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

24-1811

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.

Fig. 1. The Islands of Micronesia.

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.