1000 Beehler et al. [Auk, Vol. 124 Downloaded from https://academic.oup.com/auk/article-abstract/124/3/1000/5562655 by guest on 01 June 2020 Fr t e e. Melipotes carolae, sp. nov. (Wattled Smoky Honeyeater). Original watercolor painting by John Anderton. The Auk 124(3):1000–1009, 2007 © The American Ornithologists’ Union, 2007. Printed in USA. A NEW SPECIES OF SMOKY HONEYEATER (MELIPHAGIDAE: MELIPOTES) FROM WESTERN NEW GUINEA Bruce M. Beehler,1,2,6 Dewi M. Prawiradilaga,3 Yance de Fretes,4 and Neville Kemp4,5 1 Abstract.—We describe a new species of smoky honeyeater from the Foja Mountains, an isolated range in western New Guinea (Papua Province, Indonesia). A common inhabitant of montane forest and forest-edge in this little-known north coastal range, this species possesses a fleshy pendant suborbital wattle, unique in the genus Melipotes, among other characteristics that distinguish it from all congeners. This fleshy wattle provides a morphological link between Melipotes and the monotypic genus Macgregoria, an alpine inhabitant of the Central Ranges of New Guinea, traditionally treated as a bird of paradise (e.g., Frith and Beehler 1998) but now regarded as the sister genus to Melipotes (Cracraft and Feinstein 2000). The presence of an endemic meliphagid species in the Foja Mountains highlights the biogeographic significance and conservation importance of this geographically isolated upland forest tract, which is also home to the endemic Golden-fronted Bowerbird (Amblyornis flavifrons) and Berlepsch’s Parotia (Parotia berlepschi), a distinctive, recently rediscovered species of six-wired bird of paradise (B. M. Beehler unpubl. data). Received 7 June 2006, accepted 4 November 2006. Key words: Foja Mountains, Indonesia, Meliphagidae, Melipotes carolae, New Guinea, new species, Papua, Wattled Smoky Honeyeater. Una Especie Nueva de Melipotes (Meliphagidae) del Oeste de Nueva Guinea Resumen.—Describimos una especie nueva de Meliphagidae de las montañas Foja, una cadena aislada del oeste de Nueva Guinea (Provincia de Papua, Indonesia). Esta especie es un habitante común de los bosques montanos y los bordes de bosque de esta cadena montañosa poco conocida de la costa norte. Una de las características que distinguen a esta especie de las demás especies del género Melipotes, es que ésta presenta una carúncula carnosa colgante suborbital que es única en el género. La carúncula carnosa representa una conexión morfológica entre Melipotes y el género monotípico Macgregoria, que habita las zonas alpinas de las cordilleras centrales de Nueva Guinea y que tradicionalmente había sido tratado como un ave del paraíso (e.g., Frith y Beehler 1998), pero que ahora se considera como el género hermano de Melipotes (Cracraft y Feinstein 2000). La presencia de una especie de Meliphagidae endémica en las montañas de Foja resalta la importancia biogeográfica y para la conservación de este bosque de tierras altas aislado, en el que también habitan la 6 E-mail: bbeehler@conservation.org 1001 Downloaded from https://academic.oup.com/auk/article-abstract/124/3/1000/5562655 by guest on 01 June 2020 Melanesia Center for Biodiversity Conservation, Conservation International, 1919 M Street NW, Suite 600, Washington, D.C. 20036, USA; 2 Division of Birds, Smithsonian Institution, Washington, D.C. 20013, USA; 3 Ornithology Department, Division of Zoology, Research Centre for Biology-LIPI, Jalan Raya Bogor Km 46, Cibinong 16911, Indonesia; 4 Conservation International Indonesia, Jalan Pejaten Barat 16a, Kemang, Jakarta 12550, Indonesia; and 5 IndoPacific Conservation Alliance, c/o the Bernice P. Bishop Museum, 1525 Bernice Street, Honolulu, Hawaii 96817, USA 1002 Beehler et al. [Auk, Vol. 124 especie endémica Amblyornis flavifrons y la recientemente redescubierta ave del paraíso Parotia berlepschi (B. M. Beehler datos no publicados). a bird survey documented by museum specimens (Diamond 1985, Helgen 2005). The Foja Mountains lie just east of the Mamberamo River, and north of the Taritatu (Idenburg) River. They are entirely isolated from the Central Range as well as from all other adjacent upland areas. The range is linear (west-northwest–east-southeast) with several summits above 2,000 m, the highest reaching 2,200 m. The range is very young, composed of recently uplifted deep-sea sediments that have been sutured