Chapter I: Part 1

UNIVERSITY OF KANSAS PUBLICATIONS

MUSEUM OF NATURAL HISTORY

VOLUME 3 · 1951

EDITORS

E. RAYMOND HALL, _Chairman_
A. BYRON LEONARD
EDWARD H. TAYLOR
ROBERT W. WILSON

MUSEUM OF NATURAL HISTORY
UNIVERSITY OF KANSAS
LAWRENCE
1951

MUSEUM OF NATURAL HISTORY
UNIVERSITY OF KANSAS
LAWRENCE

PRINTED BY
FERD VOILAND, JR., STATE PRINTER
TOPEKA, KANSAS
1951

24-1811

CONTENTS

1. The Avifauna of Micronesia, Its Origin, Evolution, and Distribution.
By Rollin H. Baker. Pp. 1-359, 16 figures in text. June 12, 1951

2. A Quantitative Study of the Nocturnal Migration of Birds. By George
H. Lowery, Jr. Pp. 361-472, 47 figures in text. June 29, 1951

3. Phylogeny of the Waxwings and Allied Birds. By M. Dale Arvey. Pp.
473-530, 49 figures in text, 13 tables. October 10, 1951

4. Birds from the State of Veracruz, Mexico. By George H. Lowery, Jr.
and Walter W. Dalquest. Pp. 531-649, 7 figures in text, 2 tables.
October 10, 1951

Index, Pp. 651-681.

The Avifauna of Micronesia,
Its Origin, Evolution, and Distribution

BY

ROLLIN H. BAKER

University of Kansas Publications
Museum of Natural History

Volume 3, No. 1, pp. 1-359, 16 figures in text
June 12, 1951

University of Kansas
LAWRENCE
1951

The Avifauna of Micronesia,
Its Origin, Evolution, and Distribution

BY

ROLLIN H. BAKER

University of Kansas Publications
Museum of Natural History

Volume 3, No. 1, pp. 1-359, 16 figures in text
June 12, 1951

University of Kansas
LAWRENCE
1951

UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY

Editors: E. Raymond Hall, Chairman, A. Byron Leonard,
Edward H. Taylor, Robert W. Wilson

=Volume 3, No. 1, pp. 1-359, 16 figures in text
Published June 12, 1951=

UNIVERSITY OF KANSAS
Lawrence, Kansas

PRINTED BY
FERD VOILAND, JR., STATE PRINTER
TOPEKA, KANSAS
1951

22-8131

The Avifauna of Micronesia,
Its Origin, Evolution, and Distribution

By

ROLLIN H. BAKER

CONTENTS

PAGE

INTRODUCTION 5

DESCRIPTION OF MICRONESIA 5
Climate 8
Soils 9
Surface water 9
Vegetation 10

GAZETTEER OF THE ISLANDS OF MICRONESIA 11
Mariana Islands 11
Palau Islands 13
Caroline Islands 14
Marshall Islands 15

ORNITHOLOGICAL EXPLORATION IN MICRONESIA 16

CHECK-LIST OF THE BIRDS OF MICRONESIA 21

DISCUSSION OF THE AVIFAUNA 28
Oceanic Birds 28
Inshore Oceanic Birds 29
Offshore and Pelagic Oceanic Birds 30
Faunal Components 30
Migratory Shore Birds 32
Original Homes of the Shore Birds that Visit Micronesia 32
Routes of Migration 34
Populations of Shore Birds in Micronesia 37
Land and Fresh-Water Birds 42
Polynesian Component 44
Melanesian Component 44
Moluccan and Celebesian Components 45
Philippine Component 45
Palearctic Component 46
Speciation 48
Time of Colonization 50
Factors Causing Dispersal 52
Analysis of Speciation 55

CONSERVATION OF THE AVIFAUNA OF MICRONESIA 58

THE FUTURE OF ORNITHOLOGICAL RESEARCH IN MICRONESIA 60

METHODS AND ACKNOWLEDGMENTS 60

ACCOUNTS OF THE KINDS OF BIRDS IN MICRONESIA 63

SUMMARY AND CONCLUSIONS 340

BIBLIOGRAPHY 343

FIGURES IN TEXT

FIGURE PAGE

1. The islands of Micronesia. 6

2. The Mariana Islands. 12

3. The Palau Islands. 13

4. The Caroline Islands. 15

5. The Marshall Islands. 16

6. Routes of migration used by shore birds in the Pacific. 34

7. Avifaunal regions of the central Pacific. 43

8. Faunal areas from which Micronesian birds have been derived. 46

9. Routes of dispersal of rails in the Pacific area. 120

10. Variation in length of culmen of _Sterna s. sumatrana_. 159

11. Geographic distribution of _Thalasseus bergii_. 164

12. Geographic distribution of _Gygis alba_ in the Pacific. 177

13. Distribution and dispersal of _Ptilinopus porphyraceus_. 190

14. Distribution and dispersal of _Gallicolumba_ in the Pacific. 204

15. Distribution and dispersal of _Acrocephalus_ in the Pacific. 260

16. Distribution and dispersal of _Myzomela_ in the Pacific. 316

INTRODUCTION

Birds in Micronesia comprise the most outstanding animal life of the islands, as far as vertebrates are concerned. No fewer than 206 kinds, belonging to 37 families and 91 genera have been found there. Although this number upon first consideration may seem large, actually any large land mass in the same latitude has many more kinds of birds than does Micronesia. In this connection it is pertinent to recall that the islands of Micronesia are oceanic and have apparently been formed independently of any continental land mass. Thus, animal life found on these islands has reached them by overseas migration, either by some passive means or by individual effort. Zoogeographers have had some difficulty in explaining the presence of snails and other nonflying animals on isolated oceanic islands. Crampton, in his studies of the land snails of the genus _Partula_ at Guam and Saipan (1925:10), writes, "Despite the geological difficulties, the biological findings strongly support the view that the dominant process in this part of Oceania has been one of subsidence and of insular dissection." Although there exists today some question as to how certain forms of life have reached these remote dots of land, the ornithologist has not been much in doubt as to the actual _means_ of arrival of birds. With the exception of six kinds of birds which are definitely known to have been introduced by man, the birds have apparently reached these islands by flying there from somewhere else. The ornithologist is, therefore, concerned with learning _from where_, _by what route_, _when_, and _why_ the various species of birds came and _how_ they have become established on these islands of Micronesia. These birds exist in small populations; often less than 100 individuals of one kind may be found on a small island. How have such small numbers had the ability to survive and what environmental adaptations have occurred, are two additional questions which confront the student of Micronesian birds.

DESCRIPTION OF MICRONESIA

The vast expanse of the Pacific Ocean is dotted with numerous islands, most of which are concentrated in the central and western part and are known collectively as Oceania. Within Oceania three divisions are popularly recognized: Melanesia, Polynesia, and Micronesia. According to Krieger (1943:6), the Micronesia islands include the Mariana, Palau, Caroline, Marshall, and Gilbert islands; they may take in also the Volcano, Bonin, and Ellice islands (from the standpoint of anthropology). Zoogeographically, according to Wallace (1876), Micronesia is to be included in the Polynesian Subregion of the Australian Region. Mayr (1941a:193), on the basis of the distribution of birds, ranks Micronesia as one of the four subdivisions of the Polynesian Subregion, and includes within Micronesia the Palau, Caroline, Mariana, Marshall, and Gilbert islands. Except in the discussion of distribution, this report does not treat of the avifauna of the Gilbert Islands, which straddle the equator south of the Marshall Islands. This report is concerned only with the birds in the Mariana, Palau, Caroline, and Marshall islands formerly mandated to Japan, and with the birds of the island of Guam, which is a possession of the United States.

The word Micronesia is, of course, derived from the Greek words _mikros_ meaning small and _nesos_ meaning island, and, as shown in figure 1, this term is appropriate, for the islands of this area are small. For the most part they are too small even for inclusion on standard-sized maps of the world. There are thousands of these islands in an area some 2,400 miles long from east to west and some 1,200 miles broad from north to south. All of the islands of Micronesia are oceanic islands; that is to say, they have never been connected to the Asiatic continent or to other land masses by means of land bridges.

