
other series. The only localities where it was common were Okasa and
Mengino, where the other two species were absent. It was outnumbered
by M. nigra at Bomai and by M. versteri at Awande, and was missing
(squeezed out—see p. 34) on Mt. Karimui. M. longicauda and Coracina
morio were the only two bird species (both of them characteristic of
hill forest) which were present at Bomai (3,250 ft) but absent at
Karimui Patrol Post (3,650 ft).
Both at Okasa and at Mengino Terborgh observed M. longicauda
in trees bearing small fruits.
VOICE. At Okasa Terborgh heard this species delivering a “‘sine-
wave’ song similar to that of M. nigra (Fig. 42), suggesting that voice
is not the isolating mechanism between these two sibling species. The
calls include a high-pitched twittering, and incessant sibilant notes.
Melanocharis versteri virago (Stresemann)
Fan-tailed Berrypecker
NATIVE NAMES. Fore: oowaki. Gimi: utupi.
SPECIMENS EXAMINED. Awande: 2 ¢, 1 9,1 imm. ¢@ (15-19 June 1965).
Mt. Michael: I ¢, 1 9, 1 imm. ¢ @-12 July 1964). Mit. Karimur Zone 2: 1 4;
Zone 3: 1 ©; 2 im. 63 Zone 4 1 43 Zone 5: 1 45 2)°Os Zone 6. a
624 2, ) umm. ¢; Zone 7: 1 6, 29; Zone 8:2 6,2) 931 mmm: 4 (io Aue es
Sept. 1965).
WEIGHT. W0vad: o2 97-1377 Gi9 221.0). 10) OF 1677-20001 = 08). 6 aman
$: 12.0-16.0 (13.5 + 1.3).
WING. 10 ad. ¢: 59-64 (60.6 + 1.6). 10 9: 63-72 (68.1+1.8). 6 imm. ¢: 61-
67 (63.8 + 1.9).
TAU 10ads 3S = 06-78" (735i 3.0) 10 Oe 7=10 (6526 == 3D). Grimms soos
72 (64.6 + 5.6).
STOMACH CONTENTS. Fruit 2-5 mm in diameter (11 stomachs), fruit and
insects (two), insects (one). Most stomachs of netted birds were empty, suggesting
that digestion is rapid.
TAXONOMY. This is one of the few passerine species in which
the female is larger than the male. All females I examined have heavier
weights, and all but one have longer wings, than all adult males
examined, though the males have longer tails. Immature males in
female-like plumage are intermediate in weight and wing length be-
tween adult males and females but have the short tails of the females.
As they mature, not only must males lose weight, but also their wings
must become shorter, as also true of Serzculus aureus (Diamond, 1969).
The four immature males with the longest tails (70-72) have nearly
acquired the clear gray underparts of the adult male, though they
still have female-like olive upperparts.
Eastern Highlands males are slightly darker gray below than
maculiceps from southeastern New Guinea, less dark than nominate
versteri of the Vogelkop or Cyclops specimens included in virago, and
still less dark than meeki of the Weyland and Snow Mountains. East-
ern Highlands females are darker and grayer below, less yellow, than
402
SPECIES ACCOUNTS
maculiceps and are close to a single topotypical female virago from the
Schraderberg.
BREEDING. On Mt. Karimui testes were somewhat enlarged in
almost all adult males and in half of the immature males in female-
like plumage but were not greatly enlarged in any specimen.
DISCUSSION. Melanocharis versteri, the highest species in the
M. nigra-M. longicauda-M. verstert altitudinal sequence, is common
but quite inconspicuous within its altitudinal range in forest and
dense second-growth. On Mt. Karimui, where M. longicauda was ab-
sent, M. verstert descended to 4,500 ft and was common (4% of the
local avifauna) up to the summit at 8,165 ft. At timberline it goes out
to tree ferns in alpine grassland near the forest edge. Much of its
time was spent in the lowerstory, but in fruit trees it occasionally
ranged up to 40 ft above the ground. Females, adult males, and im-
mature males were netted with the same frequency. Once I saw an
adult male hovering at flowers.
VOICE. A variety of faint and high-pitched notes, such as a thin,
rapidly repeated “‘ts-ts. ...”; a scratchy and complaining ‘“‘dee-dee-dee’’;
a high-pitched nasal “ee’’; and an upslurred inquiring ‘“‘chee-chee-
chee”. All these notes are faint, unlikely to catch one’s attention, and
not distinctive.
Melanocharis striativentris striativentris Salvadori
Streaked Berrypecker
NATIVE NAME. Fore: oowaki.
SPECIMENS EXAMINED. Awande: 8 ¢, 4 9 (19-21 June 1964; 14-20 June
1965). Miarosa: 1 g¢ (13 June 1964). Okasa: 2 g, 1 9 (23 and 24 June 1965).
Mt. Karimui Zone 2: 1 9; Zone 4: 1 ¢,2 9 (13 Aug.-2 Sept. 1965).
WEIGHT. 12 4: 16.3-20.0 (179+1.1). 8 9: 165-22.0 (19.8 + 1.7).
WING. 12 4: 69-75 (72.2415). 8 9: 71-75 (72.9+1.1).
TAXONOMY. Females have slightly heavier weights and longer
wings than males, but the difference is much less marked than in the
case of M. verstert.
‘These specimens agree well with topotypical striativentris from the
southern side of southeastern New Guinea. The race prasina of the
northern side of southeastern New Guinea differs in having the basal
portion of the tail white, while chrysocome of the Huon Peninsula
is larger, darker ventrally, and more obscurely streaked.
BREEDING. Gonads were small in most specimens but were
slightly enlarged in one male and three females.
DISCUSSION. ‘The most noteworthy feature of this species is the
extreme local variability of its abundance despite its wide geographical
range. Ihe American Museum of Natural History has a total of 14
specimens. Ihe exhaustive collections of the three prewar Archbold
expeditions included only two specimens of this species. Gilliard
403
SPECIES ACCOUNTS
found it uncommon, Shaw-Mayer missed it, and I found it uncommon
at Mt. Karimui, Okasa, and Miarosa. At Awande, however, one mile
from Miarosa and 10 miles from Okasa, it proved to be the most
abundant bird both in 1964 and 1965, and saturated our nets located
at 6,100 ft just inside the forest near its edge and adjacent to gardens.
Gyldenstolpe also found it abundant near Nondugl. The altitudinal
range was 4,750-5,960 ft on Mt. Karimui and about 4,000-6,800 ft in
the Okapa area, overlapping both M. longicauda and M. versteri (but
not M. nigra).
Rhamphocharis crassirostris piperata (De Vis)
Spotted Berrypecker
SPECIMENS EXAMINED. Miarosa: 2 ? (13 and 17 June 1964). Mt. Michael:
1 ? (13 July 1964). Mengino: 1 imm. ¢ (15 July 1964). Mt. Karimui Zone 4: 1 ?
(28 Aug. 1965). Mt. Karimui Zone 4: 1 ? (28 Aug. 1965).
WING, lei 47 ie 7
TAIL. Lamm. @: 71. 1 2: 48.
EXPOSED CULMEN. I amm:. 4: 165. 1 2: 16.0:
TAXONOMY. All specimens are in the spotted female or female-
like plumage.
DISCUSSION. This is a rare species, but it nevertheless turned up
at all of my midmontane collecting localities between 4,600 and 7,000
ft except Awande, albeit in minimal numbers. ‘Three specimens were
in the forest, one in second-growth at the forest edge, one in a fruiting
tree in second-growth, and all were shot between 7 and 50 ft above
the ground. No specimens were netted, whereas all four species of the
closely related genus Melanocharis were often netted.
Oreocharis arfaki (Meyer)
Tit Berrypecker
NATIVE NAME. Gimi: mégino.
SPECIMENS EXAMINED. Mt. Michael: 1 ¢, 2 9 (5-12 July 1964). Agotu
(Gimi territory): 1 9 (@2 July 1964). Mt. Karimui Zone 2: 1 ¢@; Zone 3: 2 9;
Zone 8: 7 4,2 9, 31mm, ¢ (17 Aug.-3' Sept. 1965).
WIGHT, 7 @2 180; 200 (@), 20-7, 214, 215, 21572 3: OF W8.0) Is78s 210s
imm. @: 18.5, 19.7, 20.0.
WING, 6 22169) 7 (4), 7b. 2595 707 7k 2 aim G95 ln
STOMACH CONTENTS. Fruit 1-7 mm in diameter.
TAXONOMY. After comparing my material with specimens from
other parts of New Guinea, including six topotypical males and two
females from the Vogelkop, I agree with Salomonsen (1960, p. 3) and
Gilliard and LeCroy (1970, p. 24) that there is too much individual
variation to recognize the race bloodi.
BREEDING. All adult males on Mt. Karimui, and one of the
immature males in female-like plumage, had the testes considerably or
ereatly enlarged.
404
SPECIES ACCOUNTS
DISCUSSION. ‘This is one of the midmontane species whose alti-
tudinal range shows local variations apparently correlated with the
presence of mossy conditions (cf., Ptiloprora guise’). On Mt. Michael,
where heavy mossing set in at 8,700 ft, Oreocharis was collected at
8,000 ft. En route from Mt. Michael to Karimui we passed through a
tongue of moss forest at about 6,000 ft near Agotu and immediately
re-encountered Oreocharis. On Mt. Karimui, where heavy mossing
began at 6,500 ft, Oreocharis descended to 4,720 ft but became common
only above 7,600 ft. In the Okapa area, where no heavy moss was
encountered as high as we collected (up to 7,000 ft), Oreocharis was
absent. It was generally in the middle- or upperstory of the forest, in
flocks, and was rarely netted. Although stomachs contained solely
fruit, I once saw an individual hanging upside-down to probe a flower.
ee 9
VOICE. A very high, drawn-out note “‘sss’” or “z-z-z’, repeated
incessantly by flocks and similar to the note of the Cedar Waxwing
(Bombycilla cedrorum, Bombycillidae) in North America.
Paramythia montium montium De Vis
Crested Berrypecker
SPECIMENS EXAMINED. Mt. Michael: 1 ¢, 1 9 (30 June and 8 July 1964).
Mt. Karimui Zone 8: 1 ¢,1 @ (5 Sept. 1965).
WEIGHT. 1 4:41. 1 eo: 43.
WING. 2 4: 97, 103. 1 9: 97.
TAIL. 2 4: 92, 94.
STOMACH CONTENTS. Fruit, 2-8 mm in diameter.
DISCUSSION. ‘This flowerpecker is common and widespread in
high altitude stunted mossy forest, from timberline down to 9,500 ft
(Mt. Michael) and on the very summit (8,165 ft) of Mt. Karimui. Most
of its altitudinal range lies above that of its smaller relative Oreocharis
arfaki. Paramythia montium feeds unsystematically on berries: it flies
into a tree with a loud wing beat, hops around vigorously, plucks a
few berries, and flies off again, leaving the tree full of uneaten berries.
It spends more time in the crowns than in the understory.
VOICE. A faint squeaky note, suggestive of the sounds made by
birdwatchers squeaking or sucking on the back of their hands to attract
birds.
ZOSTEROPIDAE: WHITE-EYES
Zosterops atrifrons tenuifrons Greenway!
Black-fronted White-eye
SPECIMENS EXAMINED. Karimui: 1 ¢, 1 ? (12 Aug. 1964). Bomai: 1 9,1?
(6 July 1965).
1 Listed as Zosterops minor in Rand and Gilliard (1967).
405
SPECIES ACCOUNTS
WEIGH E. 1 ios k0, Pe,
WING. 1 @: 58. 1 ?: 57.
TAXONOMY. ‘The black of the forehead is restricted to a small
area at the base of the bill and is much more limited than in delicatula
of the southern slopes of southeastern New Guinea, in which the
black extends beyond the eyes. In this respect my specimens agree
with the description of tenuifrons (Greenway, 1935, p. 105; perhaps
synonymous with chrysolaema) from the northern slopes of south-
eastern New Guinea, of which I have seen no topotypical material.
BREEDING. ‘The male had the testes greatly enlarged.
DISCUSSION. At Karimui and Bomai this hill forest white-eye
was uncommon (ca. 0.2% of the local avifauna) and was encountered
in groups of up to four birds in the upperstory of forest and second-
growth, sometimes in flowering trees. Z. novaeguineae lives at higher
altitudes, but the ranges may overlap slightly, since Bulmer collected
both species around 5,000 ft in Kyaka territory.
VOICE. A lengthy song with a sweet quality. The call is a high-
pitched, inquiring, sweet, upslurred “‘tswee.”
Zosterops novaeguineae wahgiensis Mayr and Gilliard
Mountain White-eye
NATIVE NAMES. Fore: tioriéba. Gimi: Ifyu.
SPECIMENS EXAMINED. Awande: 1 ¢,1 9, 1? (17 and 18 June 1965). Mt.
Michael: 1 9 (30 June 1964).
WEHIGHE. 1451207 150-120. sso:
WING. 1 (Gt Onnote TenOl:
STOMACH CONTENTS. Fruit 1-3 mm in diameter (nine stomachs); insects
(four); insects and fruit (two).