to New Guinea’s north coast by the tectonic collision of the Australian and Pacific plates (D. A. Polhemus pers. comm.). The entire massif, encompassing ~300,000 ha, is apparently unroaded and perhaps untrailed, with no villages except in the lowland fringes. Informants from Papasena and Kwerba state that local people do not enter the uplands, in part because of inaccessibility, but also because the summits are considered sacred. From 15 November through 9 December 2005, an international team organized by Conservation International (CI) and the Indonesian Institute of Sciences (LIPI) carried out a 25-day-long Rapid Assessment (RAP) of birds, mammals, amphibians, reptiles, butterflies, and plants in the western part of the Foja Mountains (Fig. 1). On 21 November 2005, the CI RAP team helicoptered into a highland peat bog, one of very few points of helicopter access to the interior of the Fojas. Within minutes of arrival at this site, members of the field team noticed a wattled honeyeater foraging unwarily in forest at the edge of the bog. It was notable for its red-orange facial skin and the pendant wattle that hangs down from each side of the face (Fig. 2). This bird represents a new species of smoky honeyeater (genus Melipotes), for which we propose the following name: Melipotes carolae, sp. nov. Wattled Smoky Honeyeater Holotype.—Museum Zoologicum Bogoriense (MZB), Cibinong, Indonesia, MZB number 30628, male (testis 4 × 6 mm), Bog Camp, Sarmi District, Papua Province, Indonesia, elevation 1,650 m, 2°34.5’S, 138°34.9’E, mist netted on Downloaded from https://academic.oup.com/auk/article-abstract/124/3/1000/5562655 by guest on 01 June 2020 For more than seven decades, New Guinea’s greatest ornithological mystery surrounded the unknown location of the native breeding haunts of the Golden-fronted Bowerbird (Amblyornis flavifrons). This species was described by Lord Walter Rothschild from a single trade skin (an adult male) obtained from an unknown location in western (then Dutch) New Guinea by indigenous collectors apparently in the employ of the plume merchant Duivenbode (Rothschild 1895, Gilliard 1969). Two additional specimens made their way to Rothschild, presumably from the same feather merchant, once the species’ novelty was determined. A general belief held that the forests that were home to this “missing” bowerbird would also support additional avian novelties, so there was considerable subsequent effort to locate the presumably montane homeland of this bowerbird in the hope of discovering an additional trove of ornithological novelties. In spite of diligent investigative field work throughout the first half of the 20th century, the mystery bowerbird eluded an array of western field ornithologists, including F. de Bruijn, Ernst Mayr, Fred Shaw-Mayer, Crown Prince Leopold, S. Dillon Ripley, Sten Bergman, Max Thompson, Phil Temple, and E. Thomas Gilliard (Mayr 1941, Gilliard 1969). Ripley (1964), in particular, was convinced that the unexplored Foja (= Gauttier) Range of northwestern New Guinea was the home of the bowerbird, but his attempt in 1960 to penetrate this range from the north via the Tor River failed because of the logistical hurdles of hiking into this remote, trackless area of difficult terrain. Later in the decade, E. Thomas Gilliard proposed to get into the Foja uplands by helicopter, but these plans were cut short by his untimely death (M. L. LeCroy pers. comm.). It was not until 1979, and the ready availability of helicopter transport, that Jared Diamond was able to carry out a pioneering observational reconnaissance of the Foja Mountains (followed by a repeat visit in 1981). He succeeded in locating the “lost” bowerbird and provided an initial characterization of the range’s montane avifauna (Diamond 1982, 1985). However, because Diamond collected no specimens for museum study, the Foja Range has remained the most significant mountain massif in New Guinea without July 2007] New Honeyeater from Western New Guinea 1003 28 November 2005, prepared by B. M. Beehler, field catalogue number BBFM-54. Diagnosis.