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A New Species of Hawk-Owl Ninox from North Sulawesi, Indonesia PDF

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Preview A New Species of Hawk-Owl Ninox from North Sulawesi, Indonesia

THE WILSON BULLETIN A QUARTERLY JOURNAL OF ORNITHOLOGY Published by the Wilson Ornithological Society VOL. Ill, NO. 4 DECEMBER 1999 PAGES 457-630 Wilson Bull., 111(4), 1999, pp. 457-464 A NEW SPECIES OF HAWK-OWL NINOX FROM NORTH SULAWESI, INDONESIA PAMELA RASMUSSEN* C. — ABSTRACT. A distinctive new species of hawk-owl, Ninox ios, is described from a specimen collected in m 1985 in forest at 1120 in Bogani Nani Wartabone (then Dumoga-Bone) National Park, North Sulawesi, Indonesia. It was previously identified as a rufous morph of the Ochre-bellied Hawk-Owl, N. ochracea. Ninox ios is small, predominantly bright chestnut, and lacks facial patterning; it has pink orbital skin, yellow irides, triangular whitish scapular spots, a finely banded and relatively long tail, unusually short, slender tarsi that are feathered for most of their length, and weak claws. Its relationships within the genus Ninox are unclear; it differs in several morphological characters from all other species. Because Ninox ios is only known from one specimen, its distribution and conservation status are unknown; nothing is known of its ecology, but it probably occurs primarily at higher elevations than N. ochracea. Received 14 Dec. 1998, accepted 5 May 1999. For many years two endemic species of the daal netted an almost entirely bright rufous genus Ninox were thought to occur on the cen- Ninox (Frontispiece) in Bogani Nani Warta- tral Indonesian island of Sulawesi. Of these, bone (then Dumoga-Bone) National Park, the Speckled Hawk-Owl (Ninox punctulata) North Sulawesi, Indonesia (Fig. 1). He con- primarily inhabits disturbed lowland habitats cluded that this individual represented “a pre- throughout the island (White and Bruce 1986), viously undescribed rufous phase” of N. and is morphologically quite different from ochracea (Rozendaal and Dekker 1989), and other endemic Indonesian Ninox. The poorly this treatment was followed by Coates and known Ochre-bellied Hawk-Owl [A. ochracea Bishop (1997). (= perversa)] of the lowland rainforests in While working on small owls at the Na- North and Central Sulawesi (White and Bruce tional Museum of Natural History/Naturalis, 1986) is a small, fairly typical member of its Leiden (NNM, formerly Rijksmuseum van RMNH) genus (Frontispiece). Because there had been Natuurlijke Historie, in June and Oc- no indication that a third species might occur, tober 1998, I chanced to see the rufous Sula- it was a surprise when in 1985 F. G. Rozen- wesi specimen, which had been registered as RMNH 84701 but had not yet been incorpo- Was'hNiHngBton3,3D6CM2R05C60,1a1n4d, MSimcihtihgsaonniStaanteIUnnsitviteurtsiiotny, rpautrecdhaisnetobythNeNmMa.inOncoltlheectsieocnonfdololcocwaisnigonitsI Museum, East Lansing, MI 48824-0590; noted that it differed in several morphological E-mail: [email protected] features from Ninox ochracea, in addition to FRONTISPIECE. Cinnabar Hawk-Owl (Ninox ios, upper two) compared with Oehre-bellied Hawk-Owl (TV. ochracea, lower left), and Bum race of Moluccan Hawk-Owl (TV. squamipila hantu, lower right). Original watercolor painting by Ian Lewington. 