A Systematic and Ecological Study of Birds of New Guinea

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1 PEABODY MUSEUM OF NATURAL HISTORY YALE UNIVERSITY BULLETIN 19 A Systematic and Ecological Study of Birds of New Guinea BY S. DILLON RIPLEY Smithsonian Institution, Washington, D. C. NEW HAVEN, CONNECTICUT 1964

2 Printed in the United States of America Copyright Peabody Museum, Yale University New Haven, Connecticut, 1964

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4 CONTENTS LIST OF ILLUSTRATIONS vi ABSTRACT 1 INTRODUCTION, ACKNOWLEDGMENTS AND ITINERARY 2 HABITATS 5 ALTITUDINAL SHIFTS OF RANGE AND ECOLOGICAL OVERLAP AT FEEDING TREES 6 EXTENSIONS OF RANGE 7 BREEDING SEASON 7 SPECIES ACCOUNT 12 LITERATURE CITED 76 INDEX 79 PLATES I II v

5 ILLUSTRATIONS PLATES I. 1. Hollandia Bay from Holtekong looking west towards Sentani. Mrs. Ripley and Jusup Khakiaj with cassowary chick. 2. Ilaga River, 7000 feet. Dani rattan and wood bridge. 3. Dani village dancing area, 7500 feet. Casuarina and pandanus trees in background, looking southwest towards upland plateau trail to west Baliem at 9000 feet. II. 4. Nest of Lonchura tristissima. 5. Sirikena at the Moasets bivouac (560 m altitude = c ft), Lower Mamberano River. Collection of Megapodius and Talegalla eggs. 6. Lowland forest with slash cultivation of cassava and plantain in foreground. Bodim, 300 feet altitude. 7. Uhunduni hunter with stone ax, Ilaga 7500 feet. FIGURES 1. Map of the central highlands area. 2. Contour feathers from specimens of the Australian Gray Duck, Anas superciliosa pelewensis. 3. Head and bill of Rhamphomantis megarhynchns specimens. vi

6 YALE UNIVERSITY PEABODY MUSEUM OF NATURAL HISTORY BULLETIN NO. 19, 87P., 2 PLS., 3 TEXT-FIGS., 1964 A SYSTEMATIC AND ECOLOGICAL STUDY OF BIRDS OF NEW GUINEA By S. DILLON RIPLEY ABSTRACT During two trips to West New Guinea in 1954 and 1960, the author made an extensive collection of birds as well as field notes in a variety of habitats ranging from the western islands adjacent to the coast of West New Guinea, the northern coastal areas from Sarmi to Hollandia (Kotabaru), and the mountains of the interior including a newly-opened valley, the Ilaga, in the Nassau Range. Some three hundred eighty-two forms of birds were observed during a fourmonth period. Descriptions of the habitats are given. In addition discussion is made of altitudinal shifts of range and the phenomenon of crowding and overlap of related species at feeding trees. At such feeding trees under compacted situations, high intensity calls and demonstrations of aggressive behavior occur. Extensions of range are listed and data on breeding seasons are presented indicating overwhelmingly that similar species breed at two rather different seasons geographically, earlier in the calendar year in the central areas of West New Guinea, later in the calendar year in the western islands and west coast, presumably correlated with the weather and monsoonal seasons. In the species list a considerable number of field notes are included as well as native names and occasional native beliefs.

7 A SYSTEMATIC AND ECOLOGICAL STUDY OF BIRDS OF NEW GUINEA* S. DILLON RIPLEY INTRODUCTORY ACCOUNT AND ACKNOWLEDGMENTS In 1954, my wife and I made a trip into the northern Moluccan islands of eastern Indonesia. For this journey I was the recipient of a Guggenheim Fellowship, as well as a grant from the National Science Foundation (no. B-664). In addition, I received a grant from the Vose Fund of the Explorer's Club of New York. In 1960, a further summer's field work in the then Netherlands New Guinea (now West Irian) was aided by a grant from the Penrose Fund of the American Philosophical Society. To all of these distinguished organizations I am most grateful for valued aid and support. I am very grateful to the authorities of the American Museum of Natural History for permission to work with their collections. I have been in frequent consultation with Dr. Thomas Gilliard and Dr. A. L. Rand, co-authors of the New Guinea handbook, which will be published in I wish to thank Shirley Hartman for her help with two drawings and a map, as well as John Howard for photographic reproductions. New Guinea, particularly that part known now as West Irian, formerly Netherlands New Guinea and the adjacent islands, is still not well known ornithologically although expeditions have been visiting the island and reporting on collections, essentially in a taxonomically descriptive sense, since the days of A. R. Wallace in the early Eighteen Fifties. However, many areas of this rich and strongly diversified habitat still remain unexplored faunistically. The latest large effort at biological exploration was that of the Archbold Expedition of the American Museum of Natural History of , primarily into the previously unexplored highlands of the central mountain ranges near Mount Wilhelmina and the Baliem Valley. That expedition was a landmark of exploration in New Guinea, using an amphibian plane, vast numbers of porters, paramilitary assistance from the Netherlands authorities, and a staff of some seven scientists as well as assistants. Altogether a notable contribution was made to recording the fauna of the area. Several of the more important papers were published as "Results of the Archbold Expeditions," Bull. Amer. Mus. Nat. Hist., and Amer. Mus. Novitates, various dates, primarily Some three hundred and eighty forms of birds were observed and noted on that expedition. In the 1954 trip, I was concerned with observations on species of birds of Malaysian or of Papuan affinity which might coexist in the interstitial habitats of the Moluccas. In addition, I was concerned with problems of competition between species. Unfortunately, various difficulties confront a European or American today in travelling in the Moluccas. The islands are not accessible by plane, and boat service is infrequent, the only current transport being the Indo- *Published with the aid of a National Science Foundation Publication Grant, No. GN