—The new species is readily assignable to the genus Melipotes by the large circumorbital patch of bright facial skin, the overall sooty-gray plumage, and the short black bill. Each of the following four characters distinguishes M. carolae (Fig. 2A, C) from all congeners: (1) elaboration of the circumorbital skin patch into a soft, loose, and fleshy pendant wattle at the base of the jaw on each side of the face; (2) the deep red-orange (rather than orangeyellow) coloration of the facial skin patch; (3) the nonflushing condition of the circumorbital patch (see below); and (4) the dull gray throat patch, only slightly paler than adjacent upper breast feathering. The color of the circumorbital patch merits comment. Individuals of all previously known species of Melipotes can “flush” this patch a reddish or deep-orange color when agitated. Melipotes carolae differs by exhibiting a circumorbital patch that, at all times, is substantially redder than the nonflushed state of those of the other species. This was visible and obvious at a distance in all free-ranging birds observed in the field (n = 15 sightings by B.M.B. and additional reported sightings by J. M. Diamond). Of apparent significance is that M. carolae appears to exhibit a circumorbital patch whose permanent state is much like the “flushed” state of the other species of Melipotes. Our many photographs of M. carolae indicate some mottling of facial color at times, but no change in state from unflushed to flushed such as that found in ater, gymnops, and fumigatus (e.g., Fig. 2B, D). The three other species of Melipotes differ from carolae in the following ways: M. ater (Fig. 2F), which inhabits the upland forests of the Huon Peninsula (Fig. 1), is substantially larger (more than twice the mass of all other Melipotes), with a yellow circumorbital patch, satin-black plumage, white spotting on the breast, and a nonpendant caruncle-like “sub-wattle” on the lower edge of the yellow circumorbital patch. Melipotes ater is considerably more sociable and noisy than any other member of the genus (J. M. Diamond pers. comm.). Melipotes gymnops (see Coates 2001), which inhabits the upland forests of the Vogelkop and Bird’s Neck Peninsula of westernmost New Guinea (Fig. 1), has a dark throat, distinctive pale streaks on the lower breast and belly, ochre-washed undertail coverts, and a yellow circumorbital patch. Melipotes fumigatus (Fig. 2B, D), a widespread upland species of the New Guinean central cordillera, the Cyclops, Bewani, and Kumawa ranges (Fig. 1), is in most ways similar to M. carolae but exhibits an orange-yellow (not red-orange) circumorbital face patch, lacks the pendant wattle, and exhibits a distinctly paler chin patch that contrasts with the dark breast plumage. Description of holotype.—Bill and legs black. Iris dark brown. Plumage coloration (following Downloaded from https://academic.oup.com/auk/article-abstract/124/3/1000/5562655 by guest on 01 June 2020 Fig. 1. Map of New Guinea, showing ranges of all species of Melipotes and Macgregoria pulchra. 1004 Beehler et al. [Auk, Vol. 124 color terminology of Smithe 1975): crown, darker than jet black (89); mantle between jet black (89) and blackish neutral gray (82); chin and upper throat between blackish neutral gray (82) and dark neutral gray (83), underparts darker than plumbeus (78), with light neutral gray (85) edging to contour feathers on breast; scapulars and primaries paler than dusky brown (19); undertail coverts darker than smoke gray (45). Measurements of holotype.—Wing chord (unflattened): 124 mm; tarsus: 35 mm; tail: 102 mm; bill from base of skull: 25 mm; bill from feathers: 19 mm; weight: 52.5 g. Specimens examined.—Five apparently adult individuals (the holotype and four paratypes) were mist netted at the Bog Camp between 28 November and 4 December 2005. Two were prepared as study skins (MZB 30628 and 30629), and three as whole anatomical specimens in alcohol (MZB 30630, 30631, and 30632). All specimens are held at the MZB in Cibinong, Indonesia. Tissue samples of all were preserved in 70% ETOH. Comparative study skins of the species ater (n = 3), gymnops (n = 11), and fumigatus (n = 29) were examined from the collections of the National Collection, Cibinong (Museum Zoologicum Bogoriense), and the National Downloaded from https://academic.oup.com/auk/article-abstract/124/3/1000/5562655 by guest on 01 June 2020 Fig. 2. (A, C) Melipotes carolae, (B, D) Melipotes fumigatus, (E) Macgregoria pulchra, and (F) Melipotes ater. July 2007] 1005 52.5 54 59, 60, 63, 65 62 – 46, 47, 49, 60 42, 48, 54 49, 58 – 25 23.5 21, 