(navigation image)
Home American Libraries | Canadian Libraries | Universal Library | Community Texts | Project Gutenberg | Children's Library | Biodiversity Heritage Library | Additional Collections
Search: Advanced Search
Anonymous User (login or join us)
Upload
See other formats

Full text of "University of Kansas publications, Museum of Natural History"

U IV 1 ^ ^td 



HARVARD UNIVERSITY 




LIBRARY 



OF THE 



MUSEUM OF COMPARATIVE ZOOLOGY 



( 



)c?-( 



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 



EB 1 1 1952 

ffAmw 

WIVEISITY 



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 AUied 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. 






FEB 1 1 1952 
HAmRO 






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



BY 



ROLLIN H. BAKER 



MUS. COV.P. 200L 
LIBRARY 

-'^'^J IS 1951 

HAHVARO 
UNIVERSITY 



University of Kansas Publications 
Museum of Natural History 

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



6ne of fifty copies printed 
on 100 per cent rag paper 



University of Kansas 

LAWBEXCK 

19S1 



UNIVERSITY OF KANSAS PUBLICATIONS 

The University of Kansas Publications, Museilm of Natural His- 
tory, are offered in exchange for the publications of learned societies 
and institutions, universities and libraries. For exchanges and in- 
formation, address the Exchange Desk, University of Kansas Li- 
brary, LAWRENCE; Kansas, U. S. A.' 

Museum of JVTatubal History. — ^E. Raymond Hall, Ch&irman, Editorial Com- 
mittee. 
This series contains contributions from the Museum 'of Natural History. 
Cited 'as Univ. Kans. Publ., Mus. Nat. Hist. 

Vol. 1. 1. The pocket gophers (genus Thomomys) of Utah, By Stephen D. 
Durrani. Pp. 1-82, 1 figure in text. August 15, 1946. 

2. The systematic status of Eiuneces pluvialis Cope, and noteworthy 
records of other amphibians and reptiles from Kansas and Okla- 
homa. By Hobart M. Smith. Pp. 85-89. August 15, 1946. 

3. The tadpoles of Bufo cognatus Say. By Hobart M. Smith. Pp. 
93-96, 1 figure in text. August 15, 1946. 

4. Hybridization between t-R?o species of garter snakes. By Hobart M. 
Smith. Pp. 97-100. August 15, 1946. 

5. Selected records of reptiles and amphibians from Kansas. By John 
Breukelman and Hobart M. Smith. Pp. 101-112. August 15, 1946. 

6. Kyphosis and other variations in soft-shelled turtles. By Hobart 
M. Smith. Pp. 117-124. July 7, 1947. 

7. Natural history of the prairie vole (Mammalian genus Microtus). 
By E. W. Jameson, Jr. Pp, 125-151, 4 figures in text. October 6, 

• 1947. 

8. The postnatal development 9f two broods of great homed owla 
(Bubo virginianus) . By Donald F. Hoffmeister and Henry W. 
Setzer. Pp. 157-173, 5 figures in text. October 6, 1947. 

9. Additions to the list of the birds of Louisiana. By George H. 
LoVery, Jr. Pp. 177-192. November 7, 1947. 

10. A check-list of the birds of Idaho. By M. Dale Arvey. Pp. 193- 
216. November 29, 1947. 

11. Subspeciation in pocket gophers of Kansas. By Bernardo Villa-R. 
and E. Raymond Hall. Pp. 217-236, 2 figures in text. November 
29, 1947. 

12. A new bat (Genus Myotis) from Mexico. By Walter W. Dalquest 
and E. Raymond Hall. Pp. 237-244, 6 figures in text. December 
10, 1947. 

13. Tadarida femorosacca (Merriam) in Tamaulipas, Mexico. By 
Walter W. Dalquest and E. Raymond Hall. Pp. 245-248, 1 figure 
in text. December 10, 1947. 

14. A new pocket gopher (Thomomys) and a new spiny pocket mouse 
(Liomys) from Michoacan, Mexico. By E. Ra3Tnond Hall and 

s Bernardo Villa-R. Pp. 249-256, 6 figures in text. July 26, 1948. 

15. A new hylid frog from eastern Mexico. By Edward H. Taylor. 
Pp. 257-264, 1 figure in text. August 16, 1948. 

(Continued on inside of back cover.) 



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



BT 



ROLLIN H. BAKER 



MUS. COMP. ZOOL 
LIBRARY 

JUN 18 1951 

HAHVARO 
UNIVERSITY 



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 

1931 



Univ'ersity of Kansas PrBLicATioNS, 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 

(3) 



University of Kansas Publs., Mus. Nat. Hist. 



PAGE 



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 Palaii 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 de- 

rived 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 Gallicolumha 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, ac- 
tually 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 diffi- 
culties, 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 ques- 
tion 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 ac- 
tual 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 addi- 
tional 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 Mi- 
cronesia. 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 

(5) 



6 University of Kansas Publs., Mus. Nat. Hist. 

the standpoint of anthropology). Zoogeographically, according to 
Wallace (1876), Micronesia is to be included in the Polynesian Sub- 
region of the Australian Region. Mayr (1941a: 193), on the basis 
of the distribution of birds, ranks Micronesia as one of the four sub- 
divisions of the Polynesian Subregion, and includes within Micro- 
nesia 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. 



ISO. 



f. 



■/ i 






MARIANA : 
ISLANDS; 



CAROLINE ISLANDS 



125 250 500 MILES 

II I I 



•WAKE 



-%, 



■^ 



^-^o 



^0.. 



<^^ ■ ■{ <^^ ..- \ 



V GILBERT 
■..-ISLANDS 



3i: 



ife" 



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 Mi- 
cronesia 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. 



Bakek: The Avifauna of Micronesia 7 

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 \vath 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, Vol- 
canoes, Marianas, Yap, Palaus, and others continuing southward 
into Melanesia. These are characterized by igneous rocks of andes- 
itic nature. 

To the eastward of the arcuate islands in Micronesia, are numer- 
ous 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 vol- 
canic 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 expos- 
ures consisting of coral rock, the island is known as a coral atoll (ex- 
ample, 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 con- 
sist 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 ap- 
proximately 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 con- 
sidered from a political standpoint as part of the Caroline Islands. 
As shown in figure 3, the Palau Islands are a chain of islands ap- 
proximately 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 east- 
ward, extend in an east-west direction for approximately 1,700 



8 University of Kansas Publs., Mus. Nat. Hist. 

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, Ponape 
has 145 square miles, and Babelthuap has 143 square miles. Asun- 
cion, in the northern Marianas, has the highest elevation, rising as 
an almost perfect cone to a height of 2,923 feet; Ponape 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 in- 
formation 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 Oc- 



Baker: The Avifauna of Micronesia 9 

tober 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 eastvi^ard, the winds of the summer are usually light and 
variable. In the Marshall Islands, the northeast trade winds pre- 
dominate 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 fre- 
quently 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. Vol- 
canic 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-vol- 
canic soils occur on some "high" islands where coral rock and vol- 
canic rock have become mixed in the decomposition process which 
forms soil. In parts of the Marianas and elsewhere, unwise prac- 
tices 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 de- 
veloped 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, pro- 
ducing suitable habitat for certain rails, gallinules and ducks. On 
the "low" islands in the Marshalls, natural surface pools are rare. 



10 University of Kansas Publs., Mus. Nat. Hist. 

Vegetation 

The "high" islands of Micronesia support a heavy cover of vege- 
tation. 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, de- 
pending 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." Con- 
cerning the "high" islands of Micronesia, Merrill (1945:210) re- 
marks that these islands are smaller and more isolated than some 
of the others in Oceania and have fewer individual species "as com- 
pared 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 ende- 
mism 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 in- 
volved 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." 



Baker: The Avifauna of Micronesia 11 

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, carbohy- 
drates, minerals, vitamins, and other essential food items for species 
of plant-eating birds, which have not become established in Micro- 
nesia. 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 simi- 
larly sized islands in the Malay region, a difference might be re- 
vealed which would partly explain why some plant-eating birds have 
not become established in Micronesia. 

GAZETTEER OF ISLANDS OF IMICRONESIA 

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 Na\'y Department 
(1943, 1944a, 1944b, and 1944c). 

M.4RIANA 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). 



12 



University of Kansas Publs., Mus. Nat. Hist. 



20 



18 



M5 



16 



URACAS . 

MAUG • 
•ASUNCION' 



AGRIHAN"^ 



PAGAN ^. 

AL AM AG AN • 
GUGUAN' 

SARIGAN. 
ANATAHAN- 

MEDINILLA' i 



SAIPAN^ 

7INIANQ 
AGIGUAN" 



ROTA 



'^ 



GUAM^O 2550 MILES 



20 



145 



Fig. 2. The Mariana Islands. 

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, Zar- 
pane). 

Saipan (also called (Saepan, St. Joseph, San Jose, 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 Pajaros, Pajaros, Urakasu). 



Baker: The Avifauna of Micronesia 



13 



Palau Islands 

The Palau Islands (also called Arrecifos, Palaos, Paleu, Pally, Paloc, Pan- 
nog, Parao, Pelew) consist of 8 large islands, 18 smaller islands, and a large 
number of minute islets, all enclosed in a single reef S3'stem. 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, 



s-oo 



134° 20- 



I34°40- 



KAYANGEL' 



•040- 



7»20 



7°00' 



BABELTHUAP 




7'W 



KOROR 
'ARAKABESAN 



URUKTHAPEL 



," ^ 



EIL MALK 



tJ 



sOGARAKAYO 
t>^ NGESEBUS 



PELELIU 



10 



20 MILES 



P 



ANGAUR 



8°oa 



7'za 



— TOO' 



I34»20' 



I34°40' 



Fig. 3. The Palau Islands. 



14 University of Kansas Publs., Mus. Nat. Hist. 

may be included with the Palau Archipelago. From the standpoint of the avian 
zoogeography, the coral islands or atolls of Kayangel, Merir, Pulo Arma, 
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). 

Ell Malk (also called Amototi, Cogeal, Irakong, Makarakaru). 

Garakayo 

Koror (also called Coror, Goreor, Kororu). 

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

Ngahad 

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, Kreian- 
gel. Moore, Ngaj angel, Ngeiangel). 

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

Pulo Anna (also called Anna, Bui, 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 Carolme Islands consist of 41 island clusters or isolated islands (exclu- 
ive 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, 
Ponape, 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, Lutke, Rukutee). 

Eauripik (also called Aurepik, Eourpyg, luripik, 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, Gurii- 
nitchi, Kabeneylon, Kapenmailang, Makarama, Pikiram, Tenuv). 

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



Baker: The Avifauna of Micronesia 15 



10 


140 145 150 


65 160 


10 


■1 1 1 

•Nqulu Foroulep' . -■. -^ > 






. WestFoyu- •P'l^'ol 








Woleo,.- .„,, '-l-amotrek .. ^-.Truk 


'* 






Eounpik - • 


•_,4 Ponape 






- 


• Pmgelop 






•' 


'^"^^ Kusce. 




5 




> Nukuoro 


5 




50 OO 200 MILES 


* Kopingomofongi 









1 1 1 


, , 





140 145 60 


155 160 



Fig. 4. The Caroline Islands. 

Lamotrek (also called Lamorsu, Lamureck, Lamutrik, Low, Namotik, Namo- 
tikku, Manochikku, Namurrek, Swede). 

Lukunor (also called Leraarafat, Lougoullos, Lougounor, Luganor, Lugunor, 
Lugunoz, Mortlock, Namonefeng, Rukunoru, Youngwilliam). 

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

Ngulu (also called Angogul, 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, Sail- 
rocks, Tucks Reef). 

Ponape (also called Ascension, Bonabee, Bonybay, Faloupet, Faounoupei, 
Funopet, Niponpei, Painipete, Ponapi, Piunipet, Puynipet, Quirosa, Seniavin, 
William IV). Ponape 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). 



16 



University of Kansas Publs., Mus. Nat. Hist, 





60 


165 


170 


175 


" ' 




1 

50 100 200 MILES 
1 1 1 1 


1 












»B1KAR 










BIKINI 
•■ ENIWETOK '. : 


.RONGELAP 










-- 


■.; ■• ^.•. 






10 




\ i 


LIKIEB {_^ ■ 

•.VWOTJE 
~ 'KWAJALEIN 

* ., 5 

\NAMU 

--' ELMORE ,. j^ARHNO 
MAJURO" ■" "-^^ 




10 


5 


- 


1 


^•-JALUIT OMILLE 
NAMORIK 

^EBON 

1 


- 


5 


1 


60 


165 


I7D 


«„. I7S 


1 



Fig. 5. The Marshall Islands. 

i4r/ino (also called Arno, Aruno, Auru). 

Bikar 

Bikini 

Ebon (also called Boston Atoll). 

Elmore (also called Ailinglap, Ailinglapalap, Iringlob). 

Eniwetok 

JaZmi (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, Name). 

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 evolu- 



Baker: The Avifauna of Micronesia 17 

tion and differentiation. Remains of stone walls, dikes, fences, pil- 
lars, graves, and other structures which may be found today at vari- 
ous 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 Mi- 
cronesia. 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 ascer- 
tained. 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 Mar- 
shall Islands; visits were made also to Guam and Rota and to Yap, 
Pais, Ulithi, Palau, and other island groups in western Micronesia. 
Freycinet's famous expedition in the ships "Uranie" and 'Thysi- 
cienne," visited Guam, Rota, and Tinian in 1819. Quoy and Gai- 
mard, the naturalists of the expedition, obtained birds, which were 
among the first to be described from Micronesia. These two natu- 
ralists revisited the Marianas in 1829 on board the ship "Astrolabe." 

2—8131 



18 University of Kansas Publs., Mrs. Nat. Hist. 

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 Lutke obtained birds at Kusaie in De- 
cember 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 cor- 
vinus) which have not been obtained since his time. His specimens 
were deposited in St. Petersburg. He was one of the most compe- 
tent 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 "Zelee" 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 Ponape, 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 representa- 
tive of Godeffroy and Sons at Yap in 1869 and obtained birds. Per- 
haps the most famous collector in this period was Johann Kubary. 
He went to Ponape at the age of nineteen and traveled in Micro- 
nesia 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) re- 
ported 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. 



Baker: The Avifauna of Micronesia 19 

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 approx- 
imately one year, 1899-1900, at Jaluit in the Marshalls and obtained 
records of birds. In 1899, Brandeis, on board the German ship 
"Kaiseriand" visited many of the islands in the Marshalls and re- 
corded 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 Decem- 
ber, 1931, with William F. Coultas as collector. Although experienc- 
ing some difficulty and being restricted somewhat in his travels by 
the Japanese officials, he managed to obtain collections at Ponape 
(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 



20 University of Kansas Publs., Mus. Nat. Hist. 

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 Ponape 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 Rhipidiira nifijrons 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, in- 
cluding Kittlitz, Lesson, Bonaparte, and Pelzeln. In the period from. 
1861 to 1880, thirty-four birds were newly named, mostly by Hart- 
laub and Finsch, from the collections which the Godeffroy Museum 
obtained through the efforts of Kubary, Tetens, Peters, and Hein- 
sohn. 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. 



Baker: The Avifauna of Micronesia 



21 



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. 



Years 


No. of 
descriptions 


Years 


No. of 
descriptions 


1821-1830 


8 
8 
4 
5 
11 
23 


1881-1890 

1891-1900 


9 


1831 1 40 


10 


1841-1850 


1901-1910 


4 


1851-1860 


1911-1920 


10 


1861-1870 


1921-1930 


15 


1871-1880 


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 Micro- 
nesia belong to 91 genera of 37 families of 13 orders. In the fol- 
lowing list, nonresident birds are marked with an *; birds intro- 
duced by man are marked with a f. 

Class AVES— birds 

PAGE 

Order Procellariiformes — albatrosses, petrels, and allies 
Family Diomedeidae — albatrosses 

Diomcdia nigripes Audubon* _ Black-footed Albatross 63 

Family Procellariidae — petrels and shearwaters 

Puff inus pad ficMs chlororhT/nchus Lesson Wedge-tailed Shearwater.. 64 

Puffinus pacificus cuncaliis SaWin Wedge-tailed Shearwater.. 65 

Puffinus tenuirostris (Temminck)* Short tailed Shearwater 66 

Puffinus nativitatus Streets Christmas Shearwater 66 

Pxiffinu-Fi Iherminieri dichrous Finsch 

and Hartlaub Dusky Shearwater 66 

Pterodroma rostrata rostrata (Peale)* Tahiti Petrel 69 

Pterodroma hypoletica hypoleuca Salvin Stout-billed Gadfly Petrel. . 70 



22 



University of Kansas Publs., Mus. Nat. Hist. 



Order Pelecaniformes — tropic birds, boobies, cormorants, frigate 

birds and allies 

Family Phaethontidae — tropic birds 

PAGE 

Phaethon aethereus mesonauta Peters* Red-billed Tropic Bird. .. . 70 
Phaethonnibricauda rothschildi (Mathev/s) Red-tailed Tropic Bird.... 71 
Phaethon 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 

Egretla intermedia intermedia (Wagler)* Plumed Egret 82 

Demigretta sacra sacra (Gmelin) Reef Heron 84 

Nycticorax nyclicorax nycticorax 

(Linnaeus)* Black-crowned Night 

Heron 87 

Nycticorax caledonicus pelewensis Mathew Rufous Night Heron 87 

Gorsachius goisagi (Temminck)* Japanese Bittern 89 

Gorsachius melanolophus melanolophv^ 

(Raffles)* Malay Bittern 90 

I xobrychus 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 ou^taleti 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 



Baker: The Avifauna of Micronesia 23 

Order Falconiformes — vultures, hawks, falcons 

Family Accipitridae — hawks, harriers, and allies ^^^^ 

Accipiter soloensis (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 Graelin* Peregrine Falcon 105 

Order Galliformes — megapodes, pheasants, and allies 

Family Megapodidae — megapodes 

Megapodius laperouse senex Hartlaub Micronesian Megapode 106 

Megapodius laperouse laperouse Gaimard Micronesian Megapode. .. . 109 

Family Phasianidae — quails, pheasants, and allies 

Coturnix chinensis lineata (Scopoli)t Painted Quail 113 

Gallus gallu^ (Linnaeus)t Red Jungle Fowl 114 

Phasianus colchicus Linnaeus t 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 jasciata (Raffles)* Malay Banded Crake 120 

Rallina eurizono'ides eurizono'ides 

(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 

Porphyria porphyria 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 cur onicms GmeVm* Ring-necked Plover 135 

Charadrius alexandrinv^ nihonensis 

Deignan * Kentish Plover 135 



24 



University of Kansas Publs., Mus. Nat. Hist. 