8 INTRODUCTION 3 nesian government-run boats called Nocemo. Occasionally an official government vessel may make a tour of some of the islands. The capital of the North Moluccas is Ternate, where my friend, Alting Abid Din Shah, Sultan of Tidore, was at that time governor. The Kapala Daerah, his official title, was kindness itself, and our trip was really made feasible by his intervention and aid. Our gratitude to him and his wife is unending. The fact that I had begun to speak the Indonesian language in 1937 on my first trip to New Guinea and its neighboring islands was of the greatest importance. Our subsequent travels would have been impossible without our knowing the local language. Food and equipment of all sorts is difficult in these outer islands, and our expedition was not aided by the unexplained disappearance of all the food we had purchased at premium prices in Java on the way out. Even rice is imported into these islands with difficulty from the Greater Sunda Islands and was always in threatened short supply. We had initially planned to concentrate on Halmahera but this was made impossible by the activities of guerilla bands of the Dar 'ul Islam movement, with the result that we spent only a few days near Djailolo, making observations already reported upon (1959C). In the end we spent five weeks on Batjan, climbing Mount Sibela (6000 ft) in the process. This proved to be a difficult feat as even the inhabitants of Labuha, Batjan's capital, were unable to direct us to a route by which the only real mountain of the island could be ascended. A British botanist named Allston had recently preceded us and had been in the end foiled by a welter of misdirections. Our efforts were successful only when we abandoned our first route and proceeded to the bay of Waiaua from which deer-hunter trails climb up to the eastern hogback which forms one of the spurs of Sibela. Our work in the Moluccas, which had been sanctioned by the Museum Zoologicum Bogoriense, was greatly aided by one of their late mantris (assistants), Saan, who proved a loyal and devoted companion. Our final month's work was spent on Misool, an island then in Dutch hands, to visit which we had to proceed via Makassar as a port of exit, to Dilli in Portuguese Timor and there catch the monthly KPM mail and cargo boat running from Singapore to New Guinea. In Sorong, which I had last visited in 1938, we were treated with the greatest consideration by the government officials from the Assistant Resident Mr. van Bodegom down, as well as by my old friend, Ong Tjoean, one of the chief Chinese merchants of the town. Here I was reunited with Jusup Khakiaj, an assistant who had worked for me and for the Academy of Natural Sciences from 1937 to 1939 and who had survived the last war totally unchanged except for gray hair. Jusup continued to work for me into 1955 collecting on Misool, Waigeu, the Ajoe Islands and Kofiau (1959B), and again in 1960 on our return. The month's work on Misool in 1954 was made notable by the discovery of four species of megapodes on the same small island, a fact of which I had not been aware during an earlier visit in 1937, nor had other naturalists ever come across these birds. In 1960 my wife, three children and I returned to New Guinea for yet another session of field work in the Hollandia area. This visit was made possible by the generous cooperation of Dr. Victor de Bruijn, the Advisor to the Governor on tribal and Papuan affairs, and also with the aid of the Christian and Missionary Alliance aim the Missionary Aviation Fellowship. The Netherlands government officials from the Governor, Dr. P. J. Platteel, on down, were all cordiality itself and our summer was made tolerable by the kindnesses shown to our children in the small Government Hotel at Dock V.

9 4 SYSTEMATIC AND ECOLOGICAL STUDY OF NEW GUINEA BIRDS My first hope had been to continue studies of the megapodes, and we commenced operations by a field trip to the Tami River and later to Holtekong, both sites in the eastern reaches of Hollandia Bay, a vast complex of sub-bays, river mouths, tidal mangrove and beaches. The lateness of the season and the failure of my tape recording equipment as well as camera difficulties seriously impaired these observations. The lowland brush turkey, Talegalla jobiensis, in this area is heavily preyed upon by Varanus lizards who perhaps follow up human trails as well as locating the nest mounds on their own. Eggs that I attempted to keep under observation were stolen by "soa-soa," as these reptiles are called in Indonesian, and all I could do was make observations on nest mounds. The incessant rain and lack of adequate water-proofing were as much a hazard to mechanical equipment in the New Guinea lowlands in 1960 as they had ever been. Our second visit was to a forestry development suggested to us by the Bureau for Economic Affairs. This was a lumber camp called Bodim, some 100 kilometers south of the north coast where the Tor River enters the sea east of Sarmi. We ascended the Tor for some 80 kilometers by double "prahu" powered by outboard motors to its junction with the Boe (Boefareh on some maps). Both rivers arise in the Gauttier and Foja Mountains. Some 20 kilometers up the Boe there is a small lumber depot and a trail leading 8 kilometers to the north into low foothills and dense stands of primary forest, dominated by dammar (A gat his) trees. Here we camped at an altitude of 300 feet above sea level for three weeks observing birds. Unfortunately the megapodes, said to be plentiful in the area, had been virtually wiped out by local hunting pressure due to the labor influx for the lumber work. Our final visit was to the Ilaga valley, an extension of the east-west mountain valley system cradled in the Nassau range, just ten miles east northeast of the Carstenz peaks. The Ilaga river flows through this valley at an altitude of approximately 7000 feet above sea level, while cultivation of sweet potatoes, sugar cane, taro, potatoes, tomatoes and edible grasses extends up along the slopes to approximately 7700 feet, making this one of the highest cultivated areas in New Guinea. The population consists of about 6000 Western Danis and 2000 Uhundunis, the latter occupying villages on the southern slopes of the valley. The first outside contact with these people had been by missionaries in 1954, but it was not until 1956 that an invitation had come from the Danis for missionaries to enter the valley. The actual settlement by two missionary families, the Gordon Larsons and the Ellenbergers, had come two years later. Our stay in the valley would have been impossible without the generous welcome of these families and the two nurses, Miss Charlotte Schon and Miss Elizabeth Little, who had recently joined them. On the flight by Cessna monoplane up to the Ilaga, piloted by Miss Elizabeth Green, the remarkable Missionary Aviation Fellowship pilot, we were suddenly offloaded due to a shutting down of the clouds in the mountains at a small strip, Bokindini, at 4500 feet altitude, where we were graciously put up for two nights by Mr. and Mrs. Garnet Erickson of the Australian Evangelical Mission until Miss Green could fly in again and take us on to the Ilaga. To all these kindly friends our thanks are due, and we often wonder, now that political events have changed the future of the Papuans so drastically, what their future will be in these strange mountains. During these travels we managed to make notes, observe and collect some three hundred and eighty-two forms of birds. Short publications on some of these species have already appeared in Postilla of the Peabody Museum, 1957 and 1959, as well as in the Proceedings of the XIIth Internat. Ornith. Congress, Helsinki, 1958, published in In addition, I have quoted from an