Geologists and oceanographers have shown (see descriptions by Hobbs, 1945), that islands of Micronesia are of two general types: arcuate and strewn. The Pacific Ocean is surrounded by rising mountain ranges which are arranged in elongated, near-circular arcs, which form an extended series of scallops. In the western Pacific these sweeping arcs extend into the ocean, where the mountain ranges project upward from the bottom of the sea with only the crests showing above the waves to point out, in dotted outline, the position of the mountains. The easternmost of these arcs is marked by the islands of the Aleutians, Kuriles, Japan, Izo, Bonins, Volcanoes, Marianas, Yap, Palaus, and others continuing southward into Melanesia. These are characterized by igneous rocks of andesitic nature.

To the eastward of the arcuate islands in Micronesia, are numerous and irregularly distributed islands, making up all of the central and eastern Carolines and the Marshalls, which are known as strewn islands. Strewn islands mark the places of former volcanoes or volcanic peaks. If these volcanic peaks have been completely drowned and are now marked by a series of low islands edged by a protecting reef formed by coral growth enclosing a lagoon and with all exposures consisting of coral rock, the island is known as a coral atoll (example, Ulithi Atoll). Some of the coral exposures lack lagoons; they are known merely as coral islands (example, East Fayu). Some atolls become elevated by geologic activity and the lagoons may dry out or drain. The accumulation of guano of oceanic birds and the residue of fish and other organisms in the area of the lagoon remains as a rich phosphate deposit; these raised atolls have been called phosphate islands (example, Fais). Other strewn islands consist of igneous rocks which are exposed above the surface of the ocean. These are known as "high" or volcanic islands and may occur as a single mountain rising out of the ocean (example, Kusaie), or be partly drowned and surrounded by a coral reef (example, Truk). The igneous rocks found on these strewn islands are basaltic in nature.

The Mariana Islands consist of a chain of volcanic islands approximately 450 miles long. As shown in figure 2, there are 14 single islands and one group of three islands (Maug), from Uracas in the north to Guam in the south. The Palau Islands which are situated in the easternmost part of Micronesia have often been considered from a political standpoint as part of the Caroline Islands. As shown in figure 3, the Palau Islands are a chain of islands approximately 120 miles long from north to south. Sonsorol, Tobi, Merir, Pulo Anna, and Helen Island occur to the southward of the Palaus and may be considered as part of the Carolines or as part of the Palaus. The Palaus together with the Carolines, to the eastward, extend in an east-west direction for approximately 1,700 miles. The Palaus and Carolines include (as shown in figures 3 and 4) 37 atolls, 34 banks, 11 coral islands without lagoons, 2 uplifted phosphate islands, 4 volcanic islands, and the Palau chain. The Marshall Islands to the extreme eastward extend approximately 700 miles from north to south and, as shown in figure 5, contain 29 atolls and five coral islands without lagoons. No volcanic exposures occur in the Marshall Islands.

There is a total land surface of approximately 846 square miles in the islands of Micronesia. The Palaus and Carolines have 525 square miles, the Marianas 247 square miles, and the Marshalls 74 square miles of land surface. Guam has the largest land surface of any of the islands of Micronesia with 225 square miles, Ponapé has 145 square miles, and Babelthuap has 143 square miles. Asuncion, in the northern Marianas, has the highest elevation, rising as an almost perfect cone to a height of 2,923 feet; Ponapé reaches a height of 2,579 feet above the sea level. The volcanic islands are known as "high" islands, and the coral atolls are known as "low" islands. The coral islands usually rise but a few feet above sea level.

CLIMATE

In Micronesia there are two seasons: a wet summer and a dryer winter. Temperatures rarely go above 90° F. and rarely below 70° F. Rainfall in the Marianas averages approximately 85 inches per year, in the Palaus approximately 150 inches, in the Carolines it ranges from 129 to 185 inches, and in the Marshalls it goes up to 160 inches. The humidity is excessive, the average annual mean of relative humidity for selected islands in Micronesia being between 82 and 86 percent. The relative humidity is lower in the western Carolines and the Palaus, than in other parts of Micronesia.

The Mariana Islands lie between the area of the Asiatic monsoon and the belt of the northeast trade winds. At Saipan from November until March or April, winds usually are easterly or northeasterly and are strong and steady since the northeast trades and the winter monsoon reinforce each other. In April and May the directions of the winds shift toward the southeast, and they become weaker and more variable. In this period there may be some easterly winds in addition to the predominating southeasterly winds. Detailed information is not available on the winds which occur in the Marianas north of Saipan, but at Pagan easterly winds probably prevail from May to July and westerly winds prevail in the remainder of the year. The Carolines lie in the belt of alternating northeast trade winds and southwest monsoons. The northeast trades begin in October and prevail until May or June. The southwest monsoon occurs from May to October and may be felt as far east as Truk. To the eastward, the winds of the summer are usually light and variable. In the Marshall Islands, the northeast trade winds predominate from about December to April, especially in the northern part of the Marshalls. In summer, winds are variable and weak; periods of calm may occur. Typhoons and squalls occur most frequently in the spring and summer in Micronesia. Some of the severe typhoons are known to engulf entire islands, as did the one at Woleai in 1907.

SOILS

The soils of the islands of Micronesia have been derived from volcanic materials or from depositions of coralline limestone. Volcanic soils occur on the "high" islands of Micronesia. In many places, especially on the islands of the northern Marianas there is little soil; there are large areas of bare igneous rock, because the islands are geologically of relatively recent origin and little erosion has occurred. On islands where volcanic rocks have decomposed, the resulting soil may have a top layer of humus. The richest soils of the islands are along drainage areas and in alluvial deposits.

Coralline soils result from the decomposition of limestone, coral fragments, shells, and sand, and are overlain by some humus. Where the layer of humus is deep, the fertility is greatest. Coralline-volcanic soils occur on some "high" islands where coral rock and volcanic rock have become mixed in the decomposition process which forms soil. In parts of the Marianas and elsewhere, unwise practices of burning and overgrazing have allowed extensive erosion to occur, resulting in reduced fertility of the soil. On the island of Yap certain sedimentary rocks are exposed which are thought to have been elevated from the ocean bottom. Soils at Yap which have developed from this rock are considered more fertile than soils of coralline origin, although the fertility there also is dependent on the depth of the layer of humus.

SURFACE WATER

There is little fresh water on the coral atolls, but brackish marshes are present on some islands, and many of these marshes are used for the cultivation of taro by the natives. Some volcanic islands, on the other hand, possess small streams and fresh water lakes, producing suitable habitat for certain rails, gallinules and ducks. On the "low" islands in the Marshalls, natural surface pools are rare.

VEGETATION

The "high" islands of Micronesia support a heavy cover of vegetation. Typically the lowlands and stream courses are covered with dense jungle vegetation, and the slopes and higher hills are covered with grasses and brush. The vegetation of the "low" coral atolls and islands is, by comparison, much less dense. Many shorelines are covered with scant grasses and shrubs and the interior in many places is dominated by coconut, betel palms, breadfruit, papaya, and pandanus. References to papers dealing with plants in the islands of the Pacific may be obtained in Merrill (1945), who (1945:207) writes, "Botanically, the low islands are very uninteresting and monotonous. The flora of one is usually quite the same as that of another, although these islands and islets may be separated by many hundred and in some cases several thousand miles. The native vegetation may be scanty or reasonably well developed, depending on the size of the island, the quality of its soil, and whether or not it is permanently inhabitated." Of the vegetation on the "high" islands of the Pacific area, Merrill (1945:209) comments that the vegetation "is well developed, particularly within the forested areas, but for these high islands within the Pacific basin as a whole, the number of endemic genera is relatively small and most of them have definite relationships with those of Malaysia." Concerning the "high" islands of Micronesia, Merrill (1945:210) remarks that these islands are smaller and more isolated than some of the others in Oceania and have fewer individual species "as compared with what one finds on islands of a similar size located within limits of the Malay Archipelago. Thus with all of the islands under Japanese mandate, and including a number of high, but at the same time relatively small islands, less than 1,300 different species are known, of which 230 manifestly represent purposely or accidentally introduced ones. This relatively small flora includes representatives of approximately 620 genera in 192 families.... Specific endemism is relatively high, for approximately 460 species are confined to the islands within the area under consideration. The generic endemism is very low; about seven endemic genera only are involved for the whole group." The figures for endemism of plants are comparable to those for birds. Of endemic birds there are 5 genera, 35 species, and 73 subspecies. The total number of species of birds known from Micronesia is only 206 as compared with 1,300 plants. Yamada (1926:966) writes that the number of species of plants that Micronesia has in common with Japan may be due to the influence of the "Japan Stream."