TAXONOMY. Mayr and Gilliard (1951, pp. 14-15) and Gylden-
stolpe (1955, p. 176) recognized three races of Z. novaeguineae, two of
them (wahgiensis and shawmayert) new, within an area 45 miles in
extent containing no distributional barriers to the species. Examination
of the unique type of shawmayeri shows that it 1s indistinguishable from
wahgiensis, as Mees (1961) suspected and as Rand and Gilliard (1967,
p. 591) and Mayr (1967, p. 309) recently agreed. All Eastern Highlands
material collected to date belongs to a single thinly differentiated race
wahgiensis, which differs from crissalis in the slightly darker, more
ereen, less yellow back and the more gray, less powder-brown abdomen.
My specimens differ from Nondugl wahgiensis mainly in having a
slightly brighter yellow throat.
BREEDING. The gonads of all specimens were small.
DISCUSSION. Zosterops novaeguineae occurs in flocks in the
crowns and middlestory, occasionally the understory, of forest at about
5,000-7,500 ft. It is somewhat commoner in disturbed habitats (casu-
arina stands, bushes of second-growth).
406
SPECIES ACCOUNTS
VOICE. The common call note is a sweet, down-slurred “tsyew”
with a sharp attack and suggestive of a Lonchura mannikin. ‘This note
is faint, but the pooled sound of many members of a flock calling at
once is loud. There is also a short, dry, upslurred, trilled call mote
similar to that of Zosterops lateralis of New Zealand and Z. hypoxantha
of New Britain. The lengthy, sweet song of whistles and upslurs
suggests the song of Saxicola caprata in quality.
ESTRILDIDAE: WAXBILLS and MANNIKINS
Erythrura trichroa sigillifera (De Vis)
Blue-faced Parrot-Mannikin
SPECIMENS EXAMINED. Karimui: 1 ¢ (6 Aug. 1965). Mt. Karimui Zone 8:
lL @ (7 Sept. 1965),
WEIGHT. 2 4: 15:8; 15.0:
WING. 2. 6: 62, G2.
TAU, 2 63747, 48.
EXPOSED CULMEN. 2 ¢: 11.5, 12.0.
STOMACH CONTENTS. Dry granular vegetable matter, possibly bamboo seeds.
BREEDING. ‘Testes were considerably enlarged in both specimens.
DISCUSSION. ‘This mannikin lives in forest and second-growth
at 3,500-9,200 ft and is solitary and uncommon, except in thickets of
wild bamboo, where it is numerous. Like Gallicolumba beccarit, it
becomes locally superabundant when the bamboo produces seed.
When disturbed, it flies up with a loud whirring of the wings.
VOICE. The call is a very rapid descending series of 2-5 thin,
unvoiced notes, similar in quality to the song of Peltops montanus
or to light spiccato playing on the violin.
Erythrura papuana Hartert
Papuan Parrot-Mannikin
NATIVE NAME. Fore: kukusia.
SPECIMENS EXAMINED. Awande: 1 ¢,1 9, 2 ? (19 June 1964; June 14-17,
1965). Okasa: 1 9 ? (22 June 1965).
WEIGHS. I s@2518s. “hoe 73. 1 Oe I. “bers US.5;
WENG. 1 °g2 ao. “TD Ot 67. 140" fe 67. Zork 05; GS,
TAM, J 4:48. 1 9:50, 1° 7: 87. 2 >: 3% 49.
BAPOSsED GULMEN. LL g27i5. 1°93 12.5, 1 oO Py 130. 222 130, 139;
STOMACH CONTENTS. Dry, granular vegetable matter.
TAXONOMY. ‘These five specimens, plus two specimens from the
Vogelkop (including the type) and 10 from southeastern New Guinea
in the American Museum of Natural History, were compared critically
with the available large series of the very similar E. trichroa sigillifera
to evaluate diagnostic differences. ‘The only difference in color is that
the chin has a blue patch of significant extent in adult males of E.
407
SPECIES ACCOUNTS
papuana, lacking (usually) or present but very small (infrequently)
in adult males of E. trichroa sigillifera (and in females of both species).
The stouter bill of adult males of E. papuana is visible at a glance.
Measurements (‘Table 19) show that all or almost all E. papuana have
longer wings, longer bills, and heavier weights than all or almost all
E. trichroa sigillifera. Colors of the soft parts in life were the same for
my specimens of both species (bill black, eye brown, legs pale flesh-
brown).
DISCUSSION. ‘Three specimens were netted in second-growth near
the forest edge and one shot in the forest at 6,200 ft near Awande.
The fifth specimen was shot at 4,000 ft in the Okasa forest nine miles
from Awande.
The only other records of this rare species are from four localities
in southeastern New Guinea (Aroa River, Kotoi, Mt. Tafa, Mt. Albert-
Edward), from the Wissel Lakes of western New Guinea, and from the
Arfak and ‘Tamrau Mountains of the Vogelkop. The distributional
relationship with the widespread and much commoner E. trichroa
sigillifera can be described approximately as checkerboard allopatry.
In the Okapa area (4,000 and 6,200 ft) I obtained only E. papuana,
while elsewhere in the Eastern Highlands (3,500-9,200 ft: Karimui,
Mt. Karimui, Mt. Kubor, Mt. Wilhelm, Mt. Hagen, Mt. Giluwe,
Wahgi Divide Mountains, Weiga, Schraderberg, Baiyer Valley, Kaironk
Valley) other collectors and I obtained only E. trichroa. In southeastern
New Guinea the First Archbold Expedition collected E. papuana on
the east slope of Mt. Tafa, E. trichroa on the west slope of Mt. Tafa and
at three other localities. At the Wissel Lakes and at Bon Kourangen
in the ‘l'amrau Mountains only E. papuana was taken. ‘There are only
two instances of local sympatry: both species were recorded at Mayr’s
TABLE 19
MEASUREMENTS AND WEIGHTS OF Erythrura papuana AND E. trichroa sigillifera
Wing Exposed Culmen Weight
E. trichroa
sigillifera
17 8 59 (2), 61 (2), 62 (7), LOK) y 1ie(8); LE (1), 122); 1316);
Garg), 0% 12.0 (7) 14(2); 153), LO(a
17 (2); 18a)
139 60 (5), 61, 62 (4), LTOKS); Wil (2), 121), SS) ae),
63, 64 (2) 12.0 (7) 1b(2); 16(), Wey
18 (1)
E. papwuana
92 65, 66, 67 (5), 68, 69 12-5, 1300, 13:51(2),9 180; 18.5, 19.0, 921005
14:0'(2), 14.5°(2) 24.0
49 67 (2), 68 (2) 12.5 (2)y. WO; 185 h73; 1870,20:0) 2100
4? 65, 66, 67, 68 13.0 (4) 1710, W815, 190) 2100
a
408
SPECIES ACCOUNTS
locality of Siwi in the Arfak Mountains of the Vogelkop; and on Mt.
Albert-Edward in southeastern New Guinea I found E£. trichroa abun-
dant at 6,000-9,200 ft, E. papuana at 5,000-6,000 ft. No differences in
habitat preference, altitudinal preference, or behavior have been noted,
except for the altitudinal segregation on Mt. Albert-Edward. Both
forms inhabit forest and the forest edge from about 3,000 to 7,000 ft
(E. papuana) or 9,000 ft (E. trichroa). Unlike the situation among the
Geospiza finches of the Galapagos Islands, the difference in bill size
of these mannikins is evidently not sufficient to permit local overlap.
E. papuana is endemic to New Guinea and presumably is the older
invader, while E. trichroa extends from Celebes to Micronesia. ‘The
relict distribution of E. papuana suggests that its congener is in the
process of replacing it and that it has been able to hold out only at a
few scattered localities.
Lonchura grandis grandis (Sharpe)
Great-billed Mannikin
SPECIMENS EXAMINED. Okasa: 1 9 (26 June 1965).
WEIGHT. 16.
WING. 56.
TAXONOMY. The specimen has a duller back, a duller and more
yellow-edged tail, and paler brown flanks than nominate grandis from
southeastern New Guinea. However, it can be matched by some speci-
mens of grandis in the first two respects and is approached by some
grandis in the third respect. The race ernesti has the edges of the
rectrices still paler, duller, yellower, and less ochraceous, while heurni
has the back richer chestnut.
DISCUSSION. In 1964 I observed a flock of five of this species at
4,200 ft in the extensive grasslands surrounding the Okasa forest. In
1965 L. spectabilis, the widespread Lonchura of the midmontane grass-
lands in the Eastern Highlands, was the common form at Okasa, and
only the single specimen of L. grandis was obtained. Mixed flocks con-
taining both species were never observed.
Examination of the distribution of Lonchura mannikins in the mid-
montane grasslands of New Guinea proves interesting. Extensive mid-
montane grasslands probably came into existence only within the past
millenium, as a result of native agriculture, and must originally have
been confined to very narrow strips along a few of the larger lakes and
rivers, and perhaps landslide areas. A total of eight species of Lonchura
may now be found in the midmontane grasslands in various parts of
New Guinea. Of these eight, two are localized species (L. vana and
L. teerinki) endemic to this habitat; five are lowland species (one, L.
caniceps, confined to southeastern New Guinea; the other four—L.
tristissima, L. grandis, L. castaneothorax, and L. spectabilis—wide-
spread) that appear to have spread upwards to the new habitats as
409
SPECIES ACCOUNTS
they were created; and the eighth, L. montana, an alpine species that
has spread downward. Apart from Okasa in the Eastern Highlands
(L. grandis and L. spectabilis) and Mafalu in southeastern New Guinea
(L. grandis and L. caniceps), each midmontane locality supports only
a single species, but the identity of the locally successful colonist varies
in an unpredictable checkerboard fashion over New Guinea. An
extreme example of this irregularity occurs in the Herzog Mountains,
where four different colonists occur at four different localities within
35 miles: L. tristissima in the Snake River Valley, L. castaneothorax
at Mumena Creek, L. grandis at Biolowat, and L. spectabilis at Wau.
The details for the rest of New Guinea, proceeding east to west, are
as follows: southeastern New Guinea, L. caniceps (plus L. grandis at
Mafulu); Herzog Mountains, cited in previous sentence; Huon Penin-
sula, L. tristissima at some localities, L. spectabilis at others; Adelbert
Mountains, L. tristissima; Eastern Highlands, L. spectabilis (plus L.
grandis at Okasa); ‘Vorricelli Mountains, L. tristisstma; Baliem Valley
of the Snow Mountains, L. teerinki; Ilaga Valley of the Snow Moun-
tains, L. montana; Wissel Lakes, L. castaneothorax; Weyland Moun-
tains, L. tristissima; Anggi Lakes of the Vogelkop, L. vana; other
localities in the Vogelkop, L. tristissima. As discussed on p. 40, this
checkerboard pattern probably reflects the facts that the midmontane
grasslands provide islands of suitable habitat for the Lonchura man-
nikins, that establishment of the first successful colonist prevents the
establishment of later potential colonists, and that it is in part a
random matter which species arrives first.
Lonchura spectabilis wahgiensis Mayr and Gilliard
and L. s. gajduseki Diamond
New Britain Mannikin
NATIVE NAMES. Fore: shui. Daribi: buba.
SPECIMENS EXAMINED. L. s. wahgiensis: Awande, 1 9,1 imm. ¢ (19 Aug.
1964; June 17, 1965); Mengino, 1 ? (16 July 1964). L. s. gajduseki: Karimui, 8 ¢,
4 9,4 imm. 2, 1 juv. ? (3-17 July 1965).
WEIGHT. L. s. wahgiensis: 1 imm. @, 9.3. L. s. gajduseki: 8 @, 10.5-12.7
(7 30:7); 3,9 10.3; 11.0, EL; 4 tom. ?; 9:5; WL (2), Libs 1 juve P70:
WING. L. s. wahgiensis: 1 imm. 6,50. L. s. gajduseki: 6 ¢, 49, 50, 51 (3), 52;
4 ©, 49, 50 (2), 51; 1 imm. ?, 51.
TAXONOMY. L. s. wahgiensis is the widespread Lonchura of the
midmontane grasslands west at least to ‘Tari and east at least to the
Okapa area, with the underparts whitish (sometimes pale buff in
adults and medium buff in immatures). L. s. gajyduseki is a distinctive
form confined to the Karimui Basin, with the underparts medium
buff in both adults and immatures (see Diamond, 1967a, p. 15, for
diagnosis).
410
SPECIES ACCOUNTS
BREEDING. The testes of all adult L. s. gajduseki were much
enlarged, indicating synchronized breeding activity in the dry season.
DISCUSSION. Flocks of up to 30 individuals are seen between
3,000 and 8,000 ft in the Eastern Highlands at any locality where there
are large expanses of medium-tall grassland. Unlike L. tristissima,
which sometimes perches in bushes and trees at the forest edge or in
second-growth, L. spectabilis seems quite unable to utilize forest edge
habitats. My general impression of New Guinea Lonchura species
(with the possible exception of L. tristissima) is that they are among
the last grassland birds to colonize a new garden or airstrip carved out
of the forest and appear only when the available area of grassland is
substantial, possibly because of their inability to use the forest edge
and their need to associate in large flocks.