10 HABITATS 5 unpublished note by Philip Temple of the New Zealand New Guinea Expedition of On the conclusion of our trip, Jusup Khakiaj stayed on in the New Guinea highlands in the Baliem Valley with members of the Harvard Peabody Museum anthropology expedition. His collection was made sporadically, during his spare time, but enabled us to secure a number of new records and additional species in the period up until early 1962, when the expedition was concluded. Subsequently Miss Denise O'Brien, a Yale graduate student in Anthropology, obtained some birds for us in the Swart Valley in HABITATS The lowland habitats of New Guinea where our work was undertaken consist of the dense closed forest on Misool, and similar areas along the Tami River in eastern Geelvink Bay. A good description of the beach, secondary and lowland forest associations is in Gibbs (1917) as well as the summary by Archbold, Rand and Brass (1942), and Lam (1945). Coastal forest consists of stands of mangrove, interspersed with casuarina along the beaches, and a wealth of Pandanus species Inland on marshy, wet ground. Here and there, especially near Hollandia and round Lake Sentani, but also along the Tor river banks in the ox-bow bends, there are stands of grassland, Imperata or "lalang" and Saccharum as well as lesser sedges, grasses and ground orchids, Spathoglottis. Inland along the meandering rivers and back from them as far inland as Bodim, we found ourselves essentially in climax evergreen forest. Along the banks of the Tami and Tor rivers, swampy species of trees occurred, breadfruits, Artocarpus, pandanus, Hibiscus, occasional pockets of enclosed marsh with sago palm, Metroxylon. The river edges also included occasional Barringtonia, Timonius, and Syzygium, which, with small Ficus species was alive with flocks of parrots and lories in June and more particularly in July. The high forest at Bodim was dominated by Agathis, Polygonum, Intsia, Ficus of many species and Pittosporum. Lower substage trees included Eugenia species, many palms including Licuala, (on whose leaves I found a nest of Ptilinopus) Borassus, Cyrtostachys and of course the climbing Calamus (associated with a nest of Lonchura) and Korthalsia. Here and there on the ridges above Bodim an occasional oak tree occurred, but we did not work in the mid-stage forests from 1500 feet up. The Baliem valley and the Lower Ilaga valley are almost totally deforested. Here fields and grassland are interspersed with small areas of cut-over scrub. Casuarina trees dominate the landscape often near villages, and the low shrubbery consists of bracken, Pteridium, various rhododendrons, Grevilea, Euphorbia, Vaccinium, Rubus, ground orchids, grasses, Saccharum, Pennisetum, Phragmites, Imperata, and Sorghum. The cultivated plants in the Ilaga were sweet potato, pandanus, "sirih," taro, sugar cane, and recently introduced tomatoes and irish potatoes. Gourds also are grown, Lagenaria, to be dried and used as penis sheaths. In the upper Ilaga valley we penetrated into dense beech forest, Nothofagus, full of tree ferns, mosses and epiphytes, including more Calamus and climbing bamboos. At the higher levels above 9000 feet, we occasionally broke into open marshy flat areas of moorland, the floor of peat containing tussocks of sedge, Gahnia, and peat-stained streamlets and pools. Here there were edges of bracken fern, and low vacciniums and rhododendrons. The open areas of forest near these swampy patches were dominated by Podocarpus and occasional Libocedrus.

11 6 SYSTEMATIC AND ECOLOGICAL STUDY OF NEW GUINEA BIRDS Above 9500 feet lies the heath and moor zone with low alpines, peatmosses and a rich variety of epiphytes. This alpine zone forms an undulating plateau, uninhabited, but crisscrossed by overland trails running north and south to connect Uhunduni or Dani settlements, and east and west from the grand valley of the Baliem in the east, west to the feeder valleys of the Wissel Lakes. ALTITUDINAL SHIFTS OF RANGE AND ECOLOGICAL OVERLAP AT FEEDING TREES In mountainous tropical environments, related bird species are isolated from each other ecologically in a variety of interesting and effective ways. One of these is the rather strict stratification that occurs by altitude. Many authors have commented on the zonation of species in this manner, and Rand (1942) drew attention to the major or minor physical variations in size between populations of the same species or subspecies which could be observed while travelling upwards into the montane zones of New Guinea. Such physical differences could only mark the occurrence of genetic isolate populations in which any such change, or Bergmann-like effect, would be preserved. Everything that we have ever learned of ranges of birds in New Guinea tends to reinforce this general observation. Occasionally, however, observations can be made of bird species wandering or temporarily shifting range. Some recent publications such as those of Slud (1960) have evinced cases of this kind in the New World tropics. Frith (1957) has published on feeding movements of pigeons in Australia, as has Smythies (1960) for Borneo. Chapin (1954) has mentioned altitudinal feeding movements of starlings, Onychognathus, on Mount Ruwenzori, as has Moreau to me (in litt.) on Mount Kenya, Onychognathus and Poeoptera. Thus it was extremely interesting for me to come across cases where known highland species were entirely out of their normal range, competing for food with closely related, normally ecologically separated species. The first case occurred on Batjan Island at Gandasuli, altitude 350 feet above sea level, during the period September 26-October 1. The local albizzia trees (Albizzia molucca) were in flower. Although Myzomela obscura simplex, the common lowland small honeyeater was normally in evidence in these trees, we also collected the similar-sized Myzomela dibapha batjanensis in the same single tree during the six-day period. Otherwise the highland honeyeater, dibapha, was found only on the Sibela ridge at 5000 feet and above. Zosterops atriceps and Dicaeum erythrothorax were also feeding in the same tree, but in greatly reduced numbers. They were quiet and unobtrusive, Dicaeum making its ticking calls only as it left the tree in flight. However, the two Myzomela species were very noisy, making loud buzzing notes and calling incessantly, as I have described in an earlier paper (1959A). Whereas I pointed out in that paper that obscura was widely scattered at a low density over the lowlands of Batjan, and appeared to be highly aggressive when in contact with similar-sized sunbird species, here was an occasion where aggression was temporarily suspended for securing food. The result seemed to be achieved in a series of darts and dashes through the feeding tree, accompanied by noisy buzzings. In contrast to the silent yet intent behavior of feeding species of honey creepers, Diglossa and Conirostrum. in competition with each other for food in the Ecuadorian Andes, described by Moynihan (1963), these species were very noisy. Constant movement and displacement, one individual by another, was the pattern, accompanied by advertising or warning notes. Moynihan notices the absence of overt short-range disputes between interspecific contacts. In contrast, I noted a high degree of activity with