Many land birds in Micronesia depend directly on the plant life for food. Possibly the soil (including its mineral content), upon which the plants themselves depend for development of fruits and other edible parts, may offer a limiting factor to the distribution of birds in Micronesia. Possibly the fruits and other edible parts of plants do not provide the necessary amounts of proteins, carbohydrates, minerals, vitamins, and other essential food items for species of plant-eating birds, which have not become established in Micronesia. Possibly some species of plant-eating birds have reached Micronesia but have failed to establish themselves because of some dietary deficiency caused by poverty of the soils on which the plants grow. If a comparison were made of soils and of the food values of fruits of plants in both the islands of Micronesia and similarly sized islands in the Malay region, a difference might be revealed which would partly explain why some plant-eating birds have not become established in Micronesia.

GAZETTEER OF ISLANDS OF MICRONESIA

In the following list the name in current usage for each island or island group in Micronesia is followed by other names which have been used. There is no attempt made to list the names of the small islands of each atoll or those of the myriads of small islets that lie offshore from the larger volcanic islands. Collections have not been made on most of the smaller islands. For the few on which a species has been collected, the islet is adequately described in the account of the particular species concerned. The reader may refer to Brigham (1900) for a listing of the islands of the Pacific Ocean. Most of the islands included in the following list may be located on the map of Micronesia as shown in figures 2, 3, 4, and 5. These listings follow in order of arrangement those in the Civil Affairs Handbooks, published by the United States Navy Department (1943, 1944a, 1944b, and 1944c).

MARIANA ISLANDS

The Mariana Islands (also called Ladrone, Marianne, Marian) consist of 14 single islands and one group of three islands. The Marianas are all "high" or volcanic islands. The islands, shown in figure 2, are listed as follows:

_Agrihan_ (also called Agrigan, Arijan, Francisco Xavier, Granger,
Gregus, Grigan, San Francisco Javier).

_Agiguan_ (also called Agaigan, Agiigan, Agiguan, Agigwan,
Aguigan, Aguijan, Aguyan, Guigan, Saint Ange, Santa Angel).

_Alamagan_ (also called Almagan, Aramagan, Concepcion).

_Anatahan_ (also called Anatagen, Anatajen, Anataxan, San
Joaquin).

_Asuncion_ (also called Asonson, Assongsong, Assumption).

_Guam_ (also called Guaham, Guahan).

_Guguan_ (also called Guguwan, Guugwan, Piedras, San Felipe, St.
Philippe).

_Maug_ (also called Mang, Mangs, Mauga, Monjas, Mougu, Saint
Laurent, San Lorenzo, Tunas).

_Medinilla_ (also called Bade, Bird, Farallon de Medinilla,
Rocher).

_Pagan_ (also called Pagon, Paygan, St. Ignace, San Ignacio).

_Rota_ (also called Luta, St. Anne, Santa Ana, Sarpan, Satpana,
Suta, Zarpane).

_Saipan_ (also called (Saepan, St. Joseph, San José, Saypan,
Siepan, Serpan, Seypan).

_Sarigan_ (also called St. Charles, San Carlos, Sariguan,
Sarigwan).

_Tinian_ (also called Bona Vista, Buenavista, Temean, Tenian,
Tiniamou).

_Uracas_ (also called Guy Rock, Farallon de Pájaros, Pájaros,
Urakasu).

PALAU ISLANDS

The Palau Islands (also called Arrecifos, Palaos, Paleu, Pally, Paloc, Pannog, Parao, Pelew) consist of 8 large islands, 18 smaller islands, and a large number of minute islets, all enclosed in a single reef system. The northern islands (Babelthuap and Koror) are of volcanic origin; the southern islands (Peleliu and others) are of coralline formation. Angaur, to the south of Peleliu, may be included with the Palau Archipelago. From the standpoint of the avian zoogeography, the coral islands or atolls of Kayangel, Merir, Pulo Anna, Sonsorol, and Tobi are also included. The principal islands, shown in figure 3, are listed below:

_Arakabesan_ (also called Ngarekobasang).

_Aurapushekaru_ (also called Aburashokoru, Auluptagel, Oluksakel,
Oropu-shakaru).

_Babelthuap_ (also called Babeldzuap, Babel Taob, Babelthouap,
Baberthaob, Baberudaobu, Babldaob).

_Eil Malk_ (also called Amototi, Cogeal, Irakong, Makarakaru).

_Garakayo_

_Koror_ (also called Coror, Goreor, Kororu).

_Malakal_ (also called Amalakell, Malaccan, Marakaru, Nanalake).

_Ngabad_

_Ngesebus_ (also called Guadokusu).

_Peleliu_ (also called Pelelew, Periryu, Pililer, Peliliu, Uler).

_Urukthapel_ (also called Cape, Kuapasungasu, Ngurukdapel, Ulugeang,
Uruk-taaburu, Uruktapi).

Included with the Palau group because of proximity and relationships of the avifauna are the following:

_Angaur_ (also called Angauru, Angyaur, Ngaur, Ngeour, N'Yaur).

_Kayangel_ (also called Kadjangle, Kajanguru, Kazyanguru, Kianguel,
Kreiangel, Moore, Ngajangel, Ngeiangel).

_Merir_ (also called Marir, Meliel, Meriel, Meriru, Pulo Marier, Warren
Hastings).

_Pulo Anna_ (also called Anna, Bul, Bur, Current, Paola, Pul, Puru,
Wull).

_Sonsorol_ (also called St. Andrew, San Andreas, Sonesor, Songosor,
Sonseron, Sonsol, Sonsoru, Tschontil).

_Tobi_ (also called Codopuei, Johnstone, Kadogubi, Lectobis, Lord North,
Nevil, Togobei, Tokobei).

CAROLINE ISLANDS

The Caroline Islands consist of 41 island clusters or isolated islands (exclusive of submerged coral reefs). These are of coral formation. They are atolls or single islands except for Yap, which is of sedimentary rock, and Kusaie, Ponapé, and Truk, which are of volcanic rock. The principal islands are shown in figure 4 and are listed as follows:

_East Fayu_ (also called Fajo, Faliao, Lutké, Rukutee).

_Eauripik_ (also called Aurepik, Eourpyg, Iuripik, Kama, Low,
Yorupikku, Yuripik).

_Fais_ (also called Astrolabe, Feis, Feys, Fuhaesu, Huhaesu,
Tromelin, Woaje).

_Faraulep_ (also called Faraulip, Faroilap, Fattoilap, Foroilap,
Furaarappu, Gardner, Huraarappu).

_Ifalik_ (also called Evalook, Faloc, Furukku, Hurukku, Ifalouk,
Ifelug, Two Sisters, Wilson).

_Kapingamarangi_ (also called Bakiramarang, Constantine,
Greenwich, Guriinitchi, Kabeneylon, Kapenmailang, Makarama,
Pikiram, Tenuv).

_Kusaie_ (also called Arao, Armstrong, Experiment, Hope, Kusai,
Kuschai, Kushai, Kuthiu, Oualan, Quollen, Strong, Teyoa, Ualan,
Walang).

_Lamotrek_ (also called Lamorsu, Lamureck, Lamutrik, Low, Namotik,
Namotikku, Manochikku, Namurrek, Swede).

_Lukunor_ (also called Lemarafat, Lougoullos, Lougounor, Luganor,
Lugunor, Lugunoz, Mortlock, Namonefeng, Rukunoru, Youngwilliam).

_Namonuito_ (also called Anonyma, Baxos de San Bartolomeo, Bunkey,
Las Hermanas, Livingstone, Lost Jardines, Lutké, Namenwita, Olol,
Omun, Onon, Ororu, Remp, Ueito, Ulul).

_Ngulu_ (also called Angegul, Anolul, Goulou, Kurru, Lamoliao,
Lamoliork, Lamuliur, Lamuniur, Matelotas, Ngilu, Ngoli, Ngolog,
Spencer Keys, Ulu).