PAGE 

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 viadagascariensis (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 gJarcola Linnaeus * Wood Sandpiper 142 

Actitus hypoleucos lA\nnSie\xs* Common Sandpiper 143 

Heteroscelus brevipes (Vieillot) * Gray-tailed Tattler 144 

Heteroscelus incanus (Gmelin) * Amer. Wandering Tattler. . 145 

Arenaria interprcs 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 siibminuta (Middendorff) * Least Sandpiper 152 

Erolia melanotos (Vieillot) * Pectoral Sandpiper 152 

Erolia acuminata iiloTsfield) * 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 argcntatus vegae Palmen * Herring Gull 154 

Chlidonias leucoplerus (Temminck) * White-winged Black Tern. . 155 

Sterna hirundo longipennis "Nor draann* 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 juscata oahuensis Bloxham Sooty Tern 161 

Sterna albijrons sinensis Gmelin * Least Tern 161 

Thalasseiis bergii pelecanoides (King) Crested Tern 162 

'•'■' Procelsterna cerulea saxatilis 

W. E. Fisher * Blue-gray Tern 164 

•- Anoiis stolidus pileatus (Scopoli) Common Noddy 165 

Anoi'is tenuirostris marcusi (Bryan) White-capped Noddy 170 

' Gygis alba Candida (Gmelin) White Tern 174 

' Gygis alba pacifica (Lesson) White Tern 180 



Baker: The Avifauna of Micronesia 25 

Order Columbiformes — pigeons, doves, and allies 
Family Columbidae — pigeons and doves 

PAGE 

C olumba livia Gmelin t Blue Rock Pigeon 182 

Ptilinopus porphyraceus ponapensis 

(Finsch) Crimson-crw'd Fruit Dove, 182 
Ptilinopus porphyraceus hemsheimi 

(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 toumsendi (Wetmore) Micronesian Pigeon 194 

Ducula oceanica oceanica (Lesson and 

Garnot) Micronesian Pigeon 195 

Ducula oceanica ratakensis (Takatsukasa 

and Yamashina) Micronesian Pigeon 197 

Streptopelia hitorquata dusumieri 

(Temminck)t Philippine Turtle Dove.. . . 198 
Gallicolumba canijrons (Hartlaub and 

Finsch) Palau Ground Dove 201 

Gallicolumba xanthonura xanthonura 

(Tcmminck) 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) Ponape 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 podarginm (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 



26 University of Kansas Publs., Mus. Nat. Hist. 

Order Apodiformes — swifts and hummingbirds 
Family Apodidae — swifts 

PAGE 

Collocalia inexpectata pelewensis Mayr Edible Nest Swiftlet 221 

Collocalia inexpectata bartschi Meams 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 cinnamomiyia 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 macrocercu^ harterti S. Baker t 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 

(Miiller) * Chinese Blue Rock Thrush, 248 

Turdus obscurus obscurus Gmelin * Dusky Thrush 248 



Baker: The Avifauna of Micronesia 



27 



Family Sylviidae — Old World warblers 



PAGE 



Psamathia annae Hartlaub and Finsch 
Acrocephalus luscinia liLScinia (Quoy and 

Gaimard) 
Acrocephalus luscinia syrinx (Kittlitz) 
Acrocephalus luscinia yamashinae 

(Takatsukasa) 
Acrocephalus luscinia nijoi (Yamashina) 

Family Muscicapidae — Old World flycatchers 



Palau Bush-warbler 249 



Nightingale Reed-warbler. . 251 
Nightingale Reed-warbler. . 254 



Nightingale Reed-warbler. 
Nightingale Reed-warbler. 



Rhipidura rufifrons uraniae Oustalet 
Rhipidura rufifrons saipancnsis Hartert 
Rhipidura rufifrons mariae R. H. Baker 
Rhipidura rufifrons versicolor Hartlaub 

and Finsch 
Rhipidura rufifrons kubaryi Finsch 
Rhipidura lepida Hartlaub and Finsch 
Metabolus rugensis (Hombron and 

Jacquinot) 
Monarcha godcfjroyi Hartlaub 
Monarcha takatsukasae (Yamashina) 
Myiagra oceanica erythrops 

Hartlaub and Finch 
Myiagra oceanica freycineti Oustalet 
Myiagra oceanica oceanica Pucheran 
Myiagra oceanica pluto Finsch 
Muscicapa narcissina narcissina 

Temminck * 
Muscicapa griseisticta (Swinhoe) * 

Colluricincla tenebrosa (Hartlaub and 
Finsch 

Family Artamidae — wood-swallows 
Artamus leucorhynchus pelewensis Finsch 

Family Sturnidae — starlings 

Aplonis opacus opacus (Kittlitz) 
Aplonis opacus ponapensis Takatsukasa 

Yamashina 
Aplonis opacus angus Momiyama 
Aplonis opacus kurodai Momiyama 
Aplonis opacus orii (Takatsukasa and 

Yamashina) 
Aplonis opacus guami Momiyama 
Aplonis opacus aeneus (Takatsukasa and 

Yamashina) 
Aplonis pelzelni Finsch 
Aplonis corvinus (Kittlitz) 
Sturnu^ philippensis (Forster)* 
Sturnus cineraceus Temminck* 



Rufous-fronted Fantail . 
Rufous-fronted Fantail. 
Rufous-fronted Fantail. 



256 
257 



261 
262 
263 



Rufous-fronted Fantail 264 

Rufous-fronted Fantail 265 

Palau Fantail 266 

Truk Monarch 269 

Yap Monarch 272 

Tinian Monarch 274 

Micronesian Broadbill 275 

Micronesian Broadbill 277 

Micronesian Broadbill 279 

Micronesian Broadbill 280 

Narcissus Flycatcher 282 

Chinese Gray-spotted 

Flycatcher 282 

Palau Morning Bird 282 

White-breasted Wood- 
swallow 284 

Micronesian Starling 286 

Micronesian Starling 288 

Micronesian Starling 289 

Micronesian Starling 291 

Micronesian Starling 292 

Micronesian Starling 293 

Micronesian Starling 297 

Ponape Mountain Starling, 299 
Kusaie Mountain Starling. . 301 

Violet-backed Starling 302 

Ashy Starling 302 



28 



University of Kansas Publs., Mus. Nat. Hist. 



Family Meliphagidae — honey-eaters 

Cleptornis marchei (Oustalet) 
Myzomela cardinalis rubratra (Lesson) 
Myzomela cardinalis dichromata Wetmore 
Myzomela cardinalis major Bonaparte 
Myzomela cardinalis safjordi Wetmore 
Myzomela cardinalis kurodai Momiyama 
Myzomela cardinalis kobayashii 
Momiyama 

Family Zosteropidae — white-eyes 
Zosterops conspicillata conspicillata 

(Kittlitz) 
Zosterops conspicillata saypani Dubois 
Zosterops conspicillata rotensis 

Takatsukasa and Yamashina 
Zosterops conspicillata semperi Hartlaub 
Zosterops conspicillata owstoni Hartert 
Zosterops conspicillata takatsukasai 

Momiyama 
Zosterops conspicillata hypolais 

Hartlaub and Finsch 
Zosterops cinerea cinerea (Kittlitz) 
Zosterops cinerea ponapensis Finsch 
Zosterops cinerea finschii (Hartlaub) 
Rukia palauensis (Reichenow) 
Rukia oleaginea (Hartlaub and Finsch) 
Rukia ruki (Hartert) 
Rukia sanfordi (Mayr) 

Family Ploceidae — weaver-finches 

Erythrura trichroa trichroa (Kittlitz) 
Erythrura trichroa clara Takatsukasa 

and Yamashina 
Erythrura trichroa pelewensis Kuroda 
Lonchura nigerrima minor (Yamashina) 
Lonchura punctulata cabanisi (Sharpe)t 



PAGE 

Golden Honey-eater 302 

Cardinal Honey-eater 304 

Cardinal Honey-eater 307 

Cardinal Honey-eater 307 

Cardinal Honey-eater 309 

Cardinal Honey-eater 312 

Cardinal Honey-eater 313 

Bridled White-eye 31G 

Bridled White-eye 318 

Bridled White-eye 319 

Bridled White-eye 320 

Bridled White-eye 321 

Bridled White-eye 322 

Bridled White-eye 323 

Micron. Dusky White-eye, 326 
Micron. Dusky White-eye, 327 
Micron. Dusky White-eye, 328 
Palau Greater White-eye.. 330 
Yap Greater White-eye... 331 
Truk Greater White-eye... 332 
Ponape Greater White-eye, 333 

Blue-faced Parrot-finch 336 

Blue-faced Parrot-finch 337 

Blue-faced Parrot-finch.... 338 

Black-faced Weaver-finch . . 339 

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 sys- 
tem used by Mayr (1945a). 

Oceanic Birds 
Oceanic birds found in Micronesia belong to the following fam- 
ilies: Diomedeidae, Procellariidae, Phaethontidae, Pelecanidae, 



Baker: The Avifauna of Micronesia 



29 



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, Phae- 
thontidae). As shown in table 2 there are 30 kinds of oceanic birds 
in Micronesia, 18 kinds that are resident and 12 kinds that are re- 
garded 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 
kinds 


Nonresident 
kinds 


Dinme.ddn, 



4 
1 
2 
3 
1 


2 
1 

2 
2 


1 


PuMnus 


1 


Pterodroma 


1 


Phaethon . 


1 


Sula 





Fregata 


1 


Larus 


I 


Chlidonias 


1 


Sterno, 


4 


Thalasseiis 





Procelsterna 


1 


Atious 





GvQZS 










Inshore Oceanic Birds 

The inshore zone, according to Wynne-Edwards (1935:240), ''ex- 
tends from high-water mark to a maximum of four or five miles out 
to sea, including islands and reefs within sight of shore." In Micro- 
nesia the majority of the Laridae occur in this zone including such 
residents as Sterna sumatrana, S. anaetheta, Thalasseus bergii, 
Anoils stolidus, A. tenuirostris, Gygis alba. These birds, especially 
*S. anaetheta, Thalasseus, and Anoils, may venture into the offshore 
zone. Visitors to Micronesia include several terns which probably 



30 University of Kansas Publs., Mus. Nat. Hist. 

normally range in the inshore (as well as in offshore) zones, such as 
Childonias leucopterus and Steima 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 extend- 
ing to the continental edge ; however, in Micronesia where small is- 
lands 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 argen- 
tatus), 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 {Puffi- 
nus and Pterodroma), and possibly the tropic birds (Phaethon). 

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 de- 
posits 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.), 



Baker: The Avifauna of Micronesia 31 

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 deposi- 
tion by birds with a population similar to that found there today. 

The oceanic avifauna of Micronesia contains birds which are ap- 
parently 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 (espe- 
cially Palearctica) reach the northern and western edges of Micro- 
nesia as winter visitors. These include Larus argentatus, Chlidonias 
leucopterus, and Sterna hirunxlo. Another northern gull, Larus ridi- 
bundus, has been reported in the Marianas. 

One bird of the North and Northcentral Pacific, Diomedea ni- 
gripes, 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 Micro- 
nesia, either as visitors or residents, include Puffinus tenuirostris, P. 
nativitatis, Pterodroma rostrata, P. hypoleuca, Sterna lunata, and 
Procelstema 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 terms. 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. Ihermi- 
nieri, Phaethon, Sula, Fregata, Sterna anaethetus, S. fuscata, Anoiis 
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. 



32 



University of Kansas Publs., Mus. Nat. Hist. 



Migratory Shore Birds 

Twenty-eight species of shore birds of the families Charadriidae 
and Scolopacidae have been recorded from Micronesia, and one 
other of the family Phalaropodidae apparently occurs in the area, 
making a total of 29 kinds. From the entire Southwest Pacific, 
Mayr (1945a :28-47) lists 31 species and subspecies of shore birds 
and mentions six other species which may occur there. Thus, of a 
possible 37 kinds of shore birds in this large area (which includes 
Micronesia), 29 are present in the islands of Micronesia. For pur- 
poses of discussion, shore birds are here placed in one of two groups: 
regular visitors or uncommon visitors. A regular visitor is one 
which has been recorded in the literature or in unpublished field 
accounts as being frequently observed in Micronesia in periods of 
migration. An uncommon visitor is one which has been infrequently 
observed in Micronesia. Of the 28 kinds of shore birds recorded 
from Micronesia, 17 are classed as regular visitors and 11 are 
classed as uncommon visitors. 

Original Homes of the Shore Birds that Visit Micronesia 

The shore birds which are known to visit Micronesia breed in the 
Northern Hemisphere. Table 3 summarizes the data concerning the 



Table 3. Breeding and Wintering Grounds of the Species of Migr.atory 

Shore Birds in Micronesia 

Part A. Location of breeding grounds 



Class 


Number 


Circum- 
polar* 

5 
2 


Asiatic 


American 


Regular visitors 


17 
11 


10 

8 


2 


Uncommon visitors 


1 






Totals 


28 


7 


18 


3 







Part B. Location of wintering grounds 



Class 


Number 


Circum- 
tropical 


Asiatic 


American 


Oceanic 


Regular visitors .... 
Uncommon visitors. . 


17 
11 


2 

1 


13 

8 


1 
2 


1 



Totals 


28 


3 


21 


3 


1 







•Denotes birds which breed on both American and Asiatic sides of the Pacific Ocean. 



Baker: The Avifauna of Micronesia 33 

breeding and wintering areas of these birds. As shown in part A 
of table 3, 18 of the 28 species which visit Micronesia come from 
Asiatic breeding grounds. Seven have circumpolar breeding ranges 
and three (two are regular visitors) come from American breeding 
grounds. As shown in part B of table 3, 21 of the 28 waders have 
their winter ranges on the Asiatic side of the Pacific with eastern 
extensions to Micronesia and other parts of Oceania. Of the 7 re- 
maining species, the winter ranges of three are circumtropical ; the 
winter range of a fourth is restricted to Oceania; and the winter 
ranges of the remaining three (two classed as uncommon visitors) 
are American. 

Bryan and Greenway (1944:109-115) record 14 species of shore 
birds from the Hawaiian Islands, One of these, Himantopus himan- 
topus knudsoni, is a resident, probably of New World origin, accord- 
ing to Mayr (1943:56). The others, listed in table 7, include three 
species unknown in Micronesia. One of these, Phalaropus fulicarus, 
apparently winters at sea off the west coast of South America. The 
other two species {Charadrius vociferus vociferus and Gallinago 
delicata) are classified by Bryan and Greenway as "accidental" and 
"occasional" visitors from North America. The ten species com- 
mon to both the Hawaiian Islands and Micronesia include seven 
whose breeding grounds are circumpolar, two whose breeding grounds 
are in Arctic America and one whose breeding ground is in Arctic 
Asia. The winter ranges of these ten species include four which are 
circumtropical, three which are Asiatic, one which is restricted to 
Oceania, and only two which are American. 

The ability of the shore birds to migrate almost as well over water 
as over land may explain their spread into Oceania. The likelihood 
that shore birds, when migrating may have ventured to ]\Iicronesia 
and Polynesia initially from the Asiatic side of the Pacific is strongly 
suggested by the data given in the paragraph above. Also, on the 
Asiatic side of the Pacific there are large numbers of islands, which 
form several archipelagoes extending from Kamchatka south to 
Malaysia. Once accustomed to migrating along these chains of 
islands from the Arctic to Australia, birds would probably have to 
make only minor adjustments to extend the breadth of their migra- 
tory routes eastward into the islands of the Pacific Ocean. In con- 
trast, on the Pacific coast of North America there are few coastal or 
offshore islands and there is a vast area of open water separating the 
Hawaiian Islands from the American mainland. Probably the vast- 
ness of this area of water offers little stimulus to birds to expand 

3—8131 



34 



University of Kansas Publs., Mus. Nat. Hist. 



their migratory ranges westward, and in part accounts for the small 
North American contingent in the population of shore birds of the 
Central Pacific. Some North American shore birds do visit the Pa- 
cific. The brisk trade winds from the northeast might be an aid to 
the birds in their flights from Nearctica to Hawaii. 

The long flight now made by shore birds going from the Aleutians 
to the Hawaiian Islands may have commenced as a gradual expan- 
sion from the west, or perhaps such a route was initiated by birds 
flying northward through the Hawaiian Chain to the Arctic in mi- 
grating to their breeding grounds, and then later returning via the 
same route to reach their wintering grounds. 

Routes of Migration 

The small and isolated islands of Oceania might, upon first inspec- 
tion, seem to offer but little attraction to shore birds. Hesse, Allee 
and Schmidt (1937:172, 173) point out that the "open southeastern 
Pacific" being least supplied with water from land sources, which is 
an important means of fertility, is known to have one of the poorest 
faunas found anywhere in the oceans. However, there are extensive 
tidal flats, especially on the leeward sides of the islands, and these 




Fig. 6. Routes of migration used by shore birds in the Pacific area. From 
west to east these are: The Asiatic-Palauan Fly way, the Japanese-Marianan 
Flyway, the Nearctic-Hawaiian Flyway. 



Baker: The Avifauna of Micronesia 35 

flats apparently afford extensive feeding grounds for these birds. 
Also, the absence of competition from resident birds as well as the 
virtual absence of predatory animals (native man and his domesti- 
cated animals excepted) are other factors which may help to make 
the islands attractive wintering grounds for shore birds. 

Only a few birds have been banded in the Pacific, and the knowl- 
edge which comes from the recovery of banded birds gives but little 
aid to the student of movements of birds in the Pacific. The prob- 
able flyways for migratory shore birds there have to be deduced from 
sight records, data from specimens collected, known stations of 
breeding and wintering (summarized by Peters, 1934:234-293), and 
from a study of maps of the region. Analysis of information from 
the above-mentioned sources indicates that there are three routes 
taken by shore birds which migrate from Micronesia to and from 
their northern breeding grounds (see figure 6) : (1) Asiatic-Palauan 
Flyway; (2) Japanese-Marianan Flyway; (3) Nearctic-Hawaiian 
Flyway. 

1. Asiatic-Palauan Flyway. For shore birds, there appears to 
be a migration route extending almost due south from the Riu Kiu 
and the Japanese islands to the Palau Islands. Some birds may 
migrate via the Philippines and others may pass to the east of the 
Philippines. This route is considered to be distinct from that used 
by birds which follow the Asiatic Coast and coastal islands, because 
the Palau Islands are situated approximately 600 miles east of the 
Philippines. Moreover, there are fewer species — only 20 recorded 
from the Palaus as compared with the number recorded from islands 
closer to the mainland of Asia. Delacour and Mayr (1946:68-74) 
list 46 species of shore birds from the Philippines; the Hand-list of 
Japanese Birds (Hachisuka et al, 1942) lists 34 species from the 
Riu Kiu Islands. 

The information available indicates that migrant shore birds 
which utilize this flyway move east into the Carolines (examples, 
Tringa nebularia, Charadrius leschcnaultii) ; however, the recording 
of 20 species from the Palaus as compared with only 12 species in 
the western Carolines (table 4) indicates that this spread eastward 
may not be very pronounced. Migrants in autumn probably move 
from the Palaus in a southerly direction toward the New Guinea 
area. Eight species of shore birds which reach the Palaus (and 
adjacent islands in the western Carolines), are not recorded from 
other parts of Micronesia. Species which apparently utilize the 
Asiatic-Palauan Flyway are listed in table 5. 