The presence of two endemic midmontane races of Lonchura
spectabilis raises some evolutionary problems (see pp. 76-77). These
problems are less acute in the case of L. s. wahgiensis, which differs
only slightly from L. s. mayri of the lowlands and now has a large
midmontane range, extensive parts of which have probably been under
cultivation for at least several centuries. ‘This timespan and the slight
racial differences make it plausible that wahgiensis could have evolved
in the last few centuries. An alternative hypothesis is that it may be
much older and may have eeked out a precarious existence along the
edge of the Wahgi and other rivers prior to extensive native agri-
culture (cf., Rand and Brass, 1940, p. 377; Rand, 1941b).
L. s. gajduseki is more distinct and inhabits a much smaller range
with a well-known recent history. Information given by natives sug-
gests that the Karimui Basin has been inhabited somewhat less than a
century. Originally the natives’ diet was mainly sago, and dependence
on gardens began only with the termination of intertribal warfare
and the introduction of the sweet potato in the 1950's. Having flown
and walked over much of the basin, I find it impossible to imagine
where L. s. gajduseki could have maintained itself before garden
agriculture. The streams traversing the basin are confined to narrow
gorges. Although I have been in 10 of these gorges, I have yet to meet
a Lonchura or to see riparian vegetation that seemed to offer promise
of supporting L. s. gajdusekt. Either this race must have slowly evolved
in some part of the basin not yet discovered—an unattractive pos-
sibility because of the circumscribed size of the basin and the large
areas of grassland necessary for L. spectabilis—or else it must have
evolved in a very short time, perhaps as little as 15 years. This latter
possibility cannot be dismissed. The studies of Johnston and Selander
(1964) on Passer domesticus in North America show that rapid morpho-
logical changes can occur in populations of species colonizing new
habitats. ‘The variants of wahgiensis in which the adult has pale buff
underparts may have served as the starting material. Finally, the
411
SPECIES ACCOUNTS
selection pressures at Karimui (due to heavy rainfall throughout the
year) may have been unusually severe, as seen by the general tendency
towards dark plumage expressed most strikingly in the melanistic
population of Accipiter novaehollandiae.
VOICE. A faint, sweet upslur.
Oreostruthus fuliginosus hagenensis Mayr and Gilliard
Crimson-sided Mountain Grass Finch
STOMACH CONTENTS. Seeds (two stomachs), fruit (one), insects (one).
DISCUSSION. ‘This uncommon high-altitude grass finch was col-
lected in a forest glade on Mt. Hagen by Gilliard and in alpine grass-
land on Mt. Giluwe by Shaw-Mayer, at 8,500-11,500 ft.
412
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419
INDEX
Citations of all significant page references for each New Guinea bird discussed
are provided both under the generic name and under the specific name. Boldface
type is used for the first page of the main account of a bird that occurs in the
Eastern Highlands. Birds occurring in the Eastern Highlands are also cited under
the English name, the subspecific name, and any scientific name used by Rand and
Gilliard (1967) but replaced in the present work. Such citations refer only to the
first page of the main account.
Acanthiza murina, 96, 221, 222, 224, 295,
227
Accipiter buergersi, 42, 43, 47, 104, 108
fasciatus, 35, 45, 72, 78, 107, 249
melanochlamys, 35, 85, 108
novaehollandiae, 42, 43, 45, 56, 85, 105,
107, 108
poliocephalus, 35, 42, 43, 58, 85, 105,
107, 108
Aceros plicatus, 60, 85, 193
Acrocephalus arundinaceus, 73, 233, 235
Actitis hypoleucos, 120
acuminata, Calidris, 79, 121
Erolia, 121
adamsoni, Coracina, 202
adolphinae, Myzomela, 21, 36, 47, 74, 76,
80, 358, 359
Aegotheles albertisii, 39, 41, 43, 177, 178
archboldi, 39, 41, 47, 177
bennettii, 51, 54, 56, 176
insignis, 43, 60, 178
wallacii, 176
Aepypodius arfakianus, 37, 113
affinis, Milvus, 103
Ailuroedus buccoides, 27, 51, 54, 55, 64,
85, 89, 347
crassirostris, 27, 52, 54, 55, 64, 203,
347, 348
alaris, Gallicolumba, 137
alba, Egretta, 99
Lyte, 36, 74, 173
alberti, Lophorina (Pteridophora), 12,
20, 31, 46, 64, 321, 339
Pteridophora, 314, 320, 321, 339
albertisii, Aegotheles, 39, 41, 43, 177, 178
Drepanornis, 309, 310, 329
Epimachus (Drepanornis), 63, 320, 321,
329
Gymnophaps, 56, 58, 132
albifrons, Henicophaps, 59, 139
albigularis, Pachycephalopsis, 270
albispecularis, Heteromyias, 24, 37, 262,
263, 270
421
albogularis, Conopophila, 350
Eurostopodus, 79, 179
albolimbata, Rhipidura, 29, 38, 62, 69,
239, 240, 242, 244, 246, 248
albonotata, Meliphaga, 369
Poecilodryas, 62, 262, 263, 265
alboscapulatus, Malurus, 73, 76, 78, 84,
85, 165, 166, 214, 216, 217, 234, 260
Alcyone azurea, 183
alecto, Monarcha, 254
Myiagra, 74, 254, 268
Alisterus chloropterus, 52, 59, 157
Amalocichla incerta, 37, 87, 208
sclateriana, 18, 20
Amaurornis olivaceus, 51, 54, 74, 78, 84,
118
Amblyornis macgregoriae, 22, 53, 64, 91,
342
subalaris, 22, 47, 343
amboinensis, Macropygia, 35, 43, 59, 74,
78, 133, 135, 136, 137
amethystina, Cnemophilus (Loria), 49
amethystinus, Cnemophilus (Loria), 322
anais, Mino, 22
analoga, Meliphaga, 37, 38, 42, 44, 65,
361, 362, 363, 364, 365, 366, 368, 371,
372, 373, 374, 378
Anas gibberifrons, 72, 100
querquedula, 79, 100
superciliosa, 72, 100
Androphobus viridis, 24, 47
Anhinga rufa, 99
Anthus gutturalis, 48, 71, 205
novaeseelandiae, 73, 76, 205, 249
Anurophasis monorthonyx, 47
Aplonis cantoroides, 79, 296
metallica, 80, 296
apoda, Paradisaea, 316, 321, 340
approximans, Circus, 45, 54, 72, 78, 80,
110
archboldi, Aegotheles, 39, 41, 47, 177
Eurostopodus, 179
Petroica, 47
INDEX
Archboldia papuensis, 20, 24, 46, 48, 49,
342
ardens, Sericulus, 346
arfaki, Oreocharis, 12, 37, 65, 404, 405
Oreopsittacus, 38, 59, 86, 146, 148, 149,
150
Tyto, 173
arfakiana, Melanocharis, 18, 19, 20, 21,
25, 47
arfakianus, Aepypodius, 37, 113
Sericornis, 30, 34, 38, 54, 55, 61, 67,
217, 218, 223
Arses telescophthalmus, 253
Artamus leucorhynchus, 296
maximus, 72, 75, 78, 85, 89, 295
arthuri, Pitohui, 293
aruensis, Geoffroyus, 156
Meliphaga, 38, 42, 44, 65, 362, 363, 364,
365, 366
Monarcha, 252
arundinaceus, Acrocephalus, 73, 233, 235
Astrapia mayeri, 338
Princess Stephanie’s, 337
Ribbon-tailed, 338
stephaniae, 337
ater, Manucodia, 307, 308, 321, 325
aterrimus, Probosciger, 153
atra, Chalcopsitta, 141
Rhipidura, 38, 54, 5b, 62, 239) 240,
244, 245, 246, 250, 324
atrifrons, Zosterops, 52, 405
atrovirens, Lalage, 39, 196, 207
auga, Meliphaga, 36, 44, 52, 74, Dy Ws
85, 89, 355, 362, 363, 364, 365, 366,
367
aurantiifrons, Loriculus, 164
Ptilinopus, 123, 126
aurea, Pachycephala, 25
aureaus, Xanthomelus, 346
aureus, Sericulus, 346, 402
Aviceda subcristata, 54, 58, 85, 102, 239
axillaris, Monarcha, 38, 54, 62, 249, 251
azurea, Alcyone, 183
azureus, Ceyx, 51, 72, 183, 184
bastille, Meliphaga, 370
beccarii, Gallicolumba, 37, 59, 138, 407
Goura, 140
Sericornis, 217, 224
Bee-eater, Rainbow, 191
belfordi, Melidectes, 35, 39, 46, 48, 49,
377, 387, 388, 389, 391, 392, 393
bella, Charmosyna, 146
bellus, Ptilinopus, 125
bennettii, Aegotheles, 51, 54, 56, 176
Casuarius, 37, 53, 54, 55, 97
422
benecora, Falco, 35,72, 78; 85, 90) 112
Berrypecker, Black, 398
Crested, 405
Fantailed, 402
Midmountain, 401
Spotted, 404
Streaked, 403
Tit, 404
bimaculatus, Peneothello, 37, 52, 62, 262,
263, 268
Bird, Cicada, 199
Bird of Paradise, Blue, 341
King, 337
King of Saxony, 339
Lawes’ Six-wired, 332
Lesser, 341
Loria’s, 322
Magnificent, 335
Multi-crested, 320
Queen Carola’s Six-wired, 332
Raggiana, 340
Superb, 330
Yellow-breasted, 323
Bittern, Black, 100
Forest, 99
bivittata, Petroica, 48, 262
Blackbird, New Guinea, 209
blamivilliis Peltops, 275 375. (OL 745. sp;
236, 239
bloodi, Epimachus, 328
bonapartii, Todopsis, 216
Bowerbird, Archbold’s, 342
Golden, 346
Lauterbach’s, 346
MacGregor’s, 342
boyeri, Coracina, 36, 45, 54, 60, 197, 198,
201
brachyrhyncha, Rhipidura, 38, 62, 239,
240, 242, 243, 245
brehmii, Psittacella, 43, 59, 158, 159, 160,
161
brevicauda, Paradigalla, 306, 316, 321,
327
brevipes, Heteroscelus, 120
Tringa, 79, 120
brevirostris, Drymodes, 206
bruijni, Grallina, 72, 261, 299
Pomareopsis, 299
bruijnii, Drepanornis, 309, 310
Epimachus (Drepanornis), 321, 329, 330
Micropsitta, 153
brunneiceps, Piohui, 286
buccoides, Ailuroedus, 27, 51, 54, 55, 64,
85, 89, 347
buergersi, Accipiter, 42, 43, 47, 104, 108
Sericornis, 220
INDEX
Butcherbird, Black, 301
Black-headed, 300
Buzzard, Long-tailed, 103
Cacatua galerita, 59, 154, 155, 156
Cacomantis castaneiventris, 30, 35, 53,
60, 165, 167, 168
pyrrhophanus, 30, 73, 76, 79, 90, 165,
166
variolosus, 30, 35, 52, 74, 78, 85, 164,
166, 167, 168, 170, 216
caeruleiceps, Trichoglossus, 142
caeruleogrisea, Coracina, 43, 48, 53, 60,
197, 202
caerulescens, Eupetes, 27, 211
caeruleus, Elanus, 72, 102
caledonicus, Nycticorax, 99
Calidris, acuminata, 79, 121