12 RANGE AND BREEDING SEASON 7 frequent short chasings indicating overt aggression. This behavior parallels that observed in captive waterfowl (unpublished). A second case occurred at Bodim on the Boe River where a cluster of three myrtaceous flowering trees attracted honeyeaters and lorikeets during the period July 29-August 2. Here Myzomela eques primitiva was the common lowland honeyeater, but we managed also to secure a single Myzomela nigrita meyeri, not common at all at this low altitude. Here also we found three species of lorikeets, Charmosyna pulchella rothschildi, Charmosyna r. rubronotata and Charmosyna placentis ornata, feeding in the same tree. Again, as on Batjan, we were attracted by the high intensity of buzzing calls and parakeet shrieks made by the birds as they fed on the nectar of the tree flowers. It was not continuous. The noise rose and fell as flocks of lorikeets left the tree or arrived, and it broke out at intervals as birds crowded together. There was considerable competition for perch space by from two to seven lorikeets at a time crowding onto a favorable feeding post. The honeyeaters, Myzomela, tended to dart about with buzzings and squeakings in the tree as much as they had been doing on Batjan. A few larger nectar-eating parrots also fed in the trees, but were not identified as well as two larger honeyeaters, Xanthotis chrysotis and polygramma. Competition within a micro-habitat seemed to be occurring between three species of lorikeets, one of which was collected several hundred miles west of its known range, and another of which, Charmosyna pulchella, is normally a more montane species (see later discussion, p ). A third case is that of the two Alpine Lorikeets, Neopsittacus musschenbroekii and AT. pullicauda, found in the same feeding trees in the Ilaga. Presumably these species have a wide overlap in altitudinal range between meters (7250^ 9500 feet) where they coexist. The smaller sibling occupies the higher range. EXTENSIONS OF RANGE An additional subject concerns extensions of range. It seems obvious that peaceful conditions in the Baliem Valley and north at lower levels towards Archbold Lake have created new environments. Increased agriculture has resulted in new habitats for water birds and birds of open country, not previously collected above a few thousand feet. As in other tropical countries such as Ceylon (1946), changing habitats, the creation of plantations, and the destruction of forests allow these species to exploit new altitudinal ranges. The following species are here recorded from the Baliem valley or other highlands for the first time: Pelecanus conspicillatus Hydranassa picata Egretta alba modesta Egretta intermedia plumifera Nycticorax caledonicus hilli Aplonis cantoroides Haliastur indus girrenera (7500 ft) Monarcha rubiensis Columba vitiensis halmaheira Opopsitta d. diopthalma Cacatua galerita triton (Jalimo, Baliem?) Halcyon s. sancta (up to 7500 ft) Accipiter cirrhocephalus papuanus Philemon novaeguineae jobiensis Porphyrio poliocephalus melanotus BREEDING SEASON The following list includes all those bird species for which we obtained some evidence of a breeding cycle from gonadal activity of specimens dissected. Where a question mark follows the record it refers to the presumption of breeding from the fact that only males possessed enlarged gonads. In some cases only young

13 8 SYSTEMATIC AND ECOLOGICAL STUDY OF NEW GUINEA BIRDS juvenile or nestling birds were collected. In other cases the only record is the notation made by my assistant, Khakiaj, on a specimen label, where I, myself, was not present at the time of dissection. The latter records refer essentially to March or April birds from the Baliem. In a number of cases males only showed enlarged testes and from the onset of moult we assumed that gonadal regression had set in and that actual nesting had just concluded. Here I have made an entry for the month to indicate some activity and an entry followed by a question mark to indicate the supposition that breeding had occuired in the preceding month. The data following show conclusively that in the central areas of the island where we collected, Hollandia, Bodim and the mountains, the breeding season is in the southeast trade wind season, May to October, which is on average, the dry season with only thirty-six per cent of the annual rainfall. A few species give evidence of having two breeding seasons (two ducks, a megapode, two warblers, a thickhead, honeyeater, flowerpecker and white-eye). The evidence is inconclusive, but at least suggestive. In contrast to this the western island breeding season of extreme western New Guinea, Sele Strait, Misool, Waigeu, Kofiao, etc., and one or two references to adjacent Batjan Island in the Moluccas, is overwhelmingly in the months October to January, namely the interim period between the end of the southeast trade wind season (which is rougher with more storms in the western islands like Misool) and the beginning of the rather uncertain northwest monsoon which traditionally begins in November, but often in the western area, does not become really rainy until February. Thus it would appear that most species have young well on the wing and are entering the completion of the moult cycle by the onset of the rainy season. The only exception appears to be the starlings in which breeding activity in the western areas apparently occurs as early as August (Aplonis mysolensis). Gyldenstolpe (1955B) records breeding starlings collected by Bergman in August and September. One of the rails also, Gymnocrex plumbeiventris, appears to breed as early as September. Otherwise the trend appears unmistakable. x = Central New Guinea, i.e. Hollandia, Tami, Bodim, Nassau, Baliem jf: = Western New Guinea, Arar, Misool, Waigeu o = Batjan Podiceps ruficollis tricolor Dupetorflavicollisgouldi Threskiornis moluccus Anas gibberifrons gracilis Anas superciliosa pelewensis Salvadorina waigeuensis Megapodius freycinet freycinet Megapodius freycinet affinis Megapodius wallacei Talegalla cuvieri Talegalla j. jobiensis Coturnix chinensis novaeguineae Rallus pectoralis captus J F M A M J J A S 0 N D x? x? X? X? X X X X? X? X? X? X? X X? X X I J I F I M I A I M I J I J. A I S I Q I N -D #?