_Nukuoro_ (also called Dunkin, Matakema, Menteverde, Nugoru,
Nukor, Nukuor).

_Pikelot_ (also called Bigali, Biguela, Coquille, Lydia, Pigela,
Pigerotto, Pigouelao, Pik, Pyghella).

_Pingelap_ (also called Macaskill, Musgrave, Pelelap, Piigerappu,
Punlap, Sailrocks, Tucks Reef).

_Ponapé_ (also called Ascension, Bonabee, Bonybay, Faloupet,
Faounoupei, Funopet, Niponpei, Painipete, Ponapi, Piunipet,
Puynipet, Quirosa, Seniavin, William IV). Ponapé is the largest
island of the Senyavin Islands.

_Truk_ (also called Djuk, Hogoleu, Hogolu, Hoguleu, Lugulus, Ola,
Rough, Ruck, Ruk, Torakku, Tuck, Ugulut). The Truk group includes
approximately 100 islands.

_Ulithi_ (also called Mackenzie, Mogmog, Mogumogu, Mokomok,
Ouluthy, Uluthi, Uluti, Urushi).

_West Fayu_ (also called Faiyao, Fajahu, Faliau, Huiyao, West
Faiu).

_Woleai_ (also called Anagai, Mereyon, Oleai, Ouleyai, Thirteen
Islands, Uala, Ulea, Uola, Ulie, Wolea).

_Yap_ (also called Eap, Guap, Heap, Jap, Ouap, Uap, Wuap, Yappu).

MARSHALL ISLANDS

The Marshall Islands consist of 29 atolls and 5 coral islands without lagoons arranged in two chains, the Ralik and the Radak chains, which extend in a northwesterly to southeasterly direction. No volcanic rocks are exposed in these islands. The principal islands shown in figure 5 are as follows:

_Ailuk_ (also called Ailu, Fisher, Krusenstern, Tindall, Watts).

_Arhno_ (also called Arno, Aruno, Auru).

_Bikar_

_Bikini_

_Ebon_ (also called Boston Atoll).

_Elmore_ (also called Ailinglap, Ailinglapalap, Iringlob).

_Eniwetok_

_Jaluit_ (also called Bonham, Taluit).

_Kwajalein_

_Likieb_ (also called Likiep).

_Majuro_ (also called Arrowsmith, Mezyuro).

_Mejit_

_Maloelab_

_Mille_ (also called Mulgrave).

_Namorik_

_Namu_ (also called Musquillo, Namo).

_Rongelap_

_Wotje_ (also called Romanzov, Wotze, Wozzie).

ORNITHOLOGICAL EXPLORATION IN MICRONESIA

The Micronesian islands were first explored and colonized by a a people who came from Malaysia. It is thought that these people spread into the Palau, Caroline, Mariana, Marshall, and Gilbert islands as a single wave of migration. Following this occupation, the people apparently underwent a normal process of cultural evolution and differentiation. Remains of stone walls, dikes, fences, pillars, graves, and other structures which may be found today at various islands in Micronesia were constructed by the ancestors of the islanders of the present day. It is thought by archeologists that the Polynesians moved eastward into the Pacific islands by way of Micronesia. The date of this wave of migration is thought to have been approximately 1200 A. D. What kinds of birds may have been exterminated by this earliest of human colonization cannot be ascertained. Edible species, particularly megapodes, rails, and pigeons, probably were eliminated or reduced in numbers, as is indicated by later discussions.

The first Europeans to visit Micronesia, as far as the present writer can ascertain, left no accounts of the birds significant for the study here reported upon. Magellan, on his trip around the world, was the leader of the first party of Europeans who touched at Guam; this was on March 6, 1521. Rota, Agiguan, Saipan, and Tinian were also discovered by this Portuguese sea captain in the service of the king of Spain. Eltano, one of Magellan's lieutenants, revisited the Pacific and stopped at Rota in 1524. After the voyage of Magellan, other seafarers, mostly in the service of Spain, visited the Micronesian islands. The Caroline Islands were apparently first observed by the Portuguese captain, Diego de Rocha, in 1526. Loyasa and Saavdera, both Spaniards, visited the Marshall Islands in 1526 and 1529, respectively.

One of the first travelers to record observations on the bird life was Henry Wilson. Wilson was captain of the schooner "Antelope" which became grounded on a reef in the Palau Islands in August, 1783. He lived with the islanders while the ship was being repaired and kept a journal of his observations (Wilson, 1788). Wilson also visited several other islands in western Micronesia. Adelbert von Chamisso (1821), as naturalist with the Russian expedition in the ship "Rurick," made observations of the animal life in Micronesia in 1817 and 1818. Under the command of Otto von Kotzebue, this Russian expedition made the first detailed exploration of the Marshall Islands; visits were made also to Guam and Rota and to Yap, Fais, Ulithi, Palau, and other island groups in western Micronesia. Freycinet's famous expedition in the ships "Uranie" and "Physicienne," visited Guam, Rota, and Tinian in 1819. Quoy and Gaimard, the naturalists of the expedition, obtained birds, which were among the first to be described from Micronesia. These two naturalists revisited the Marianas in 1829 on board the ship "Astrolabe." Scientific results of both of these expeditions (Quoy and Gaimard, 1824-'26 and 1830-'35) include texts and plates dealing with the birds obtained.

The French expedition in the corvette "La Coquille" visited Kusaie in June, 1824. Lesson (1829) wrote the zoology of this trip. Kittlitz (1836) of the expedition which sailed in the corvette "Le Seniavine" commanded by Lutké obtained birds at Kusaie in December and January, 1827-'28, at Guam in March, 1828, and at Lukunor and other islands of the Carolines. At Kusaie, Kittlitz found a rail (_Aphanolimnas monasa_) and a starling (_Aplonis corvinus_) which have not been obtained since his time. His specimens were deposited in St. Petersburg. He was one of the most competent of the early naturalists; his writings contain accounts of habits as well as descriptions and are accompanied by colored plates. The expedition which sailed on the "Astrolabe" and the "Zélée" in 1827-'40 under the command of Dumont d'Urville visited the Caroline Islands. The naturalists, Hombron and Jacquinot, obtained birds at Truk, including the interesting flycatcher, _Metabolus rugensis_, which they described (1841). The "Novara," in the course of its voyage around the world (1857-'59) visited the Caroline Islands in 1858. Birds were recorded from Ponapé, Lukunor and other islands by Pelzeln in his account of the birds of the expedition (1865).

In the years following the middle of the Nineteenth Century, Godeffroy and Sons, of Hamburg, opened branches of its trading firm in Micronesia. Representatives of the company including Heinsohn and Peters, who were ship captains, obtained collections of birds at Palau and Yap. These were deposited in the Godeffroy Museum at Hamburg and reported on by Hartlaub and Finsch (Hartlaub, 1868; Hartlaub and Finsch, 1868a and 1872). Tetens became representative of Godeffroy and Sons at Yap in 1869 and obtained birds. Perhaps the most famous collector in this period was Johann Kubary. He went to Ponapé at the age of nineteen and traveled in Micronesia for many years for Godeffroy and Sons. He obtained birds at many of the islands of the Carolines, spending fourteen months at Truk. In 1873, one of his collections of some 200 birds was lost in a shipwreck. Hartlaub and Finsch, (Hartlaub and Finsch, 1872; Finsch, 1876a) described much of his material; Nehrkorn (1879) reported on nests and eggs which he obtained. Hartlaub and Finsch (1868b) also reported on birds obtained at Palau by Doctor Semper, which were deposited in the museum at Altona. Otto Finsch (1880b, 1880d, 1881b, 1881c) traveled in Micronesia about 1880, observing birds in the eastern Carolines and in the Marshalls.

One of the largest collections from Micronesia was made by Alfred Marche in the Marianas. He arrived there on April 22, 1887, and stayed until May, 1889. He obtained approximately 732 specimens of birds, nests, and eggs at Guam, Rota, Tinian, Saipan, Pagan, and Alamagan, which were deposited in the Paris Museum and reported on by Oustalet (1895-'96). Shortly after Marche's visit, Japanese collectors in the hire of Alan Owston, a professional collector of Yokahama, obtained birds in the Marianas and at Truk in the years 1894-'97. These went to the Rothschild collection at Tring and were reported on by Hartert in 1898 and 1900.