ruficollis, 79, 121
Caliechthrus leucolophus, 60, 171
callopterus, Alisterus, 157
Caloenas nicobarica, 122
Campochaera sloetii, 24, 54, 60, 195, 199
caniceps, Lonchura, 40, 409, 410
cantoroides, Aplonis, 79, 296
Capella hardwickii, 121
megala, 12]
capensis, Tyto, 36, 73, 76, 173, 174
caprata, Saxicola, 74, 76, 78, 206, 407
Caprimulgus macrurus, 73, 178
captata, Melanocharis, 401
captus, Rallus, 116
carbonarius, Dicrurus, 302
carolae, Lophorina (Parotia), 22, 39, 48,
321, 332, 333
Parotia, 311, 312, 321, 332
carunculata, Paradigalla, 316, 321
caspica, Motacilla, 205
cassicus, Cracticus, 36, 54, 74, 75, 78, 85,
300
Cassowary, Double-wattled, 98
Dwarf, 97
castaneiventris, Cacomantis, 30, 35, 53,
60, 165, 167, 168
castaneothorax, Lonchura, 409, 410
castanonotus, Eupetes, 27, 36, 37, 61, 210,
213, 241, 273
Causarius bennettii, 37, 53, 54, 55, 97
casuarius, 37, 39, 54, 55, 98
unappendiculatus, 39
casuarius, Casuarius, 37, 39, 54, 55, 98
Catbird, Green, 347
White-eared, 347
cenchroides, Falco, 79, 90, 111
centralis, Heteromyias, 270
423
Centropus menbeki, 36, 54, 60, 139, 172
phasianinus, 36, 73, 172, 173
cervicalis, Otidiphaps, 139
Psittaculirostris, 150
cervinicaudus, Epimachus, 329
Ceyx, azureus, 51, 72, 183, 184
lepidus, 60, 183
Chaetorhynchus papuensis, 63, 303
Chaetura novaeguineae, 180
Chalcites lucidus, 169
meyerii, 169
ruficollis, 169
chalconota, Ducula, 24, 27, 131, 132
Chalcophaps stephani, 37, 54, 59, 137
Chalcopsitta atra, 141
duivenbodei, 141
scintillata, 59, 140
superspecies, 50, 151
chalybatus, Manucodia, 62, 85, 89, 307,
308, 321, 324, 327
Charmosyna josefinae, 47
multistriata, 25
papou, 29, 86, 145, 146, 147, 149
placentis, 29,954; 55; 59) 86,146, 447,
152
pulchella, 29, 54; 55, 59; 36, 1467924
wilhelminae, 25, 147
Chat, Pied, 206
chinensis, Coturnix, 73, 76, 115
Excalfactoria, 115
Chlamydera lauterbachi, 73, 346
chloronota, Gerygone, 25, 36, 227
chloroptera, Chalcopsitta, 140
Melanocharis, 398
chloropterus, Alisterus, 52, 59, 157
Chrysococcyx lucidus, 169, 170
malayanus, 169, 170
meyer, 20,60, 169
ruficollis, 169, 170
chrysogaster, Gerygone, 36, 61, 225, 226,
227
chrysogenys, Oreornis, 47
chrysomela, Monarcha, 37, 38, 54, 62,
252
chrysoptera,
335
Neositta, 348, 349
chrysotis, Xanthotis, 374
Cicinnurus regius, 306, 310, 311, 321,
337
cinerea, Gerygone, 36, 61, 225, 227, 257
Motacilla, 79, 205
cinereus, Poliolimnas, 117
Pycnopygius, 65, 378, 379
cinnamomeus, Ailuroedus, 347
Lophorina . (Diphyllodes),
INDEX
Circus approximans, 45, 54, 72, 78, 80,
110
spilonotus, 45, 72, 109, 110
Cisticola exilis, 73, 91, 235, 236
clamosa, Rhipidura, 51, 241
Climacteris leucophaea, 18, 19, 20, 270
placens, 18
Clytoceyx rex, 60, 184, 186
Clytomyias insignis, 61, 87, 91, 216
Cnemophilus macgregorii, 22, 47, 49, 64,
67, 318, 320, 322
loriae, 22, 31, 49, 64, 320, 321, 322
Cockatoo, Palm, 153
Sulphur-crested, 154
Collocalia esculenta, 36, 43, 60, 72, 75, 78,
85, 180, 181, 182
hirundinacea, 36, 43, 72, 75, 78, 85, 88,
180, 181
papuensis, 182
vanikorensis, 181]
whiteheadi, 43, 72, 180, 182
Colluricincla harmonica, 36, 74, 274, 286
megarhyncha, 36, 53, 63, 87, 89, 274,
284
Columba vitiensis, 133
comrii, Manucodia, 307, 308, 321
Coracina boyeri, 36, 45, 54, 60, 197, 198,
201
caeruleogrisea, 43, 48, 53, 60, 197, 202
lineata, 17, 18, 20, 21, 25, 47, 197
longicauda, 197, 203, 204
melaena, 36, 45, 51, 60, 197, 198
montana, 29, 33, 43, 60, 196, 197, 199,
200, 202
morio, 43, 45, 197, 199, 402
novaehollandiae, 79, 197, 203
papuensis, 36, 74, 185, 197, 204, 205
Sclusticeps, 29), 33, 30, 4, 45, 60, 196,
197, 198, 199, 200, 201, 202
tenuirostris, 36, 74, 197, 199, 200
Cormorant, Little Black, 98
Little Pied, 98
coromandelianus, Nettapus, 72, 101
coronulatus, Ptilinopus, 39, 40, 123, 125,
128
Corvus orru, 249
tristis, 63, 304
Coturnix chinensis, 73, 76, 115
Coucal, Greater, 172
Pheasant, 173
Cracticus cassicus, 36, 54, 74, 75, 78, 85,
300
quoyi, 36, 63, 300, 301
Crake, Marsh, 116
Spotless, 117
White-browed, 117
424
Craspedophora magnifica, 333
crassirostris, Ailuroedus, 27, 52,
64, 203, 347, 348
Rhamphocharis, 404
Crateroscelis murina, 10, 27, 28, 29, 30,
35. 36, 07, ol, Glo 69) 875 2062 229;
230, 232, 233, 243, 248
misronuta, 25; 29, 35,37, ol, 220.0231
robusta, 10, 23, 27, 28, 29, 30; 35,36,
37, Ol, G9, 229; 230; 2315252
cristatus, Pitohui, 37, 274, 286, 293
Crow, Bare-faced, 304
cruentata, Myzomela, 37, 64, 86, 358, 359,
360
cryptoleucus, Peneothello, 22, 47
Cuckoo, Chestnut-breasted Brush, 168
Fan-tailed Brush, 166
Golden Bronze, 169
Gray-breasted, 164
Meyer’s Bronze, 169
Mountain Bronze, 169
Oriental, 164
Cuckoo-shrike, Golden, 195
Cuculus saturatus, 79, 164
cuicui, Microeca, 51
cuvierl, Talegalla, 113
cyanocephala, Todopsis, 74, 216
cyanoleuca, Myiagra, 79, 254
cyanus, Peneothello, 22, 36, 37, 62, 69, 87,
262, 263, 266, 267, 271
549);
Dacelo gaudichaud, 54, 74, 78, 85, 185,
186
leachii, 187
novaeguineae, 187
Daphoenositta miranda, 24, 49, 348, 349
Darter, Austalian, 99
dauma, Oreocincla, 207
Zoothera, 37, 207
decora, Paradisaea, 316, 321
desmarestii, Psittaculirostris, 39, 54, 59,
81, 85, 89, 141, 150, 151
devisi, Rhipidura, 243
devittatus, Domicella, 145
Dicaeum geelvinkianum, 37, 52, 65, 86,
350, 398
dichrous, Pitohui, 45, 46, 63, 76, 78, 251,
274, 286, 289, 292
Dicrurus hottentottus, 54, 74, 78, 85, 156,
302, 303
diminuta, Cisticola, 235
dimorpha, Uroglaux, 24
diophthalma, Opopsitta, 52, 152
Diphyllodes magnificus, 306, 310, 311,
Sid, dl6, 320) 321, 335
respublica, 306, 310, Sil; Sis, 321
INDEX
Dollarbird, 192
Domicella hypoinochroa, 144, 145
lory, 51, 59, 144, 145, 154, 204, 398
dominica, Pluvialis, 79, 120
doriae, Megatriorchis, 24, 85, 109
dorsalis, Pachycephala, 283
Dove, Amboina Cuckoo, 133
Beautiful Fruit, 124
Beccari’s Ground, 138
Black-billed Cuckoo, 135
Great Cuckoo, 136
Least Fruit, 126
Little Coroneted Fruit, 125
Magnificent Fruit, 129
Ornate Fruit, 127
Pink-spotted Fruit, 128
Red-throated Ground, 137
Stephan’s Ground, 137
Superb Fruit, 124
White-breasted Fruit, 125
White-breasted Ground, 139
Drepanornis albertisii, 309, 310, 329
bruijnii, 309, 310
Drongo, Mountain, 303
Spangled, 302
Drymodes superciliaris, 25, 37, 61, 87,
206
ducalis, Lophorina (Astrapia), 337
Duck, Black, 100
Ducula chalconota, 24, 27, 131, 132
pinon, 130, 133
rufigaster, 27, 45, 58, 131, 132
zoeae, 45, 58, 85, 89, 132, 133
duivenbodei, Chalcopsitta, 141
dumontii, Mino, 54, 74, 78, 193, 297
Dupetor flavicollis, 100
Eagle, New Guinea, 109
Red-backed Sea, 104
edwardi, Porzana, 117
edwardsii, Psittaculirostris, 39, 51, 151,
153
Egret, Plumed, 99
Egretta alba, 99
intermedia, 99
Elanus caeruleus, 72, 102
elisabeth, Halcyon, 189
eques, Myzomela, 37, 44, 51, 54, 64, 358,
359
Epimachus albertisii, 63, 320, 321, 329
bruijnii, 321, 329, 330
fastosus, 22, 31, 47, 63, 163, 309, 310,
314, 321, 328
meyeri, 22, 31, 49, 163, 309, 310, 314,
321, 328
erebus, Turdus, 209
425
ernesti, Falco, 110
Erolia acuminata, 121
ruficollis, 121
erythrogaster, Pitta, 37, 60, 86, 193, 194
erythropleura, Ptiloprora, 22, 47, 385, 386
erythrothorax, Domicella, 144
Erythrura papuana, 17, 21, 39, 41, 47,
407
trichroa, 39, 41, 65, 407, 408
esculenta, Collocalia, 36, 43, 60, 72, 75,
78, 85, 180, 181, 182
Eudynamis parva, 170
scolopacea, 37, 54, 80, 167, 170, 171
Eugerygone rubra, 61, 96, 256
Eulacestoma nigropectus, 63, 294
Fuipetes caerulescens, 27, 211
castanonotus, 27, 36, 37, 61, 210, 213,
241, 273
High Mountain, 212
leucostictus, 27, 36, 37, 61, 211, 212
Midmountain, 210
Eurostopodus albogularis, 79, 179
archboldi, 179
mystacalis, 179
papuensis, 179
Eurystomus orientalis, 45, 74, 78, 79,
85, 90, 192, 297
Excalfactoria chinensis, 115
excelsa, Psittacella, 160
excitus, Cacomantis, 166
exiguus, Anthus, 205
exilis, Cisticola, 73, 91, 235, 236
facialis, Meliphaga, 371
Falco berigora, 35, 72, 78, 85, 90, 112
cenchroides, 79, 90, 111
peregrinus, 110
severus, 35, 111
Falcon, Brown, 112
Peregrine, 110
fallax, Monarcha, 249
Fantail, Black, 245
Chestnut-bellied, 245
Dimorphic Rufous, 243
Gray-breasted Rufous, 241
Sooty Thicket, 240
White-breasted Thicket, 241
White-eared, 246
White-throated, 247
fasciatus, Accipiter, 35, 45, 72, 78, 107,
249
fastosus, Epimachus, 22, 31, 47, 63, 163,
309, 310, 314, 321, 328
feminina, Lophorina (Astrapia), 337
ferrugineus, Pitohui, 45, 51, 54, 63, 274,
286, 292, 293
INDEX
festetichi, Opopsitta, 152
Finch, Crimson-sided Mountain Grass,
412
finschi, Paradisaea, 341
flaviceps, Campochaera, 195
flavicollis, Dupetor, 100
flavigaster, Microeca, 257
flavirictus, Meliphaga, 25, 362, 363, 364,
365, 366, 373
flaviventer, Machaerirhynchus, 27, 62,
254, 255, 256
Meliphaga, 44, 51, 65,
374, 376
flavogrisea, Pachycare, 62, 212, 241, 272
278
flavovirescens, Microeca, 51, 54, 55, 62,
87, 255, 257, 258
flavum, Oedistoma, 356
Flowerpecker, Red-capped, 398
Flycatcher, Black and White Thicket,
263
Black-breasted Flatbill, 256
Black-throated Thicket, 265
Ground Thicket, 270
Lowland Peltops, 236
Mountain Peltops, 238
Olive Microeca, 258
Red-backed, 256
River, 261
Satin, 254
Shining, 254
Slaty Thicket, 267