36 University of Kansas Publs., Mus. Nat. Hist. 

2. Japanese-Marianan Flyway. Shore birds from Asiatic, and 
probably Aleutian and Alaskan, breeding grounds may follow the 
Asiatic Coast or the adjacent island chains southeast to the Japa- 
nese Archipelago. From there some of the birds apparently fiy 
south through the Bonin and Volcano islands to the Marianas, from 
where they may spread in fanlike fashion to the southeast, south 
and southwest, even reaching to the Palau Islands (example, Heter- 
oscelus incanus) . The number of species of shore birds recorded 
from the Marianas (see table 4) is greater than that found in the 
Carolines, but it must be remembered that more intensive investiga- 
tions have been made by ornithologists in the Marianas, which 
might account for the recording of more species (especially strag- 
glers, such as Gallinago gallinago) . Species which apparently use 
this flyway are named in table 6. 

3. Nearctic-Hawaiian Flyway. Shore birds from breeding 
grounds in western Canada, Alaska, the Aleutians, the Bering Sea 
area, and probably northeastern Asia may fly in a southerly direc- 
tion along a broad front to the Hawaiian Islands. This flyway is 
probably the one which supplies to central and eastern Oceania the 
largest wintering populations of shore birds. From the Hawaiian 
Islands birds may fly directly south through the scattered islands 
to southern Polynesia, or they may fly in a southwesterly direction 
and reach the Marshall Islands. The shore birds which visit tlie 
Marshall Islands apparently move south through the Gilbert, Ellice 
and other more southern island groups rather than west into the 
Carolines as exemplified by the fact that Numenius tahitiensis, a 
characteristic migrant through the Marshalls from the Hawaiian 
Islands, is rarely found west of the Marshall Islands in Micronesia. 
Species which apparently use this flyway are listed in table 7. 

Flyways additional to the three suggested above may be utilized 
by some shore birds on their southward (and northward) migrations. 
Species reaching Wake and the Marcus Islands may fly directly 
south from the islands of the North Pacific. Bryan (1903:115, 116) 
lists four species of shore birds from Marcus {Erolia acuminata, 
Heteroscelus incanus, Pluvialis dominica, Arenaria interpres). 

Table 4. List of Species of Shore Birds Known From Five Geographical 

Areas of Micronesia 

Western Central Eastern 

Palaus Carolines Marianas Carolines Carolines Marshalls 

Number of species 20 12 17 11 10 10 



Baker: The Avifauna of Micronesia 



37 



Table 5. Shore Birds Which May Use the Asiatic-Palauan Flyway 



Uncommon? Visitors 
Charachius dubius curonicus 
Charadrius alexandrinus 
Calidris tenuirostris 
Erolm jerruginea 
Erolia suhminuta 
Limicola jalcineUus sibirica 



Regular Visitors 
Pluvialis dominica fulva 
Charadrius mongolus stegmanni 
Charadrius leschenaultii 
Numenius phaeopus variegaius 
Numenius madagascariensis 
Limosa lapponica baueri 
Tringa nebularia 
Tringa glareola 
Actitis hypoleucos 
Heteroscelus brevipes 
Arenaria i. interpres 
Gallinago megala 
Erolia minuta ruficoUis 
Erolia acuminata 

Table 6. Shore Birds Which May Use the Japanese-Marianan Flyway 



Regular Visitors 
Pluvialis dominica julva 
Charadrius m.ongolus stegmanni 
Numenius phaeopus variegatus 
Limosa lapponica baueri 
Actitis hypoleucos 
Heteroscelus brevipes 
Heteroscelus incanus 
Arenaria i. interpres 
Gallinago megala 
Crocethia alba 
Erolia acuyninata 

Table 7. Shore Birds Which May Use the Nearctic-Hawaiian Flyway 



Uncommon? Visitors 

Sqimtarola squatarola 

Numenius tahitiensis 

Nu m e n ins madagascariensis 

Tringa glareola 

Gallinago gallinago gallinago 

Erolia minuta ruficoUis 



Regular Visitors 
Pluvialis dominica julva* 
Numenius tahitiensis* 
Heteroscelus incanus* 
Arenaria i. interpres* 
Crocethia alba* 
Phalaroptis julicarius 
Phalaropus lobatus*? 



Uncommon? Visitors 
Squatarola squatarola* 
Charadrius hiaticula semipalmatus^ 
Charadrius r\ vocijerus 
Limosa lapponica baucii* 
Tringa mclanolerica*^ 
Gnllinago delicala 
Erolia melanotos* 
Erolia acuminata* 



•Indicates species which are found in Micronesia. 

tindicates species not recorded from the Hawaiian Islands; see Brvan and Greenway (1944: 
109-115). 

Populations of Shore Birds in Micronesia 

Although shore birds have been observed in Micronesia on many 
occasions, actual counts of numbers of individuals of the different 
birds have rarely been made. Kubary, Finsch, Marche, Scale and 
other early collectors and observers record some data of this kind 



38 



University of Kansas Publs., Mus. Nat. Hist. 



as have the Japanese investigators in later times. William Coultas 
of the Whitney South Sea Expedition obtained considerable infor- 
mation of this nature at Guam, Saipan, Kusiae, Ponape, and the 
Palaus, but it is unpublished. His records were made in fall, winter 
and spring, when migrants were present in large numbers and these 
observations offer evidence that many of the migrants are com- 
paratively numerous, especially in the Carolines, throughout the 
winter months. McElroy's observations made on his trip for 
NAMRU2 to Truk in December, 1945, offer further evidence of this. 

Table 8. Populations of Migratory Shore Birds Seen at Guam in 1945 

























CO 


























03 
















-(— 


.f.— 








3 








•I 

s 


00 


§ 

o 


d 


=2 


CO 

=1, 




CO 




"2 


03 




•5 


o 


c 


s 


Q. 


s 


!~. 


s 


Sr 




*-< 


tn 




s 




^ 


^ 


m 


■(?» 




o 


<u 










-§ 


so 


55, 

2 


o 
a. 


=0 


« 

"*^ 
lij 


■to 


a. 

a. 


53 


-% 

yn 


O 

d 


O 

d 




•?* 






-c 


CO 






c 


s 


r^ 


^ 


^ 




e 


« 


<i> 




o 


O 


g 


CO 




OJ 


, ^ 


, ^ 




S 


S 


s 


."S 








O 


g 


TS 


oj 


c3 




^ 


« 






■♦.A 






jt 


■« 






*i 






.<; 




o 


15 


1) 






s^ 


^ 


O 


O 




Q, 


o 


:^ 


^ 


&: 


^ 


^ 


^q 


'^ 


P 


H 


H 


March 11 


X 

10 

X 
X 




















X 

13 

X 

X 


1 


March 17 




1 

X 




2 

X 












3 


March 19 








X 




^ 


April 24 








1 


April 26 
















1 






1 


1 


May 19 










3 


?. 










5 


1 


May 21 










4 












4 


1 


May 26 










X 


?. 










X 


1 


June 1 






1 

X 

1 

12 
2 
















1 

X 

1 

14 
3 
2 
2 


1 


June 6* 








X 










X 


4 


June 11 








1 


June 12 








2 












? 


June 22 














1 

2 


2 


June 30 


















1 


July 7 






9. 














1 


July 8 


3 




X 




1 












,x 


3 


July 16* 


6 




3 


3 


4 












17 


4 


July 19 


X 




X 




X 












X 


3 


July 24* 


10 




6 




3 




?. 




3 




29 


fi 


July 26 






8 
















8 


1 


August 2 






X 
















X 


2 


August 3 






1 
















1 


1 


August 6* 






6 




1? 












IS 


2 


September 29 


X 




X 












X 




X 


2 


October 3* 


X 
X 








X 

X 












X 

X 


2 


October 10 




X 




2 






X 




4 


October 11 


















2 

X 




2 

X 


1 


October 23* 


X 




X 




X 


1 


1 




5 


October 24 


X 




















X 


1 

























X Observed but numbers not recorded. 

* Observations made on beach at Agfayan Bay area. 

t Figures based on identified skins. 



Baker: The Avifauna of Micronesia 39 

None of the above workers, however, obtained very much informa- 
tion on comparative numbers of species. 

Tables 8, 9, and 10 present the writer's findings on populations of 
migratory shore birds in Micronesia in 1945. At Guam, as shown 
in table 8, the records for March, April and early May are few, 
owing to a limited amount of field observation. Beginning in late 
May and until October 24 a greater amount of time was spent in 
the field and more regular records were obtained. No observations 
were made by the author at Guam in the period from August 11 to 
September 25. The dates marked with an asterisk are those on 
which observations were made on the extensive tidal flats at Ag- 
fayan Bay and vicinity. These fiats, at low tide, present excellent 
feeding grounds for waders and in 1945 were undisturbed by parties 
of service personnel, because the area was "off-limits." 

Table 8 shows that Pluvialis dominica, Numenius phaeopus, and 
Heteroscelus spp. were the shore birds most frequently found at 
Guam in this period. Pluvialis dominica was the most numerous of 
the three species. Of Heteroscelus there was approximately equal 
representation of H. incanus and H. brevipes as indicated by speci- 
mens collected. These birds were not identified to species in the 
field. 

Although records were made only infrequently in the spring mi- 
gration, such information as was obtained indicates that the popu- 
lations were largest in March and early April. On April 24, Pluvi- 
alis dominica was the only bird observed on beaches and in upland 
openings. On April 26, a single Limosa lapponica was recorded. On 
May 15, no shore bird was seen on a trip along several beaches. In 
late May and early June, single individuals of Heteroscelus were 
found. Of this genus, those collected in May were in nuptial plum- 
age, and those collected in June were in winter plumage and prob- 
ably should be classed as non-migrants. Numenius phaeopus was 
occasionally recorded beginning in early June, but waders were 
totally absent from beaches at Agfayan Bay and vicinity on June 
18 and 19. Few shore birds were seen in early August. In late 
September, birds, especially Pluvialis dominica, Numenius phaeopus, 
and Heteroscelus spp., were numerous. These species were numer- 
ous until October 24, when observations were discontinued. 

Of the 17 species of migratory shore birds recorded from the 
Mariana Islands, eight were identified. Of these eight, three species, 
Limosa lapponica, Actitis hypoleucos, and Charadrius mongolus, 
were found on only one occasion. Never more than four species 



40 



University of Kansas Publs., Mus. Nat. Hist. 



were identified on a single field trip. These data give an idea of 
the lack of variety of species that may be observed on Micronesian 
islands. 

Table 9. Populations of Migratory Shore Birds Seen at Ulithi Atoll 

IN 1945 





Island and Date 


Species 


Potangeras 


Fas- 
sarai 


Mange- 
jang 


Pau 


Losiep 




Aug. 
14 


Aug. 
15 


Aug. 
16 


Aug 
17 


Aug. 
19 


Aug. 
20 


Aug. 
21 


Aug. 
22 


Pluvialis 

dominica 






6 


5 


4 




10 

X 

1 
2 
6 

30 


5 


Charadrius 

mongolus 






2 


Numenius 

phaeopus 

Acititis 

hypoleucos 


1 


4 




1 






2 






2 


Heteroscelus 

snn 










2 


1 


3 


H. incanus * 










2 


Crocethia 

alba 












5 


















Total No. of 

Individuals .... 
Total No. of 

Species 


1 
1 


4 
1 


6 

1 


6 
2 


6 
2 


1 
1 


49 
6 


21 
6 



X Observed but numbers not recorded. 
* Figures based on identified skins. 

Table 9 lists the shore birds seen at Ulithi Atoll, Caroline Islands, 
on eight field excursions in the period from August 14 to August 22, 
1945. Of seven species of shore birds known to visit the atoll, six 
were taken in this period. As observed at Guam, Pluvialis dominica 
and Numenius phaeopus were the species most frequently found. 
Heteroscelus was seen on three occasions; those collected were iden- 
tified as H. incanus. Most of the shore birds were seen at Pau and 
Losiep, islands unoccupied by man. Similar tidal flats are present 
at most of the other small islands in the atoll, but these islands 
(Asor, Fallalop, Potangeras, Fassarai and Mangejang were visited) 
were occupied by small detachments of service personnel or by na- 
tives, which may have tended to keep many of the shore birds away. 
At the more populated islands of Asor and Fallalop, no shore birds 



Baker: The Avifauna of Micronesia 



41 



were seen. Almost as many species were recorded at Ulithi on the 
eight field trips as were found by the author at Guam in eight 
months of observations. 

Table 10. Populations of Migr.atory Shore Birds Seen at the Palau Islands 

IN 1945 





Peleliu 


Angaur 


Species 


August 


September 


Sept. 




24 


28 


1 


6* 


8* 


9t 


16* 


20* 


21 


Pluvialis dominica .... 
Charadrius mongolu . . . 


X 




X 


X 

X 
X 
X 


25 
25 
25 
30 

1 
3 


...... 


20 
5 
5 

20 

3 


X 
X 
X 
X 

15 


X 

X 


C. leschenaultii 








X 


Numenius phaeopus . . 








X 


N. madagascariensis . . . 








Limosa lapponica 












Tringa nebidaria 




6 










T. alareolat 














1 


Actitis hypoleucos 












2 








Heteroscelus sp 








X 

3 


75 

2 

20 


X 

2 


X 


X 


H brevipesX 








Arenaria inter pres 












Capella megala 
















4 


Calidris tenuirostris .... 










15 
50 




20 
50 






Erolia minuta 


.... 






X 


X 


X 


E. acuminatiX 








3 


E ferruainea t 








1 












Limicola falcinellusX ■ ■ ■ 
















1 


Unidentified 


.\ 






X 


X 




X 


X 


X 










Total number of 

individuals 

Total number of 

species 


X 

1 


6 
1 


X 

3 


X 

7 


271 + 
10 


3 

2 


129 + 
9 


X 

7 


X 

10 







* Observations made on beaches at Akarakoro Point, Peleliu. 
t Observations made at fresh water ponds. 
X Observed but numbers not recorded. 
t Figures based on identified skins. 

Table 10 presents field counts at the Palau Islands in the period 
from August 24 to September 21, 1945. Of 20 species of shore birds 
known from the Palaus, 17 species were collected or identified on 
this trip. It was apparent that the fall migration was at its height 
at this time. Birds were numerous at inland openings and ponds, 
air field strips, and on the extensive tidal flats at Akarakoro Point. 
The latter area is between Peleliu and the adjacent island of 
Ngesebus to the north. Several observations were made at this area 
(as indicated by the dates marked with asterisks in the table) ; on 
September 8, 271+ shore birds were counted; on September 16, 



42 University of Kansas Publs., Mus. Nat. Hist. 

129+ were counted. Six species were observed to be abundant. 
The majority of the birds found at these beaches were in small 
flocks which consisted of several birds of one or more species. 

The birds observed at Angaur on September 21 were seen at sev- 
eral fresh and brackish ponds. Four species {Tringa glareola, Erolia 
acuminata, Limicola falcinellus, Gallinago megala) , which were not 
taken on the tidal flats or elsewhere at Peleliu, were found at these 
ponds. 

The abundance, and more especially the variety, of shore birds at 
the Palau Islands during this period was in marked contrast to the 
smaller and less diversified populations of shore birds in rather simi- 
lar insular environments at Ulithi and Guam. These differences 
offer support for the supposition that the Asiatic-Palauan Migratory 
Shore Bird Flyway is distinct from the Japanese-Marianan Migra- 
tory Shore Bird Flyway. 

Land and Fresh-water Birds 

The land and fresh-water avifauna of Micronesia consists of 147 
kinds of birds. Of these, 37 kinds are non-residents, 104 kinds are 
residents, and 6 kinds have been introduced by man. The 104 resi- 
dent birds include 98 kinds (94 percent) which are found only 
within the confines of Micronesia. Included in these 98 kinds which 
are restricted to Micronesia are 5 endemic genera, 31 endemic spe- 
cies and 76 endemic subspecies. 

Gulick (1932: 407, 413) stresses that the fauna and flora of the 
oceanic islands may be "disharmonic" (he uses Easter Island as his 
example) and says, "It is evident that mature groups of islands will 
attain an internal harmony, from the standpoint of the systematist. 
But this harmony, instead of reflecting the pre-existing harmony of 
some continental source (as in the case of the continental islands or 
land-bridge remnants) will be recognizably derivable by descent 
from a quite limited number of original importations, at the start 
distinctly miscellaneous and 'disharmonic'." Analysis of the land 
and fresh-water avifauna of Micronesia supports Gulick's view. 

As mentioned previously, the islands of Micronesia, from the 
zoogeographical viewpoint, have been regarded as a part of the 
Polynesian Subregion of the Australian Region. Mayr (1941a: 192) 
defines the Polynesian Subregion as comprising "all the tropical and 
subtropical islands of the Pacific Basin which indicate by their im- 
poverished fauna that they have had no recent continental connec- 
tion (after early Tertiary) and which derived the major part of 



Baker: The Avifauna of Micronesia 



43 



their fauna directly or indirectly from the Papuan Region or jointly 
from Australia and the Papuan Region." As based on the distribu- 
tion of the resident avifauna, Mayr (1941a: 193) subdivides the 
Polynesian Subregion into the following districts: Micronesia ("in- 
cluding Palau, the Marianne, Caroline, Marshall, and Gilbert is- 
lands") ; Central Polynesia ("including Fiji, Tonga, Samoa, Phoe- 
nix, Ellice, Union islands, and a number of small islands, such as 
Rotuma, Fotuna, Keppel, Niue, Niouafu, and Uvea") ; Eastern 
Polynesia ("all the islands east of 165°W") ; and Southern Mela- 
nesia ("including the Santa Cruz group, Banks Islands, New Hebri- 
des, Loyalty Islands, and New Caledonia"). He considers that the 
Hawaiian Islands, Solomon Islands, and possibly New Caledonia 
are bordering districts to the Polynesian Subregion. Figure 7 show^s 




Fig. 7. Divisions of the islands of part of the Pacific Basin from the 
standpoint of the distribution of land birds and fresh-water birds: (1) Mi- 
cronesia; (2) Hawaii; (3) Central Polynesia; (4) Eastern Polynesia; (5) 
Southern Melanesia; (6) Melanesia. 



the divisions of the islands of the Pacific Basin from the standpoint 
of the distribution of the land and fresh-water birds. I have placed 
the Gilbert and Marshall islands in the Central Polynesian rather 
than in the Micronesian District. For purposes of discussion in this 
report, however, I am considering the Marshalls to be a part of 
Micronesia. The birdlife of the Bonin and Volcano islands north- 
ward of the Marianas is regarded as having its closest affinities to 
the Japanese avifauna. The Papuan or Melaneslan Subregion of 
the Australian Region includes the districts of New Guinea and 



44 University of Kansas Publs., Mus. Nat. Hist, 

Northern Melanesia, including the Bismarck Archipelago, the Ad- 
miralty Islands, and the Solomon Islands. 