24.5, 26, – 21, 22, 23, 25 20(2), 22(3) 21,24,24.3,25.5 20, 22(4), 24, 24.5 18, 21(2), 22(3), 24 19(2), 20(2), 21, 22 22.5, 23.5 109, 118, 119, 122, 123(2) 105, 107, 108, 110(2), 111, 114 104, 107, 113,114, 115(2),117, 121 108, 109, 112, 109, 114 100, 108, 109, 121(2) 103, 108, 106, 107, 113, 115 109, 119 Male Female Male Female Male Female Unsexed 35 30.5 33, 33.5, 34, – M. carolae 102 95 99(2), 100, – M. fumigatus goliathi 29(3), 30, 34.5 103, 108, 109, 111, 113 29(6), 30, 31 95, 101, 103(2), 104, 108(2), M. f. fumigatus 27, 28(2) , 29, 33.5,34(2),35 91, 93, 94 96,100,101, 28(2), 29(3), 31,34 93(2),96, 103, 106,109 M. gymnops 27(3) 94, 97, 111 26, 27(2), 28(3), 91(2), 96(2), 97, 99 29, 32 93.5, 100 Mass (g) Culmen (mm) Tail (mm) Tarsus (mm) Wing chord (mm) 124 112.5 123, 125(2), 125.5 The genus Melipotes is an important member of the upland forest bird fauna of New Guinea Male Female Unsexed Remarks Downloaded from https://academic.oup.com/auk/article-abstract/124/3/1000/5562655 by guest on 01 June 2020 Museum of Natural History, Smithsonian Institution, Washington, D.C. The measurements of wing, tail, tarsus, and bill for M. carolae did not differ substantially from those for its putative sister form, M. fumigatus (Table 1). Distribution, ecology, and behavior.—Recorded only above 1,150 m elevation in the Foja Mountains of western (Indonesian) New Guinea, M. carolae was found by the field team to be common in forest interior and forest-edge. It was seen repeatedly foraging unwarily in the middle and upper levels of the vegetation, especially at plants producing small fruits. The Foja Mountains support a large expanse of continuous, closed, humid tropical submontane forest. From the forest physiognomy, we estimate that this mountain range receives what appears to be a moderate amount of rainfall annually (about 3–4 m per annum, with a dry season that apparently peaks in August, according to local informants). The type locality is a sphagnum bog ~500 m in diameter surrounded by mossy forest of moderate stature (20–25 m) dominated by gymnosperms (e.g., Dacrydium, Dacrycarpus) and montane angiosperm taxa (e.g., Nothofagus, Lithocarpus). The montane vegetation of this isolated mountain massif exhibits the Massenerhebung Effect—thus, the forest at 1,650 m in the Foja Mountains is similar to a forest of higher elevation in New Guinea’s central cordillera. Melipotes carolae exhibited behavior typical of the genus Melipotes (Beehler et al. 1986, Coates 1990), with one exception: the individuals encountered on 15 occasions during the field trip were silent. We never heard or recorded this species making a sound. We encountered no evidence of nesting or other reproductive behavior during the field period (late November–early December). The two dissected individuals had nonbreeding gonads. Etymology.—The specific epithet honors Carol Beehler, wife of the senior author, acknowledging her long and unstinting support and her personal commitment to biodiversity studies in New Guinea. The English name highlights the most distinctive feature of this novel species. Table 1. Measurements for specimens of three species of Melipotes (sample sizes > 1 in parentheses). New Honeyeater from Western New Guinea 1006 Beehler et al. polytypic–biological species of Mayr (1963) but that we consider geographically well-delimited and morphologically well-defined allotaxa as species rather than subspecies. Figure 3 supports the divergence of the Melipotes lineage from a Macgregoria-like sister lineage, with attendant character evolution from a large-bodied black montane songbird with a fleshy orbital wattle to a smaller sooty songbird with a circumorbital patch of bare skin. In this regard, Melipotes ater appears to be basal within Melipotes and the gymnops–fumigatus–carolae species-group appears to be derived. Within the latter clade, fumigatus and carolae appear to be sister forms with many shared derived characters. We thus see an interesting pattern of geographic differentiation in three distinct steps. First, there is a Central Cordillera–Huon Peninsula split (proto-Macgregoria–proto-Melipotes basal lineages). Second, we postulate a colonization of the Central Cordillera range and Bird’s Head Uplands by basal Melipotes from the Huon Peninsula, followed by east–west differentiation of this lineage into a