14 x = Central New Guinea, i.e. Hollandia, Tami, Bodim, Nassau, Baliem = Western New Guinea, Arar, Misool, Waigeu o = Batj an RANGE AND BREEDING SEASON 9 JlFlMlA M JlJ A S Q N D Porzana tabuensis edwardi x? x? Poliolimnas cinereus minimus x? Gymnocrex p. plumbeiventris # Amaurornis olivaceus frankii o Porphyrio poliocephalus melanotus x? Irediparra gallinacea novae-guineae x? x? Sterna s. sumatrana x Ptilinopus s. superbus x? Ptilinopus pulchellus decorus x x Ptilinopus coronulatus trigeminus #? # Ptilinopus coronulatus quadrigeminus x? x? x? x Ptilinopus iozonus humeralis Ptilinopus iozonus jobiensis x Ptilinopus rivoli bellus x Ptilinopus viridis salvadorii x Ptilinopus aurantiifrons x Megaloprepia magnifica septentrionalis x? x? x Ducula m. myristicivora # Ducula bicolor o o Gymnophaps a. albertisii x Reinwardtoena reinwardtsi griseotincta x x Henicophaps a. albifrons x Gallicolumba r. rufigula # # Gallicolumba r. septentrionalis x Goura victoria beccarii x? Chalcopsitta a. atra # Trichoglossus h. haematodus # Lorius I. lory # Lorius I. viridicrissalis x Charmosyna papou goliathina x Micropsitta keiensis chloroxantha #? # # Probosciger aterrimus goliath Alisterus chloropterus moszkowskii x x? Psittacella lorentzi x Cacomantis pyrrophanus excitus x Cacomantis castaneiventris arfakianus x #? Podargus papuensis x #? #? Podargus ocellatus # Caprimulgus macrurus schillmolleri o Caprimulgus macrurus yorki x? Collocalia vanikorensis grand x? Hemiprocne mystacea confirmata o Alcyone azurea lessonii #? Merops philippinus salvadorii x Aceros plicatus ruficollis Pitta erythrogaster macklotii $ Pitta sordida novae-guineae Cracticus c. cassicus x? ^ J F M A M J J A S 0 N D

15 lo SYSTEMATIC AND ECOLOGICAL STUDY OF NEW GUINEA BIRDS x = Central New Guinea, i.e. Hollandia, Tami, Bodim, Nassau, Baliem # = Western New Guinea, Arar, Misool, Waigeu o = Batjan J F M A M J J A S O N D Cradieus q. quoyi #? #? # Oriolus szalayi x x # Dicrurus hottentotus carbonarius # Artamus maximus x x Aplonis cantoroides x x Aplonis m. mysolensis #? Aplonis m. metallica x x # Mino dumontii violaceus x x Corvus tristis x Corvus o. orru $ Loria I. loriae x? x? Macgregoria pulchra carolinae x? x Manucodia jobiensis rubiensis x Ptiloris m. magnificus x Seleucides melanoleuca auripennis x Drepanornis bruijnii x Cicinnurus regius similis x Diphyllodes magnificus chrysopterus x x Paradisea m. minor x x Paradisea m. pulchra # Archboldia p. papuensis x? Chlamydera cerviniventris x Coracina papuensis melanolora o Lalage a. atrovirens Crateroscelis robusta sanfordi x Pomatostomus i. isidori x x x # Saxicola caprata belensis x Malurus alboscapulatus aida x x x Todopsis c. cyanocephala # # # Clytomyias insignis oorti x? x Megalurus timoriensis macrurus x? x Cisticola exilis diminuta x? Sericornis s. spilodera x? Sericornis n. nouhuysi x? Sericornis perspicillatus x x Sericornis papuensis burger si x? x? Gerygone cinerea x Gerygone c. chrysogaster x? x? Gerygone chloronota cinereiceps # Gerygone magnirostris affinis x Gerygone ruficollis insperata x? x x Peltops montanus x? Rhipidura t. threnothorax x x # Rhipidura r. rufidorsa # # Rhipidura a. albolimbata x x Rhipidura rufiventris gularis x # Rhipidura leucophrys melaleuca x ' ' o Monarcha alecto chalybeocephalus J F I M A M J J A S 0 N D

16 x = Central New Guinea, i.e. Hollandia, Tami, Bodim, Nassau, Baliem # = Western New Guinea, Arar, Misool, Waigeu o = Batjan RANGE AND BREEDING SEASON 11 J F M[ A M -J J A S O N D Monarcha guttula x x # # Monarcha chrysomela melanonotus $ jf Monarcha t. telescopthalmus Monarcha insularis x x x Machaerirhynchus n. saturatus x Microeca flavovirescens cuicui x Microeca papuana x Poecilodryas albonotata griseiventris x x Peneothello sigillatus quadrimaculatus x Peneothello cyanus atricapilla x x Pachycephala pectoralis balim x? x? Pachycephala schlegelii viridipectus x x Pachycephala lorentzi x Myiolestes megarhynchus misoliensis # Myiolestes megarhynchus hybridus x Pitohui kirhocephalus uropygialis $ Pitohui kirhocephalus meyeri x x Pitohui ferrugineus heurni x Nectarinea jugularis frenata $ if x Timeliopsis g. griseigula x x Glycichaera f. fallax x x Myzomela e. eques Myzomela e. primitiva x? x Myzomela r. rosenbergii x? x? x x Toxorhamphus n. novae-guineae x? x $ Toxorhamphus i. iliolophus x Melipotes fumigatus goliathi x? x Melidectes nouhuysi x? x Xanthotis chrysotis meyeri x? x Xanthotis polygramma kuehni # Meliphaga aruensis sharpei x x # Meliphaga a. analoga # x x # Ptiloprora per striata lorentzi x? x Pycnopygius cinereus marmoratus x? Philemon meyeri x? x Philemon n. novaeguineae # Philemon n. jobiensis x x Dicaeum pectorale ff Dicaeum geelvinkianum centrale x? x Melanocharis n. nigra Melanocharis n. unicolor x? x x Paramythia montium alpinum x Zoster ops f. fuscicapilla x? x? x? Lonchura tristissima calaminoros x? x Lonchura spectabilis mayri x? x? x Lonchura teerinki x? Lonchura monticola x? x J F M A M J J A S 0 N D