457 458 THE WILSON BULLETIN • Vol. Ill, No. 4, December 1999 FIG. 1. Map of Wallacea showing collection locality of the holotype of the Cinnabar Hawk-Owl (Ninox ios), other locations mentioned in the text, and approximate ranges within the region of other species of Ninox and subspecies of W. scutulata, which occurs throughout the region. CINNABAR HAWK-OWL the obvious color differences. Subsequent Ninox ios sp. nov. mensural analyses of series of all species of — RMNH Ninox have confirmed the distinctness of the Holotype. 84701, adult male rufous Sulawesi specimen (an adult in good (Frontispiece), according to the label collected m condition) in many characters. Although sev- in a forested valley at 1 120 at Clark’s camp eral Ninox species from other areas are typi- (Hill 1440), east-central Bogani Nani Warta- cally rufous, morphism (and thus true rufous bone National Park, North Sulawesi, Indone- morphs) appears to be unknown for any Ni- sia (ca 0°40' N, 123° 0' E) by F. G. and C. nox, and in any case most of the differences M. Rozendaal the night of 5-6 April 1985 (the are structural and thus would not be related to label date of 7 April presumably indicates date morph. Despite the fact that only one rufous of death). Label data: “Completely ossified specimen is known thus far from Sulawesi, skull”, “weig—ht 78 g”. A there is no reason to believe that any of its Diagnosis. small, lightly built, nearly several novel character states are aberrant, and uniformly rich chestnut hawk-owl with a rel- there can be no reasonable doubt that it rep- atively long tail and narrow pointed wings, lax resents a new species. feathering, no facial pattern, mostly feathered Rasmussen • A NEW SPECIES OF HAWK-OWL 459 short slender tarsi, and rufous, narrowly dark- the patterning of its breast feathers, which barred wings and tail. have a light rufous (vs dark brown) area sur- Compared with all flying states of Ninox rounding the whitish shafts. = ochracea [n 20 (three of which are fully The Philippine Hawk-Owl {Ninox philip- grown juveniles); 6 males, 4 females, 10 un- pensis) superspecies (sensu Dickinson et al. sexed], N. ios is much smaller in most dimen- 1991, but see Collar and Rasmussen 1998) is sions (Table 1, Fig. 2), but has a relatively composed of several dark brown to brown- longer tail and rictal bristles. Its wing, while and-ocher forms that are either barred or shorter than that of N. ochracea, is narrower streaked below. None of the taxa included in and more pointed (Fig. 3). Ninox ios has a TV. philippensis can be described as warmer- much shorter, shallower bill and smaller nares toned than rufescent brown. All have much than N. ochracea. It has short, slender tarsi heavier claws and relatively shorter tails (Fig. that are mostly feathered on both surfaces, 2A) than TV. ios, from which they also differ whereas N. ochracea has longer, stout tarsi in wing shape (Fig. 3). One form, TV. [philip- that are largely unfeathered on the anterior pensis] mindorensis (see Frontispiece), is (acrotarsal) side and are virtually unfeathered somewhat similar in overall size and tarsal on the posterior (plantar) side, with numerous feathering to TV. ios than is any other taxon, stiff bristles over the unfeathered areas. The including TV. ochracea (Fig. 2C), but not in new species has relatively sparse, fine rufous plumage or the above-mentioned shape char- bristles on the extreme lower tarsi and on its acters. slender toes (although the bristles are heavier All taxa of the paraphyletic Moluccan and longer on the hallux), while N. ochracea Hawk-Owl {Ninox squamipila\ split provi- has more