western–Bird’s Head form (gymnops) and a central range vicariant (fumigatus). Finally, we propose a colonization of the north coastal ranges by M. fumigatus, followed by isolation and differentiation of the Foja Mountain isolate into M. carolae. We suggest that speciation of carolae in the Foja Mountains is a product of the geographic extent and height of the range and its Fig. 3. Cladogram of the Macgregoria + Melipotes lineage. Downloaded from https://academic.oup.com/auk/article-abstract/124/3/1000/5562655 by guest on 01 June 2020 (Salomonsen 1967; Beehler et al. 1986; Coates 1990, 2001). Virtually all upland forests in New Guinea support a single species of Melipotes. In each montane habitat, the Melipotes tends to be one of the most common and visible members of the local montane forest community. Systematics.—A comprehensive systematic review of the genus must await analysis of tissue samples of M. carolae, which currently remain in Indonesia. Therefore, we here offer some preliminary thoughts on systematics and biogeography of the lineage. First, using molecular analysis, Cracraft and Feinstein (2000) recently demonstrated that the distinctive subalpine frugivore Macgregoria pulchra (“Macgregor’s Bird of Paradise” in Frith and Beehler 1998) is the sister form to Melipotes. We provide a provisional cladogram for the Macgregoria + Melipotes clade, based on external morphology and naturalhistory traits (Fig. 3 and Appendices 1 and 2). We did not believe it necessary to run the small data set through a cladistic algorithm (e.g., PAUP) because only a single instance of homoplasy appears (transformation of character state 8.1 to 8.0 for M. gymnops). In addition, note that we included basal-derived characters that unite both the Macgregoria and Melipotes clades. These are not informative in terms of tree generation but provide additional evidence of the monophyly of the two lineages (cf. Cracraft and Feinstein 2000). We note that the species concept employed here approximates the [Auk, Vol. 124 July 2007] New Honeyeater from Western New Guinea Guinean mountain ranges, Diamond (1982, 1985) provided the initial indication of the biological importance of the Foja Mountains in a regional context. Our discovery of an endemic meliphagid in the Foja Mountains and rediscovery of Parotia berlepschi (the “lost” six-wired bird of paradise; Kleinschmidt 1897, B. M. Beehler unpubl. data) further confirm the faunal distinctiveness of these mountains and provide strong evidence for their longterm biotic isolation (Fig. 1). Although the Foja Range is often united with other outlying northern New Guinean mountain ranges (especially the Cyclops, Bewani, Torricelli, and Prince Alexander ranges) as part of a single biogeographic unit (e.g., Stattersfield et al. 1998, Helgen 2005), our results offer new evidence of the Fojas’ faunal uniqueness compared with even these closely associated ranges. This is also indicated by the discovery of new endemic species of butterflies, mammals, frogs, and plants during our recent Foja survey (van Mastrigt 2006, S. J. Richards unpubl. data). Given that we conducted surveys over relatively limited elevational spans (100–900 m and 1,150–1,850 m, with attention primarily focused at 100, 350, and 1,650 m) and in only one part of the range, we believe that further field work may reveal additional exciting ornithological discoveries in the Fojas. It is apparent from Diamond’s work and ours that the end of the calendar year is the nonbreeding “quiet” season for birds. Field work undertaken in August– September may thus encounter more avian song and breeding activity and potentially reveal certain retiring species that have been overlooked by Diamond and us. Our field work in the Foja Mountains further validates Diamond’s (1985) suggestion that these mountains represent a significant biodiversity resource that merits large-scale conservation. The Fojas are currently designated as a wildlife sanctuary (suaka margasatwa), but this remains in essence a “paper park.” We suggest that the entire mountain range merits a management plan and that additional research should be invested in delineating the distribution of all globally restricted-range species inhabiting the Fojas. In addition, and perhaps more importantly, Conservation International is working with indigenous