White-faced, 260
White-rumped Thicket, 268
White-throated Thicket, 270
White-winged Thicket, 266
Yellow-breasted Flatbill, 254
Yellow-footed Microeca, 259
Yellow Microeca, 259
Yellow Thicket, 263
foersteri, Melidectes, 387, 388
forbesi, Rallicula, 22, 23, 37, 52, 117
Fowl, Scrub, 112
frater, Monarcha, 36, 37, 38, 62, 250,
255, 2/9
freycinet, Megapodius, 37) LZ
Friarbird, Meyer’s, 380
New Guinea, 380
Frogmouth, Great Papuan, 175
Little Papuan, 175
frontalis, Hirundo, 90, 195
fulgidus, Psittrichas, 155
fulginosa, Rhipidura, 239, 244
fuliginosus, Oreostruthus, 18, 48, 71,
412
fulva, Pluvialis, 120
69) 389; 21
fulvigula, Timeliopsis, 64, 350
fumigatus, Melipotes, 31, 38, 65, 397
fuscata, Pseudeos, 80, 141, 144, 145, 148
Sterna, 80, 122
fuscicapilla, Zosterops, 47
fuscirostris, Talegalla, 39, 58, 112, 114
fuscus, Melidectes, 45, 48, 387, 388, 389
gajduseki, Lonchura, 410
galatea, Tanysiptera, 184, 186, 190
galerita, Cactua, 59, 154, 155, 156
Gallicolumba beccarii, 37, 59, 138, 407
jobiensis, 37, 42, 45, 137, 139
rufigula, 37, 42, 45, 51, 137
Gallinago hardwickii, 79, 121
megala, 79, 121
Gallinula tenebrosa, 72, 91, 119
gaudichaud, Dacelo, 54, 74, 78, 85, 185,
186
geelvinkianum, Dicaeum, 37, 52, 65, 86,
350, 398
geislerorum, Ailuroedus, 347
geoffroyi, Geoffroyus, 54, 59, 156, 157,
303
Geoffroyus geoffroyi, 54, 59, 156, 157,
303
simplex, 59, 80, 156
Gerygone chloronota, 25, 36, 227
chrysogaster, 36, 61, 225, 226, 227
cinerea, 36, 61, 225, 227, 257
magnirostris, 36, 74, 225, 226, 228
olivacea, 225, 226
palpebrosa, 36, 61, 87, 225, 227, 228
ruficollis, 36, 37, 54, 61, 73, 76, 225,
226, 228, 257, 396
gibberifrons, Anas, 72, 100
gigantea, Melampitta, 47
giluwensis, Turnix, 115
girrenera, Haliastur, 104
giulanettii, Phylloscopus, 235
Glareola isabella, 79, 121
godmani, Psittaculirostris, 150
goldiei, Psitteuteles, 80, 144, 146
goliath, Probosciger, 153
goliathi, Melipotes, 397
goliathina, Charmosyna, 145
Goose, White-quilled Pygmy, 101
Goshawk, Australian, 107
Buerger’s, 104
Black-mantled, 108
Gray, 105
New Guinea Gray-headed, 107
White, 105
gouldi, Circus, 110
Dupetor, 100
426
INDEX
Goura scheepmakeri, 39, 140
superspecies, 50
victoria, 22, 39, 51, 140
gracilis, Anas, 100
Meliphaga, 365
Grallina bruijni, 72, 261, 299
grandis, Lonchura, 73, 409
Oreopsittacus, 148
Grassbird, Tawny, 233
Graybird, Australian, 203
Black, 198
Black-bellied, 200
Black-hooded, 204
Gray's, 198
Miiller’s, 199
Papuan, 204
Rufous-underwing, 201
Stout-billed, 202
Grebe, Little, 98
grisea, Coracina, 204
griseiceps, Pachycephala, 29, 30, 63, 274,
275, 280
griseoceps, Microeca, 54, 55, 257, 258,
259
griseiventris, Poecilodryas, 265
egriseotincta, Reinwardtoena, 136
guilielmi, Paradisaea, 306, 316, 321, 342
mise Piiloprora, 125 13; 22, 27, 31, 34,
47, 49, 65, 86, 279, 372 384, 386, 387,
405
gularis, Rhipidura, 247
gulielmitertii, Opopsitta, 54, 59, 151
gutturalis, Anthus, 48, 71, 205
guttula, Monarcha, 38, 62, 89, 251
guttaticollis, Ailuroedus, 347
guttatus, Sericornis, 217
Gymnocorvus tristis, 304
Gymnophaps albertisii, 56, 58, 132
haematodus, Trichoglossus, 52, 59, 74,
78, 81, 86, 89, 142, 144, 145, 146,
147, 148, 151, 352, 396
hagenensis, Oreostruthus, 412
Peneothello, 266
Halcyon macleayii, 36, 184, 188, 189, 190
megarhyncha, 27, 36, 45, 54, 55, 60, 91,
168, 184, 186, 187, 188, 190, 387
sancta, 36, 79, 91, 188, 189, 190, 191
torotoro, 27, 36, 54, 55, 60, 168, 184,
186, 187, 189, 190, 191, 387
Haliastur indus, 72, 78, 80, 104
hallstromi, Psittacella, 163
halmaheira, Columba, 133
hardwickii, Capella, 121
Gallinago, 79, 121
harmonica, Colluricincla, 36, 74, 274, 286
427
Harpyopsis novaeguineae, 58, 109
Harrier, Spotted March, 109
Swamp, 110
harterti, Monarcha, 253
hattamensis, Pachycephalopsis, 22, 47
Hawk, Brown, 112
Crested, 102
Doria’s, 109
heliosylus, Zonerodius, 99
Hemiprocne mystacea, 72, 75, 78, 85, 182
Hen, Common Scrub, 112
Purple Swamp, 119
Henicopernis lognicauda, 72, 103, 104
Henicophaps albifrons, 59, 139
henkii, Monarcha, 253
Heron, Great White, 99
Nankeen Night, 99
Pied, 99
Heteromyias albispecularis, 24, 37, 262,
263, 270
Heteroscelus incanus, 120
Hieraaetus morphnoides, 17, 18
hilli, Nycticorax, 99
hirundinacea, Collocalia, 36, 43, 72, 75,
78, 85, 88, 180, 181
Hirundo nigricans, 79, 90, 195
tahitica, 72, 90, 195
Hobby, Oriental, 111
Honeyeater, Black, 359
Black-backed Streaked, 386
Black-throated, 376
Brown, 379
Brown-backed Streaked, 384
Gray-bellied, 356
Long-bearded, 390
Long-billed, 351
Marbled, 378
Meek’s Streaked, 384
Mountain Straight-billed, 350
Obscure, 377
Pygmy, 357
Sooty, 389
Spotted, 375
Tawny-breasted, 374
Hornbill, Papuan, 193
horsfieldi, Cuculus, 164
hottentottus, Dicrurus,
156, 302, 303
hunsteini, Lophorina (Diphyllodes), 335
hyperythra, Pachycephala, 29, 30, 35, 37,
38; Ol,03,;. 211; 274, 275, 278; 280,
281, 282, 283, 285
Rhipidura, 29, 36, 38, 62, 239, 240, 242,
244, 245, 247, 248
hypoinochoa, Domicella, 144, 145
54, 74, 78, 85,
INDEX
hypoleuca, Pachycephala, 282
Poecilodryas, 262, 263, 264
hypoleucos, Actitis, 120
Tringa, 79, 120
Ifrit, Blue-capped, 213
Ifrita kowaldi, 61, 96, 213, 256
iliolophum, Oedistoma, 44, 64, 86, 352,
356
iliolophus, Toxorhamphus, 352, 356
incanus, Heteroscelus, 120
incerta, Amalocichla, 37, 87, 208
Coracina, 199
incincta, Halcyon, 189
incondita, Pseudeos, 14]
inconspicua, Gerygone, 228
indus, Haliastur, 72, 78, 80, 104
insignis, Aegotheles, 43, 60, 178
Chalcopsitta, 141
Clytomyias, 61, 87, 91, 216
insperata, Gerygone, 228
intermedia, Egretta, 99
intermedius, Trichoglossus, 52, 142
interposita, Megaloprepia, 129
iozonus, Ptilinopus, 123, 126
isabella, Glareola, 79, 121
Stiltia, 121
isidori, Pomatostomus, 210
ixoides, Pycnopygius, 37, 54, 65, 379
jamesii, Manucodia (Phonygammus), 326
javanica, Mirafra, 73, 194
jobiensis, Ducula, 130
Gallicolumba, 37, 42, 45, 137, 139
Nalegalla, 39; 58, 12, i142
josefinae, Charmosyna, 47
jugularis, Nectarinia, 355
kaporensis, Ptilinopus, 127
karimuiensis, Myzomela, 358
keiensis, Micropsitta, 26
keraudrenii, Manucodia (Phonygammus),
52, 54, 63, 67, 308, 321, 326
Phonygammus, 307, 321, 326
kerstingi, Macropygia, 133
Kestrel, Nankeen, 111
Kingfisher, Azure, 183
Common Paradise, 190
Dwarf, 183
Forest, 189
Hook-billed, 184
Lowland Yellow-billed, 187
Mountain Yellow-billed, 188
Sacred, 190
Shovel-billed, 184
428
kirhocephalus, Pitohui, 22,
274, 286, 290, 292, 293
Kite, Black, 103
Black-winged, 102
Brahminy, 104
klossi, Pachycephala, 276
kodonophonos, Pitohui, 293
Koel, Black-capped, 170
Indian; 171
White-crowned, 171
Kookaburra, Gaudichaud’s, 185
kowaldi, Ifrita, 61, 96, 213, 256
kuboriensis, Daphoenositta, 348
kutubu, Malurus, 214
45, 54, 63,
Lalage atrovirens, 39, 196, 207
leucomela, 39, 60, 196
superspecies, 196
lamonti, Synoicus, 114
Lanius schach 74, 76, 80, 85, 295
Larius roratus, 59, 155
Lark, Singing Bush, 194
Torrent, 299
latipennis, Lophorina, 330
lauterbachi, Chlamydera, 73, 346
lawesi, Lophorina (Parotia) 22, 37, 39,
44, 47, 48, 63, 321, 332
Parotia, 311, 312, 313, 321, 332
leachii, Dacelo, 187
lepidus, Ceyx, 60, 183
lessonii, Ceyx, 183
leucolophus, Caliechthrus, 60, 171
leucomela, Lalage, 39, 60, 196
leucophaea, Climacteris, 18, 19, 20, 270
leucophrys, Poliolimnas, 117
Rhipidura, 36, 74, 78, 107, 239, 242, 248
leucops, Irepellasia; 3, 154, 162) 875 2575
258, 260
leucorhynchus, Artamus, 296
leucosomus, Accipiter, 105
leucostephes, Melidectes, 387, 389
leucostictus, Eupetes, 27, 36, 37, 61, 211,
212
leucostigma, Pachycephala, 31, 42, 54, 63,
85, 274, 275
Rhagologus, 275
leucothorax, Rhipidura, 36, 51, 54, 74,
78, 85, 239, 240, 241, 242
lineata, Coracina, 7, Us 20, Sle 2on4 7s
197
Loboparadisea sericea, 24, 31, 55, 64, 318,
321, 323
Lonchura, castaneothorax, 409, 410
caniceps, 40, 409, 410
grandis, 73, 409
montana, 18, 23, 410
INDEX
monticola, 18, 20, 23
Spectabilis; 40> 56, 73, 76, 77, 78, 8D,
409, 410
teerinki, 47, 77, 409, 410
tristissima, 40, 409, 410, 411
vana, 77, 409, 410
Longbill, Slaty-chinned, 353
longicauda, Coracian, 197, 203, 204
Henicopernis, 72, 103, 104
Melampitta, 213
Melanocharis 24, 27, 33, 34, 45, 53,
400, 401, 403, 404
Lophorina superba, 31, 44, 53, 54, 64, 91,
305, 306, 311, 312, 321, 330, 336, 392
Lophorina (Astrapia) mayeri, 22, 34, 39,
47, 48, 314, 321, 332, 338, 390
nigra, 314, 321
rothschildi, 314, 321
splendissima, 22, 47, 314, 321, 339
stephaniae, 22, 34, 39, 47, 48, 49, 63,
314, 321, 332, 337, 338
Lophorina (Cicinnurus) regia, 31, 44, 51,
54, 64, 86, 321, 337
Lophorina (Diphyllodes) magnifica, 31,
44, 52, 56, 64, 85, 86, 89, 320, 321,
335, 337
respublica, 321
Lophorina (Parotia) carolae, 22, 39, 48,
321, 332, 333
lawesi, 22, 37, 39, 44, 47, 48, 63, 321,
332
sefilata, 321, 312, 313, 332
wahnesi, 306, 311, 312, 313, 314, 321,
332
Lophorina (Pteridophora) alberti, 12, 20,
31, 46, 64, 321, 339
Lophorina (Ptiloris) magnifica, 37, 64,
86, 320, 321, 333
Lophorina (Seleucidis) melanoleuca, 321
lophotis, Meliphaga, 375
lorentzi, Pachycephala, 47, 282
Loria loriae, 305, 318, 321, 322
loriae, Cnemophilus (Loria), 22, 31, 49,
64, 320, 321, 322