At the turn of the Twentieth Century, several ornithologists were visiting Micronesia. Alvin Seale (1901) obtained a collection of birds at Guam in the summer of 1900 which was deposited in the Bernice P. Bishop Museum in Honolulu. The U. S. Fish Commission steamer "Albatross" visited Micronesia from August, 1899, to March, 1900; birds obtained by the expedition were reported on by Townsend and Wetmore (1919). Paul Schnee (1901) spent approximately one year, 1899-1900, at Jaluit in the Marshalls and obtained records of birds. In 1899, Brandeis, on board the German ship "Kaiserland" visited many of the islands in the Marshalls and recorded birds. William Safford (1905) resided at Guam in the early part of this century and reported on the bird life in the course of his studies of the botany and native life. Bartsch (Mearns, 1909) also obtained a small collection of birds at Guam, this is in the United States National Museum.

In the first World War when the Japanese gained a mandated control over the islands of Micronesia, the Japanese ornithologists promptly visited the area, obtained collections, and published works concerning the birds. In 1922, Momiyama and Kuroda prepared a list of the birds of Micronesia. The work was published under the auspices of the Ornithological Society of Japan. Subsequent editions appeared in 1932 and 1942.

The Whitney South Sea Expedition of the American Museum of Natural History visited Micronesia from October, 1930, to December, 1931, with William F. Coultas as collector. Although experiencing some difficulty and being restricted somewhat in his travels by the Japanese officials, he managed to obtain collections at Ponapé (October 26, 1930, to January 1, 1931), Kusaie (January 15 to June 11, 1931), Guam (June 24 to August 30, 1931), Saipan and Tinian (September 1 to 26, 1931), and Palau (October 2 to December, 1931). Many of the species which he obtained are represented by large series of fine skins. Only part of his collections have been reported on by Mayr and his associates.

Other than the work of Coultas and that of the Japanese, there was little ornithological work done in the period between the two world wars, probably, at least in part, because of the "iron curtain," which Japan had thrown about her mandate. Bryan (1936) did visit Guam in the middle 1930's and published an account of the birds in the newspaper, Guam Recorder.

When the Micronesian islands were taken by the American forces in 1944, personnel attached to various units made observations on the bird life. The first reports, published or unpublished, were from the Marshalls, which were taken at the beginning of the campaign. Gleise, Genelly, Wallace, and others made contributions. In the Marianas considerably more observing and collecting were done by service personnel including Marshall, Stott, Borror, Strophlet, Buss, Watson, Arvey, Downs, and others. Marshall (1949) obtained also a collection of birds in the Palaus in 1945. The Laboratory of Mammalogy, United States Naval Medical Research No. 2, to which I was attached, collected at Guam (January to October, 1945), at Rota (October 17 to November 2, 1945), at Ulithi (August 11 to 23, 1945), at Palau (August 24 to September 24, 1945), and at Truk (November 24 to December 18, 1945). Following the end of the war, Harvey I. Fisher visited Micronesia and obtained a collection of birds at Yap, which is to be reported on in the near future. Larry P. Richards obtained 33 birds at Ponapé and 4 at Truk in the period from August 28, 1947, to February 10, 1948.

Descriptions of birds in Micronesia began with the naming of _Halcyon c. cinnamomina_ in 1821; the most recent description is that of _Rhipidura rufifrons mariae_ in 1946. In all, 131 descriptions have designated type localities in Micronesia. Table 1 lists the dates (on the basis of ten-year intervals) when names of birds (synonyms or otherwise) were proposed. In the period from 1821 to 1860, twenty-five birds were made known to science by the earliest workers, including Kittlitz, Lesson, Bonaparte, and Pelzeln. In the period from 1861 to 1880, thirty-four birds were newly named, mostly by Hartlaub and Finsch, from the collections which the Godeffroy Museum obtained through the efforts of Kubary, Tetens, Peters, and Heinsohn. Nineteen original descriptions were published from 1881 to 1900, principally by Oustalet and Hartert, who studied the material of Marche and Owston, respectively. From 1901 to 1910, only four birds were described, but from 1911 to 1940, forty-seven descriptions were published, mostly by the Japanese following World War I. From 1931 to 1940, the number of known birds was increased by the efforts of Mayr, who studied the material of the Whitney South Sea Expedition. From 1941 to date only two original descriptions have appeared--only one was postwar. Except for possible undescribed subspecies in the northern Marianas, I think that the heyday of the taxonomist in ornithology in Micronesia is over. The field of avian ecology in Micronesia has barely been scratched.

TABLE 1. COMPILATION OF THE DATES (ON THE BASIS OF TEN-YEAR INTERVALS) WHEN ORIGINAL DESCRIPTIONS OF BIRDS OF MICRONESIA APPEARED.

============+==============+============+==============
| No. of | | No. of
YEARS | descriptions | YEARS | descriptions
------------+--------------+------------+--------------
1821-1830 | 8 | 1881-1890 | 9
1831-1840 | 8 | 1891-1900 | 10
1841-1850 | 4 | 1901-1910 | 4
1851-1860 | 5 | 1911-1920 | 10
1861-1870 | 11 | 1921-1930 | 15
1871-1880 | 23 | 1931-1940 | 22
| 1941-1949 | 2
------------+--------------+------------+--------------

CHECK-LIST OF THE BIRDS OF MICRONESIA

The 206 kinds of birds of 150 full species known to occur in Micronesia belong to 91 genera of 37 families of 13 orders. In the following list, nonresident birds are marked with an *; birds introduced by man are marked with a [+].

Class AVES--birds

PAGE

Order PROCELLARIIFORMES--albatrosses, petrels, and allies

Family Diomedeidae--albatrosses

_Diomedia nigripes_ Audubon* Black-footed Albatross 63

Family Procellariidae--petrels and shearwaters

_Puffinus pacificus chlororhynchus_
Lesson Wedge-tailed Shearwater 64
_Puffinus pacificus cuneatus_ Salvin Wedge-tailed Shearwater 65
_Puffinus tenuirostris_ (Temminck)* Short tailed Shearwater 66
_Puffinus nativitatus_ Streets Christmas Shearwater 66
_Puffinus lherminieri dichrous_ Finsch
and Hartlaub Dusky Shearwater 66
_Pterodroma rostrata rostrata_ (Peale)* Tahiti Petrel 69
_Pterodroma hypoleuca hypoleuca_ Salvin Stout-billed Gadfly Petrel 70

Order PELECANIFORMES--tropic birds, boobies, cormorants, frigate
birds and allies

Family Phaëthontidae--tropic birds

_Phaëthon aethereus mesonauta_ Peters* Red-billed Tropic Bird 70
_Phaëthon rubricauda rothschildi_
(Mathews) Red-tailed Tropic Bird 71
_Phaëthon lepturus dorotheae_ Mathews White-tailed Tropic Bird 72

Family Sulidae--boobies and gannets

_Sula dactylatra personata_ Gould Masked Booby 75
_Sula sula rubripes_ Gould Red-footed Booby 75
_Sula leucogaster plotus_ (Forster) Brown Booby 76

Family Phalacrocoracidae--cormorants

_Phalacrocorax melanoleucus melanoleucus_
(Vieillot) Little Pied Cormorant 78

Family Fregatidae--frigate birds or man-o'-war birds

_Fregata minor minor_ (Gmelin)* Pacific Man-o'-War 79
_Fregata ariel ariel_ (Gray) Least Man-o'-War 80

Order CICONIIFORMES--herons, storks, and allies

Family Ardeidae--herons and bitterns

_Butorides striatus amurensis_ Schrenck* Amur Green Heron 81
_Bubulcus ibis coromandus_ (Boddaert)* Cattle Egret 82
_Egretta intermedia intermedia_
(Wagler)* Plumed Egret 82
_Demigretta sacra sacra_ (Gmelin) Reef Heron 84
_Nycticorax nycticorax nycticorax_
(Linnaeus)* Black-crowned Night Heron 87
_Nycticorax caledonicus pelewensis_
Mathew Rufous Night Heron 87
_Gorsachius goisagi_ (Temminck)* Japanese Bittern 89
_Gorsachius melanolophus melanolophus_
(Raffles)* Malay Bittern 90
_Ixobrychus sinensis_ (Gmelin) Chinese Least Bittern 93
_Ixobrychus eurhythmus_ (Swinhoe)* Shrenck's Least Bittern 93
_Dupetor flavicollis flavicollis_
(Latham)* Black Bittern 94