The resident land and fresh-water birds of Micronesia have been 
derived from several sources. Studies of these birds and their 
closest relatives in adjacent areas indicate that the avifauna has 
been derived from five different sources: Polynesia, Melanesia, the 
Moluccas and Celebes, Philippines, and Palearctica. 

Polynesian Component 

Aphanolininas monasa (extinct ?), Ptilinopus porphyraceus, and 
Ducula oceanica are the only species of birds which have reached 
Micronesia directly from Polynesia. There are in Micronesia, as 
Mayr (1941b: 204) points out, eight species "which are members 
of typically Polynesian species or genera" and six species which are 
either Papuan or Polynesian. The relationships between Polynesian 
and Micronesian birds is evident, but insofar as the pathways of 
colonization are concerned the majority of these Micronesian species 
listed by Mayr have come from elsewhere than Polynesia and the 
birds of these two areas are thought to have arisen from common 
ancestors. Aphanolimnas, Ptilinopus, and Ducula apparently in- 
vaded Micronesia from Central Polynesia via the Marshall Islands 
through a rather continuous chain of islands and atolls. Aphano- 
limnas is known only from Kusaie in the extreme eastern part of 
the Carolines while Ptilinopus and Ducula are known from the 
Marshalls, Carolines, and Palaus. 

Melanesian Component 

The Papuan or Melanesian Region (New Guinea, Bismarck Archi- 
pelago, Solomon Islands) has supplied to Micronesia its greatest 
number of endemic land and fresh-water residents. Fifty kinds of 
birds belonging to the following species reached Micronesia from 
Melanesia: Nycticorax caledonicus, Megapodhis laperouse, Ptilino- 
pus roseicapillus, Gallicoluniba xanthonura, G. canijrons, Caloenas 
nicobarica, Halcyon cinnamomiyia, Trichoglossus rubiginosus, Col- 
localia inquieta, Edolisoma temdrostre, Rhipidura rufijrons, Metab- 
obis regensis, Monarcha godefjroyi, M. takatsukasae, C olluricincla 
tenebrosa, Aplonis opacus, A. pelzelni, A. corvinus (extinct ?), Clep- 
tornis marchei, Myzomela cardinalis (probably by way of Southern 
Melanesia), Rukia palauensis, R. oleaginea, R. ruki, R. sanfordi, 
Erythrura trichroa. The colonization of Micronesia by these species 
has probably extended over a considerable period of time. Megapo- 
dius, Trichoglossus, and Aplonis corvinus may represent older coloni- 



Baker: The Avifauna of Micronesia 45 

zations which have become well differentiated from the ancestral 
forms; Nycticorax, Myzomela, and Erythrura may have become 
established later and have had "less time" to become modified from 
the ancestral forms. Birds from Melanesia have reached Micronesia 
probably by direct flight to the Caroline Islands. Aided by favorable 
winds which blow from the southwest, south and southeast during 
the period from May to November, birds, particularly the young of 
the year, might conceivably be blown in the direction of the Caro- 
lines, where 57 percent of the birds derived from Melanesia reside. 
The Palaus are populated with 15 percent, the Marianas with 28 
percent, and the Marshalls (lacking "high" islands) with none; these 
may be secondary colonizations from the Carolinas excepting 
Ptilinopus, Megapodius, Gallicolumba canifrons, Cleptornis, and 
Colluricincla. The Marshall Islands have received no avian compo- 
nents from Melanesia. The absence of "high" islands in the Mar- 
shalls and the possible inability of birds accustomed to life on the 
luxuriant islands of Melanesia to become established on relatively 
barren atolls are logical reasons for this. Instead of New Guinea 
itself, the outlying islands of Melanesia (Bismarck Archipelago, 
Solomons, Southern Melanesia) probably have been the principal 
"taking-off" places for birds invading Micronesia. 

MOLUCCAN AND CeLEBESIAN COMPONENTS 

Birds which reached Micronesia by way of the islands of Celebes 
and the Moluccas may have been derived originally from Melanesia. 
The following birds appear to have used this route: Poiyhyrio por- 
phyrio, probably Halcyon chloris, Rhipidura lepida, Mijiagra oce- 
anica, Zosterops conspicillata, and Z. cinerea. These birds appar- 
ently became established initially in the Palaus; Porphyrio and 
Rhipidura lepida have not been recorded elsewhere in Micronesia, 
but Myiagra and the two species of Zosterops have spread to the 
Carolines and Marianas, although not into the Marshall Islands. 
Wind from the southeast in summer and fall has probably been a 
factor aiding these colonizations. The population of Gallinula chlor- 
opus resident at Palau may also have arrived by this route. 

Philippine Component 

Ten of the kinds of birds of Micronesia have come from or by way 
of the Philippine area. These are known principally from the Palaus 
and the Marianas and include: Rallus philippinus, R. owstoni, Po- 
liolimnas cinereus, Caprimulgus indicus, Corvus kubaryi, Psamathia 
annae, Artamus leucorhynchus, possibly Lonchura nigerrima, and 



46 



University of Kansas Publs., Mus. Nat. Hist. 



Collocalia inexpectata. The Philippines may have been the actual 
point of dispersal of the birds (example, Psamathia) , or may have 
been used as a stepping stone to Micronesia by birds coming from 
Melanesia (examples, Rallus and Artamus), by birds from Malaysia 
(example, Collocalia), and by birds from Asia (example, Caprimul- 
giis) . Two birds of this component have reached the islands of east- 
ern Micronesia. A subspecies of Lonchura nigerrima is endemic at 
Ponape, and a subspecies of Poliolimnas cinereus occurs on several 
islands in the Carolines and has even been recorded at Bikini in the 
Marshall Islands. Three species are known only from the Palaus; 
two are known only from the Marianas. 

Palearctic Component 

Birds of Micronesia which have been derived directly from Pale- 
arctica are Gallinula chloropus guami, Otus podarginus, Asia jlam- 
meus, Acrocephalus luscinia and Anas oustaleti. Apparently Galli- 
nula, Asia, and Acrocephalus arrived in Micronesia by way of the 
chain of islands from Japan southward to the Bonins, Volcanoes, 
and Marianas. Otus reached Palau from Asia, possibly by way of 
the Philippines. The smallness of the representation of this com- 
ponent may result partly from lesser ability of the northern birds 




Fig. 8. Faunal areas from which the resident land birds and fresh-water 
birds of Micronesia have been derived. (1) Palearctica; (2) PhiHppines; (3) 
Moluccas and Celebes (Malaysia) ; (4) Melanesia (New Guinea and north- 
ern Melanesia) ; (5) Polynesia. 



Baker: The Avifauna of Micronesia 47 

to adapt themselves to, and to establish themselves on, the semi- 
tropical and tropical islands of Micronesia as compared with birds 
from Melanesia where the climate and ecologic conditions resemble 
more closely those found in Micronesia. Evidence supporting this 
possibility is the large number of Palearctic residents in the Bonin 
and Volcano islands as compared with fewer in the Marianas; the 
Bonins and Volcanoes are less tropical and more temperate in 
climate. 

Table 11 lists the birds concerned, by faunal areas from which 
the birds have been derived and shows the number of kinds of birds 
which are present as a result of these colonizations. There is some 
overlap in the numbers since some endemics may be found in more 
than one area in Micronesia. Figure 8 shows the faunal areas from 
which the endemic land and fresh-water birds of Micronesia have 
been derived. Melanesia (Papua) supplied 52 percent of this popu- 
lation. Birds reaching Micronesia by way of the Moluccas and 
Celebes include 21 percent of the total population. The Philippines 
have supplied 10 percent; Polynesia, 9 percent; and Palearctica, 8 
percent. This population of endemic land birds and fresh-water 
birds has seemingly evolved from 46 colonizations, of which 27 
have been derived from Melanesia, 6 from the Philippines, 5 from 
the Moluccan and Celebean areas, 5 from Palearctica, and 3 from 
Polynesia, 

The Palaus have received a large part of their avifauna from the 
west (Moluccas, Philippines, Palearctica) . Their Melanesian com- 
ponent is mostly the result of secondary colonization from the Caro- 
lines. The Carolines have received a greater share of their land 
birds and fresh-water birds from Melanesia and a smaller share 
from Polynesia. The Marshalls are definitely associated with the 
Polynesian element. The Marianas exhibit a considerable amount 
of secondary colonization from other Micronesian islands, as well as 
some unique components from the Philippines, Melanesia, and Pale- 
arctica. Thus, the number of endemics in Micronesia provides little 
information concerning the actual number of successful coloniza- 
tions by birds from other areas. Many of the endemics probably 
have resulted in this way: Individuals of an endemic subspecies 
flew to another island and there underwent further differentiation, 
producing another endemic subspecies. Such secondary colonization 
probably is going on now. 

This analysis of the avifauna shows that Micronesia, with the 
exception of the Marshall Islands (and the Gilbert Islands), has 



48 



University of Kansas Publs., Mus. Nat. Hist. 



but little affinity to Polynesia. It has greater affinity, from the 
zoogeographical standpoint, with the Papuan Region (Melanesia). 

Table 11. Avifaunal Components Which Make Up the Endemic Resident 
Land and Fresh-water Bird Population of Micronesia 



Faunal Component 


Palau 


Western 

and central 

Carolines 


Eastern 
Carolines 


Marianas 


Marshalls 


Polynesian 


2 

11 
6 
6 
2 


3 

14 

3 

2 

1 


5 
16 
4 
2 
2 




12 
7 
4 
5 


3 


Melanesian 

Moluccan-Celebean . 
Philippine 




1 


Palearctic 









Totals 


27 


23 


29 


2S 


4 







Speciation 

Of the 104 native fresh- water birds and land birds which are res- 
ident in Micronesia, only 7 kinds or 6.5 percent remain undifferen- 
tiated from populations elsewhere. These birds are Phalacrocorax 
melanoleucus, Pandion haliaetus, Demigretta sacra, Ixobrychus 
sinensis, Anas poecilorhyncha, and possibly Lonchura punctulata 
(may be an introduction by man). Another bird, Gallinula chlo- 
ropus, a resident at Palau, may or may not be distinct from the 
gallinule of Malaysia, G. c. orientalis. Of the 104 resident birds, 97 
kinds or 93.5 percent have become differentiated and can be sepa- 
rated taxonomically from populations elsewhere. Of the kinds of 
birds which are found only in Micronesia, there are 5 endemic genera 
(16 percent), 31 endemic species (32 percent) and 76 endemic sub- 
species (75 percent). If we consider the avifauna of Micronesia as 
a single element, the endemism is high as compared with that on 
larger and less isolated islands. For example, Mayr (1944a: 174) 
found 137 resident birds on Timor including 22 endemic species (16 
percent) and 67 endemic subspecies (47.5 percent). Stresemann 
(1939b: 313) found 220 species including 84 endemic species (38.2 
percent) on Celebes. Mayr (1944a: 174) also writes that on Java, 
of 337 breeding species, 16 (4.8 percent) are endemic, and on New 
Caledonia, of 68 species 19 (27.9 percent) are endemic. Speciation 
in Micronesia has not progressed much farther than that at New 



Baker: The Avifauna of Micronesia 



49 



Caledonia and not so far as at Celebes, but siibspeciation has pro- 
gressed considerably more than at the island of Timor. The avi- 
fauna of the Hawaiian Islands, as recorded by Bryan and Greenway 
(1944), has 73 resident land birds and fresh-water birds, all of which 
are endemic, including one family, 23 genera and 36 species. The 
North American night heron, Nycticorax n. hoactli, may be included 
in this list as the only resident which is undifferentiated. The devel- 
opment of full specific differentiation within the resident avifauna is 
greater in the more isolated Hawaiian chain where 49 percent of 
these birds are regarded as endemic species, while in Micronesia, 
which is less remote from other bodies of land, the specific endemism 
is only 32 percent. 

Table 12. Endemism in Families of Native Land and Fresh-water Birds 

IN Micronesia 



Family 


Residents 


Endemic 
genera 


Endemic 
species 


Endemic 
subspecies 


Total 
endemic 


Phalacrocoracidae . . . 
Ardeidae 


I 

3 

'2 

1 
2 

7 
13 
1 
2 
1 
5 
7 
3 
1 
5 
14 
1 
9 
7 
14 
5 












1* 

















1 
1 




1 
1 







1 


1 
2 
4 

1 
1 

1 
1 


1 

2 

6 



3t 

1 

6 





1 


2 
4 

11 

1 
1 
5 
7 
3 

4 
9 
1 
7 
6 

10 
4 



1 


Anatidae 


1 


Accipitridae 

Megapodidae 

Rallidae 

Columbidae 

Psittacidae 

Strigidae 

Caprimulgidae 

Apodidae 

A cedinidae 

Campephagidae. . . . 
Corvidae 




2 

6 

13 

1 
2 
1 
5 
7 
3 
1 


Sylviidae 

Muscicapidae 

Artamidae 

Sturnidae 


5 

14 
1 
9 


Meliphagidae 

Zosteropidae 

Ploceidae 


7 
14 

4 






Totals 


104 


5 


31 


76 


97 







* Aphanollmonasa is included but may be extinct. 
t Aplonis corvinus is included but may be extinct. 



Table 12 lists the families of land birds and fresh-water birds 
which have resident members as part of the avifauna of Micronesia. 
It can be observed from the table that only two families are repre- 
sented by no endemic kinds, several families are represented by one 
or two endemic kinds, and others are represented by as many as 14 

4-^131 



50 University of Kansas Publs., Mus. Nat. Hist. 

endemic kinds. Endemism has reached its greatest development in 
the families Rallidae (6), Columbidae (13), Apodidae (5), Alcedi- 
nidae (7), Sylviidae (5), Muscicapidae (14), Sturnidae (9),Meliph- 
agidae (7), and Zosteropidae (14). Generic endemism is greatest in 
the Sylviidae where one endemic genus occurs among 5 endemic spe- 
cies and subspecies (20 percent), in Rallidae one in 6 (17 percent), 
in Meliphagidae one in 7 (14 percent). Specific endemism is great- 
est in Psittacidae and Corvidae where the single representative of 
each family in Micronesia is considered specifically distinct (100 
percent), in Megapodidae and Strigidae one in 2 (50 percent), in 
Muscicapidae and Zosteropidae 6 in 14 (43 percent) in Sylviidae 2 
in 5 (40 percent), in Rallidae 2 in 6 (33 percent), in Sturnidae 3 in 9 
(33 percent) in Columbidae 4 in 13 (31 percent). Subspeciation 
within species which are endemic in Micronesia has occurred in 8 
families, occurring within two species in each of the families Colum- 
bidae and Zosteropidae and once in each of the families Megapo- 
didae, Apodidae, Alcedinidae, Sylviidae, Muscicapidae, and Stur- 
nidae. 

In summary, the families of land and fresh-water birds found in 
Micronesia which have the greatest number of endemic forms are 
Muscicapidae (14), Zosteropidae (14), Columbidae (13), and Stur- 
nidae (9). Speciation has occurred in the single representative of 
the families Psittacidae {Trichoglossus rubiginosus) and Corvidae 
{Corvus kubaryi). Where family representation is large, speciation 
has occurred most frequently, as in the Muscicapidae (6 in 14 = 43 
percent), in the Zosteropidae (6 in 14 = 43 percent), and in the Co- 
lumbidae (4 in 13 = 31 percent). Subspeciation has occurred in 8 
families, in two species in the Columbidae and Zosteropidae and in 
one species in each of 6 other families. 

Time of Colonization 

Previously (and in the accounts of the species to follow), com- 
ments are made concerning the subjects of from where and by what 
route the various kinds of birds have arrived at Micronesia. The 
problem of when these birds arrived is a difficult and usually un- 
answerable one. Although geology provides some evidence on the 
relative age of the islands, and although deposits of bird guano on 
now elevated coral islands show that oceanic birds have inhabited 
these islands for a long time, there is no evidence to show the time 
of the first colonization by land birds. No fossil remains of land 
birds or fresh-water birds have been found in Micronesia. The 
relative extent of differentiation in color and structure, which has 



Baker: The Avifauna of Micronesia 51 

taken place between different birds, offers one means for estimating 
the relative length of residence in the area, provided all other fac- 
tors are equivalent. Concerning the birds of the Galapagos, Lack 
(1947:113) writes "That Darwin's finches are so highly differen- 
tiated suggests that they colonized the Galapagos considerably 
ahead of the other land birds." Evidence from this source actually 
is of little value, because the speed of evolution is unknown and its 
rate may be different in different species, even though they live 
under the same circumstances. Dobzhansky (1941) says that evo- 
lution is a modification of the genetic equilibrium, which, if true, 
may not result in similar manifestations in different kinds of birds 
living under the same conditions of life. Relative antiquity of the 
birds might be ascertained by measuring their ecologic adaptations. 
The Guam Rail {RalliLS oavstoni) and the Micronesian White- 
browed Rail {Poliolimnas) can be examined in this way. R. owstoni 
has the ability to live in both brackish and fresh water swamps, as 
well as in the scrub and grass of the uplands and in the virtually 
barren, rocky areas in the dense jungles. Poliolimnas, on the other 
hand, appears to be restricted to swampy areas in Micronesia. If 
the swampy areas were removed this rail probably would become 
extinct. R. owstoni appears to have been resident in Micronesia 
longer than Poliolimnas. However, ability to live in a variety of 
habitats might be acquired by R. owstoni in a relatively short time. 
Another possibility is that the birds, which are less differentiated 
from their ancestral stocks, may be less differentiated because of 
suppression of newly evolved characters by dilutions, which result 
from interbreeeding with new birds, which may be arriving at irreg- 
ular intervals from the ancestral home. Interbreeding of the resi- 
dent population with newcomers may overshadow any modifications 
which might have appeared as a result of insular isolation, especi- 
ally modifications which have little adaptive significance. One 
would suspect, from their modifications, that Rallus owstoni, Meta- 
bolus rugensis, Corvus kubaryi, and other endemic forms have ex- 
perienced less of this "dilution," than such birds as Rallus philippen- 
sis pelewensis, Artamus leucorhynchus pelewensis, Myzomela cardi- 
nalis, and others. Murphy (1938) mentions this "dilution" effect 
in his discussion of "strong" and "weak" subspecies among warblers 
of the Marquesas. He writes that "strong" subspecies maj'' develop 
if the birds are present on islands which are upwind from islands 
containing related subspecies. The wind acts to block interisland 
migration in these weak-flyers. On the other hand, "weak" sub- 
species may show the effect of "dilution," being situated on islands 



52 University of Kansas Publs., Mus. Nat. Hist. 

downwind from islands containing related subspecies. The direction 
of the wind acts to aid the weak flyers to move to the downwind 
islands and continually ''dilute" the resident subspecies. Similar 
examples can be cited for Micronesian birds. Hesse, Allee, and 
Schmidt (1937:87) write, "Endemism on islands is most frequent 
in forms for which the difficulty of reaching the island is most ex- 
treme, so that new increments of the parent form are unlikely to 
follow." 