17 LIST OF THE SPECIES CASUARIIDAE, CASSOWARIES Casuarius unappendicwlatus philipi Rothschild A fine adult of this species, the face and upper throat very blue, and with a high casque, was watched in damar forest at Bodim on July 28. The bird came quite silently out onto a trail, its slightly rocking gait as the head remains still and the neck carries back during each step, giving it an extremely dignified air. After perhaps three minutes pause on the trail, it apparently heard something and bounded off into heavy rattan-shrouded forest, as silently as a deer, only a faint crack or two marking its passing. A chick perhaps eighteen inches high was purchased from Papuan villagers in July at Holtekong. The bird is beginning to get black feathers interspersed among the brown now in July a year later. Its color pattern corresponds exactly to that of a juvenile taken by me in West New Guinea in In the Ilaga, the Western Dani and Uhunduni people believe that the few heavy wing quills of the mountain cassowary are breasts and that these birds suckle their young. They believe that there are no males among cassowaries, the females are parthenogenetic and produce eggs by copulating with a stick. PODICIPEDIDAE, GREBES Podiceps ruficollis tricolor (Gray) A series of four specimens from Lake Sentani and the Baliem Valley seem identical with a female taken in northern Celebes in October in breeding condition. Two October males are similarly listed as having enlarged gonads. A single adult was solicitously protecting a brood of well-grown young on the lagoon lake at Mosso, July 2. Wing $ 97, 97.5, 98, $ 96; soft parts: iris red, bill and feet blackish. Podiceps novaehollandiae novaehollandiae (Stephens) Rand (1942, p. 426) points out quite rightly that the Australian dabchick should be kept specifically separate from the Papuan ruficollis as it apparently overlaps. A series of four birds from Wamena and the Baliem River includes a juvenile specimen taken in February just moulting into the first winter or first basic plumage. The stripes are broken into isolated blackish feathers on the throat and sides of the neck. The crown is dull brown and the sides of the lower neck are rufous. Wing # 101, 107, 9 100, juv. $ 100.5; weight $ 200, juv. $ 207 grams. Mayr in Hartert (1930, p. 112) records a male of the Little dabchick (tricolor) from Lake Sentani, taken in September as weighing 165 g. PELICANIDAE, PELICANS Pelecanus conspicillatus Temminck An Australian pelican was killed on the Ilaga River at an altitude of 6800 feet in early August, 1959, as it perched in a very large casuarina tree. The photograph of the bird taken by Mr. Larson was unequivocal. This is a first record for the highlands. PHALACROCORACIDAE, CORMORANTS, ANHINGAS Phalacrocorax melanoleucos melanoleucos (Vieillot) Two specimens from Poee on Lake Sentani were taken in September and October. 12

18 LIST OF THE SPECIES 13 A single bird was seen on the small lagoon lake at Mosso. The little pied cormorant also occurs in the Baliem. Wing $ 239, o 224. Grand Valley Dani name, "waluem." Anhinga rufa papua Rand A pair in non-breeding condition were taken at Poee in October. The bill of these birds was given on the label as "orange," the legs "gray." Wing $ 340, $ 330. This species is shy, and specimens are hard to obtain. It is undoubtedly eagerly sought for as food by the fishing people of the lake. ARDEIDAE, HERONS Notophoyx novaehollandiae austera, subsp. nov. TYPE: $ ad. YPM no , collected March 2, 1961, at Wamena, 5000 feet, Baliem Valley, Netherlands New Guinea, by Jusup Khakiaj. DIAGNOSIS: from typical novaehollandiae of Australia this form differs by being noticeably darker in tone on the crown of the head, in the coloration of the neck plumes especially on the lower neck and upper breast, on the flanks and thighs, and on the back and rump. From nana Amadon (1942A) of New Caledonia this form differs by being larger, as in novaehollandiae, and also of course darker. MEASUREMENTS: wing $ 316 (worn), 324, 327, 328, 335, 340 (type), $ 320, 328, o 300, 318 (worn), 327, 332, 334. REMARKS: birds collected in February and March tended to have worn primaries, although this varies, as the type is in good condition. This series, all from the Baliem Valley agrees with birds taken by the Archbold Expedition of 1938 in the American Museum of Natural History collection in being noticeably dark. Birds from Papua and the southeast coast are also darker than Australian birds although less noticeably so. The large series taken by Khakiaj, however, demonstrates clearly that New Guinea birds are darker than those of Australia. RANGE: New Guinea in the central mountains (Baliem Valley), the south and southeast. GENERIC POSITION: Bock (1956) has united Notophoyx with Ardea as he calls it a dwarf Ardea resembling especially Ardea pacifica in color patterns and plumes. While sympathetic to his attempt to point out the relationships of the heron family, I feel that novaehollandiae is very dwarfish indeed for an Ardea and that proportionally also its legs are rather short for that genus. It seems wiser to leave the species as monotypic for the time being in the genus Notophoyx until some clearer anatomical evidence is forthcoming that it is as close to Ardea as assumed. Grand Valley Dani name, "mugugo." Hydranassa picata (Gould) Two specimens, a male and female, were collected at Wamena in October and November, the first record for this lowland species in the Baliem Valley. Wing 218, $ 220; oilmen $ 59, $ 65. Jusup, my assistant, has marked on the label that the local Grand Valley Dani name for this species is "wamena," a curious coincidence as the nearby village has a similar name. These small plumed herons might well be combined in Hydranassa as Bock (1956) has suggested. Egretta alba modesta (Gray) A pair were secured at Poee and a male at Wamena. This was the commonest species seen here and there in the lowlands wading in the large river shallows or on sand banks, or near the lake shore. The Wamena bird is a new record for the Baliem Valley. Wing $ 353, 374, $ 371. Iris orange yellow, bill yellow. Grand Valley Dani name, "gut."