Order ANSERIFORMES--ducks, geese, swans, and allies

Family Anatidae--ducks, geese, and swans

_Anas oustaleti_ Salvadori Marianas Mallard 94
_Anas poecilorhyncha pelewensis_
Hartlaub and Finsch Australian Gray Duck 98
_Anas querquedula_ Linnaeus* Garganey Teal 100
_Anas crecca crecca_ Linnaeus* European Teal 100
_Anas crecca carolinensis_ Gmelin* Green-winged Teal 100
_Anas acuta acuta_ Linnaeus* Pintail 101
_Anas acuta tzitzihoa_ Vieillot* Pintail 101
_Anas penelope_ Linnaeus* Widgeon 102
_Anas clypeata_ Linnaeus* Shoveller 102
_Aythya fuligula_ (Linnaeus)* Tufted Duck 103
_Aythya valisineria_ (Wilson)* Canvas-back 103

Order FALCONIFORMES--vultures, hawks, falcons

Family Accipitridae--hawks, harriers, and allies

_Accipiter soloënsis_ (Horsfield)* Chinese Goshawk 104
_Accipiter virgatus gularis_ (Temminck
and Schlegel)* Asiatic Sparrow Hawk 104
_Pandion haliaetus melvillensis_
Mathews Osprey 105

Family Falconidae--falcons and caracaras

_Falco peregrinus japonensis_ Gmelin* Peregrine Falcon 105

Order GALLIFORMES--megapodes, pheasants, and allies

Family Megapodidae--megapodes

_Megapodius lapérouse senex_ Hartlaub Micronesian Megapode 106
_Megapodius lapérouse lapérouse_ Gaimard Micronesian Megapode 109

Family Phasianidae--quails, pheasants, and allies

_Coturnix chinensis lineata_
(Scopoli)[+] Painted Quail 113
_Gallus gallus_ (Linnaeus)[+] Red Jungle Fowl 114
_Phasianus colchicus_ Linnaeus[+] Ring-necked Pheasant 115

Order GRUIFORMES--cranes, rails, and allies

Family Rallidae--rails, gallinules, and coots

_Rallus philippensis pelewensis_ (Mayr) Banded Rail 116
_Rallus owstoni_ (Rothschild) Guam Rail 118
_Rallina fasciata_ (Raffles)* Malay Banded Crake 120
_Rallina eurizonoïdes eurizonoïdes_
(Lafresnaye)* Philippine Banded Crake 121
_Aphanolimnas monasa_ (Kittlitz) Kusaie Black Rail 121
_Poliolimnas cinereus micronesiae_
Hachisuka White-browed Rail 123
_Gallinula chloropus_ subsp. near
_orientalis_ Horsfield Gallinule 126
_Gallinula chloropus guami_ Hartert Gallinule 127
_Porphyrio porphyrio pelewensis_ Hartlaub
and Finsch Purple Swamphen 129
_Fulica atra atra_ Linnaeus* Common Coot 131

Order CHARADRIIFORMES--shorebirds, gulls, and auks

Family Charadriidae--plovers, turnstones, and allies

_Squatarola squatarola_ (Linnaeus)* Black-bellied Plover 131
_Pluvialis dominica fulva_ (Gmelin)* Pacific Golden Plover 132
_Charadrius hiaticula semipalmatus_
Bonaparte* Semipalmated Plover 134
_Charadrius dubius curonicus_ Gmelin* Ring-necked Plover 135
_Charadrius alexandrinus nihonensis_
Deignan* Kentish Plover 135

_Charadrius mongolus stegmanni_
Stresemann* Mongolian Dotteral 135
_Charadrius leschenaultii_ Lesson* Large Sand Dotteral 137

Family Scolopacidae--snipe, sandpipers, and allies

_Numenius phaeopus variegatus_
(Scopoli)* Whimbrel 137
_Numenius tahitiensis_ (Gmelin)* Bristle-thighed Curlew 139
_Numenius madagascariensis_ (Linnaeus)* Long-billed Curlew 140
_Limosa lapponica baueri_ Naumann* Pacific Godwit 140
_Tringa nebularia_ (Gunnerus)* Greenshawk 141
_Tringa melanoleuca_ (Gmelin)* Greater Yellow-legs 142
_Tringa glareola_ Linnaeus* Wood Sandpiper 142
_Actitus hypoleucos_ Linnaeus* Common Sandpiper 143
_Heteroscelus brevipes_ (Vieillot)* Gray-tailed Tattler 144
_Heteroscelus incanus_ (Gmelin)* Amer. Wandering Tattler 145
_Arenaria interpres interpres_ (Linnaeus)* Turnstone 147
_Gallinago megala_ Swinhoe* Marsh Snipe 149
_Gallinago gallinago gallinago_ (Linnaeus)* Common Snipe 150
_Crocethia alba_ (Pallas)* Sanderling 150
_Calidris tenuirostris_ (Horsfield)* Asiatic Knot 151
_Erolia minuta ruficollis_ (Pallas)* Little Stint 151
_Erolia subminuta_ (Middendorff)* Least Sandpiper 152
_Erolia melanotos_ (Vieillot)* Pectoral Sandpiper 152
_Erolia acuminata_ (Horsfield)* Sharp-tailed Sandpiper 152
_Erolia ferruginea_ (Pontoppidan)* Curlew Sandpiper 153
_Limicola falcinellus sibirica_ Dresser* Broad-billed Sandpiper 154

Family Phalaropidae--phalaropes

_Phalaropus lobatus_ (Linnaeus)* Northern Phalarope 154

Family Laridae--gulls and terns

_Larus argentatus vegae_ Palmén* Herring Gull 154
_Chlidonias leucopterus_ (Temminck)* White-winged Black
Tern 155
_Sterna hirundo longipennis_ Nordmann* Black-billed Com. Tern 155
_Sterna sumatrana sumatrana_ Raffles Black-naped Tern 156
_Sterna lunata_ Peale Spectacled Tern 160
_Sterna anaetheta anaetheta_ Scopoli Bridled Tern 160
_Sterna fuscata oahuensis_ Bloxham Sooty Tern 161
_Sterna albifrons sinensis_ Gmelin* Least Tern 161
_Thalasseus bergii pelecanoides_ (King) Crested Tern 162
_Procelsterna cerulea saxatilis_
W. E. Fisher* Blue-gray Tern 164
_Anoüs stolidus pileatus_ (Scopoli) Common Noddy 165
_Anoüs tenuirostris marcusi_ (Bryan) White-capped Noddy 170
_Gygis alba candida_ (Gmelin) White Tern 174
_Gygis alba pacifica_ (Lesson) White Tern 180

Order COLUMBIFORMES--pigeons, doves, and allies

Family Columbidae--pigeons and doves

_Columba livia_ Gmelin[+] Blue Rock Pigeon 182
_Ptilinopus porphyraceus ponapensis_
(Finsch) Crimson-crw'd Fruit
Dove 182
_Ptilinopus porphyraceus hernsheimi_
(Finsch) Crimson-crw'd Fruit
Dove 184
_Ptilinopus porphyraceus pelewensis_
Hartlaub and Finsch Crimson-crw'd Fruit
Dove 185
_Ptilinopus roseicapillus_ (Lesson) Marianas Fruit Dove 186
_Ducula oceanica monacha_ (Momiyama) Micronesian Pigeon 190
_Ducula oceanica teraokai_ (Momiyama) Micronesian Pigeon 193
_Ducula oceanica townsendi_ (Wetmore) Micronesian Pigeon 194
_Ducula oceanica oceanica_ (Lesson and
Garnot) Micronesian Pigeon 195
_Ducula oceanica ratakensis_ (Takatsukasa
and Yamashina) Micronesian Pigeon 197
_Streptopelia bitorquata dusumieri_
(Temminck)[+] Philippine Turtle
Dove 198
_Gallicolumba canifrons_ (Hartlaub and
Finsch) Palau Ground Dove 201
_Gallicolumba xanthonura xanthonura_
(Temminck) White-thrt'd Ground
Dove 203
_Gallicolumba xanthonura kubaryi_ (Finsch) White-thrt'd Ground
Dove 207
_Caloenas nicobarica pelewensis_ Finsch Nicobar Pigeon 209