profuse, heavier, mostly pale bristles sionally into at least three species by Norman (which are usually longer but sometimes worn et al. 1998) are considerably larger and heavi- down to stubs) on the tops and sides of its er-legged than TV. ios, and all differ from it stouter toes. Ninox ios has much smaller, more additionally in having whitish-barred under- slender claws that are dark for most of their parts and scapulars. Despite the above differ- length (vs large and mostly pale in N. ochra- ences, Ninox s. hantu (Frontispiece) of Bum cea). The holotype of TV. ios had pink orbital superficially resembles the much smaller TV. skin (vs blackish in TV. ochracea) and yellow ios because of its overall mfescence and re- eyes, as does TV. ochracea according to Strese- duced barring below, as well as its obscure mann (1940), who based this statement on G. facial pattern and finely barred tail. The Sum- Heinrich’s specimens [although Meyer and ba Hawk-Owl (TV. rudolfi) is large and strik- Wiglesworth (1898) mentioned a brown-eyed ingly different, with a heavily spotted crown, TV. ochracea]. The base of the bill and the cere barred underparts, and broadly banded and of TV. ios appear entirely pale (vs the basal speckled upperparts. The widespread and var- two-thirds conspicuously dark in specimens of iable Brown Hawk-Owl {Ninox scutulata) is TV. ochracea). also a much larger species, with a broadly In plumage, TV. ios differs conspicuously banded tail and large, heavily feathered tarsi. from both adults and juveniles ofTV. ochracea It is dark brown above with the underparts in its overall bright rufous coloration (vs dark heavily streaked, or nearly solid dark brown brown and yellow-ocher). Unlike all flying in TV. s. obscura of the Andamans. The nom- stages of TV. ochracea, it lacks facial pattern- inate race of the Andaman Hawk-Owl (TV. a. ing, including the whitish supercilia typical of affinis) is smaller than TV. scutulata, to which most of its relatives, and also lacks white it is otherwise quite similar, while the larger markings in the wing coverts and flight feath- Nicobar race (TV. a. isolata) is even more like ers. Less obvious distinctions from TV. ochra- some races of TV. scutulata. cea include its more triangular (vs squarer The highly varied subspecies (including a tipped) whitish scapular spots, its mainly ru- new one described from Roti Island, south- fescent rictal bristles (vs blackish with white west of Timor, Lesser Sundas; Johnstone and bases), its more narrowly barred rectrices, its Darnell 1997) usually grouped in the Southern vaguely dark-scalloped lower underparts (vs Boobook {Ninox novaeseelandiae) as well as plain ocher or somewhat brown-streaked), and the Manus Hawk-Owl (TV. meeki) are also * ' . -1' 460 THE WILSON BULLETIN • Vol. Ill, No. 4, December 1999 uc/5 2<D .ti c/5 ofl d) TD x: "OD.c0^ o (N o 0m0 r04o CroO froO frNo fSOt fOO roo 0O4n 0O4N 0O4S CNOO o CmO o CN^O5 ud) Cac3 -dC) (—N1 *§ ;:; 3.9 C(NN rsod 0^ 3.5 0sd4 dO; d 04; qrd q04 oi Q0O4 0i4n q cdn 0d0 0044 Ci. ocb OC^L -CO0C=JX3) <LC9<sj Cr+Nn1 (Cd+N1N <(+N1N dr+1- 0Oi+4n1n S0+O14 Nt+ON1 Nd+O1 dS+t1 1+/15 io+n1 O0d+N41 0rr+01od Co+Oi1 NI+*O1-* d00+401 c0+d01 N—+O1H N—+O1 <Uui 3 c3 E c/5 ji d) 'u OCJJ 3 ^ to DC V«) 'O On 00 o r- in 04 — mOS O' mfO o om os in o 00 in •o r- SO NO r- NO NO SO so »n in O' O' NO O' sO d) o — m “_O3 3o 2^ ka.. r4 d 04 sOo' rd 0d0 O—N d dOn 04 f0O4 0040 c0d4 c00d 0o4i dCO d0- 0C4O O c ^ 2 s</<31 +1 +1 m+1 +1 +1 +1 O+1 +1 +1 +1 +1 —+1 +1 +1 +1 +1 +1 +1 +1 p o NO in so 00 3d- Sdt NO NO ods Ods 0d0 0- NO 04 c d §; (N NrO4 04 0- in ,0-4H sd in in i0n4 cCdO 00 04 cd i"n d) C3 (U N,(mD 3 X £ 3 a 2 o <— ^C/5 lU 3 53 ou y s m a -5 3 in SO 0- so sO in so so so NO CO SO in NO in ^ CN CN 04 04 04 04 04 04 04 04 04 04 04 04 04 04 04 04 04 xo3: 2OX) 13 q idn O) 5.2 04 1.7 1.2 0.8 2.1 2.6 2.7 2.8 CcOd 3.9 qoi 0d4 0d- q —_o £ p c-^ 2 « l d+1 i+n1 Od+S1 o+1 +1 +1 —+1' -—+t1 rd+o1 N2+O1 0+41 +1 +1 N+O1 O+N1 0+41 N+O1 0+41 O+1 CN c(nN •NdO Om'! 