forest-dwelling communities inhabiting the fringes of the Fojas to delineate a long-term comanagement plan Downloaded from https://academic.oup.com/auk/article-abstract/124/3/1000/5562655 by guest on 01 June 2020 physical isolation. Thus, the presence of typical fumigatus in the Cyclops, Bewani, and Kumawa ranges may be evidence that these ranges are too small to foster the differentiation of a species vicariant of Melipotes (because of periodic extirpation and recolonization). Gibbs (1994) recently observed a Melipotes in the Fak Fak Mountains (Fig. 1) that exhibited a pendant facial wattle. On the basis of pale lower ventral plumage, he believed that this population was a derivative of gymnops, not fumigatus as Diamond (1985) had postulated. It is certainly worth returning to the Fak Fak Mountains to collect the honeyeater and other novelties inhabiting that undersurveyed mountain range. New Guinean biogeography.—The genus Melipotes assumes greater biogeographic significance with the discovery of M. carolae. Melipotes can be added to the list of readily speciating New Guinean montane-forest bird genera that have produced allospecies-level vicariants on several of New Guinea’s north coastal ranges (other examples include Rallicula, Melidectes, Ptiloprora, Amblyornis, Sericulus, Parotia, and Astrapia; cf. Diamond 1969, 1985). These lineages form the systematic “backbone” of the New Guinean montane forest avifauna, and five of these seven “core” genera are endemic to New Guinea’s humid uplands. This discovery confirms that the Foja Mountains are New Guinea’s third-mostsignificant north coastal range, when measured by species-level avian endemism (Diamond 1985). The Tamrau–Arfak Mountains of the Bird’s Head Peninsula of the far west support nine montane endemic bird species, the massive and high Finisterre–Saruwaged Mountains of the Huon Peninsula support four species, and the Foja three species. Like the uplands of the Huon and Bird’s Head, the Fojas harbor an endemic species of Melipotes and an endemic bird of paradise (Parotia). As with Bird’s Head, the Foja possess an endemic gardener bowerbird of the genus Amblyornis (remarkably absent from the Huon, which instead possesses a subspecies of the central cordilleran species A. macgregoriae). The Fojas also possess endemic populations–subspecies of Sericulus, Melidectes, Ptiloprora, and Rallicula (Diamond 1985; B. M. Beehler unpubl. data). Conservation significance.—On the basis of ornithological surveys in outlying New 1007 Beehler et al. 1008 for this important forest block. We believe that these communities are the local stewards of one of New Guinea’s most remarkable natural resources. Acknowledgments Literature Cited Beehler, B. M., T. K. Pratt, and D. A. Zimmerman. 1986. Birds of New Guinea. Princeton University Press, Princeton, New Jersey. Coates, B. J. 1990. The Birds of Papua New Guinea, vol. 2. Dove, Alderley, Australia. Coates, B. J. 2001. Birds of New Guinea and the Bismarck Archipelago. Dove, Alderley, Australia. Cracraft, J., and J. Feinstein. 2000. What is not a bird of paradise? Molecular and morphological evidence places Macgregoria in the Meliphagidae and the Cnemophilinae near the base of the corvoid tree. Proceedings of the Royal Society of London, Series B 267: 233–241. Diamond, J. M. 1969. Preliminary results of an ornithological exploration of the North Coastal Range, New Guinea. American Museum Novitates 2362:1–57. Diamond, J. M. 1982. Rediscovery of the Yellowfronted Gardener Bowerbird. Science 216: 431–434. Diamond, J. M. 1985. New distributional records and taxa from the outlying mountain ranges of New Guinea. Emu 85:65–91. Frith, C. B., and B. M. Beehler. 1998. The Birds of Paradise. Oxford University Press, Oxford, United Kingdom. Gibbs, D. 1994. Undescribed taxa and new records from the Fakfak Mountains, Irian Jaya. Bulletin of the British Ornithologists’ Club 114:4–12. Gilliard, E. T. 1969. The Birds of Paradise and Bower Birds. Natural History Press, Garden City, New York. Helgen, K. M. 2005. The amphibious murines of New Guinea (Rodentia, Muridae): The generic status of Baiyankamys and description of a new species of Hydromys. Zootaxa 913:1–20. Kleinschmidt, O. 1897. Beschreibung eines neuen Paradiesvogels. Ornithologische Monatsberichte 5:46. Mayr, E. 1941. List of New Guinea Birds: A Systematic and