The Project Gutenberg eBook of The Avifauna of Micronesia, Its Origin, Evolution, and Distribution
Title: The Avifauna of Micronesia, Its Origin, Evolution, and Distribution
Author: Rollin H. Baker
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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.
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).