Eupetes, 212
Loria, 305, 318, 321, 322
Loriculus aurantiifrons, 164
Lorikeet, Bat, 164
lory, Domicella, 51, 59, 144, 145, 154,
204, 398
Lory, Dusky-orange, 141
Eastern Black-capped, 145
Little Red, 146
Orange-billed Mountain, 149
Papuan, 145
Plum-faced Mountain, 148
429
Pygmy Streaked, 147
Rainbow, 142
Red-capped Streaked, 144
Western Black-capped, 144
Yellow-billed Mountain, 149
Yellow-fronted Blue-eared, 147
Yellow-streaked, 140
lucidus, Chalcites, 169
Chrysococcyx, 169
lugubris, Melampitta, 24, 37, 213
Lycocorax pyrrhopterus, 316, 317, 321
Macgregoria pulchra, 18, 20, 24, 315, 317,
321
macgregoriae, Amblyornis, 22, 53, 64, gl,
342
macgregorii, Cnemophilus, 22, 47, 49, 64,
67, 318, 320, 322
Machaerirhynchus flaviventer, 27, 62, 254,
255, 256
nigripectus, 27; 62) 2147255, 256
macklottii, Pitta, 193
macleayii, Halcyon, 36, 184, 188, 189, 190
Macropygia amboinensis, 35, 43, 59, 74,
78, 133, 135, 136, 137
Nigrirostris, 35, 45, 59, 86, 134, 135,
139, 172
macrorhina, Melidora, 45, 60, 184, 186
macrurus, Caprimulgus, 73, 178
maculosa, Turnix, 73 76, 115
madaraszi, Psittacella, 22, 43, 49,
159, 160, 161, 162, 163
mafulu, Malurus, 73, 214
magnifica Craspedophora, 333
Lophorina (Diphyllodes), 31, 44, 52,
56, 64, 85, 86, 89, 320, 321, 335, 337
Lophorina (Ptiloris) 37, 64, 86, 320,
321, 333
Megaloprepia, 54, 58, 86, 126, 129,
132
magnificus, Diphyllodes, 306, 310, 311,
313, 316, 320, 321, 335
Ptilons; S11; 312; 313, 315, 320, 321,
333
magnirostris, Gerygone, 36, 74, 225, 226,
228
major, Neopsittacus, 149
Psittacella, 161
malaynaus, Chrysococcyx, 169, 170
Malurus alboscapulatus, 73, 76, 78, 84,
85, 165, 166, 214, 216, 217, 234, 260
Mannikin, Blue-faced Parrot, 407
Great-billed, 409
New Britain, 410
Papuan Parrot, 407
Manucode, Crinkle-collared, 324
158,
INDEX
Manucodia ater, 307, 308, 321, 325
chalybatus, 62, 85, 89, 307, 308, 321,
324, 327
comri, 307, 308, 321
jobiensis, 307, 321
Manucodia (Phonygammus) keraudrenii,
52, 54, 63, 67, 308, 321, 326
margaritae, Paradisaea, 53, 341
marmoratus, Pycnopygius, 378
Martin, Tree, 195
maximus, Artamus, 72, 75, 78, 85, 89, 295
mayeri, Astrapia, 337
mayeri, Lophorina (Astrapia), 22, 34, 39,
47, 48, 314, 321, 332, 338, 390
mayri, Ptiloprora, 385, 386
meeki, Halcyon, 187
Loriculus, 164
bytes 73
meekiana, Ptiloprora, 17, 25, 384
megala, Capella, 121
Gallinago, 79, 121
Megaloprepia magnifica, 54, 58, 86, 126,
129, 132
Megalurus timoriensis, 48, 72, 73, 76, 78,
85, 233, 235
Megapodius freycinet, 37, 112
megarhyncha, Colluricincla, 36, 53, 63,
87, 89, 274, 284
Halcyon, 27, 36, 45, 54, 55, 60, 71,
168, 184, 186, 187, 188, 190, 387
megarhynchus, Melilestes, 64, 89, 197,
351
Myiolestes, 284
Rhamphomantis, 24
Megatriorchis doriae, 24, 85, 109
melaena, Coracina, 36, 45, 51, 60, 197
198
melaleuca, Rhipidura, 248
Melampitta gigantea, 47
Lesser, 213
lugubris, 24, 37, 213
melanocephalus, Ailuroedus, 347
Melanocharis arfakiana, 18, 19, 20, 21, 25,
47
loneicauda, 25,27, 33; 34, 40; 53x 400;
401, 403, 404
mera, 26, 29) do, G5, 398; 401, 402, 403,
404
striativentris, 45, 52, 65, 403
versteri, 27, 29, 33, 45, 65, 108, 257,
399, 400, 401, 402, 403, 404
melanochlamys, Accipiter, 35, 85, 108
melanoleuca, Lophorina (Seleucidis), 321
Seleucidis, 314, 321
melanoleucos, Phalacrocorax, 72, 98
melanops, Coracina, 203
3
430
Melidectes belfordi, 35, 39, 46, 48, 49,
377, 387, 388, 389, 391, 392, 393
Belford’s 391
foersteri, 387, 388
fuscus, 45, 48, 387, 388, 389
leucostephes, 387, 389
nouhuysi, 20, 22, 26, 46, 47, 387, 388,
391
ochromelas, 20, 35, 39, 47, 387, 388,
389, 392, 395
princeps, 20, 22, 26, 45, 46, 47, 48, 387,
388, 390
Reichenow’s, 392
rufocrissalis, 35, 36, 38, 39, 46, 48, 49,
65, 86, 377, 387, 388, 389, 392, 393,
397
torquatus, 36, 54, 74, 76, 229, 387, 388,
392, 393, 396
Melidora macrorhina, 45, 60, 184, 186
Melilestes megarhynchus, 64, 89, 197,
351
Meliphaga albonotata, 369
analoga, 37, 38, 42, 44, 65, 361, 362,
363, 364, 365, 366, 368, 371, 372,
373, 374, 378
aruensis, 38, 42, 44, 65, 362, 363, 364,
365, 366
auga, 36, 44,52, 745,775; 78; So, 89) 3b,
362, 363, 364, 365, 366, 367
flaviventer, 44, 51, 65, 69, 89, 251, 374,
376
flavirictus, 25, 362, 363, 364, 365, 366,
373
gracilis, 365
Large Spot-breasted, 370
Mimic, 373
mimikae, 36, 38, 44, 65, 362, 363, 364,
365, 366, 368, 369, 370, 372, 373
montana, 362, 367, 368, 369
obscura, 37, 65, 377
orientalis, 38, 44, 46, 52, 53, 54, 65, 86,
362, 363, 364, 365, 366, 369, 371,
373
polygramma, 25 65, 375
Puff-backed, 366
Small Spot-breasted, 371
Southern White-eared, 367
subfrenata, 37, 65, 376
versicolor, 377
Yellow-gaped, 373
Melipotes, Common, 397
fumigatus, 31, 38, 65, 397
menbeki, Centropus, 36, 54, 60,
172
Merops ornatus, 79, 191
metallica, Aplonis, 80, 296
lags
INDEX
meyeri, Epimachus, 22, 31, 49, 163, 309,
310, 314, 321, 328
Myzomela, 51, 359
Philemon, 44, 65, 389
Timeliopsis, 350
meyerli, Chalcites, 169
Chrysococcyx, 25, 60, 169
Microdynamis parva, 170
Microeca flavigaster, 257
flavovirescens, 51, 54, 55, 62, 87, 255,
257, 258
griseoceps, 54, 55, 257, 258, 259
papuana, 62, 257, 258, 259
Micropsitta bruijnii, 153
keiensis, 26
pusio, 26, 153
migrans, Milvus, 72, 79, 103, 104
Milvus migrans, 72, 79, 103, 104
mimikae, Meliphaga, 36, 38, 44, 65, 362,
363, 364, 365, 366, 368, 369, 370,
372, 373
Mino dumontii, 54, 74, 78, 193, 297
anais, 22
minor, Paradisaea, 39, 50, 51, 74, 171,
316, 321, 340, 341, 342
Zosterops, 405
minutus, Numenius, 79, 120
Mirafra javanica, 73, 194
miranda, Daphoenositta, 24, 49, 348, 349
modesta, Egretta, 99
Pachycephala, 29, 30, 38, 43, 49, 53,
274, 275, 282
Psittacella, 22, 158, 159, 160, 161, 162,
163
monacha, Pachycephala, 283
Monachella muelleriana, 72, 261, 299
Monarch, Black, 249
Black and Yellow, 252
Black-winged, 250
Frilled, 253
Spot-winged, 251
Monarcha alecto, 254
axillaris, 38, 54, 62, 249, 251
chrysomela, 37, 38, 54, 62, 252
frater, 36, 37, 38, 62, 250, 255, 279
guttula, 38, 62, 89, 251
telescophthalmus, 62, 253
monorthonyx, Anurophasis, 47
montana, Coracina, 29, 33, 43, 60, 196,
197, 199, 200, 202
Lonchura, 18, 23, 410
Meliphaga, 362, 367, 368, 369
montanus, Megalurus, 233
Peltops, 27, 37, 61, 74, 75, 85, 102, 194,
237, 238, 401, 407
monticola, Lonchura, 18, 20, 23
431
montium, Paramythia, 65, 67, 405
Moorhen, Dusky, 119
Rufous-tailed, 118
morio, Coracina, 43, 45, 197, 199, 402
morphnoides, Hieraaetus, 17, 18
Motacilla cinerea, 79, 205
muelleri, Rhipidura, 245
muelleriana, Monachella, 72, 261, 299
multistriata, Charmosyna, 25
murina, Acanthiza, 96, 221, 222, 224, 225,
227
Crateroscelis, 10, 27, 28, 29, 30, 35, 36,
37, 51, 61, 69, 87, 206, 229, 230,
232, 233, 243, 248
musschenbroekii, Neopsittacus, 59, 144,
146, 148, 149, 150
Myiagra alecto, 74, 254, 268
cyanoleuca, 79, 254
Myiolestes megarhynchus, 284
Myna, Yellow-faced, 297
mystacalis, Eurostopodus, 179
mystacea, Hemiprocne, 72, 75,
182
Myzomela adolphinae, 21, 36, 47, 74, 76,
80, 358, 359
cruentata, 37, 64, 86, 358, 359, 360
eques, 3/7, 44, dil, 54,64, 3585359
Mountain Red-headed, 359
nigrita, 37, 44, 51, 64, 358, 359
Red, 359
Red-collared, 360
Red-spotted, 358
rosenbergii, 31, 36, 38, 64, 86, 88, 358,
359, 360, 377
sclateri, 359
78; 38D,
nanus, Ptilinopus, 25, 43, 54, 58, 123, 126,
129
nea, Colluricincla, 284
Nectarinia jugularis, 355
sericea, 54, 64, 74, 75, 78, 85, 350, 355,
398
Neopsittacus musschenbroekii, 59, 144
146, 148, 149, 150
pullicauda, 146, 149
Neositta chrysoptera, 348, 349
Nettapus coromandelianus, 72, 101
neumanni, Manucodia (Phonygammus)
326
nicobarica, Caloenas, 122
Nightjar, Archbold’s, 179
White-tailed, 178
White-throated, 179
nigra, Lophorina (Astrapia), 314, 321
Melanocharis, 26, 29, 33, 65, 398, 401,
402, 403, 404
>
INDEX
nigrescens, Pitohui, 46, 63, 67, 274, 289,
290, 291
nigricans, Hirundo, 79, 90, 195
nigripectus, Machaerirhynchus,
214, 255, 256
nigrirostris, Macropygia, 35, 43, 59, 86,
134, 135, 139, 172
nigrita, Myzomela, 37, 44, 51, 64, 358,
359
nigropectus, Eulacestoma, 63, 294
nigrorufa, Crateroscelis, 25, 29, 35, 37, 91,
229, 231
nigroviridis, Sericornis, 25, 217
Ninox novaeseelandiae, 175
theomacha, 60, 174
niveifrons, Pachycephala, 283
nobilis, Otidiphaps, 56, 139
Notophoyx picata, 99
nouhuysi, Melidectes, 20, 22, 26, 46, 47,
387, 388, 391
Sericornis, 38, 42, 43, 48, 61, 87, 166,
217, 219, 222, 224, 233
novaeguineae, Chaetura, 180
Coturnix, 73
Dacelo, 187
Falco, 90
Harpyopsis, 58, 109
Philemon, 44, 65, 74, 80, 298, 327, 375,
380
Pitta, 194
Toxorhamphus, 22, 352, 353
Zosterpos, 47, 49), 309>, 406
novaehollandiae, Accipiter, 42, 43, 45, 56,
85, 105, 107, 108, 412
Coracina, 79, 197, 203
Podiceps, 72, 98
novaeseelandiae, Anthus, 73, 76, 205, 249
Ninox, 175
nuditarsus, Collocalia, 182
Numenius minutus, 79, 120
Nuthatch, Pink-faced, 348
Nycticorax caledonicus, 99
nympha, Tanysiptera, 25
Dis 102,
obscura, Meliphaga, 37, 65, 377
Pachycephala, 275
obscurior, Pachycephala, 277
obscurus, Oreornis, 377
ocellatus, Podargus, 43, 60, 175
ochrogaster, Ceyx, 183
ochromelas, Melidectes, 20, 35, 39, 47,
387, 389, 392, 395
Oedistoma iliolophum, 44, 64, 86, 352,
356