Order PSITTACIFORMES--lories and parrots

Family Psittacidae--lories, parrots, and allies

_Trichoglossus rubiginosus_ (Bonaparte) Ponapé Lory 211

Order CUCULIFORMES--cuckoos, plantain-eaters

Family Cuculidae--cuckoos, anis, and allies

_Cuculus canorus telephonus_ Heine* Common Cuckoo 213
_Cuculus saturatus horsfieldi_ Moore* Oriental Cuckoo 214
_Eudynamis taitensis_ (Sparrman)* Long-tailed New
Zealand Cuckoo 214

Order STRIGIFORMES--owls

Family Strigidae--owls

_Otus podarginus_ (Hartlaub and Finsch) Palau Scops Owl 215
_Asio flammeus flammeus_ (Pontoppidan)* Short-eared Owl 217
_Asio flammeus ponapensis_ Mayr Short-eared Owl 218

Order CAPRIMULGIFORMES--goatsuckers and allies

Family Caprimulgidae--goatsuckers

_Caprimulgus indicus jotaka_ Temminck
and Schlegel* Jungle Nightjar 219
_Caprimulgus indicus phalaena_ Hartlaub
and Finsch Jungle Nightjar 219

Order APODIFORMES--swifts and hummingbirds

Family Apodidae--swifts

_Collocalia inexpectata pelewensis_ Mayr Edible Nest Swiftlet 221
_Collocalia inexpectata bartschi_ Mearns Edible Nest Swiftlet 222
_Collocalia inquieta inquieta_ (Kittlitz) Carolines Swiftlet 224
_Collocalia inquieta rukensis_ Kuroda Carolines Swiftlet 225
_Collocalia inquieta ponapensis_ Mayr Carolines Swiftlet 226

Order CORACIIFORMES--kingfishers, rollers, and allies

Family Alcedinidae--kingfishers

_Halcyon cinnamomina cinnamomina_ Swainson Micronesian
Kingfisher 227
_Halcyon cinnamomina pelewensis_ Wiglesworth Micronesian
Kingfisher 229
_Halcyon cinnamomina reichenbachii_
(Hartlaub) Micronesian
Kingfisher 230
_Halcyon chloris teraokai_ Kuroda White-collared
Kingfisher 233
_Halcyon chloris orii_ Takatsukasa and
Yamashina White-collared
Kingfisher 235
_Halcyon chloris albicilla_ (Dumont) White-collared
Kingfisher 235
_Halcyon chloris owstoni_ Rothschild White-collared
Kingfisher 237

Family Coraciidae--rollers

_Eurystomus orientalis connectens_
Stresemann* Dollar Bird 238

Order PASSERIFORMES--perching birds

Family Hirundinidae--swallows

_Hirundo rustica gutteralis_ Scopoli* Eastern Barn Swallow 239

Family Campephagidae--cuckoo-shrikes

_Edolisoma tenuirostre monachum_
(Hartlaub and Finsch) Cicada Bird 239
_Edolisoma tenuirostre nesiotis_
(Hartlaub and Finsch) Cicada Bird 241
_Edolisoma tenuirostre insperatum_ (Finch) Cicada Bird 242

Family Dicruridae--drongos

_Dicrurus macrocercus harterti_ S. Baker[+] Black Drongo 244

Family Corvidae--crows, magpies, and jays

_Corvus kubaryi_ Reichenow Marianas Crow 244

Family Turdidae--thrushes

_Luscinia calliope calliope_ (Pallas)* Siberian Rubythroat 248
_Monticola solitaria philippensis_ (Müller)* Chinese Blue Rock
Thrush 248
_Turdus obscurus obscurus_ Gmelin* Dusky Thrush 248

Family Sylviidae--Old World warblers

_Psamathia annae_ Hartlaub and Finsch Palau Bush-warbler 249
_Acrocephalus luscinia luscinia_ (Quoy and
Gaimard) Nightingale
Reed-warbler 251
_Acrocephalus luscinia syrinx_ (Kittlitz) Nightingale
Reed-warbler 254
_Acrocephalus luscinia yamashinae_
(Takatsukasa) Nightingale
Reed-warbler 256
_Acrocephalus luscinia nijoi_ (Yamashina) Nightingale
Reed-warbler 257

Family Muscicapidae--Old World flycatchers

_Rhipidura rufifrons uraniae_ Oustalet Rufous-fronted
Fantail 261
_Rhipidura rufifrons saipanensis_ Hartert Rufous-fronted
Fantail 262
_Rhipidura rufifrons mariae_ R. H. Baker Rufous-fronted
Fantail 263
_Rhipidura rufifrons versicolor_ Hartlaub
and Finsch Rufous-fronted
Fantail 264
_Rhipidura rufifrons kubaryi_ Finsch Rufous-fronted
Fantail 265
_Rhipidura lepida_ Hartlaub and Finsch Palau Fantail 266
_Metabolus rugensis_ (Hombron and Jacquinot) Truk Monarch 269
_Monarcha godeffroyi_ Hartlaub Yap Monarch 272
_Monarcha takatsukasae_ (Yamashina) Tinian Monarch 274
_Myiagra oceanica erythrops_ Hartlaub and
Finch Micronesian Broadbill 275
_Myiagra oceanica freycineti_ Oustalet Micronesian Broadbill 277
_Myiagra oceanica oceanica_ Pucheran Micronesian Broadbill 279
_Myiagra oceanica pluto_ Finsch Micronesian Broadbill 280
_Muscicapa narcissina narcissina_ Temminck* Narcissus Flycatcher 282
_Muscicapa griseisticta_ (Swinhoe)* Chinese Gray-spotted
Flycatcher 282
_Colluricincla tenebrosa_ (Hartlaub and
Finsch) Palau Morning Bird 282

Family Artamidae--wood-swallows

_Artamus leucorhynchus pelewensis_ Finsch White-breasted
Wood-swallow 284

Family Sturnidae--starlings

_Aplonis opacus opacus_ (Kittlitz) Micronesian Starling 286
_Aplonis opacus ponapensis_ Takatsukasa and
Yamashina Micronesian Starling 288
_Aplonis opacus angus_ Momiyama Micronesian Starling 289
_Aplonis opacus kurodai_ Momiyama Micronesian Starling 291
_Aplonis opacus orii_ (Takatsukasa and
Yamashina) Micronesian Starling 292
_Aplonis opacus guami_ Momiyama Micronesian Starling 293
_Aplonis opacus aeneus_ (Takatsukasa and
Yamashina) Micronesian Starling 297
_Aplonis pelzelni_ Finsch Ponapé Mountain
Starling 299
_Aplonis corvinus_ (Kittlitz) Kusaie Mountain
Starling 301
_Sturnus philippensis_ (Forster)* Violet-backed
Starling 302
_Sturnus cineraceus_ Temminck* Ashy Starling 302

Family Meliphagidae--honey-eaters

_Cleptornis marchei_ (Oustalet) Golden Honey-eater 302
_Myzomela cardinalis rubratra_ (Lesson) Cardinal Honey-eater 304
_Myzomela cardinalis dichromata_ Wetmore Cardinal Honey-eater 307
_Myzomela cardinalis major_ Bonaparte Cardinal Honey-eater 307
_Myzomela cardinalis saffordi_ Wetmore Cardinal Honey-eater 309
_Myzomela cardinalis kurodai_ Momiyama Cardinal Honey-eater 312
_Myzomela cardinalis kobayashii_ Momiyama Cardinal Honey-eater 313