Employing the criteria mentioned above, the birds of Micronesia 
can be tentatively divided into four groups as regards the relative 
time when they arrived at the islands: 

1. Birds of ancient colonizations which reached certain individual 
islands, became modified, and dispersed no farther. Examples are 
Aphanolimnas, Rallus omstoni, Aplonis corvinus, Metabolus rugen- 
sis, and Corvus kubaryi. 

2. Birds of ancient colonizations which reached or dispersed 
through a number of islands but are now restricted to relatively 
few islands. Examples are Ducula oceanica, Ptilinopus porphy- 
raceus, Megapodius laperouse, Asio fiaymneus, and Acrocephalus 
luscinia. 

3. Birds of ancient, or possibly more recent, colonizations which 
initially reached or subsequently dispersed to many of the islands 
of Micronesia possessing habitat suitable for them. Examples are 
Myzomela cardinalis, the two species of Halcyon, Aplonis opacus, 
and Zosterops conspicillata. 

4. Birds of rather recent colonizations, which may have reached 
only a few islands and are relatively unmodified from their parental 
stocks. Examples are Artamus leucorhynchus, Caprimulgits indicus, 
Poliolimnas cinereus, and Nycticorax caledoniciis. 

Factors Causing Dispersal 

Darlington (1938:274) in discussing the origin of the fauna of 
the Greater Antilles uses the term "over-water dispersal" in refer- 
ring to the spread of terrestrial animals across water. He is against 
the use of the term "accidental dispersal" since many factors besides 
accident are involved. He contends, as do others, that certain forms 
of organisms, owing to their "nature and behavior" cross water bar- 
riers more successfully than others. These observations may be 
applied to the "over-water dispersal" of birdlife to the islands of 
Micronesia. Certain groups of birds are more evident in Micronesia 
than others. Certain groups of birds which are found on other 
islands of the Pacific basin are found in Micronesia only in small 



Baker: The Avifauna of Micronesia 53 

numbers or may not be represented; Mayr (1945a: 284) writes, 
"Remarkable is the almost complete absence of parrots and honey- 
eaters, the small number of pigeons and the absence of such wide- 
spread genera as Lalage, Turchis, and Pachycephala." The absence 
of some species and the presence of others produces the character- 
istic insular effect termed "disharmonic" by Gulick (1932:407), as 
compared with the continental area or island which derived its avi- 
fauna by way of a land bridge. One would think from looking at 
table 12 that members of the families Rallidae, Columbidae, Musci- 
capidae, Sturnidae, and Zosteropidae were the most successful colo- 
nizers in Micronesia on the basis of the number of successful coloni- 
zations (not necessarily on the number of endemics developed from 
a single colonization). Of these families, Sturnidae and Zosteropidae 
and possibly Columbidae contain species which often move in flocks. 
Furthermore, these families as well as the Muscicapidae feed on 
either fruits, seeds, or insects, any one of which is a type of food 
which might "give out" suddenly, stimulating a migratory behavior 
within the birds. From a flock embarking seaward in "search" of 
more food, a part or even all of the birds might survive in a chance 
flight to an isolated island in Micronesia. If a flock containing both 
males and females reaches an island, the species has a good chance 
of becoming established. Evidence that such a rapid colonization by 
flocks of birds can take place is found in the remarkable colonization 
of New Zealand by Zosterops lateralis from the Australian area. 
The bird was first seen as a winter migrant in New Zealand in 1856 
and records of nestings were obtained at North Island in 1862, ac- 
cording to Oliver (1930:489). In the case of rails there is no evi- 
dence that they move in flocks; however, they are among the most 
successful colonizers and are on many of the oceanic islands in the 
tropical and subtropical oceans. Representatives of several species 
of the family Rallidae have invaded Micronesia and have success- 
fully established 6, or possibly 7, "colonies." 

Darlington (1938:274) further writes that "it is no accident that 
some islands, because of their nature and position, the direction of 
winds and currents, and the nature of the neighboring land, receive 
more organisms than other islands do." Semper (1881:294) writes 
that the distribution of flying creatures "must be in a great degree 
dependent on the direction and strength of atmospheric currents." 
These statements are applicable to the history of the avifauna of 
Micronesia. The Caroline Islands, for example, present a "broad 
front" for wanderers from the Melanesian islands. As mentioned 



54 University of Kansas Publs., Mus. Nat. Hist, 

previously, the prevailing winds in the late spring, summer, and 
early fall are from the south, southwest, and southeast and would 
favor bird flight to the northward towards the Carolines. In addi- 
tion, the breeding season of many of the birds in Melanesia is from 
November to February, and in the spring and summer, restless 
young birds seeking living space might fly seaward and aided by the 
winds fly northward towards Micronesia. Adults, which may have 
well-established home territories, may be less likely to attempt such 
a movement. 

One could conclude from the above discussion that the Micrones- 
ian islands, especially the Carolines, might be well populated with 
a large variety of birds from Melanesia, a scant 500 or more miles 
away. As it turns out, there are only a few islands in this extensive 
archipelago possessing proper vegetation, fresh water, and other 
qualities which make them capable of supporting the land and fresh- 
water birds of Melanesia. The few islands which have these qual- 
ities are the so-called "high" islands, including the entire Mariana 
chain, the Palaus, and four widely separated islands in the Caro- 
lines: Yap, Truk, Ponape, and Kusaie. The other islands of Micro- 
nesia are "low" coral islands, which often lack fresh water and have 
a meager variety of fruits, insects and other foods. Thus, if birds 
do reach Micronesia but arrive at the atolls instead of the "high" 
islands, these birds may be doomed. It is noteworthy that the Mi- 
cronesian islands are small compared with the Solomons, Fijis, and 
others. The smaller the island, the fewer the number of ecologic 
niches and the fewer the kinds of birds present. 

Mayr (1941b:215) writes that the distance from the nearest 
land mass and the climatic conditions are important factors con- 
trolling dispersal. With regard to the degree of remoteness of the 
islands, table 13 lists the number of resident land and fresh-water 
birds present in the Palaus and the "high" islands of the Carolines. 
Also, the approximate distance from the nearest large land mass 
and the area in square miles are given. There is some correlation 
between the distance from the nearest land mass and the number 
of resident land birds and fresh-water birds. For example, Palau, 
with 32 resident birds, is only 410 miles from the nearest land mass 
whereas Kusaie, with only 11 resident birds, is 720 miles from the 
nearest land mass. The comparative size of the land mass must 
also be taken into account, as shown by the fact that the large 
island of Ponape contains more kinds of birds but is more remote 
from large land masses than either Yap or Truk. 



Baker: The Avifauna of Micronesia 



55 



Table 13. Correlation Between Number of Resident Land and Fresh-water 
Birds and Distance From Large Land Masses of "High" Islands of Mi- 
cronesia 



Island 


No. 

of 

birds 


Approximate 

distance from 

nearest land 

mass (statute 

miles) 


Nearest land mass 


Area in 

square 

miles 


Palau 


32 


410 


Approximately equal distance from 
Mindanao, Morotai, New Guinea 


171 


Yap 


13 


580 


New Guinea 


83 


Truk 


17 


525 


New Ireland 


50 


Ponape. . 


20 


630 


New Ireland 


145 


Kusaie . . . 


11 


720 


Malaita (Solomons) 


42 



Climatic factors are important in the dispersal of bird life; Micro- 
nesia, where the climate is tropical to subtropical, is better suited 
for colonization by birds from the tropics (Melanesia) than by birds 
from the temperate or cold climates (Palearctica). The climatic 
factor may be one of the principal reasons why birds from Pale- 
arctica make up only a small part of the avifauna of Micronesia. 

Analysis of Speciation 

The process of speciation within insular populations has been 
discussed by many authors. Hesse, Allee, and Schmidt (1937:517) 
list the motives for differentiation as, "Special character of insular 
faunae rests on the conditions common to all islands — isolation, 
freedom from competition, space restriction, and special insular 
climates." This combination of characteristics is seldom found 
elsewhere in nature, and as Murphy (1938:357) points out, an 
island is the nearest approach to a ''man-controlled laboratory." 
Isolation of small populations is probably the most influential factor 
in the process of speciation in insular organisms. Lack (1947:134) 
writes that "in all organisms the isolation of populations is an es- 
sential preliminary to the origin of new species." Buxton (1938: 
265) also stresses this point with regard to the formation of species 
of insects in Samoa and emphasizes that evolution may occur more 
quickly in small populations. When mutations appear in such small 
and isolated populations, they have a greater chance to become fixed 
than do mutations in less restricted populations in a larger land 
mass, where such a mutation might be lost by the swamping effects 



56 University of Kansas Publs., Mus. Nat. Hist. 

of outbreedings. In addition, Wright (1931 and elsewhere) suggests 
the possibility of change by accidental elimination and recombina- 
tion of hereditary characters in micropopulations. This mechanism 
could well be a factor in Micronesian bird populations, many of 
which possess no more than a few hundred individuals. Huxley 
(1938:256) emphasizes that "accidental" mutations may be per- 
petuated in small, isolated groups. It might be added that such 
changes might be either advantageous or disadvantageous to the or- 
ganism concerned. Huxley (1938:263) states also that geographic 
isolation may promote nonadaptive differentiation, which may be 
caused by "colonization by a random sample" or by subsequent 
"preservation of nonadaptive mutations in numerically small iso- 
lated groups." Mayr (1942b :237) cites the importance of the 
"founder" principal for reduced variability in small populations. He 
points out that if the "founders" of the population carried with 
them only "a very small proportion of the variability of the parent 
population," one would expect to see divergence from the ancestral 
stock. 

Freedom from competition, especially interspecific strife, is an 
important factor in differentiation ; this is especially true in the early 
period of colonization. Lack (1947:113) points to the absence of 
food competitors, especially in the initial period of colonization, as 
an important influence in the evolution of Darwin's finches at the 
Galapagos Islands. Once a population has become established and 
"adjusted" to a given environment on a small island, intraspecific 
competition might bring about adaptative selection. Subsequent 
colonists might be eliminated by the competition brought about by 
these previously adapted organisms, especially if both organisms 
were adapted for life in the same ecologic niche. Space restriction 
may be important in such Micronesian birds as Rhipidura and 
Myiagra, which appear to possess recognizable territories. A new 
colonist entering the territory of one of these birds might be forced 
out. This competition might not play such an important part among 
birds, which live in flocks and do not range in closely guarded terri- 
tories; birds in this group include some pigeons, starlings, and white- 
eyes. 

Freedom from the pressure of predation probably exerts a direct 
influence on formation of species. Aside from a few migrant hawks 
and two kinds of resident owls, most of the avifauna feeds on vege- 
table and invertebrate foods. The large lizard Varanus may be 
classed as the only native predator on many of the islands. Man has 
been responsible for the introduction of rats, house cats, and other 



Baker: The Avifauna of Micronesia 57 

mammals, which may be destructive to birds. Thus, before the ad- 
vent of man the factor of predation may not have been of great con- 
sequence. As mentioned previously, nonadaptive modifications may 
be perpetuated where the ''weeding-out" process by predation is not 
an influence. Flightless rails have apparently developed in the 
absence of predation. 

The absence of the pressure of predation should remove a certain 
amount of control on the population turn-over. As Hesse, Allee, and 
Schmidt (1937:521) write, a characteristic of the faunas of oceanic 
islands is the fact that they are distinguished by the occurrence of 
"disproportionately developed taxonomic groups in which one or a 
few basic types have undergone adaptative radiation and come to fill 
unduly large proportions of the population as compared with condi- 
tions that obtain on neighboring continents." Lack (1947:114) 
writes, "that the absence of predators may well have accelerated the 
adaptative radiation" in the Galapagos finches. In Micronesia, the 
starling {Aplonis opacus) dominates much of the available habitat on 
some of the Caroline atolls, and even on "high" islands, where other 
land birds are present. There appears to be no tendency towards 
selective adaptations occurring, or towards ecologic isolation. 

Available data indicate that the life spans of individual birds in 
Micronesia may be short. For example, it was obvious on many of 
the islands visited by the NAMR.U2 party that starlings (Aplonis 
opacus) in immature jilumage outnumbered starlings in adult plum- 
age, although it is pdssible that immature plumages are retained 
longer in these island birds than in others. Similar observations were 
made by Coultas, who noted the ratio of birds in immature plumage 
to birds in adult plumage at Kusaie to be 5 to 1. If the life span is 
shorter in these insular forms as compared with that of the ancestral 
stocks, the higher annual population turn-over would allow for the 
speed of genetic changes to be accelerated. 

The origin of species by hybridization between different kinds of 
organisms has been a subject of frequent discussion. Lack (1947: 
100) concludes that it is improbable that hybridization has played 
an important part in the origin of new kinds of birds. Nevertheless, 
the absence of sufficient mates in the confines of a small island prob- 
ably stimulates the crossbreeding between two species of birds. 
Fertile offspring of such a cross might conceivably account for some 
of the populations, the origins of which are puzzles to present day 
taxonomists. Such Micronesian forms as Metabolus and Cleptornis 
could conceivably have been derived in such a manner. Yamashina 
(1948) has described the origin of Anas oustaleti as a result of by- 



58 University of Kansas Publs., Mus. Nat. Hist. 

bridization between A. platyrhynchos and A. poecilorhyncha. It 
might be difficult to explain every case of the formation of other 
insular species on the basis of the effects of isolation and paucity- 
alone. However, Mayr (1942b:236) includes the development of 
questionable and unusual kinds of insular forms in a general state- 
ment: "The potentiality for rapid divergent evolution in small pop- 
ulations explains also why we have on islands so many dwarf or 
giant races, or races with peculiar color characters (albinism, mela- 
nism), or with peculiar structure (long bills in birds), or other pe- 
culiar characters (loss of male plumage in birds)." 

Nutrition may be also a factor influencing speciation in bird life. 
The types of food plants (coconut, papaya, breadfruit, pandanus, 
etc.) might be similar on a Micronesian island and on a continental 
island in the Philippine region; however, the value of these plants 
as foods might vary and might reflect differences in mineral content 
of the soils. For example, if the soils on an island lack, or by leach- 
ing out have lost, sufficient amounts of potassium and other ele- 
ments, plants may store foods, not as proteins, but possibly as carbo- 
hydrates, simple sugars, or alkaloids. Whether nutritional influences 
might have a selective effect on the bird populations, has not been 
ascertained. 

In summary, it may be said that genetic change altering the 
phenotypic expression of avian characteristics is no more apt to 
happen in insular populations than in continental populations but 
genetic change may have a greater chance of being perpetuated in 
small insular populatians where isolation, limited competition, free- 
dom from the selective influences of predation, and other factors 
exert influences. 

CONSERVATION OF THE AVIFAUNA OF MICRONESIA 

The islands of Micronesia are small and their occupation by man 
often produces serious effects on the endemic animal life of the 
islands. The vulnerability of insular bird populations is well at- 
tested by the fact that the majority of birds, which have become 
extinct in the past two hundred years, have been insular forms. Two 
birds in Micronesia, the Kusaie Rail {Aphanolimnas) and the Ku- 
saie Mountain Starling (Aplonis corvinus) , are known to be either 
extinct or so rare that they have not been taken since the time of 
Kittlitz, who visited the island of Kusaie in December, 1827, and 
January, 1828. Other birds {Anas oustaleti, Caloenas nicobarica, 
Megapodius I. laperouse, and Metabolus rugensis) have become 
redused in numbers and may be threatened with extermination. 



Baker: The Avifauna of Micronesia 59 

Nelson (1921:270-274) has described the following agencies de- 
structive to island life of the Pacific: fire, volcanic eruptions, tidal 
waves, hurricanes, clearing of the land, introduction of domestic 
animals and grazing, introduction of wild animals and birds. Mayr 
(1945c) also presents a discussion of conservation problems in these 
islands. 

Fire is a serious hazard to island life, especially to the land birds. 
It destroys both food and cover, these two habitat requirements 
being most essential to the birds. The firing of open lands to improve 
grazing conditions was a practice which persisted in the Marianas 
during the time of the Spanish. This practice has declined, but the 
resultant vegetational changes and erosion have adversely affected 
the avifauna. Tidal waves and hurricanes (typhoons) are occasion- 
ally of such intensity as to flood low coral atolls. Such events are 
damaging to, or might even exterminate populations of land birds 
(Aplonis, Acrocephalus and others), and prevent colonizations which 
might otherwise occur. Clearing of the land for agricultural use 
probably has affected the avifauna, especially on the island of 
Tinian where much of the island has been placed in cultivation. The 
occurrence of domestic stock, especially feral hogs and cats, has 
affected the birds. Hogs, apparently, have been in the islands for a 
long time. The English privateer, Lord Anson, visited Tinian in 
October, 1742, and noted a large number of hogs present at that 
time. At Guam, in 1945, the NAMRU2 party found both hogs and 
cats moving freely in all parts of the island. Stomachs of cats exam- 
ined showed that they had been feeding principally on rodents. 

Introduction of wild animals and plants have not been so extensive 
as in the Hawaiians or other islands. There have been at least five 
importations of land birds to Micronesia as well as several mam- 
mals, other vertebrates and invertebrates. The effect of these estab- 
lished colonies on the native bird life has not been studied. 

The late world war has brought changes to the population of bird 
life in Micronesia. The author (1946b) has elsewhere described 
some of the effects of the bombing, invasion, and occupation of small 
islands. Some islands, like Peleliu, suffered severely from bombing 
and invasion operations. Some islands, especially smaller ones like 
Kwajalein and Ulithi, were partly or almostly entirely cleared of 
vegetation by occupation forces. Other effects were caused by ''rec- 
reational" shooting of birds by garrison forces; introductions of pests 
in materials unloaded; and pest control by clearing, draining, and 
spraying with DDT and other insecticides to the detriment of 
inoffensive species. 



60 University of Kansas Publs., Mus. Nat. Hist. 

It is obvious that a well-planned program of conservation should 
be placed in operation to insure survival of the endemic avifauna of 
Micronesia. 

THE FUTURE OF ORNITHOLOGICAL RESEARCH 

IN MICRONESIA 

Collections of birds have been made at most of the major islands 
of Micronesia, and it is thought that there are but few if any un- 
named birds in the region. The distribution of several species has 
not been completely investigated, especially those land birds 
(Dvciila, Ptilinopus, and Aplo7ns) which inhabit coral atolls in the 
Carolines and Marshalls. The bird life of the nortliern Marianas is 
also incompletely known. Continued observations in the Microne- 
sian islands will increase our knowledge of the kinds of migratory 
shore birds and migratory land birds which reach the island as win- 
ter visitors. Further information is needed concerning the breeding 
activities of sea birds in Micronesia, especially in the Marshalls and 
Carolines. 