Faunal List of the Birds of New Guinea and Adjacent Islands. American Museum of Natural History, New York. Mayr, E. 1963. Animal Species and Evolution. Belknap Press of Harvard University Press, Cambridge, Massachusetts. Ripley, S. D. 1964. A systematic and ecological study of birds of New Guinea. Yale Peabody Museum Bulletin 19:1–87. Rothschild, W. 1895. A new species of bower bird. Novitates Zoologicae 2:480. Salomonsen, F. 1967. Meliphagidae. Pages 338–450 in Check-List of Birds of the World, vol. 12 (R. A. Paynter, Jr., Ed.). Museum of Comparative Zoology, Cambridge, Massachusetts. Smithe, F. 1975. Naturalist’s Color Guide. American Museum of Natural History, New York. Downloaded from https://academic.oup.com/auk/article-abstract/124/3/1000/5562655 by guest on 01 June 2020 We thank the Indonesian Institute of Sciences (LIPI), the Local Forestry Offices (BKSDA) in Papua, the Department of Forests, Directorate General of Forest Protection and Nature Conservation, and the Ministry of Interior, for permission to conduct fieldwork in Indonesian New Guinea. We gratefully thank D. Darnaedi, Director of the Research Center for Biology of LIPI, for his strong support of the scientific collaboration between the Conservation International team and his Indonesian scientific team. We thank Conservation International Indonesia for abundant assistance and support of our field expedition, especially J. Supriatna, I. Budiningrum, and the Mamberamo field team. Our field work in the Foja Mountains could not have proceeded without the hospitality and partnership of the people of Kwerba and Papasena villages, especially P. I. Tawani and P. T. Kawena. HeliMission made its helicopter available to us, without which our montane survey could not have taken place. We salute S. Richards, co-leader of this expedition, for all that he did to ensure its success. We thank R. C. Banks, J. M. Diamond, K. Helgen, M. LeCroy, R. Pasquier, P. Rasmussen, D. Wilcove, and two anonymous reviewers for technical and editorial suggestions that improved this paper. The project was generously supported by grants from the Swift Foundation, the Gordon and Betty Moore Foundation, the National Geographic Society, and the Global Environment Project Institute. [Auk, Vol. 124 July 2007] 1009 New Honeyeater from Western New Guinea Stattersfield, A. J., M. J. Crosby, A. J. Long, and D. C. Wege. 1998. Endemic Bird Areas of the World. Birdlife International, Cambridge, United Kingdom. van Mastrigt, H. 2006. New (sub)species of Delias (Hübner, 1819) from the Foja Appendix 1 1. Facial wattle: 0 = present. 2. Wattle type: 0 = semicircular and ocular, 1 = circular and facial, 2 = circular and with pendant extensions. 3. Wattle color, at rest: 0 = yellow-orange, 1 = orange-red. 4. Body size: 0 = large, 1 = medium. 5. Contour plumage background color: 0 = black, 1 = gray. 6. Ventral plumage pattern: 0 = plain, 1 = pale spotted–streaked, 2 = mottled–scalloped. Associate Editor: N. S. Sodhi 7. Undertail coverts plumage pattern: 0 = dark, 1 = ochre. 8. Chin plumage: 0 = dark, 1 = distinctly paler, contrasting breast; 2 = slightly paler than breast. 9. Wing patch: 0 = present, 1 = absent. 10. Sociability: 0 = sociable, 1 = solitary. 11. Vocalization: 0 = vocal, 1 = mildly vocal, 2 = silent. 12. Wing sound production: 0 = present, 1 = absent. 13. Bill morphology: 0 = short and straight. 14. Diet: 0 = highly frugivorous. Note: Characters 1, 13, and 14 are included to demonstrate characters linking Macgregoria with Melipotes (see Cracraft and Feinstein 2000). Appendix 2. Character matrix for the Macgregoria–Melipotes lineage. Character number Character 1 2 3 4 5 6 7 8 9 10 11 12 13 14 Facial wattle Wattle type Wattle color Body size Contour plumage color Ventral plumage pattern Undertail coverts Chin plumage Wing patch Sociability Vocalization Wing sound production Bill morphology Diet Macgregoria pulchra Melipotes ater Melipotes gymnops 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 1 1 0 0 1 0 0 0 1 0 1 1 1 1 0 1 1 1 1 0 0 Melipotes Melipotes fumigatus carolae 0 1 0 1 1 2 0 1 1 1 1 1 0 0 0 2 1 1 1 2 0 2 1 1 2 1 0 0 Downloaded from https://academic.oup.com/auk/article-abstract/124/3/1000/5562655 by guest on 01 June 2020 Character Matrix for Melipotes and Macgregoria Mountains, Papua, Indonesia (Lepidoptera: Pieridae). Futao 51:11–18.