pygmaeum, 37, 44, 64, 80, 86, 352, 355,
357, 358
432
olivacea, Gerygone, 225, 226
olivaceus, Amaurornis, 51, 54, 74, 78, 84,
118
olivascentior, Amalocichla, 208
Opopsitta diophthalma, 52, 152
gulielmitertii, 54, 59, 151
oorti, Glytomyias, 216
Sericornis, 222
Oreocharis arfaki, 12, 37, 65, 404, 405
Oreocincla dauma, 207
oreophilus, Cacomantis, 164
Oreopsittacus arfaki, 38, 59, 86, 146, 148,
149, 150
Oreornis subrenata, 376
obscurus, 377
chrysogenys, 47
Oreostruthus fuliginosus, 18, 48, 71, 412
orientalis, Eurystomus, 45, 74, 78, 79, 85,
90, 192, 297
Meliphaga, 38, 44, 46, 52, 53, 54, 65,
86, 362, 363, 364, 365, 366, 369,
371, 373
Oriole, New Guinea, 297
Oriolus szalayi, 37, 63, 74, 78, 85, 297, 382
ornatus, Merops, 79, 191
Ptilinopus, 29, 30, 42, 43, 123, 126, 127,
129
orru, Corvus, 249
Orthonyx temminckii, 18, 19, 20
Otidiphaps nobilis, 56, 139
Owl, Barn, 173
Grass, 174
Papuan Boobook, 174
Sooty, 173
Owlet-nightjar, Collared, 176
Large, 178
Mountain, 177
Wallace’s, 176
Pachycare flavogrisea, 62, 212, 241, 272,
278
Pachycephala aurea, 25
griseiceps, 29, 30, 63, 274, 275, 280
hyperythra, 29, 30, 35, 37, 38, 51, 63,
ON, QA, Bib, 278, 280, 28ly 282,
283, 285
leucostigma, 31, 42, 54, 63, 85, 274, 275
lorentzi, 47, 282
modesta, 29, 30, 38, 43, 49, 53, 274,
275, 282
monacha, 283
rufinucha, 43, 44, 63, 274, 275, 277, 283
rufiventris, 36, 74, 78, 85, 211, 274, 278,
283, 286
schilegelit, 29) 30) 015 95, 68; 44, 68,274;
DIOn wily ZOuy 2olbe zoe, 200
INDEX
soror, 25, 29, 34, 38, 44, 52, 54, 63, 211,
274, 276, 278, 280, 281
tenebrosa, 20, 25, 46, 274, 284
Pachycephalopsis hattamensis, 22, 47
poliosoma, 22, 36, 37, 52, 54, 62, 69, 89,
262, 263, 266, 270
pacificus, Eurystomus, 79, 90, 192
pallida, Psittacella, 158
palpebrosa, Gerygone, 36, 61, 87, 225,
227, 228
palustris, Porzana, 116
papou, Charmosyna, 29, 86, 145, 146, 147,
149
papua, Anhinga, 99
papuana, Erythrura, 17, 21, 39, 41, 47,
407
Microeca, 62, 257, 258, 259
papuanus, Falco, 111
papuensis, Archboldia, 20, 24, 46, 48, 49,
342
Chaetorhynchus, 63, 303
Collocalia, 182
Coracina, 36, 74, 185, 197, 204, 205
Eurostopodus, 179
Podargus, 43, 175
Sericornis, 30, 32, 38, 43, 61, 88, 217,
218, 220, 223, 224
Tyto, 174
Zoothera, 207
Paradigalla brevicauda, 306, 316, 321, 327
carunculata, 316, 321
Short-tailed, 327
superspecies, 46
Paradisaea apoda, 316, 321, 340
decora, 316, 321
guilielmi, 306, 316, 321, 342
minor, 39, 50, 51, 74, 171, 316, 321, 340,
341, 342
raggiana, 22, 39, 50, 64, 74, 78, 301,
316, 321, 340, 341, 342
rubra, 316, 321
rudolphi, 22, 47, 48, 53, 306, 316, 321,
341
Paramythia montium, 65, 67, 405
Parotia carolae, 311, 312, 321, 332
lawesi, 311, 312, 313, 321, 332
sefilata, 312, 313, 321, 332
wahnesi, 306, 311, 312, 313, 314, 321,
332
Parrot, Blue-collared, 156
Brehm’s Tiger, 158
Edward’s Fig, 151
King William’s Fig, 151
Large Fig, 150
Madarasz’s Tiger, 163
Modest Tiger, 163
433
Mountain Pygmy, 153
Painted Tiger, 160
Papuan King, 157
Red-cheeked, 156
Red-sided Eclectus, 155
Two-eyed Fig, 152
Vulturine, 155
parva, Eudynamis, 170
Microdynamis, 170
pectoralis, Larius, 155
Rallus, 36, 73, 116
pelewensis, Anas, 100
Peltops blainvillii, 27, 37, 61, 74, 85, 236,
239
montanus, 27, 37, 61, 74, 75, 85, 102,
194, 237, 238, 401, 407
Peneothello bimaculatus, 37, 52, 62, 262,
263, 268
cryptoleucus, 22, 47
cyanus, 22; 36, 37, 62, 69,87, 262; 263,
266, 267, 271
sigillatus, 12, 37, 48, 62, 262, 263, 266,
267, 320
peregrinus, Falco, 110
periophthalmicus, Monarcha, 250
perlatus, Ptilinopus, 29, 30, 42, 43, 54, 58,
12a 26 128
perneglecta, Pachycephala, 280
perspicillatus, Sericornis, 30, 33, 38, 43,
695 217, 219; 221,222, 223
perstriata, Ptiloprora, 22, 27, 34, 385, 386
Petroica archboldi, 47
bivittata, 48, 262
Phalacrocorax melanoleucos, 72, 98
sulcirostris, 98
phasianinus, Centropus, 36, 73, 172, 173
Philemon meyeri, 44, 65, 380
novaeguineae, 44, 65, 74, 80, 298, 327,
375, 380
philippensis, Rallus, 36, 72, 116
Phonygammus keraudrenii, 307, 321, 326
Phylloscopus trivirgatus, 37, 54, 61, 214,
299, 235
picata, Notophoyx, 99
picta, Psittacella, 49, 158, 159, 160
Pigeon, D’Albertis’, 132
Magnificent Ground, 139
Pinon Imperial, 130
Purple-tailed Imperial, 131
Red-breasted Imperial, 131]
Scheepmaker’s Crowned, 140
Victoria Crowned, 140
White-capped Ground, 139
White-throated, 133
Wompoo, 129
Zoe Imperial, 132
INDEX
pinon, Ducula, 130, 133
piperata, Rhamphocharis, 404
Pipit, Alpine, 205
Richard’s, 205
Pitohui, Black, 291
Black-headed, 289
Crested, 293
cristatus, 37, 274, 286, 293
dichrous, 45, 46, 63, 76, 78, 251, 274,
286, 289, 292
ferrugineus, 45, 51, 54, 63, 274, 286,
292, 293
kirhocephalus, 22, 45, 54, 63, 274, 286,
290, 292, 293
nigrescens, 46, 63, 67, 274, 289, 290, 291
Rusty, 291
Variable, 286
Pitta, Black-headed, 194
Blue-breasted, 193
erythrogaster, 37, 60, 86, 193, 194
sordida, 37, 194
placens, Climacteris, 18
Poecilodryas, 24, 25, 37, 54, 62, 87, 262,
263, 266, 268
placentis, Charmosyna, 29, 54, 55, 59, 86,
146, 147, 152
plagosus, Chrysococcyx, 79, 169, 170
Platalea regia, 79, 100
plicatus, Aceros, 60, 85, 193
Plover, Golden, 120
plumbea, Ptiloprora, 18, 25
plumifera, Egretta, 99
Pluvialis dominica, 79, 120
Podargus ocellatus, 43, 60, 175
papuensis, 43, 175
Podiceps novachollandiae, 72, 98
Poecilodryas albonotata, 62, 262, 263, 265
hypoleuca, 262, 263, 264
placens,.24,-25, 57,104, 62, °8/, 262, 263;
266, 268
poliocephalus, Accipiter, 3p, 42, 45, 68,
85, 105, 107, 108
Turdus, 20, 48, 71, 209
Poliolimnas cinereus, 117
poliopsa, Coracina, 198
poliopterus, Toxorhamphus, 22, 36, 64,
89, 251, 352, 353, 356
poliosoma, Pachycephalopsis, 22, 36, 37,
52, 54, 62, 69, 89, 262, 263, 266, 270
polycryptus, Accipiter, 107
polygramma, Meliphaga, 25, 65, 375
Xanthotis, 375
polygrammica, Lalage, 196
polyphonus, Melidectes, 396
Pomareopsis bruijni, 299
434
Pomatostomus isidori, 210
temporalis, 210
Porphyrio porphyrio, 72, 91, 119
porphyrio, Porphyrio, 72, 91, 119
Porzana pusilla, 72, 116
tabuensis, 20, 46, 72, 85, 117
Pratincole, Australian, 121
princeps, Melidectes, 20, 22, 26, 45, 46,
47, 48, 387, 388, 390
prionurus, Cacomantis, 166
Probosciger aterrimus, 153
propinquus, Centropus, 173
Pseudeos fuscata, 80, 141, 144, 145, 148
Psittacella brehmii, 43, 59, 158, 159, 160,
161
madaraszi, 22, 43, 49,
161, 162, 163
modesta, 22, 158, 159, 160, 161, 162,
163
picta, 49, 158, 159, 160
Psittaculirostris desmarestil, 39, 54, 59,
81, 85, 89, 141, 150, 151
edwardsii, 39, 51, 151, 153
superspecies, 50
Psitteuteles goldiei, 80, 144, 146
Psittrichas fulgidus, 155
Pteridophora alberti, 314, 320, 321, 339
Ptilinopus aurantiifrons, 123, 126
coronulatus, 39, 40, 123, 125, 128
iozonus, 123, 126
nanus, 25, 43, 54, 58, 123, 126, 129
ornatus, 29, 30, 42, 43, 123 126, 127,
L538, 1595, 1605
129
perlatus, 29; 30; 42, 43,545 58, 125, 126;
128
pulchellus, 39; 40, 42, 43, 58, 88, 123,
124, 125, 128
rivoli, 30; 58, 122, 1235, 125, 128, 129
superbus, 42, 43, 45, 86, 88, 96, 123,
124, 126, 128
Vantais. 1225123
Ptiloprora erythropleura, 22, 47, 385, 386
piiset, M2 Woy 222, Oly Ot. eee Oe
86, 279, 372 384, 386, 387, 405
mayri, 385, 386
meekiana, 17, 25, 384
perstriata, 22, 27, 34, 385, 386
plumbea, 18, 25
Ptiloris magnificus, 311, 312, 313, 315,
320) 8212333
pulchella, Charmosyna, 29, 54, 55, 59, 86,
146, 324
pulchellus, Ptilinopus, 39, 40, 42, 43, 58,
88, 123, 124, 125, 128
pulcher, Eupetes, 210
INDEX
pulchra, Macgregoria, 18, 20,
O17, 021
pullicauda, Neopsittacus, 146, 149
purpureoviolaceus, Manucodia (Phony-
gammus), 326
pusilla, Porzana, 72, 116
pusio, Micropsitta, 26, 153
Pycnopygius cinereus, 65, 378, 379
ixoides, 37, 54, 65, 379
stictocephalus, 25
pygmaeum, Oedistoma, 37, 44, 64, 80, 86,
352, 355, 357, 358
pyrrhophanus, Cacomantis, 30, 73, 76, 79,
90, 165, 166
pyrrhopterus, Lycocorax, 316, 317, 321
24, Sd,
Quail, Black-backed Bustard, 115
Brown, 114
Chinese, 115
King, 115
Red-backed, 115
querquedula, Anas, 79, 100
quoyi, Cracticus, 36, 63, 300, 301
raggiana, Paradisaea, 22, 39, 50, 64, 74,
78, 301, 316, 321, 340, 341, 342
Rail, Banded Land, 116
Forbes’ Chestnut, 117
Lewin, 116
Red-necked, 118
Slate-breasted, 116
Rallicula forbesi, 22, 23, 37, 52, 117
rubra, 22, 47
Rallina tricolor, 54, 118
Rallus pectoralis, 36, 73, 116
philippensis, 36, 72, 116
regia, Lophorina (Cicinnurus), 31, 44, 51,
54, 64, 86, 89, 321, 337
Platalea, 79, 100
regius, Cicinnurus, 306, 310, 311, 321, 337
Reinwardtoen reinwardtsi, 59, 135, 136
reinwardtsi, Reinwardtoena, 59, 135, 136
respublica, Diphyllodes, 321
Lophorina (Diphyllodes), 321
rex, Clytoceyx, 60, 184, 186
Lophorina (Cicinnurus), 337
Rhagologus leucostigma, 275
Rhamphocharis crassirostris, 404
Rhamphomantis megarhynchus, 24
Rhipidura albolimbata, 29, 38, 62, 69,
239, 240, 242, 244, 246, 248
atra, 38, 54, 55, 62, 239, 240, 244, 245,
246, 250, 324
brachyrhyncha, 38, 62, 239, 240, 242, 243,
245
fuliginosa, 239, 244
435
hyperythra, 29, 36, 38, 62, 239, 240,
949, 244, 245, 247, 248
leucophrys, 36, 74, 78, 107, 239, 242,
248
leucothorax, 36, 51, 54, 74, 78, 85, 239,
240, 241, 242
rufidorsa, 61, 239, 240, 241