19 !4 SYSTEMATIC AND ECOLOGICAL STUDY OF NEW GUINEA BIRDS Egretta intermedia plumifera (Gould) Taken at Wamena this is a new record for the species at this altitude in New Guinea. March birds have long plumes. Wing $ 291, 2 296, o 302, 303. Iris orange yellow, bill yellow. Egretta garzetta nigripes (Temminck) A male from Poee has a wing measurement of 269. This species was also seen occasionally along the rivers. Nycticorax caledonicus hilli Mathews Adults were taken at Poee and in the Baliem Valley, a new altitude record for the species in New Guinea. A juvenile example was secured November 16 on Misool, evidently a bird of the preceding breeding season. Wing $ 296, $ 296, o 277, 295; soft parts: juvenile iris, yellow, ocular skin yellowish green, bill yellowish green, oilmen and tomia black, feet yellowish green. Zonerodius heliosylus (Lesson) A female in non-breeding condition was shot on a small tree overlooking a small swampy clearing at Bodim near the Tor River on July 27. I have seen this rare forest bittern only twice in New Guinea, both times in trees overlooking very small streams, or forest pools. Soft parts: iris yellow, facial skin green, bill upper mandible black, lower mandible grayish flesh, feet blackish green, tarsus distally black, basally yellowish green. Dupetor flavicollis gouldi (Bonaparte) An adult, an immature and a juvenile just out of the nest were taken. The immature bird collected at Poee in October differs from the adult in being rather rusty brown on the upper surface, the wings being a paler grayish bluish rather than the blackish tone of the wings of the adult. The throat and neck feathers are heavily washed with dark brown with rufous and brownish rather than blue-black spotting. There is an indistinct chestnut brownish sub-ocular streak extending back to the ear coverts. The under surface is dull mauvish gray-brown, the ends of the feathers of the abdomen being dark isabelline. The thigh feathers are cinammon buff. The nestling or post-nestling is very blackish on the upper surface and below brownish black, the feathers of the chin, throat and abdomen being finely edged with pale brown. Soft parts (including the nestling): iris yellow, bill black, lower mandible greenishbrown, feet dark brown. Wing $ 209, $ (im.) 215. THRESKIORNITHIDAE, IBISES Threskiornis moluccus moluccus (Cuvier) A male was shot while incubating two white eggs in the crown of a coconut palm at Arar on Sele Strait on December 1. The bird had enlarged gonads. Unfortunately I did not realize that this bird was actually incubating when I destroyed it, as the nest, while distinguishable, was still of flimsy construction and appeared to me to be an incipient rather than a real nest. No mate was seen during our stay of nearly a week at the nearby village where we camped. Soft parts: iris brown, bill black, facial skin and nape patches rosy pink, feet mauve, joints black, tarsi pink. Wing 375, bill (culmen) 171. ANATIDAE, DUCKS Dendrocygna guttata Schlegel A female taken at Poee in October was the only bird encountered. Wing 210. Tadorna radjah radjah (Lesson) Radjah Shelducks were present on the Tor River and on the Tami, usually seen in

20 LIST OF THE SPECIES 15 small parties of five or six, perhaps adults and well-grown young of the year. A female was collected at Kabare on Waigeu in October; iris brown, bill and feet creamy white. Wing 287. Anas gibberifrons gracilis Buller Apparently common along the Baliem River near Wamena and the adjacent streams and lakes, the Gray Teal is apparently a breeding bird in the central highlands, as three males and one female taken in October are indicated as having enlarged gonads. Two of the males with enlarged gonads show heavy wear on the primaries, indicating that they are perhaps about to go into moult, the eclipse or pre-basic plumage so far as is known at present indistinguishable from the breeding plumage. An immature female was taken in October in the prebasic or juvenal plumage, marked by dappled small spots uniformly on the underparts. Wing $ 200, 202, 207; (im.) 193, 196 (2), o 197, 201, 203. Grand Valley Dani name, "walo-walo." Anas superciliosa pelewensis Hartlaub and Finsch Twelve males, five females and a number of birds of undetermined sex were taken on the Baliem River. February, March, August, September and October birds are listed as having enlarged gonads, an indication that nesting was in prospect. February specimens are in moult. In addition three September and October birds show heavy wear on the primaries and a third October specimen appears to have freshly moulted primaries which have shorter projecting tips beyond the ends of the secondaries than in the rest of the series, Wing a 231 (2), 233, 238, 239, 241, 244 (2), 245, 2 220, 222, o 228, 227, 231, 233, 237, 238, 243. Four immature birds were taken, all characterized by short wing measurements (under 220) and by having very broad buffy tips to the feathers of the abdomen. This would appear to be part of a prebasic or juvenal plumage and rather easily recognizable in museum skins, less so in live birds on the wing, although to the experienced eye such young birds often have an appearance of youth in their demeanor and flight. These young birds were taken in March and September. One young bird is extremely pale and washed out in appearance and heavily worn. A few darker feathers of the basic plumage are appearing spottily on the under surface from breast to vent. Another small February female is also very worn and the undertail coverts and abdominal feathers are so fluffy as to indicate the downy quality of the juvenal. This bird is in juvenal plumage, and here again the breast and abdominal feathers are distinctive. The three types may be diagrammed as follows then in Anas super ciliosa pelewensis (breast feathers only): juvenal basic or first non-nuptial alternate or nuptial Figure 2. Contour feathers from three specimens of the Australian Gray Duck, Anas superciliosa pelewensis, showing juvenal, immature of the first year, and adult plumage.