0044 rd c0d4 CO C0O0 0oi0 0O4s cd »c—d o _C/5 d) x: *CONJ &-c- doa:; 1 ."dcno/)5 WoSM>i', x--EO;w OCIrdTSN1) m(idNn 0Cod44 m1OOs' soro- <0io4n1 0Ow4N1 rOwo1 wO1 l—wo1i 00i44n1 m00q00401 dC0O01 iCinOn1 qoO00S00 iNC004nOO4 scpd1 00Cd/-4^O'1V qN0cr-4pd1s 3r- “QOJ r- On N—O O—n 00 m ^r- m^ ^ NO S"O so NO 00 NO NO ON d 0u E^ ? "" a ir)'r'--00O000N00^O'O'Ot^0N—I'sTroiriO (ifnj\ V r" — r-iOu-istr<S—'O—-(NCNricNCNVO—"OOrO CL ^ +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 c/5 m — ^cNO(Nmr<^staNOt^t^—'f^'Ooor^oo — O d — — m s ro n O—' lO 0\ On 00 On 0c0 fs 00 lo (N 'sj' On OJ <u s '5 > *5 c u <u Co-* •-! ^'^.SJ Or—n'NCONOr;or'im—vocNO 04——<' 0—4r(OSrOOONN^lO^Ntr^N4fONOld0O0NN——O 04 (N r' in I d ^ s u ° M <£ _ c Q- "5 ^ (/) Co -uO 3 +1-^-2 +1 C/5 CO S C E S3 ^ o 1) ^ o wj sw: —c c3/5 £ c/5 U_JL_l SViJ UaB« jacj &C0 ddc/))5 2toc2oc2ot^2od2oct2oc2odt2ot2c: —o 3-Cc:Elij cUs3/i5 0cd)0 S(d3U> CSQ ?.S <cM/u5 E DE « ^C/5C/5C/5C/5C/5C/5C/5C/5 -ocC/5 cD/5 s s 3 ^ 2 -^rl^^i^»r)'Or-ooc^ c/5 ,o § 2 U < Qi cucuCua<(xa<cu(XCL a! s s D Rasmussen • A NEW SPECIES OF HAWK-OWL 461 Culmen from skull (mm) Tarsus length (mm) FIG. 2. Bivariate scatter plots (measurements in mm) for Ninox ios (filled circles), N. ochracea, N. “squam- ipila” (sensu White and Bruce 1986), and N. philippensis (sensu Dickinson et al. 1991): A. culmen vs tail length; B. auricular vs rictal bristle length; and C. tarsus length vs extent of unfeathered tarsus. For A, the main Philippine taxon groups are treated separately, while for B and C they are combined. 462 THE WILSON BULLETIN • Vol. Ill, No. 4, December 1999 - 60 sell (1977) notation, in which the first number and letters refer to the hue, the number pre- ceding the slash is the value or lightness, and the last number is the chroma or saturation. The holotype was directly compared with 11 specimens of N. ochracea (including the ho- lotype) at NNM, and a series of photographs of it was compared to specimens from other museums. Front ofhead from base ofbill through cen- ter of forecrown and including supercilia, uni- form rich chestnut (5YR 5/8); center of crown 5YR through mantle slightly darker (close to 4/6); rictal bristles fairly long (maximum 24 mm), profuse, and dark chestnut, somewhat blackish near tips; auriculars with fairly long FIG. 3. Shortfalls from wingpoint of each primary distally extended barbs (total length of longest (PI = outer primary; shortfalls are distance of tip of feather 23 mm) that are paler basally (5YR 5/ each primary from longest primary in folded wing) for 10) and grade to black near the tips; chin and Ninox ios, N. ochracea, N. [philippensis] spilocephala, throat paler chestnut (5YR 6/8) than forehead. and N. [/?.] spilonota. Ninoxp. philippensis and similar Sides of neck and breast, back, rump, and races are virtually identical in pattern ofprimary short- falls to N. [p.] spilocephala and thus are not shown uppertail coverts are all approximately the separately, while all bar-bellied populations in the N. same