The systematic status of most of the birds in Micronesia is already 
established. It is hoped that the present account advances our 
knowledge of the methods of colonization. Although these funda- 
mental investigatons have been nearly completed in Micronesia the 
field of avian ecology has been relatively untouched. In the past, 
expeditions have visited Micronesia with the aim of obtaining within 
a short time collections of the animal life as large and as represent- 
ative as possible. Many of the collectors made few or no field notes 
on the bird life; some, like Finsch, Kubary, Marche, and Coultas, 
made valuable observations on the habits of the birds. Intensive 
ecological researches may be accomplished more thoroughly by res- 
ident investigators, who can devote full time to such pursuits. 

METHODS AND ACKNOWLEDGMENTS 

My own opportunity to study the bird life of Micronesia came as 
a member of the scientific staff of the Laboratory of Mammalogy 
of United States Naval Medical Research Unit No. 2 (NAMRU2) 
in the late war. The primary duty of this laboratory was to obtain 
examples of the vertebrate fauna for examination for ectoparasites 
by the Laboratory of Acarology and to preserve specimens for iden- 
tification. As a result sizeable collections of mammals, birds, and 
other vertebrates were obtained. In addition, ecological data were 
obtained (as time permitted), especially as an aid in studying the 
distribution of ectoparasites which affected man. In 1945, I spent 



Baker: The Avifauna of Micronesia 61 

eleven months in Micronesia; for most of this time I was stationed 
at Guam, the headquarters of the Unit, although one month was 
spent in the Palau Islands, two weeks were spent at Ulithi Atoll, 
and short stop-overs were made at Eniwetok and Kwajalein atolls. 
Other members of the laboratory staff visited Rota and Truk islands. 

Subsequent to the field studies in the Pacific, I was sent to Wash- 
ington and spent approximately eight months at the United States 
National Museum studying the collections of birds and preparing 
several reports for publication. In this period other material was 
studied, both in the United States National Museum and at the 
American Museum of Natural History, New York, and the litera- 
ture dealing with the birds of Micronesia was explored and a bibli- 
ography of Micronesian birds was prepared. At the University of 
Kansas, I continued the bibliographic work, borrowed and studied 
some specimens, and completed accounts of the avifauna of Micro- 
nesia. 

Under the account of each bird, all known references in the litera- 
ture, which mention the scientific name of the bird and its distribu- 
tion in Micronesia, are listed. The references are arranged as fol- 
lows: (1) citation to the original description, and (2) citations to 
names in literature in the order of their first appearance. AVhen a 
name is a pure synonym, it may be recognized as such by the fact 
that the type locality is given immediately following the citation. 
In compiling these references the writer made use of the invaluable 
work by Wiglesworth (1891) and of Utinomi's "Bibliographica Mi- 
cronesia," made available through the translation by Fisher (1947). 
The arrangement of the families follows that of Peters (1931-1945) 
and Wetmore (1940). 

Specimens examined are designated as to collection in which cata- 
logued by the following abbreviations: USNM, the United States 
National Museum ; AMNH, the American Museum of Natural His- 
tory; MCZ, the Museum of Comparative Zoology; and KMNH, the 
University of Kansas Museum of Natural History. Average and 
extreme measurements of specimens are usually listed in tables; 
unless otherwise indicated, measurements are in millimeters, and 
are of adult specimens. The wings have been measured by flatten- 
ing them on a ruler. Weights are in grams. Unless otherwise in- 
dicated, descriptions of the birds have been written by the author. 
Descriptions of shore birds are not given; for these the reader may 
refer to Mayr (1945a :28-47) where characters useful for identifi- 
cation of the birds in the field also are given. The writer is espe- 



62 University of Kansas Publs., Mrs. Nat. Hist. 

cially indebted to Dr. Ernst Mayr for making available the descrip- 
tions of Micronesian birds made by Miss Cardine Bogert, especially 
those dealing with color of the irides, feet, and bill. Color terms 
in quotation marks refer to those in Ridgway (1912). 

In dealing with insular forms the criterion of intergradation as 
indicative of subspecies cannot be applied as it can in kinds of birds 
on the mainland which have geographically continuous distributions. 
Instead, degree of difference in combination with geographic position 
plus other factors such as degree of variation in the geographic races 
of the same species or a related species on continental areas are used 
in deciding whether two closely related kinds are subspecies or full 
species. Many kinds of birds in the islands are modified but little 
from island to island (examples, Rhipidura rufifrons, Aplonis opacus, 
Ducula oceanica, and Myzomela cardinalis), and can be treated as 
subspecies. Others show much variability from island to island and 
it is uncertain whether they should be treated as subspecies or as 
separate species (examples, Myiagra oceanica, Zosterops cinerea, 
Rukia, and possibly Acrocephalus luscinia). Decisions on generic 
status are equally difficult to make. In many cases the experience 
and judgment of the taxonomist may be the only criteria by which 
he can decide whether a bird is different enough to be considered as 
a distinct genus. This "human element" has caused some disagree- 
ment. Knowing whether the bird is to be considered as a distinct 
genus or instead merely as a species may not be as important as 
knowing its correct phylogenetic relationship. The circumstance 
that variation in these insular birds is in general less predictable 
than in mainland birds adds, I think, to the pleasure inherent in the 
classification of the variations. 

First, I thank Commodore Thomas N. Rivers (MC) USNR, then 
commanding oSicer of NAMRU2, for the opportunity to join the 
Unit, for his interested cooperation in seeing that the plans for field 
trips were successful, and for his thoughtfulness in obtaining for me 
the orders for duty at the United States National Museum subse- 
quent to our field investigations. Greatly appreciated also is the 
help rendered by my former colleagues of NAMRU2, including Dr. 
David H. Johnson, Dr. George W. Wharton, Dr. Aaron B. Hard- 
castle, Mr. Odis A. Muennik, Mr. L. P. McElroy, Mr. Charles 0. 
Davison, Mr. Merle H. Markley, Mr. Walter L. Necker, Dr. Wilbur 
G. Downs, Dr. Bernard V. Travis, and Mr. E. W. Coleman. Other 
personnel, then stationed in Micronesia, who contributed data used 
in this report include: Dr. Joe T. Marshall, Jr., (who generously 



Baker: The Avifauna of Micronesia 63 

loaned some of the specimens taken by him in Micronesia), Dr. C. 
K. Dorsey, Dr. George Hensel, Mr. Tom Murray, Dr. Irwin O. 
Buss, Mr. James 0. Stevenson, Dr. Wilfred D. Crabb, Mr. Herbert 
Wallace, and Dr. M. Dale Arvey. Authorities of the United States 
National Museum provided generously for using the collections 
there, and I am especially grateful to Dr. Alexander Wetmore, Dr. 
Herbert Friedman, and Mr. Herbert G. Deignan for their coopera- 
tion and assistance. Doctor Wetmore kindly made available many 
of the birds collected at Bikini during the atomic bomb experiments. 
Dr. Robert Cushman Murphy, Dr. Ernst Mayr, and Dr. Dean 
Amadon of the American Museum of Natural History made avail- 
able the collections in their charge. Doctor Murphy allowed me to 
examine some of the heretofore unstudied collections of sea birds of 
the Whitney South Sea Expedition. Doctor Mayr generously helped 
me with taxonomic and evolutionary problems and made available 
to me some of his owti unpublished taxonomic notes, the unpublished 
field accounts of Mr. William F. Coultas and a partly completed 
manuscript on the birds of Micronesia by Miss Cardine Bogert. Mr. 
James L. Peters generously loaned specimens from the Museum of 
Comparative Zoology. The use of unpublished field notes made by 
Mr. Larry P. Richards at Ponape and Truk in 1947 and 1948 is also 
gratefully acknowledged. I am grateful also to my colleagues at 
the Museum of Natural History of the University of Kansas and 
would single out for special mention Dr. E. Raymond Hall who gave 
critical assistance with the manuscript, Drs. Edward H. Taylor and 
Herbert B. Hungerford who made helpful suggestions, and Mrs. 
Virginia Cassell Unruh who drew the distributional maps. 

ACCOUNTS OF THE KINDS OF BIRDS OF MICRONESIA 

Diomedea nigripes Audubon 
Black-footed Albatross 

Diomedea nigripes Audubon, Ornith. Biog., 5, 1839, p. 327. (Type locality, Pacific 
Ocean, lat. 30°44'N., long. 146°W.) 

Diomedea fidiginosa Oustalet, Le Nat., 1889, p. 261 (Mariannes). 

Diomedea nigripes Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris (3). 8. 1896, p. 51 
(Agrigan); Hartert, Novit. Zool., 5, 1898, p. 68 (Marianne); Seale, Occ. Papers Bernice 
P. Bishop Mus. 1, 1901, p. 22 (Marianas); Safford, Osprey, 1902, p. 70 (Mariannes); 
idem. The Plant World, 7, 1904, p. 268 (Guam?); Kuroda, in Momiyama, Birds Micro- 
nesia, 1922, p. 32 (Marriane); Peters, Check-list Birds World, 1, 1931, p. 43 (Mar- 
shalls); Hand-list Japanese Birds, rev., 1932, p. 188 (Marianas); Hand-list Japanese 
Birds, 3rd ed., 1942, p. 210 (Marianas) ; Mayr, Birds Southwest Pacific, 1945, p. 5 
(Marshalls). 

Geographic range. — North Pacific Ocean. Breeds on islands northwest of 
Hawaii. In Micronesia : Mariana Islands — Agrigan. 

Characters. — A large oceanic bird with sooty-brown coloration; darker on 



64 University of Kansas Publs., Mus. Nat. Hist. 

nape, wings and tail; lighter on forehead, sides of head, and abdomen; area 
surrounding bill whitish; tail whitish at base; bill dark reddish-brown; feet 
black. 

Remarks. — This albatross has been recorded from waters near the 
Mariana Islands. Quoy and Gaimard (1824:145) observed "alba- 
tross" between the Mariana and the Hawaiian Islands. The only 
actual specimens obtained from the islands were reported on by 
Oustalet (1896:51). These were eight Black-footed Albatrosses 
which were taken on the coast of Agrigan by Marche in December, 
1888, and January, 1889. Oustalet gives the following measure- 
ments: total length, 680-785; wing, 485-525; tail, 180-225; tarsus, 
80-90; culmen, 108-125. The specimens are apparently in the Paris 
Museum. 

Peters (1931:43) lists the Marshall Islands as part of the range 
of D. nigripes. 

In the period of the late war Gleise (1945:221) observed eight 
Short-tailed Albatrosses (D. albatrus Pallas) "off Saipan." Speci- 
mens of D. albatrus have not been taken in Micronesia. According 
to Austin (1948b:32) this albatross "is now virtually extinct," and 
this record may be questioned. 

Puffinus pacificus chlororhynchus Lesson 
Wedge-tailed Shearwater 

Puffinus chlororhynchiis Lesson, Traite d'Omith., 8, 1931, p. 613. (Type is from 
Shark's Bay, West Australia.) 

Puffinus sphenurus Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, 
p. 299 (Mortlock). 

Puffinus chlororhynchus Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 
1890-1891 (1891), p. 80 (Luganor?); Godman, Monogr. Petrels, pt. 2, 1908, p. 88 
(Carolines); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 33 (Luganor or Ruk?). 
Puffinus pacificus chlororhynchus Hand-list Japanese Birds, rev., 1932, p. 187 
(Lukunor or Truk?, Kusaie) ; Hand-list Japanese Birds, 3d ed., 1942, p. 209 (Lukunor 
or Truk?, Kusaie). 

Geographic range. — Breeds at the Seychelles, Australia, Lord Howe, Nor- 
folk, and other islands in the Australian area. Ranges throughout most of the 
warmer parts of the Indian and Pacific oceans. In Micronesia : Mariana Islands 
— Guam; Caroline Islands — Lukunor or Truk?, Kusaie. 

Characters. — A large shearwater with long wedge-shaped tail; upper parts 
sooty-brown with crown, neck, and wings darker and forehead paler; under 
parts paler than upper parts; bill dark; feet flesh-colored. 

Remarks. — This shearwater was taken by Kubary either at 
Lukunor or at Truk in the Caroline Islands. At a later date, appar- 
ently between 1922 and 1932, the Japanese recorded the bird at 
Kusaie. In using this subspecific name, I am following the Hand- 
list of Japanese Birds (Hachisuka et al., 1932:187). 



Baker: The Avifauna of Micronesia 65 

At Guam on August 10, 1931, Coultas obtained a male shearwater, 
which is tentatively placed in this subspecies. Its measurements are 
as follows: wing, 290; tail, 128; exposed culmen, 39; tarsus, 47. 
Coultas (field notes) writes that he was told by natives that petrels 
nest and roost on the high cliffs behind the city of Agafia on Guam. 
At sea south of the eastern Caroline islands, Coultas obtained five 
other birds which appear to be the same as the bird from Guam. All 
specimens are in the collections of the American Museum of Natural 
History. 

Puffinus pacificus cuneatus Salvia 

Wedge-tailed Shearwater 

Puffinus cuneatus Salvin, Ibis, 1888, p. 353. (Type locality, Krusenstern Island ^ 
Ailuk, Marshall Islands, fide Fisher, Auk, 63, 1946, pp. 587-588.) 

Puffinus cuneatus Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890- 
1891 (1891), p. 80 (Krusenstern); Salvin, Cat. Birds British Mus., 25, 1896, p. 371 
(Krusenstern) ; Godman, Monogr, Petrels, pt. 2, 1908, p. 76 (Marshalls). 

Puffinus pacificus cuneatus Mathews, Birds .\ustralia, 2, 1912, p. 84 (Marshall 
Group) ; Peters, Check-list Birds World, 1, 1931, pp. 55-56 (Krusenstern) ; Hand-list 
Japanese Birds, 3d ed., 1942, p. 209 (Krusenstern); Fisher, Auk, 63, 1946, pp. 587- 
588 (Ailuk). 

Thyellodroma cuneata cuneata Mathews and Iredale, Ibis, 1915, p. 597 (Krusen- 
stern); Mathews, Syst. .^vium Australasianarum, 1, 1927, p. 113 (Marshall Group). 

Thyellodroma cuneata Oberholscr, Auk, 34, 1917, p. 474 (Krusenstern). 

Thyellodroma pacificia cuneata Mathews, Novit. Zool., 39, 1934, p. 186 (Caroline 
Islands). 

Geographic range. — Pescadores cast to the Hawaiian Islands and south to 
eastern Micronesia. In Micronesia: Marshall Islands — Ailuk. 

Remarks. — Osbert Salvin received two specimens of this shear- 
water from H. J. Snow, who got them at the Krusenstern Islands 
in 1883. In describing them, Salvin (1888: 353) comments that the 
locality is seemingly in the Marshall Islands at approximately 
10°17' N. and 190° W. This locality was confusing to Seebohm 
(1891:191) who thought it was between the Hawaiians and the 
Marshalls, while Hartert (1926:352) decided it was really Krusen- 
stern Rocks in the Hawaiian Group. To clear the matter up, Fisher 
(1946:587-588) writes that Salvin was correct and suggests that the 
name of the island should be the better established one, Ailuk, rather 
than the little used one, Krusenstern. 

P. p. cuneatus resembles P. p. chlororhynchus but is whiter on the 
underparts, especially the breast. These two subspecies are insep- 
arable according to the twenty-fourth supplement to the American 
Ornithologists' Union Check-list of North American Birds (Auk, vol. 
66,1949:281). 

5—8131 



66 University of Kansas Publs., Mus. Nat. Hist. 

PuflSnus tenuirostris (Temminck) 
Short-tailed Shearwater 

Procellaria tenuirostris Temminck, PI. Co!., livr. 99, 1835, text to pi. 587. (Type 
locality, Seas north of Japan and shores of Korea.) 

Pufinus tenuirostris tenuirostris Bryan, Guam Rec, vol. 13, no. 2, 1936, p. 15 
(Guam). 

Pufinus tenuirostris Yamashina, Tori, 10, 1940, p. 677 (Kinajon, Marshall Islands) ; 
Hand-list Japanese Birds, 3d ed., 1942, p. 210 (Kinajon, Marshall Islands). 

Geographic range. — Breeds in Tasmania, southeastern Australia, islands in 
Bass Straits, and Bounty Islands. Ranges north to the Bering Sea. In 
Micronesia: Mariana Islands — Guam?; Marshall Islands — Kinajon. 

Character. — A rather large shearwater with short, rounded tail; upper parts 
sooty brown; underparts paler and more grayish than back; throat may be 
occasionally whitish; bill lead-gray; feet grayish, browner on outer side. 

Remarks. — On migration this shearwater probably reaches most 
parts of Micronesia. It has been recently recorded by the Japanese 
at Kinajon in the Marshall Islands. Bryan (1936:15) includes 
this species as a ''chance arrival" in his list of the birds of Guam. 

Puffinus nativitatis Streets 
Christmas Shearwater 

Puffinus (Nectris) nativitatis Streets, Bull. U. S. Nat. Mus., 7, 1877, p. 29. (Type 
locality, Christmas Island, Pacific Ocean.) 

Puffinus nativiatis Salvin, Cat. Birds British Mus.. 25, 1896, p. 389 (Krusenstern) ; 
Lister, Proc. Zool. Soc. London, 1891, pp. 295-300 (Krusenstern); Godman, Monogr. 
Petrels, pt. 3, 1908, p. 153 (Marshalls). 

Geographic range. — Breeds at Wake and Laynan Islands south to Christmas, 
Phoenix, Marquesas, Tuamotu, and Austral Islands. In Micronesia: Marshall 
Islands — Ailuk. 

Characters. — Upper parts chocolate brown; underparts resemble upper parts 
but throat may be slightly grayer; bill and feet black. P. nativitatis resembles 
P. pacificus but is similar with black feet. 

Remarks. — The only specimens of this bird known from Micro- 
nesia, are those taken in the spring of 1883 by H. J. Snow at 
Krusenstern (Ailuk) in the Marshall Islands. For two birds from 
this island in the collections of the British Museum, Godman (1908: 
154) gives the following measurements: wing, 9.6 and 10.0; tail, 
3.35 and 3.4; culmen, 1.15 and 1.2; tarsus, 1.7 and 1.8; middle toe 
and claw, 2.0 and 2.1. 

Puffinus Iherminieri dichrons Finsch and Hartlaub 
Dusky Shearwater 

Puffinus dichrous Finsch and Hartlaub, Fauna Centralpolynesiens, 1867, p. 244. 
(Type locality, McKean Island, Phoenix Group.) 

Puffinus dichrous Hartlaub and Finsch, Proc. Zool. Soc. London, 1872, pp. 90, 
108 (Pelew); Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 6, 44 (Palau). 