Family Zosteropidae--white-eyes

_Zosterops conspicillata conspicillata_
(Kittlitz) Bridled White-eye 316
_Zosterops conspicillata saypani_ Dubois Bridled White-eye 318
_Zosterops conspicillata rotensis_
Takatsukasa and Yamashina Bridled White-eye 319
_Zosterops conspicillata semperi_ Hartlaub Bridled White-eye 320
_Zosterops conspicillata owstoni_ Hartert Bridled White-eye 321
_Zosterops conspicillata takatsukasai_
Momiyama Bridled White-eye 322
_Zosterops conspicillata hypolais_
Hartlaub and Finsch Bridled White-eye 323
_Zosterops cinerea cinerea_ (Kittlitz) Micron. Dusky
White-eye 326
_Zosterops cinerea ponapensis_ Finsch Micron. Dusky
White-eye 327
_Zosterops cinerea finschii_ (Hartlaub) Micron. Dusky
White-eye 328
_Rukia palauensis_ (Reichenow) Palau Greater
White-eye 330
_Rukia oleaginea_ (Hartlaub and Finsch) Yap Greater
White-eye 331
_Rukia ruki_ (Hartert) Truk Greater
White-eye 332
_Rukia sanfordi_ (Mayr) Ponapé Greater
White-eye 333

Family Ploceidae--weaver-finches

_Erythrura trichroa trichroa_ (Kittlitz) Blue-faced
Parrot-finch 336
_Erythrura trichroa clara_ Takatsukasa
and Yamashina Blue-faced
Parrot-finch 337
_Erythrura trichroa pelewensis_ Kuroda Blue-faced
Parrot-finch 338
_Lonchura nigerrima minor_ (Yamashina) Black-faced
Weaver-finch 339
_Lonchura punctulata cabanisi_ (Sharpe)[+] Phil. Nutmeg
Mannikin 340

DISCUSSION OF THE BIRDS OF MICRONESIA

Of the 206 kinds of birds found in Micronesia, 30 kinds are classed as sea birds, 29 kinds as migratory shore birds, and 147 kinds are classed as land and fresh-water birds. For purposes of discussion these birds are arranged in these three categories, following the system used by Mayr (1945a).

OCEANIC BIRDS

Oceanic birds found in Micronesia belong to the following families: Diomedeidae, Procellariidae, Phaëthontidae, Pelecanidae, Fregatidae, and Laridae. Following Wynne-Edwards (1935:240) and Murphy (1936:326), these birds may be grouped as inshore birds (Laridae and others), offshore birds (Pelecanidae, Fregatidae and others), and pelagic birds (Diomedeidae, Procellariidae, Phaëthontidae). As shown in table 2 there are 30 kinds of oceanic birds in Micronesia, 18 kinds that are resident and 12 kinds that are regarded as visitors to the area. Records of nestings are few; field work in the future probably will yield evidence that more kinds of oceanic birds are actually resident in the Micronesian islands.

TABLE 2. LIST OF RESIDENT AND NONRESIDENT OCEANIC BIRDS OF MICRONESIA

-------------------+----------+-------------
Genera | Resident | Nonresident
| kinds | kinds
-------------------+----------+-------------
_Diomedea_ | 0 | 1
_Puffinus_ | 4 | 1
_Pterodroma_ | 1 | 1
_Phaëthon_ | 2 | 1
_Sula_ | 3 | 0
_Fregata_ | 1 | 1
_Larus_ | 0 | 1
_Chlidonias_ | 0 | 1
_Sterna_ | 2 | 4
_Thalasseus_ | 1 | 0
_Procelsterna_ | 0 | 1
_Anoüs_ | 2 | 0
_Gygis_ | 2 | 0
-------------------+----------+-------------

INSHORE OCEANIC BIRDS

The inshore zone, according to Wynne-Edwards (1935:240), "extends from high-water mark to a maximum of four or five miles out to sea, including islands and reefs within sight of shore." In Micronesia the majority of the Laridae occur in this zone including such residents as _Sterna sumatrana_, _S. anaetheta_, _Thalasseus bergii_, _Anoüs stolidus_, _A. tenuirostris_, _Gygis alba_. These birds, especially _S. anaetheta_, _Thalasseus_, and _Anoüs_, may venture into the offshore zone. Visitors to Micronesia include several terns which probably normally range in the inshore (as well as in offshore) zones, such as _Childonias leucopterus_ and _Sterna hirundo_. These birds feed to a considerable extent inside the outer reefs surrounding the lagoons, coming to shore frequently in small or large groups. _Gygis alba_ probably spends considerable time on shore; stomachs examined contained fish, crustaceans and insects, indicating that they obtain some of their food ashore.

OFFSHORE AND PELAGIC OCEANIC BIRDS

Wynne-Edwards (1935:241) defines the offshore zone as extending to the continental edge; however, in Micronesia where small islands rise abruptly out of the ocean's depths, there is no useful way to separate the offshore zone from the pelagic zone. Since certain species go farther from the land than others, the two zones may be combined as a single zone extending beyond the sight of land. Birds which frequent this area beyond the inshore zone but may not range extensively at sea include _Fregata_, _Sula_, _Sterna fuscata_, _S. hirundo_, _S. anaetheta_, and others. The Herring Gull (_Larus argentatus_), which has been taken in the northern Marianas, may be classed with this group although it probably ranges widely in the open sea. Birds which spend considerable time at sea and may seldom approach land include _Diomedea nigripes_, the petrels (_Puffinus_ and _Pterodroma_), and possibly the tropic birds (_Phaëthon_).

In numbers of individuals the birds inhabiting the inshore zones are relatively more numerous than those preferring the offshore and pelagic zones, although 12 of the 18 resident kinds of oceanic birds apparently prefer the offshore zone, while only 6 kinds appear to be restricted primarily to the inshore areas.

FAUNAL COMPONENTS

The oceanic birds were probably among the earliest birds to reach the islands of Micronesia. The presence of phosphate deposits on islands (Fais, Angaur), denoting deposition of guano by oceanic birds (possibly boobies, noddies, sooty terns), indicates long time residence by these birds. A person is prone to think that these deposits must have been made by larger concentrations of oceanic birds than are found in these islands today. Whether there were actually more individuals present during the period of deposition of phosphate in the lagoons of these islands is not known, although the elevation of the lagoons (forming the raised islands of Fais and Angaur) with the resulting freshening of the water probably was a great attraction to oceanic birds, especially to those which prefer to drink fresh water. According to Leonard P. Schultz (_in litt._), the abundance of fish in the areas about these Pacific islands has been approximately the same since Pleistocene times, so that there was apparently no greater concentration of fish near these islands to attract large populations of fish-eating sea birds. Probably the time element is of sufficient magnitude to account for such deposition by birds with a population similar to that found there today.

The oceanic avifauna of Micronesia contains birds which are apparently from ancestral homes in the Palearctic Region, in the North and Central Pacific, in Polynesia, in Melanesia and Malaysia, and from homes the positions of which are uncertain because of the widespread circumtropical occurrence of the birds. There are no sea birds that are endemic in Micronesia.

Oceanic birds whose range is in the Northern Hemisphere (especially Palearctica) reach the northern and western edges of Micronesia as winter visitors. These include _Larus argentatus_, _Chlidonias leucopterus_, and _Sterna hirundo_. Another northern gull, _Larus ridibundus_, has been reported in the Marianas.

One bird of the North and Northcentral Pacific, _Diomedea nigripes_, reaches the northern Marianas where it has been taken at Agrihan. It is not unlikely that other birds of the North Pacific reach northern Micronesia as occasional visitors.

Species of oceanic birds which are restricted in their distribution to Polynesia and some adjacent islands and which range to Micronesia, either as visitors or residents, include _Puffinus tenuirostris_, _P. nativitatis_, _Pterodroma rostrata_, _P. hypoleuca_, _Sterna lunata_, and _Procelsterna cerulea_. The islands of the vast Pacific basin offer havens for many kinds of oceanic birds. Apparently there has been considerable speciation among sea birds in Polynesia, especially in its marginal areas. Micronesia has received only a small part of this avifauna.

Two terns, _Sterna sumatrana_ and _Thalasseus bergii_, have reached Micronesia, either directly or indirectly, each from a dispersion point somewhere in the Melanesian or the Malayan area. These two birds are restricted in their ranges to the western Pacific and the Indian oceans.

Many of the species of oceanic birds found in Micronesia have circumtropical ranges. These include _Puffinus pacificus_, _P. lherminieri_, _Phaëthon_, _Sula_, _Fregata_, _Sterna anaethetus_, _S. fuscata_, _Anoüs stolidus_, _A. tenuirostris_, and _Gygis alba_. Some of these kinds range along continental shores as well as in island archipelagoes. Others, like _Gygis alba_, are rarely found along the shores of continents or even at coastal islands.

MIGRATORY SHORE BIRDS