rich dark chestnut (5YR 4/8). The un- philippensis superspecies are similar to race N. p. spi- derparts appear very lightly dappled, slightly lonota. paler chestnut (5YR 5/8) than upperparts. Most breast feathers have pale shaft streaks (5YR 7/8) and pale rufous surrounding areas, larger than N. ios and are streaked or heavily some with darker dappling at sides, and feath- blotched below. None of the remaining Aus- ers of lower underparts are mostly pale rufous tralasian taxa (Papuan Boobook Owl, N. theo- with vague darker scalloping (2.5YR 5/8); un- macha; Rufous Owl, N. rufa'. Powerful Owl, dertail coverts rufescent whitish with the tips N. strenua-, or Barking Owl, N. connivens) ap- scalloped rufous (5YR 6/8). proach N. ios more closely than the above. The scapulars have large mostly triangular The other Sulawesi endemic. Speckled Hawk- whitish spots with broad dark chevron-shaped Owl (N. punctulata), and some Melanesian tips (5YR 4/4). The upper secondary coverts taxa (Bismarck Hawk-Owl, N. variegata-. New are almost uniformly rufous (5YR 6/8) and the Britain Hawk-Owl, N. odiosa; and Solomons upper primary coverts are darker (5YR 4/2). Hawk-Owl, N. jacquinoti) are strikingly dif- The remiges are faded, pale, and worn, in ferent in plumage and morphology, with short striking contrast to the fresh, richly colored tails, very heavy tarsi, and Athene plum- scapulars. The inner webs of the primaries and age pattern and toe bristles; in fact some had narrow vague dark bands of the outer webs been placed in that genus (among others) in are dark grayish brown (5YR 4/4); only the the past. The White-browed Owl (Ninox su- outer webs have broader light bands (5YR 7/ perciliaris) of Madagascar is very different 6). The base color of the secondaries is dull from other Ninox (H. F. James, pers. comm.) rufescent ochraceous (7.5YR 6/8), with fine as would be pr—edicted by its distribution. dark dusky brown bars (7.5YR 4/4). The inner Distribution. To date Ninox ios is known webs of the undersurfaces of the inner pri- only from the type locality in North Sulawesi, maries and secondaries are basally pale rufous Indonesia. It might occur at similar elevations (7.5 YR 8/6), as are the uppersurfaces of the elsewhere in the Minahasa Peninsula of North inner webs of the inner secondaries, which contrast strongly with the dark bands. The un- Sulawesi. — Description ofthe holotype. Color match- derwing coverts are solid pale rufous (7.5YR ing was done under natural light using Mun- 7/8). The uppertail surface has pale bands of Rasinussen • A NEW SPECIES OF HAWK-OWL 463 — dull rufous (SYR 5/6) that are narrow basally Habitat and elevation. Most researchers and wider distally, and about 12 narrow very have considered N. ochracea to be restricted dark brown bands (SYR 3/2) that fade out to- to the lowlands below 800 m (Stresemann ward the tip. There are no definite bands for 1939, White and Bruce 1986, Stattersfield et the terminal 20 mm. The rectrices are heavily al. 1998). More recently, Coates and Bishop worn and faded. (1997) gave the elevational range ofN. ochra- The short, slender tarsi are completely cea as up to 1780 m, but this was probably feathered with short pale cinnamon (7.SYR 7/ based on the questionable vocal records men- mm 6) pennaceous feathers to about 12 ante- tioned in Rozendaal and Dekker (1989) and riorly (measured from joint of digits 1-2 of the collection of the type of N. ios at 1120 m. mm middle toe) and posteriorly to about 6 All montane