Puffinus opisthomelas var. minor Hartlaub, Proc. Zool. Soc. London, 1867 (1868), 
p. 832 (Type locality, Pelew); Finsch, Journ. f. Ornith., 1872, p. 57 (Pelew). 



Baker: The Avifauna of Micronesia 



67 



Puffinus opisthomelas Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 9, 
118 (Pelew); Finsch, Journ. f. Ornith., 1870, p. 371 (Pelew). 

Puffinus tenebrosus Pelzeln, Ibis, 1873, p. 47, fig. 1 (Type locality, unknown = 
Pelew Islands, ex Mathews); Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 8, 
1896, p. 55 (Rota); Hartert, Novit. Zool., 5, 1898, p. 69 (Marianne); Seale, Occ. 
Papers Bernice P. Bishop Mus., 1, 1901, p. 23 (Marianas?); Safford, The Plant 
World, 7, 1904, p. 268 (Guam). 

Puffinus obscuTus Finsch, Journ. Mus. Godeffroy, 12, 1876, pp. 18, 40 (Ponape, 
Palau) ; idem, Proc. Zool. Soc. London, 1877, p. 786 (Palau) ; idem, Proc. Zool. Soc. 
London, 1877 (1878), p. 782 (Ponape) ; idem, Journ. f. Omith., 1880, pp. 295, 309 
(Ponape, Kuschai) ; idem, Proc. Zool. Soc. London, 1880, p. 577 (Ruk); idem. Ibis, 
1881, p. 109 (Kuschai); idem, Ibis, 1881, pp. 113, 115 (Ponape); Schmeltz and Krause, 
Ethnogr. Abth. Mus. Godeffroy, 1881, p. 353 (Ruk); Salvin, Ibis, 1888, p. 357 
(Pelew); Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), 
p. 79 (Ruk, Ponape, Pelew); Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 8, 1896, 
p. 54 (Saypan, Palaos); Salvin, Cat. Birds British Mus., 25, 1896, p. 382 (Carolines, 
Pelews); Hartert, Novit. Zool., 5, 1898, p. 68 (Marianne); Seale, Occ. Papers Bernice 
P. Bishop Mus., 1, 1901, p. 23 (Marianas?); Safford, Osprey, 1902, p. 70 (Marianas); 
Dubois, Syn. Avium, 2, 1904, p. 1031 (Pelew, Carolines); Godman, Monogr. Petrels, 
pt. 2, 1908, pp. 126, 127 (Pelew, Ruk, Ponape). 

Puffinus obscurus obscurus Hartert, Novit. Zool., 7, 1900, p. 10 (Ruk); Kuroda, 
in Moniiyama, Birds Micronesia, 1922, p. 32 (Saipan, Ruk, Ponape, Pelew). 

Puffinus Iherminieri minor Mathews, Birds Australia, 2, 1912, p. 70 (Pelew, Caro- 
lines). 

Puffinus assimitis minor Mathews, Syst. Avium Australasianarum, 1, 1927, p. Ill 
(Pelew). 

Puffinus Iherminieri dichrous Murphy, Aiiier. Mus. Novit., no. 276, 1927, p. 10 
(Pelews, Carolines); Peters, Check-list Birds World, 1, 1931, p. 60 (Pelew); Yama- 
shina. Tori, 7, 1932, p. 408 (Arakabesan) ; Hand-list Japanese Birds, rev., 1932, p. 
188 (Saipan, Truk, Ponape, Palaus); Hand-list Japanese Birds, 3rd ed., 1942, p. 209 
(Saipan, Truk, Ponape, Palau); Mayr, Birds Southwest Pacific, 1945, p. 10 (Caro- 
lines, Palaus) ; Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 37 (Palau). 

Alphapuffinus Iherminieri minor Mathews, Novit. Zool., 39, 1934, p. 182 (Pelew 
Islands). 

Puffinus obscura Bryan, Guam Rec, vol. 13, No. 2, 1936, p. 15 (Guam). 

Geographic range. — Known from Phoenix. Nauru, Micronesia, and south to 
the Samoan, Society, Tuamotu, and Marquesas islands. In Micronesia : Mari- 
ana Islands — Guam, Rota, Saipan; Palau Islands— Babelthuap, Koror, Araka- 
besan; Caroline Islands — Truk, Ponape, Kusaie. 

Characters. — A small shearwater with upper parts sooty-black; under parts 
white except for sides of breast grayish and under tail-coverts blackish; bill 
blackish; feet yellowish, outer toe black. 

Measurements. — Measurements of 17 adult birds (9 males, 7 females, 1 un- 
sexed) from Micronesia (Palau, Truk, Ponape. Kusaie) and 10 adult birds (6 
males, 4 females) from the Phoenix Group (Enderbury, Canton) are listed in 
table 14. 



T.-iBLE 14 


Me-asurements of Puffinus 


Iherminieri dichrous 


Locality 


Wing 


Tail 


Exposed 
cu men 


Tarsus 


Micronesia 

Phoenix 


203 (197-211) 
197 (193-203) 


83.6 (77-89) 
82.2 (79-85) 


27.9 (26-30) 
26.3 (25-28) 


38.5 (37.5-40) 
37.2 (36-39) 







68 University of Kansas Publs., Mus. Nat. Hist. 

Specimens examined. — Total number, 72 (44 males, 19 females, 9 unsexed), as follows: 
Palau Islands, AMNH — exact locality not givpn, 64 (Oct., Nov., Dec); Caroline Islands, 
AMNH — Truk, 4 (June 15, 16)— Ponape, 3 (imdated)— Kusaie, 1 (April 25). 

Nesting. — The Dusky Shearwater in Micronesia nests in holes on high, and 
usually isolated, coral cliffs. Owston's collectors, according to Hartert (1900:10), 
found a nest with one egg at Truk on June 16. The nest was in a hole four feet 
deep in the side of a cliff. The egg is white and mea.sures 42 X 35. Yamashina 
(1932a :408) records the taking of one egg at Arakabesan, Palau Islands, on 
May 26. Coultas (field notes) gives an interesting account of nesting activities 
of this shearwater at the Palau Islands. He found the bird nesting on small 
islands of the group from October to December, 1931 ; however, he states that 
the natives told him that the bird nests throughout the year. Land crabs and 
shearwaters were often found together in the same burrow. Apparently the 
adult birds did not remain in the burrow with the young during the day. At 
Kusaie, Coultas was told by the natives that the adult birds were caught by 
tying the mandibles of the young together. When the parent birds approached 
and hovered over the young birds expecting their mouths to open, the natives 
had the opportunity to strike them down with clubs. Coultas collected six 
downy nestlings at Palau in November and December. 

Remarks. — The first published account of this shearwater in 
Micronesia was apparently by Kittlitz (1858, pt. 1:358) when he 
recorded his "Schwarzlicher Sturmvogel" at Kusaie, according to 
Wiglesworth (1891a:79). Finsch (1875:44 and 1881b:113, 115) 
studied specimens taken by Tetens, Heinsohn, and Kubary at the 
Palau Islands and those taken by Kubary at Ponape. Earlier, 
Hartlaub (1868:832) used some of these specimens from the Palau 
Islands to describe his Puffinus oyisthomelas var. minor, which was 
destined to be placed in synonymy (Murphy, 1927:10). Oustalet 
(1896:54, 55) recorded specimens taken by Marche at Saipan in 
May, 1887, and at Rota in July, 1888. Oustalet referred to them as 
P. obscuras and P. tenebrosus, respectively. T. W. Gulick obtained 
undated skins at Ponape. Hartert (1900:10) reported on specimens 
taken by Owston's collectors at Truk. In 1931, Coultas with the 
Whitney South Sea Expedition took one shearwater at Kusaie and a 
series of 64 skins at the Palau Islands. He failed to find birds at 
Ponape and wrote that their scarcity there may have been due to 
persistent hunting of them by the inhabitants of the island. The 
NAMRU2 party obtained no information concerning the birds at 
Guam, Rota, or Truk, but at the Palau Islands observed shearwaters 
at sea approximately 6 miles east of Babelthuap Island on Septem- 
ber 2, 1945. 

Murphy (1927:6-15) revised the shearwaters of the Puffinus 
Iherminieri group, and recognized several subspecies. P. I. dichrous 
was assigned a range consisting of Micronesia, the Phoenix Islands, 
and Nauru Island. The breeding range of P. I. polynesiae was given 



Baker: The Avifauna of Micronesia 69 

as the Samoan, Society, Tuamotu and Marquesas islands. Color dif- 
ferences between the two subspecies are very slight, and he separated 
them on the basis of the length of the exposed culmen as follows: 
P. I. dichrous 22.6-27 (26) in P. I. polijnesiae 25.5-30 (28.9). In 
other measurements they closely resembled one another. At the time 
of his study, Murphy did not have the shearwaters from Micronesia 
collected by Coultas and actually did not have a large series from 
these islands. On studying this new material, I find the length of 
the exposed culmen of 17 adult birds from Micronesia (including 12 
from the Palaus) to be 26-30 (27.9). In comparison with Murphy's 
findings, my measurements of Micronesian birds fall almost midway 
between the measurements which he recorded as characteristic of 
P. I. dichrous (from the Phoenix Islands) and P. I. polynesiae. The 
intermediate position of the measurements of the Micronesian birds, 
together with the absence of other distinguishing characters, suggests 
that these shearwaters belong to only one subspecies which consists 
of a group of isolated and variable populations. Unless the old spe- 
cific name, obscuras of Gmelin, is revived, the name for the entire 
group in Micronesia and Polynesia would be P. I. dichrous. I agree 
with Murphy that the Bonin form, P. /. bannermani, is a well- 
defined subspecies. 

Pterodroma rostrata rostrata (Peale) 
Tahiti Petrel 

Procellaria rostrata Peale, U. S. Expl. Exp., 8, 1848, p. 296. (Type locality, Moun- 
tains about 6,000 feet on Tahiti, Society Islands.) 

Procellaria desolata Pucheran, Voy. Pole Sud, 3, 1853, p. 138 (des lies Carolines); 
Hartlaub, Journ. f. Ornith., 1854, p. 168 (Carolinen). 

Procellaria (.Aestrelata) desolata Gray, Cat. Birds Trop. Is. Pacific Ocean, 1859, 
p. 55 (Caroline Islands). 

Oestrelata rostrata Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 
1890-1891 (1891), p. 82 (Caroline Is.); Godman, Monogr. Petrels, pt. 3, 1908, p. 190 
(Caroline Is.). 

Pterodroma rostrata Kuroda, in Momiyama, Birds Micronesia, 1922, p. 33 (Caro- 
lines). 

Pterodroma rostrata subsp. (?) Hand-list Japanese Birds, rev., 1932, p. 188 (Caro- 
lines); Hand-list Japanese Birds, 3d ed., 1942, p. 210 (Carolines). 

Geographic range. — Known to breed on the Society and Marquesas Islands. 
In Micronesia: Caroline Islands — exact locality unknown. 

Characters. — A large petrel with blackish-brown plumage except for belly 
and under tail-coverts white and throat, upper breast and flanks pale brown; 
bill black; legs yellowish; feet black. This oceanic bird differs from other 
petrels and shearwaters of Micronesia by the presence of a white abdomen in 
contrast with dark plumage on upper parts, throat, and breast. 

Remarks. — A petrel which is referred to this subspecies has been 
taken once in Micronesia, by Hombron and Jacquinot in the Caro- 
line Islands. It may be pointed out that the subspecies P. r. becki 



70 University of Kansas Publs., Mus. Nat. Hist. 

Murphy is known from the sea east of the Bismarck Archipelago 
and might range into Micronesian waters. 

Pterodroma hypoleuca hypoleuca Salvia 
Stout-billed Gadfly Petrel 

Oestrelata hypoleuca Salvin, Ibis, 1888, p. 359. (Type locality, Krusenstern 
Island = Ailiik, Marshall Islands, fide Fisher, Auk. 63, 1946, pp. 587-588). 

Oestrelata hypoleuca Salvin, Cat. Birds British Mus., 25, 1896, p. 409 (Krusen- 
stern); Godman, Monogr. Petrels, pt. 3, 1908, p. 212 (Krusenstern). 

Cookilaria hypoleuca hypoleuca Mathews, Syst. Avium Australasianarum, 1, 1927, 
p. 122, (Marshall Group). 

Pttrodroma leucoptera hypoleuca Hand-list Japanese Birds, rev., 1932, p. 188 
(Marshalls); Hand-list Japanese Birds, 3d ed., 1942, p. 210 (Krusenstern); Fisher, 
Auk, 63, 1946, pp. 387-388 (Ailuk). 

Pterodroma hypoleuca hypoleuca Mayr, Birds Southwest Pacific, 1945, p. 11 
(Micronesia). 

Geographic range. — Ranges from the Bonins east to the Hawaiians and 
south to Micronesia. In Micronesia: Marshall Islands — Ailuk. 

Characters. — Upper parts grayish except for forehead whitish, crown and 
nape sooty-black; underparts whitish except for sides of breast sooty-black; 
legs and feet flesh color except for tips of toes and webs which are black. 

Remarks. — In Micronesia, this petrel is known only from the type 
locality, Krusenstern or Ailuk, Marshall Islands. Fisher (1946: 
587-588) has corrected the confusion regarding the exact position 
of this type locality. 

Phaethon aethereus mesonauta Peters 
Red-billed Tropic-bird 

Phaethon aethereus mesonauta Peters, Occ. Papers Boston Soc. Nat. Hist., 5, 
1930, p. 261. (Type locality. Swan Key, Almirante Bay, Panama.) 

Phaeton aethereus Finsch, Ibis, 1880, pp. 329, 333, (Ratak Chain, Marshalls); 
idem, Joum. f. Ornith., 1880, p. 310 (Kuschai) ; idem. Ibis, 1881, p. 109 (Kuschai); 
Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 73 
(Kushai, Marshalls) ; Ogilvie-Grant, Cat. Birds British Mus. 26, 1898, p. 457 (Kushai, 
Marshalls); Schnee, Zool. Jahrbiicher, 20, 1904, p. 390 (Marschall Inseln) ; Kuroda, 
in Momiyania, Birds Micronesia, 1922, p. 34 (Kusaie, Marshall Islands). 

Phaethon aethereus [tmesonauta] Hand-list Japanese Birds, rev., 1932, p. 187 
(Kusaie, Marshalls); Hand-list Japanese Birds, 3d ed., 1942, p. 208 (Kusaie, 
Marshall Islands). 

Geographic range. — Tropical parts of Atlantic and eastern Pacific from Cape 
Verde Islands west to Panama and Galapagos Islands. In Micronesia: Caro- 
line Islands — Kusaie; Marshall Islands — ^Ratak Chain. 

Characters. — 'Adult: A large, white sea bird with a long white tail; dorsal 
surface marked with blackish, transverse vermiculations; bill red; tarsus and 
foot flesh-colored with a yellowish hue, with plantar surface grayish. Imma- 
ture: Resembles adults but dark transverse bars are broader; crown blacker; 
bill yellow. 

Remarks. — No specimens have been examined. The Red-billed 
Tropic-bird is placed in the list of birds known from Micronesia on 



Baker: The Avifauna of Micronesia 71 

the basis of two observations by the German ornithologist, Otto 
Finsch. It has not been reported since his time, and may be con- 
sidered as an unusual record for the area. I am following the Hand- 
list of Japanese Birds (Hachisuka et al, 1942:208) in assigning 
the bird to the subspecies, P. a. mesonauta. 

Phaethon rubricauda rothschildi (Mathews) 
Red-tailed Tropic Bird 

Scaeophaethon rubricauda rothschildi Mathews, Birds Australia, 4, 1915, p. 303. 
(Type locality, Laysan and Niihau.) 

Phaeton rubricaudus Finsch, Journ. f. Ornith., 1880, p. 296 (Carolines); idem, Ibis, 
1881, p. 115 (Ponape). 

Phaeton rubricauda Finsch, Proc. Zool. Soc. London, 1880, p. 577 (Ruk) ; Wigles- 
worth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 73 (Ruk, 
Ponape, Marshalls). 

Phaeton rubricauda Ogilvie-Grant, Cat. Birds British Mus., 26, 1898. p. 451 (Caro- 
line Islands); Hartert, Novit. Zool., 7, 1900, p. 11 (Ruk); Hand-list Japanese Birds, 
rev., 1932, p. 187 (Pagan, Truk, Ponape, Marshalls). 

Scaeophaethon rubricauda Kuroda, in Momiyama, Birds Micronesia, 1922, p. 34 
(Mariannes, Ruk, Ponape, Marshalls). 

Phaethon rubricauda rothschildi Yamashina, Tori, 7, 1932, p. 406 (Pagan); idem, 
Tori, 10, 1940, p. 676 (Maug). 

Phaethon rubricaudus rothschildi Hand -list Japanese Birds, 3d ed., 1942, p. 209 
(Maug, Pagan, Truk, Ponape, Marshalls). 

Geographic range. — Bonin and Hawaiian islands south to Micronesia. In 
Micronesia: Mariana Islands— Maug, Pagan ; Caroline Islands— -Truk, Ponape; 
Marshall Islands — exact locality unknown. 

Characters.— kdwM: Long-tailed sea bird white with pinkish tint except 
for black lores and eye streak; black shafts on feathers of secondaries, flanks, 
and tail coverts; black bases on feathers of head; central tail feathers elongate 
with black shafts and bright red webs; bill orange-red with black nasal streak; 
tarsus and foot bluish-yellow, distal part blackish. Immature: Resembles 
adult but barred with black above ; bill blackish. 

Measurements. — Yamashina (1940:676) hsts the measurements for seven 
adult birds from Maug in the northern Marianas as wing 304-319 and exposed 
culmen 55-62. 

A^esfmgr.— Yamashina (1932a :406) reports the taking of one egg at Pagan 
in the Marianas on February 15, 1931. 

Remarks. — The Red-tailed Tropic Bird has been recorded from 
the Mariana, Caroline, and Marshall Islands. On the basis of our 
present knowledge it appears to be uncommon in most of Micronesia 
and may be established as a resident bird only in the northern 
Marianas, as shown by Yamashina (1932a :406 and 1940:676), 
Coultas obtained an immature male at 3° N and 158° E, which is 
at a point in the ocean south of the eastern Carolines. Possibly this 
bird and others obtained in the Carolines are representatives of the 
subspecies, P. r. melanorhynchos Gmelin, which is known from the 
Palmerston, Society and Turtle islands. 



72 University of Kansas Publs., Mus. Nat. Hist. 



Phaethon lepturus dorotheae Mathews 
, White-tailed Tropic Bird 

Phoethon lepturus dorotheae Mathews, Austr. Avium. Rec, 2, 1913, p. 7. (Type 
locality, Queensland.) 