rufiventris, 36, 62, 74, 78, 239, 240, 242,
246, 247
threnothorax, 36,
239, 240, 242
rhododendri, Anthus, 205
Riflebird, Magnificent, 333
rivoli, Ptilinopus, 30, 58, 122, 123, 125,
128, 129
Robin, Forest, 262
Northern Scrub, 206
robusta, Crateroscelis, 10, 23, 27, 28, 29,
30; 35, 36, 37, Gl, 69; 229, 230; 231,
232
rogersi, Anas, 100
roratus, Larius, 59, 155
rosenbergii, Myzomela, 31, 36, 38, 64, 86,
88, 358, 359, 360, 377
Scolopax, 121
rothschildi, Lophorina (Astrapia), 314,
32]
rubiensis, Domicella, 144
Meliphaga, 374
rubra, Eugerygone, 61, 96, 256
Rallicula, 22, 47
rubrocoronatum, Dicaeum, 398
rudolphi, Paradisaea, 22, 47, 48, 53, 306,
BM}, SAM, Bx4tI
rufa, Anhinga, 99
ruficollis, Aceros, 193
Calidris, 79, 121
Chalcites, 169
Chrysococcyx, 169, 170
Erolia, 121
Getysone; 36," 37,54, Ol, 73, 16, 225,
226, 228, 257, 396
ruficrissum, Amaurornis, 118
rufidorsa, Rhipidura, 61, 239, 240, 241
rufigaster, Ducula, 27, 45, 58, 131, 132
rufigula, Gallicolumba, 37, 42, 45, 51, 137
rufinucha, Pachycephala, 43, 44, 63, 274,
275, 277, 283
rufiventer, Eudynamis, 171
rufiventris, Pachycephala, 36, 74, 78, 85,
211, 274, 278, 283, 286
Rhipidura, 36, 62, 74, 78, 239, 240, 242,
246, 247
rufocrissalis, Melidectes, 35, 36, 38, 39,
46, 48, 49, 65, 86, 377, 387, 388, 389,
392, 393, 397
37, 38, 54, bb, 6),
INDEX
salvadorii, Aegotheles, 177
Meliphaga, 376
Pachycephala, 281
Paradisaea, 340
Salvadorina waigiuensis, 72, 89, 100
sancta, Halcyon, 36, 79, 91, 188, 189, 190,
19]
Sandpiper, Common, 120
Sharp-tailed, 121
sanfordi, Archboldia, 342
sanguineus, Cnemophilus, 320
saturata, Scolopax, 37, 120, 12]
saturatior, Eugerygone, 256
saturatus, Cuculus, 79, 164
Machaerirhynchus, 256
Saxicola caprata, 74, 76, 78, 206, 407
schach, Lanius, 74, 76, 80, 85, 295
scheepmakeri, Goura, 39, 140
schistacinus, Accipiter, 108
schisticeps, Coracina, 29, 33, 36, 43, 45,
60, 196, 197, 198, 199, 200, 201, 202
schlegelii, Pachycephala, 29, 30, 31, 35,
Je, 44,569, 274, 276, 277, 280, 281,
282, 283
schraderensis, Melidectes, 391
scintillata, Chalcopitta, 59, 140
sclateri, Myzomela, 359
sclateriana, Amalocichla, 18, 20
scolopacea, Eudynamis, 37, 54, 80, 167,
170, 171
Scolopax saturata, 37, 120, 121
sefilata, Lophorina (Parotia), 312, 313,
321, 332
Seleucidis melanoleuca, 314, 321
Semioptera wallacei, 316, 317, 321
septentrionalis, Gallicolumba, 137
sericea, Loboparadisea, 24, 31, 55, 64,
38, 321, 323
Nectarinia, 54, 64, 74, 75, 78, 85, 350,
355, 398
Sericornis arfakianus, 30, 34, 38, 54, 55,
Oly 67, 217, 218). 223
beccarii, 217, 224
Beccari’s, 224
Buff-faced, 219
Gray-green, 218
Large Mountain, 222
nigroviridis, 25, 217
nouhuysi, 38, 42, 43, 48, 61, 87, 166,
INT, 219) 222, 224, 253
Pale-billed, 217
Papuan, 220
papuensis, 30, 32, 38, AP Gile ogy 2 kits
218, 220, 223, 224
Perplexing, 224
436
perspicillatus, 30, 33, 38, 43, 69, 217,
219: 221, 222,223
spilodera, 30) 54, 5,01, 2175 210
virgatus, 217, 224
Sericulus aureus, 346, 402
severus, Falco, 35, 111
shawmayeri, Casuarius, 97
Shrike, Schach, 295
Shrike-thrush, Gray, 286
Rufous, 284
Shrike-tit, Wattled, 294
Sicklebill, Black, 328
Black-billed, 329
Brown, 328
sigillatus, Peneothello, 12, 37, 48, 62, 262,
263, 266, 267, 320
sigillifera, Erythrura, 407
similis, Lophorina (Cicinnurus), 337
simplex, Geoffroyus, 59, 80, 156
Sittella, Papuan, 348
sloetii, Campochaera, 24, 54, 60, 195, 199
smaragdina, Ducula, 131
Snipe, Japanese, 121
Marsh, 121
solitarius, Ceyx, 183
somu, Domicella, 144
sordida, Pitta, 37, 194
soror, Pachycephala, 255529734, 025, 0ct4,
52, 54, 63, 211, 274, 276, 278, 280, 281
spectabilis, Chalcopsitta, 141
Tonchura, 40; 56, 735 76, 77, (ese,
409, 410
Spilodeta, Sericornis, 30) 54, 555 Ol 217;
219
spilonotus, Circus, 45, 72, 109, 110
spilothorax, Circus, 109
splendissima, Lophorina (Astrapia), 22,
47, 314, 321, 339
Spoonbill, Royal, 100
Starling, Metallic, 296
Singing, 296
stenozona, Aviceda, 102
stephani, Chalcophaps, 37, 54, 59, 137
stephaniae, Astrapia, 337
Lophorina (Astrapia), 22, 34, 39, 47,
48, 49, 63, 314, 321, 332, 337, 338
Sterna fuscata, 80, 122
stictocephalus, Pycnopygius, 25
Stiltia isabella, 121
Stint, Red-necked, 121
stonii, Ailuroedus, 347
strenua, Coracina, 202
stresemanni, Epimachus, 328
Lanius, 295
Sericornis, 222
INDEX
striativentris, Melanocharis, 45, 52, 65,
403
stictocephalus, Pycnopygius, 25
suavissima, Opopsitta, 151
subalaris, Amblyornis, 22, 47, 343
Coracina, 201
subaurantia, Pachycare, 272
subcristata, Aviceda, 54, 58, 85, 102, 239
subcyaneus, Peneothello, 267
subfrenata, Meliphaga, 37, 65, 376
Oreornis, 376
subpallida, Pachycare, 272
sulcirostris, Phalocrocorax, 98
Sunbird, Black, 350
superba, Lophorina, 312
superbus, Ptilinopus, 42, 43, 45, 86, 88,
96, 123, 124, 126, 128
superciliaris, Drymodes, 25, 37, 61, 87,
206
superciliosa, Anas, 72, 100
Swallow, Greater Wood, 295
Pacific, 195
Swift, Mustached, 182
New Guinea Spine-tailed, 180
Swiftlet, Glossy, 180
Mountain, 181
Whitehead’s, 182
Synoicus ypsilophorus, 73, 76, 78, 85,
114
szalayi, Oriolus, 37, 63, 74, 78, 85, 297,
382
tabuensis, Porzana, 20, 46, 72, 85, 117
tachycrypta, Colluricincla, 286
tahitica, Hirundo, 72, 90, 195
Talegalla cuvieri, 113
fuscirostris, 39, 58, 112, 114
jobiensis, 39, 58, 112, 114
Tanysiptera galatea, 184, 186, 190
nympha, 25
tappenbecki, Colluricincla, 284
Tattler, Gray-tailed, 120
Teal, Garganey, 100
Gray, 100
Salvadori’s, 100
teerinki, Lonchura, 47, 77, 409, 410
telescophthalmus, Arses, 253
Monarcha, 62, 253
temminckii, Orthonyx, 18, 19, 20
temporalis, Pomatostomus, 210
tenebricosa, Tyto, 36, 173
tenebrosa, Gallinula, 72, 91, 119
Pachycephala, 20, 25, 46, 274, 284
tenuifrons, Zosterops, 405
tenuirostris, Coracina, 36, 74, 197, 199,
200
437
terborghi, Aegotheles, 176
Tern, Sooty, 122
theomacha, Ninox, 60, 174
thomasi, Melidectes, 392
Thornbill, New Guinea Mountain, 224
threnothroax, Rhipidura, 36, 37, 38, 54,
55, 61, 239, 240, 242
Thrush, Island, 209
Lesser New Guinea, 208
White’s Ground, 207
Timeliopsis fulvigula, 64, 359
timoriensis, Megalurus, 48, 72, 73, 76, 78,
85, 233, 235
Todopsis cyanocephala, 74, 216
torotoro, Halcyon, 27, 36, 54, 55, 60, 168,
184, 186, 187, 189, 190, 191, 387
torquatus, Melidectes, 36, 54, 74, 76, 229,
387, 388, 392, 393, 396
Toxorhamphus iliolophus, 352, 356
novaeguineae, 22, 352, 353
poliopterus, 22,36, G4, 989) Zoi,” aa2,
353, 356
Tregellasia leucops, 53, 54, 62, 87, 257,
258, 260
Trichoglossus haematodus, 52, 59, 74, 78,
81, 86, 89, 142, 144, 145, 146, 147,
148, 151, 352, 396
trichroa, Erythrura, 39, 41, 65, 407, 408
tricolor, Rallina, 54, 118
Triller, Black-browed, 196
Varied, 196
Tringa brevipes, 79, 120
hypoleucos, 79, 120
tristis, Corvus, 63, 304
Gymnocorvus, 304
tristissima, Lonchura, 40, 409, 410, 411
triton, Cacatua, 154
trivirgatus, Phylloscopus, 37, 54, 61, 214,
229, 235
Trumpetbird, 326
Turdus poliocephalus, 20, 48, 71, 209
Turkey, Black-billed Brush, 112
Brown-collared Brush, 112
Wattled Brush, 113
Turnix maculosa, 73, 76, 115
Tyto alba, 36, 74, 173
capensis, 36, 73, 76, 173, 174
tenebricosa, 36, 173
umbrosa, Ptiloprora, 384
unappendiculatus, Casuarius, 39
uniformis, Chlamydera, 346
Uroglaux dimorpha, 24
vana, Lonchura, 77, 409, 410
vanikorensis, Collocalia, 181
INDEX
variolosus, Cacomantis, 30, 35, 52, 74, 78,
85, 164, 166, 167, 168, 170, 216
versicolor, Meliphaga, 377
versteri, Melanocharis, 27, 29, 33, 45, 65,
108, 257, 399, 400, 401, 402, 403, 404
victoria, Goura, 22, 39, 51, 140
virago, Melanocharis, 402
virgatus, Sericornis, 217, 224
vitiensis, Columba, 133
viridis, Androphobus, 24, 47
Ptiliopus, 122, 123
Wagtail, Gray, 205
Willie, 248
wahgiensis, Elanus, 102
Lonchura, 410
Megalurus, 233
Rallus, 116
Saxicola, 206
Tregellasia, 260
Zosterops, 406
wahnesi, Lophorina (Parotia), 306, 311,
312, 313, 314, 321, 332
waigiouensis, Eurystomus, 192
waigiuensis, Salvadorina, 72, 89, 100
wallacei, Semioptera, 316, 317, 321]
wallacii, Aegotheles, 176
Warbler, Black and White Wren, 214
Black-headed Gerygone, 228
Blue Wren, 216
Golden-headed Fantail, 235
Gray Gerygone, 225
Gray-headed Gerygone, 227
Great Reed, 233
Meats 235
Lowland Mouse, 230
Midmountain Mouse, 231
Mountain Mouse, 232
Red-necked Gerygone, 228
Rufous Wren, 216
Swamp Gerygone, 228
Yellow-bellied Gerygone, 227
Wattlebird, Cinnamon-breasted, 396
weiskei, Cacomantis, 168
Whimbrel, Little, 120
Whistler, Brown-backed, 282
Dwarf, 272
Gray-headed, 280
Mottled, 275
Rufous-breasted, 281
Rufous-naped, 283
Schlegel’s, 277
Sclater’s, 276
Sooty, 284
White-bellied, 283
White-eye, Black-fronted, 405
Mountain, 406
whiteheadi, Collocalia, 43, 72, 180, 182
wilhelminae, Charmosyna, 25, 147
Woodcock, East Indian, 121
xanthogenys, Machaererhynchus, 254
Xanthomelus aureus, 346
Xanthotis chrysotis, 374
polygramma, 375
yorki, Caprimulgus, 178
ypsilophorus, Synoicus, 73, 76, 78, 85, 114
zoeae, Ducula, 45, 58, 85, 89, 132, 133
Zonerodius heliosylus, 99
zonurus, Ptilinopus, 128
Zoothera dauma, 37, 207
Zosterops atrifrons, 52, 405
fuscicapilla, 47
minor, 405
novaeguineae, 47, 49, 359, 406
438
QL694.N4 DS
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