records of N. ochracea therefore (measured from base of hallux). The toes ap- require review in light of this new species. pear to have been slender, with sparse, short Ninox ios clearly occurs in sympatry with, al- rufous bristles on the tops and sides of each though very likely at higher elevations than, toe. The claws are small, delicate, and mostly N. ochracea. — blackish but with pale bases. Molt, breeding, and ecology The holo- . The soft part colors recorded on the original type of N. ios clearly had recently molted its label are: eyes “bright yellow; pink orbital scapulars, which were bright and fresh and skin”, bill “ivory”, feet “pale whitish-yel- contrasted strikingly with the relatively dull low”. tertials and other flight feathers. The feathers Mea—surements of the holotype (by au- on the crown appeared to be worn, while those thor). Culmen (from skull) 17.9 mm; cul- of the back appeared fresh. Only 10 rectrices men (from distal edge of cere) 10.7 mm; tar- were present. Active molt ofthe flight feathers sus 22.6 mm; wing 172 mm; tail 97 mm. Total was not detected, but avoid damaging the length of prepared specimen 220 mm. See Ta- unique specimen a thorough examination was ble 1 for measurements of other characters of not attempted. The size of the label drawing 6X4 the holotype a—nd those of other species. of the largest testis (which measures Etymology. This new species is named mm) suggests a bird not completely reproduc- Ninox ios (Greek for rust) for its striking over- tively quiescent. Because nothing is known of all coloration. The specific epithet is here used the habits ofN. ios, it is possible only to spec- as a noun in apposition to Ninox, which, al- ulate that its morphology (which recalls that though usually treated as feminine, is a port- of owlet-nightjars Aegothelidae) suggests the manteau combining Nisus and Noctua. The likelihood of its preying largely upon soft- common name “cinnabar” also refers to its bodied inverteb—rates caught in flight. predominant color, which is similar to that of Systematics. The affinities of Ninox ios mercuric sulfide before prolonged exposure to are unclear; it shows many morphological dif- light. ferences from all other species, particularly in its small size, relatively long tail, narrow DISCUSSION — pointed wing, and weak tarsi and claws. Al- Voice. Not definitely known. Rozendaal though membership in the polytypic N. phi- (Rozendaal and Dekker 1989:97) mentioned lippensis superspecies might ‘^eem geograph- “disyllabic calls ascribed to [N. ochracea] re- ically plausible, the pattern ^f primary feather corded at Clark’s camp and on the summit of lengths shown by TV. ios is closer to that of TV. G.[unung] Muajat during April 1985.” Ek- ochracea than to any form of TV. philippensis. strom and coworkers (1998:39) reported “an Phylogenetic analyses will be required to un- unknown owl Ninox sp.” giving a series of derstand the rela—tionships of TV. ios. dry hoots rising and falling in pitch in dense Conservation. As only one specimen is evergreen valley forest near the eastern known, it appears likely that Ninox ios has a boundary of Lore Lindu National Park, at limited range and/or is rare. However, noctur- about 1300 m, in the northern part of central nal birds are frequently overlooked. Also, Sulawesi (J. Tobias, pers. comm.). Either of most scientific bird collecting in Sulawesi these reports might refer to Ninox ios but con- took place before mist-nets were widely avail- firming field data are required. able, and at lower elevations. Ascertaining its 464 THE WILSON BULLETIN • Vol. Ill, No. 4, December 1999 LITERATURE CITED vocalizations and calling periods will be a pre- requisite to carrying out effective surveys, A Coates, B. J. and K. D. Bishop. 