21 16 SYSTEMATIC AND ECOLOGICAL STUDY OF NEW GUINEA BIRDS An alternative feather pattern with a central V-shaped patch of buff is sketched as well. This appears to be a first basic or non-nuptial plumage bird of unknown sex. It corresponds to an eclipse bird of Kortright's description of the Black Duck (1942), but our North American Black Duck possesses this in the alternate or nuptial plumage, and the eclipse or basic plumage is plain black with the buffy fringing edge as in the adult alternate plumage of pelewensis. This subspecies is much less broadly streaked and edged with buff than typical rogersi of Australia. Perhaps in pelewensis there is a tendency for the plumage to become more immaculate and less buffy with age and succeeding prealternate moults. In connection with Amadon's review of this species (1942B) it is of interest to record that of two males from the Palau Archipelago, type locality of pelewensis, one has a weight of grams. Unfortunately the wings of both birds which measure 227, 228, are badly worn (January, April). Salvadorina waigeuensis Rothschild and Hartert A female with ovaries granular in heavy wing moult was shot on the Ilaga River on September 9 and two ducklings were taken, both of which died after one or two days of efforts to keep them alive. The ducklings are dark, mousey-black color with white lores, superocular streaks, subauricular streaks, and white underparts. There are two pairs of white body spots and a white wing edging. Soft parts of the ducklings are: iris brown, bill upper mandible blackish, nail and lower mandible pinkish, feet olive brown, splotched with dull yellow. Weight 30 g. Our adult female had the iris brown, bill upper mandible brownish orange, center of the upper mandible brown, feet splotched dull brown and yellow. Weight 520 g. The rest of our series of these birds was taken at Wamena and on the Baliem. Males are listed as in breeding condition in February and March, and contra Delacour (1956), all have white-barred tails. One immature bird in prebasic or juvenal plumage shows traces of down feathers still on the thighs and lower abdominal flanks. Even in this specimen the tail feathers are barred and indeed the upper plumage except for being somewhat "foxed" or more brownish blackish in tone is indistinguishable from the adult. However, round the face and neck prebasic warm brown feathers are being replaced by a prebasic moult into the first basic or first non-nuptial plumage of the adult. Until far more is known of these long-tailed birds it would seem unrealistic to submerge the genus Salvadorina. Grand Valley Dani name, "kunuru." Western Dani, "uuni;" Uhunduni, "lakkum." Ay thy a australis papuana, subsp. no v. TYPE: $ ad. (YPM No ), collected February 20, 1961 at Wamena, Baliem Valley, Netherlands New Guinea by Jusup Khakiaj. DIAGNOSIS: from Australian australis, these birds differ by being darker, more chocolate chestnut on the head and neck, and much darker brown on the lower abdomen, vent, thighs and flanks. In size these populations are virtually the same, though there is an indication of slightly larger measurements in australis australis. From extima (specimens from Tana, New Hebrides and New Caledonia), this form differs by being larger and much darker, the males being pronouncedly sexually dimorphic, in contrast to the small tropical island population in which the male plumage has become dulled and diminished, more like an eclipse plumage as in the tropical gray teals, gibberifrons, or the small mallard of Hawaii, the "koloa," wyvilliana. MEASUREMENTS: wing $ (214.8), 2 209, 212; tail $ (59.1), $ 55 (worn), 60; culmen $ (45.5), $ Measurements of type: wing 212.5, tail 59, culmen 44. Australian specimens in the American Museum and the Yale collections measure: wing # (218.7), $ ; tail (64.2), $ 60-65; culmen $ (47), $ Specimens of extima from Tana ($, $) wing 201, New Caledonia $ 195 (worn). Mayr's birds from the Banks Is. (1940, p. 7) measured: $ $ , $ REMARKS: AS Rand (1942, p. 431) noted, this breeding population of the New Guinea highlands is darker, more blackish, than the Australian one. In addition, it is slightly

22 LIST OF THE SPECIES 17 smaller, and in breeding plumage, if anything, richer, more pleasing in chestnut tone, approaching Madagascan innotata, but much more blackish than typical small A. nyroca of the palearctic region. Aythya australis ledeboeri (Bartels and Franck) (1938) is identical and synonymous with typical australis in size and color, and represents a small straggling population of the Australian form of the White-eyed Pochard which extends into eastern Java, the lesser Sunda Islands and occasionally to Celebes, probably reinforced from time to time by vagrant Australian birds. The Netherlands New Guinea highlands, however, seem to be the home of a small resident population extending from Lake Habbema and the Baliem Valley west to the Wissell Lakes at altitudes of 5000 feet and above. Birds are showing very worn scapulars in February and March. Our male type lacks the white spot on the chin normally found in all males of the Ferruginous ducks. ACCIPITRIDAE, HAWKS Aviceda subscristata waigeuensis Mayr Two adult males and an immature male of this subspecies were taken at Kabare on Waigeu Island in September, The character of unbarred greater underwing coverts shows up clearly in these birds. Henicopernis longicauda minimus Junge A female (?) from Wasa, Misool, with a wing length of 331, confirms the small size of this insular race on Misool, although as Gyldenstolpe points out (1955B) Batanta Island birds are as large as those of mainland New Guinea. This specimen is below Junge's minimal measurements (1937) for males ( ), and therefore is probably also a male. Henicopernis longicauda longicauda (Garnot) A male was taken August 18, 1963, in the Swart Valley. Wing 412. Swart Dani name, "ligatlok." Haliastur indus girrenera (Vieillot) Found by us from seashore to 7500 feet in the Ilaga, a new altitude record for this species. I collected an immature male in very worn plumage commencing the moult into adult plumage atop a huge casuarina in the Ilaga, September 2, Western Dani name, "mokop." Accipiter novaehollandiae leucosomus (Sharpe) Waigeu, $(=#?) wing 210, $ wing 250. Holtekong, $ (subad.) wing 246. Baliem including Kurelu, o (= $) wing 216, 217, , 246. The Waigeu Island birds are in the pale, more grayish coloration, while the Hollandia Bay and Baliem birds are in the slatey and chestnut underparts "etorques" coloration. There appears to be no altitudinal variation in size among our specimens. Only females appear to have any pronounced barring on the tail and I am inclined to feel that this is a character which disappears with age and succeeding moults. The immature female taken at Holtekong in secondary scrub forest near cultivation had the following soft parts: iris yellow, bill black, cere orange yellow, feet orange yellow. Weight 439 grams. A fledgling was taken in the Swart Valley in September. Grand Valley Dani name, "pintev Swart Dani name, "penake." Accipiter poliocephalus Gray A single fe^nale from Misool is our only specimen of this species. Wing 213; iris brown, bill black, feet reddish. Accipiter melanochlamys (Salvadori) Two specimens, an adult female and an immature male, appear to bring to twenty-

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