1997. guide to the which will be essential to establish the degree birds of Wallacea. Alderley, Queensland, Austra- of risk faced by this unique new species. lia. The only other bird species thought to be Collar, N. J. and P. C. Rasmussen. 1998. Species restricted to North Sulawesi is the poorly limits in the Ninox philippensis complex. Ostrich known Matinan Flycatcher (Muscicapa san- 69(3-4);398. fordi), which has been found only in the Du- Dickinson, E. C., R. S. Kennedy, and K. C. Parkes. 1991. The birds of the Philippines. British Orni- moga-Bone and Tentolo-Matinan mountains thologists’ Union, Tring, U.K. between 1400 and 1780 m. The fact that a Ekstrom, j., j. Tobias, and J. Robinson-Dean. 1998. species as distinctive as Ninox ios could have Forests at the edge of Lore Lindu National Park, escaped description until now clearly under- central Sulawesi. Oriental Bird Club Bull. 28:36— scores the fact that our knowledge of the avi- 39. fauna of Sulawesi is still in a rudimentary Johnstone, R. and J. Darnell. 1997. Description of a new subspecies of Boobook Owl Ninox novae- state. seelandiae (Gmelin) from Roti Island, Indonesia. ACKNOWLEDGMENTS West. Aust. Nat. 21:161—173. Meyer, A. B. and L. W. Wiglesworth. 1898. The Special thanks are due R. W. R. J. Dekker, M. S. birds of Celebes and the neighbouring islands. dHeono,gamnodedto,EanGd. RHo.zevnadnaaGlr.oSupwe,ciNmNeMn/sNaotfucroamlpiasr,atLievie- Vol. 1. R. Friedlander & Sohn, Berlin, Germany. Munsell. 1977. Munsell color charts for plant tissues. taxa were examined in museums too numerous to enu- Gretagmacbeth, New Windsor, New York. wmehriacthe tinhdainvkidGu.alEly,Barbrutowcclhioeufglhy atnhdeRfoSlwleoewti,ngA,mefro-r Norman, J. A., L. Christidis, M. Westerman, and E iDcealnawMauIrseeMuumsoefuNmatoufrNaaltuHriastlorHyi,stoNrey,wWYiolrmki;ngGt.onHe;sDs., Asp.ecRif.icHisltla.tus1o9f98t.heMCohlreicsutlmaarsdIastlaancdoHnfaiwrkm-sOtwhle Willard, Field Museum of Natural History, Chicago; Ninox natalis. Emu 98:197—208. nRa.ti;S.S.KePrninjeodnyo,anMdusDaerujmonoo,fMNuatsuerualmHZiosotlooryg,icCuimncBion-- Rozennodtaaatled,cEheGck.liasntdofR.thWe.biRr.dsJ.ofDethkekeDru.mo1g9a89-.BoAnne- goriense, Cibinong, West Java; S. L. Olson, G. R. National Park, North Sulawesi. Kukila 4:85-109. Graves, and B. M. McPhelim, National Museum of Stattersfield, a. j., M. J. Crosby, A. J. Long, and Natural History, Smithsonian Institution, Washington, D. C. Wege. 1998. Endemic bird areas of the D.C.; M. P. Walters and R. P. Prys-Jones, The Natural world. BirdLife International, Cambridge, U.K. History Museum, Tring, U.K.; G. Boenigk, Staatliches Stresemann, E. 1939. Die Vogel von Celebes. Part 1. Naturhistorische Museum, Braunschweig, Germany; J. Ornithol. 87:299-425. and E C. Sibley, Yale Peabody Museum, New Haven. Stresemann, E. 1940. Die Vogel von Celebes. Part 2. The manuscript was improved by the comments of R. J. Ornithol. 88:389-487. W. R. J. Dekker, N. J. Collar, B. M. Beehler, and R. White, C. M. N. and M. D. Bruce. 1986. The birds Hill; J. Tobias provided information; and the frontis- of Wallacea. British Ornithologists’ Union, Lon- piece was painted by 1. Lewington. don, U.K.

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