Phaeton candidus Kittlitiz. Denkw. Reise russ. Amer. Micron, unci Kamchat., 
1, 1858, p. 382 (Ualan); Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 9, 
118 (Pelew); Finsch, Journ. f. Ornith., 1872, p. 57 (Pelew) ; Hartlaub and Finsch, 
Proc. Zool. Soc. London, 1872, pp. 90, 114 (Pelew, Ualan); Finsch, Journ. Mus. 
Godeffroy, 1875, pp. 6, 47 (Palau) ; ide7n, Proc. Zool. Soc. London, 1877 (1878), 
p. 782 (Ponape); idem, Journ. f. Ornith., 1880, pp. 296, 309 (Ponape, Kuschai) ; 
idem, Proc. Zool. Soc. London, 1880, p. 577 (Ruk) ; Schmeltz and Krause, Ethnogr. 
> . . Abth. Mus., Godeffroy, 1881, pp. 281, 299, 330, 353 (Ponape, Mortlock, Nukuor, Ruk); 
Finsch, Mitth. Ornith. Ver. Wien, 1884, p. 52 (Kuschai); Wiglesworth, Abhandl. 
und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 73 (Pelew, Ruk, Luganor, 
Nukuor, Ponape, Ualan, Marshalls) ; Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, 
(3), 8, 1896, p. 62 (Agrigan, Palaos, Ruk, Kushai, Marshalls); Hartert, Novit. Zool., 
5, 1898, p. 68 (Marianne). 

Phaeton flavirostris Finsch, Ibis, 1880, pp. 329, 333 (Ratak Chain); idem. Ibis, 
1881, pp. 105, 109, 115 (Kuschai, Ponape). 

Phaethon candidus Salvador!, Ornith. Papuasia, 3, 1882, p. 426 (Pelews, Carolines, 
Marshalls); Scale, Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 23 (Agrigan); 
Safford, Osprey, 1902, p. 70 (Mariannes); Takatsukasa and Kuroda, Tori, 1, 1915, 
p. 50 (Pelew, Ponape); Uchida, Annot. Zool. Japon., 9, 1918, pp. 489, 492 (Palau). 

Phaethon lepturus Ogilvie-Grant, Cat. Birds British Mus., 26, 1898, p. 453 
(Pelew, Carolines, Marshalls); Hartert, Novit. Zool., 7, 1900, p. 10 (Ruk); Safford, 
The Plant World, 7, 1904, p. 268 (near Guam); idcrn, Contr. U. S. Nat. Herb., 9 
1905, p. 80 (northern Marianas) ; Mayr, Birds Southwest Pacific, 1945, p. 17 (Palau) ; 
Strophlet, Auk, 63, 1946, p. 535 (Guam); Borror, Auk, 64, 1947, p. 416 (Agrihan) ; 
Stott, Auk, 64, 1947, p. 524 (Saipan). 

Phaeton lepturus Schnee, Zool. Jahrbiicher, 20, 1904, p. 390 (Marschall Insein). 

Leptophaethon lepturus dorothea Mathews, Birds Australia, 4, 1915, p. 309 (Pelew). 

Phaethan lepturus Cox, Island of Guam, 1917, p. 22 (northern Marianas). 

Leptophaethon lepturus lepturus Kuroda, in Momiyama, Birds Micronesia, 1922, 
p. 33 (Agrigan, Saipan, Pelew, Ruk, Luganor, Nukuor, Ponape, Kusaie, Marshalls). 

Phaethon lepturus dorotheae Yamashina, Tori, 7, 1932, p. 407 (Ponape) ; Hand- 
list Japanese Birds, rev., 1932, p. 187 (Agrigan, Pagan, Saipan, Agiguan, Palaus, Truk, 
Luganor, Nukuor, Ponape, Kusaie, Marshalls) ; Hand-list Japanese Birds, 3d ed., 
1942, p. 209 (Agrigan, Pagan, Saipan, Agiguan, Babelthuap, Koror, Urukthapel, 
Angaur, Unusuto, Truk, Luganor, Nukuor, Ponape, Kusaie, Namorik) ; Baker, Smith- 
son. Misc. Coll., vol. 107, no. 15, 1948, p. 38 (Guam, Peleliu, Ulithi, Truk). 

Geographic range. — Island.s in the southwestern Pacific area. In Micronesia: 
Mariana Islands — Agrigan. Pagan, Saipan, Agiguan, Rota, Guam; Palau L^lands 
— Babelthuap, Koror, Urukthapel, Peleliu, Anguar, Unusuto; Caroline Islands 
Truk, Ulithi, Luganor, Nukuor, Ponape, Kusaie; Marshall Islands — Namorik. 

Characters. — Adult : White often with pinkish shade but lores and eye streak 
black; feathers of head, flanks and under tail-coverts with bases black; black 
on outer and subterminal part of inner webbing of primaries; black, sub- 
terminal coloring on scapulars and secondaries; black on shafts of elongated 
tail plumes; bill horn yellow, dark basally; tarsus dark yellow; feet blackish. 

Immature: Resembles adult but upper parts barred with black, bill black 
on terminal part. 

Meas«rer?ien/s.— Measurements of adult birds from Micronesia are given in 
table 15. 

Weights. — The NAMRU2 party recorded weights of five adult males from 
Guam as 294 (267-321) grams. 



Baker: The Avifauna of Micronesia 



73 



Table 15. Measurements of Phaethon lepturus from Micronesia 





No. 


Wing 


Tail 


Exposed 
culmen 


Tarsus 


Marianas: Asuncion, Guam. . . . 
Palaus: Peleliu 


6 
11 
11 


264 
256-287 

257 
242-270 

261 
252-271 


107 
97-117 

108 
98-122 

105 
97-114 


47 
44-50 

45 
40-49 

47 
44-49 


21 
20-21 

21 


Carolines: Ponape, Kusaie 


19-21 

21 
21-22 


Total: Micronesia 


28 


260 

242-287 


107 
97-122 


46 
40-50 


21 
19-22 



Specimens examined.— Tota] number, 37 (22 males, 10 females, 5 unsexed), as follows: 
Mariana Islands, USNM — Guam, 5 (June 11, July 21); AMNH — Asuncion, 1 (June ?); 
Palau Islands, USNM— Peleliu, 5 (Aug. 29, 31, Sept. 5, 6); AMNH— exact locality not 
given, 7 (Oct. 13, 26, Nov. 15, 23, Dec. 18); Caroline Islands, .\MNH— Ponape, 9 (Dec. 8, 9, 
undated) — Kusaie, 10 (March 1-8, April). 

Nesting. — The NAMRU2 party observed nests of the White-tailed Tropic 
Bird at Peleliu in August and September, 1945. Several nests vpere seen in 
hollows of the Australian pine (Casuarina equisetifolia) between 20 and 30 feet 
above the ground. Birds could be seen in the nest hollows because the plumes 
of their long tail usually extended well out of the entrance. One nest was 
found in a dead tree in a battle-cleared area; others were observed in jungle 
habitat. Coultas observed nesting at Ponape between November 1 and De- 
cember 30, 1930, and found nests in the tops of trees and in hollow trees; a few 
were observed in holes in cliffs. Yamashina (1932a :407) records the taking of 
one egg at Ponape on August 18, 1931. At Guam the NAMRU2 party found 
birds along the high cliffs which edge the beach. There was no evidence that 
they were nesting from May to July; nevertheless males taken in June had 
enlarged gonads. The bird is known to breed at Namorik in the Marshall 
Islands, according to the Hand-list of Japanese Birds (Hachisuka et al., 1942: 
209). 

Food habits. — The NAMRU2 party found small fish in the stomachs of 
these birds taken at Peleliu. 

Parasites.— Uchida, (1918:489, 492) records the bird lice (Mallophaga), 
Colpocephalum epiphanes and Menopon eulasius, from the White-tailed Tropic 
Bird from Palau. 

Remarks. — Birds taken in Micronesia differ only slightly from 
those from other areas in Oceania. Within Micronesia (see table 15) 
the birds from the Palaus have the shortest wing and shortest ex- 
posed culmen. 

The White-tailed Tropic Bird appears more numerously in west- 
ern and northern Micronesia than in the Marshall Islands. This dis- 
tribution may be correlated with a preference for the "high" islands; 
especially those which have rocky cliffs, including Guam, Rota, 



74 University of Kansas Publs., Mus. Nat. Hist. 

Peleliu, Angaiir, and Truk. Reports were received in 1945 that the 
birds were only infrequently seen at Ulithi, a low atoll. Stott (1947: 
524) observed birds flying into rocky crevices at Saipan on December 
18. Gleise (1945:221) also recorded the bird in the vicinity of Sai- 
pan. Borror (1947:416) reports seeing birds at Agrigan on July 29, 
August 5 and 6, 1945. Coultas (field notes) found tropic birds com- 
mon at Ponape in November and December, 1930, in forested re- 
gions and along the cliffs. He made similar observations at Kusaie 
and Palau. At Ponape and Palau, Coultas noted the use of the eggs, 
young and adults as food by the natives. At Palau the plumes are 
used in headdresses worn by the natives, the birds being taken with 
the blowgun. 

Murphy (1936:807) states that the principal enemy of the White- 
tailed Tropic Bird at Bermuda is the introduced rat [Rattus rattus). 
Introduced rats, particularly Rattus mindanensis on Guam, may 
prey on the nesting birds. Baker (1946c :404) writes that this rat is 
a good climber and may spend considerable time in trees. The rat 
was trapped also in rough coral jungle at the edge of the cliffs, where 
tropic birds, Micronesian Starlings and other species, may have been 
nesting. 

Little has been recorded concerning the post-breeding season wan- 
derings of these tropic birds in Micronesia. They seemingly spend 
considerable time at sea, but whether they move as far from their 
breeding areas as do birds in the Atlantic, as reported by Murphy 
(1936:803), Baker (1947a:253) and others, is not known. 

Murphy (1936:796) notes that the northward distribution of the 
tropic birds in the Atlantic is dependent on the warm currents of 
water. In the western Atlantic, the poleward-flowing, warm currents 
of the Gulf Stream allow for the northern extension of the range of 
these birds to Bermuda. In the eastern Atlantic, cool currents flowing 
toward the equator restrict the northern range. The same condition 
prevails in the eastern Pacific where warm current flowing toward 
the pole enable the birds to range north to the Bonins and other 
islands. 

The three species of tropic birds known from Micronesia overlap 
very little in their ranges in this area. The White-tailed Tropic Bird 
has become firmly established in the western part of Micronesia, but 
there are only a few records from the extreme eastern part. The 
Red-tailed Tropic Bird appears to be resident only in the northern 
Marianas although it has been recorded in the Carolines and Mar- 
shalls. Interspecific competition may prevent considerable inter- 



Baker: The Avifauna of Micronesia 75 

mingling of breeding populations in Micronesia, or it may be that 
each species requires different ecologic conditions. 

Sula dactylatra personata Gould 
Masked Booby 

Sula personata Gould, Proc. Zool. Soc. London, 1846, p. 21. (Type locality, North 
and northeast coasts of Australia = Raine Island.) 

Sula cyanops Finsch, Ibis, 1880, p. 219 (Taluit); Wiglesworth, Abhandl. und Bar. 
Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 72 (Marshalls) ; Ogilvie-Grant, Cat. 
Birds British Mus., 26, 1898, p. 430 (Marshalls). 

Parasula dactylatra personata Kuroda, in Momiyana, Birds Micronesia, 1922, p. 35 
(Marshall Islands); Mathews, Syst. Avium Australasianarum, 1, 1927, p. 232 (Marshall 
Islands). 

Sula dactylatra personata Yamashina, Tori, 7, 1932, p. 407 (Medinilla) ; Hand-list 
Japanese Birds, rev., 1932, p. 187 (Medinilla, Marshall Islands) ; Hand-list Japanese 
Birds, 3d ed., 1942, p. 208 (Medinilla, Marshall Islands). 

Geographic range. — Central and western Pacific from the Hawaiian Islands 
south to Australia, probably also in the Indian Ocean. In Micronesia: Mari- 
ana Islands — Medinilla; Marshall Islands — Jaluit? 

Characters. — Adult: A large, white sea bird, with brown wings and tail; 
face dark blue; bill horn-colored with base orange-yellow in males and pink 
or light red in females; feet olive in males and lead gray in females. 

Immature: Resembles adult, but head, wings, tail, chin and throat dark 
brown ; some white mottling ma}^ be present on back and rump ; bill dark ; 
feet lead colored. 

Nesting. — Yamashina (1932a :407) reports the taking of 12 eggs on Febru- 
ary 19, 1931, at Medinilla Island in the Marianas. 

Remarks. — No specimen has been examined by me from the area 
reported upon. Little is known regarding the distribution of the 
Masked Booby in Micronesia. It is found on the island groups 
which surround Micronesia and future field observations probably 
will add to our knowledge of its occurrence in this area. It is known 
to be resident only in the northern Marianas. 

Sula sula rubripes Gould 
Red-footed Booby 

Sula rubripes Gould, Syn. Birds Australia, pt. 4, 1838, app., p. 7. (Type locality. 
New South Wales = Raine Island.) 

Pelecanus piscator Kittlitz, Obser. Zool., in Lutke, Voy. "Le Seniavine," 3, 1836, 
pp. 296, 299 (Lougounor ^ Lukunor) ; idem, Denkw. Reise russ. Amer. Micron, und 
Kamchat., 1, 1858, p. 351 (Lugunor). 

Dysporus piscator Hartlaub, Proc. Zool. Soc. London, 1867 (1868), p. 831 (Pelew); 
Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 9, 118 (Pelews) ; idem, Proc. 
Zool. Soc. London, 1872, p. 90 (Pelew); Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 
6, 47 (Palau). 

Sula piscatrix Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890- 
1891 (1891), p. 72 (Pelew, Luganor) ; Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, 
(3), 8, 1896, p. 64 (Rota, Palaos, Carolines); Hartert, Novit. Zoo!., 5, 1898, p. 68 
(Marianne); Safford, Osprey, 1902, p. 70 (Rota); idem, The Plant World, 7, 1904, p. 
267 (Guam); idem, Contr. U. S. Nat. Herb., 9, 1905, p. 80 (Guam); idem, Guam, 
1912, p. 19 (Guam): Cox, Island of Guam, 1917, p. 22 (Guam). 



76 University of Kansas Publs., Mus. Nat. Hist. 

Sula piscator Ogilvie-Grant, Cat. Birds British Mus., 26, 1898, p. 432 (Pelew) ; 
Seale, Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 24 (Guam). 

Piscatrix sula rubripes Kuroda, in Momiyama, Birds Micronesia, 1922, p. 34 (Pelew, 
Luganor, Rota). 

Sula sula rubripes Hand-list Japanese Birds, rev., 1932, p. 185 (Medinilla, Saipan, 
Rota, Palau, Lukunor, Likieb) ; Bryan, Guam Rec, vol. 13, no. 2, 1936, p. 15 
(Guam); Yamashina, Tori, 10, 1940, p. 676 (Maug, Bikar) ; Hand-list Japanese 
Birds, 3d ed., 1942, p. 208 (Maug, Medinilla, Saipan, Rota, Palau, Lukunor, Bikar, 
Likieb). 

Geographic range. — Indian Ocean east to central Pacific islands. In Micro- 
nesia: Mariana Islands — Maug, Medinilla, Saipan, Rota; Palau Islands — ex- 
act locality unknown; Caroline Islands — Lukunor; Marshall Islands — Bikar, 
Likieb, Bikini, Enivvetok. 

Characters. — Adult: A large sea bird with plumage of variable color, mainly 
white or partly bufT with black primaries and black-tippe'd secondaries, or 
grayish or brownish with white or grayish tail; throat blackish; face blue or 
green; bill bluish and lighter at tip; legs and feet red. 

Immature: Resembles adult, but often wholly brownish, lighter ventrally; 
bill blackish; feet yellowish red. Immature resembles that of S. leucogaster. 

Nesting. — Morrison obtained a male nestling at Bikini on May 3, 1946. 

Specimens examined. — Total number, 10 (3 males, 7 females) from Marshall Islands, 
USNM— Bikini (April 28, May 1, 2, 3). 

Remarks. — The writer saw several bircis approximately 20 miles 
east of Eniwetok on January 7, 1945. Morrison obtained a series of 
birds at Bikini in April and May, 1946. Murphy (1936:861-870) 
presents a wealth of information concerning the bird. He points 
out the need for a better understanding of the plumages of the adult 
birds and gives evidence that the birds of different colors may occur 
within the same population. He describes the Red-footed Booby as 
nesting in trees and shrubs. This type of nesting environment is 
present at many of the islands in Micronesia. 

Sula leucogaster plotus (Forster) 
Brown Booby 

Pelecanus Plotus Forster, Descr. Anim., ed. Licht., 1844, p. 278. (Type locality 
Near New Caledonia.) 

Dysporus sula Hartlaub, Proc. Zool. Soc. London, 1867 (1868), p. 831 (Pelew); 
Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 9, 118 (Pelew); idem, 
Proc. Zool. Soc. London, 1872, p. 90 (Pelew); Finsch, Journ. Mus. Godeffroy, 8, 
1875, pp. 6, 47 (Palau); idem, Proc. Zool. Soc. London, 1880, p. 577 (Ruk) ; Hartert, 
Novit. Zool., 7, 1900, p. 11 (Ruk). 

Sula fusca Fmsch, Ibis, 1880, p. 218 (Taluit). 

Sula leucogastra Salvador!, Ornith. Papuasia, 3, 1882, p. 423 (Pelew, Carolinis) ; 
Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 72 
(Pelew, Ruk, Marshalls) ; Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 8, 1896, 
p. 63 (Palaos, Mariannes, Marshalls, Carolines); Hartert, Novit. Zool., 5, 1898, p. 68 
(Marianne). 

Sula sula Ogilvie-Grant, Cat. Birds British Museum, 26, 1898, p. 436 (Asuncion, 
Pelew); Seale, Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 24 (Guam); Safford, 
Osprey, 1902, p. 66 (Mariannas); ide7n. The Plant World, 7, 1904, p. 267 (Guam); 
idem, Contr. U. S. Nat. Herb., 9, 1905, p. 80 (Guam); idem, Guam, 1912, p. 19 



Baker: The Avifauna of Micronesia 77 

(Guam); Prowazek, Die deutschen Marianen, 1913, p. 100 (Marianen) ; Takatsukasa 
and Kuroda, Tori, 1, 1915, p. 50 (Marianne); Cox, Island of Guam, 1917, p. 22 
(Guam) ; Uchida, Annot. Zool. Japon., 9, 1918, pp. 487, 493 (Sea off Mariana Islands). 
Sula leucogaster plotus Kuroda, in Momiyama, Birds Micronesia, 1922, p. 34 
(Pelew, Ruk, West Faiu, Uracas, Saipan, Marshalls) ; Yamashina, Tori, 7, 1932, p. 407 
(Medinilla); Hand-list Japanese Birds, rev., 1932, p. 185 (Uracas, Pagan, Medinilla, 
Saipan, Truk, West Fayu, Grimes, Marshalls); Hand-list Japanese Birds, 3d ed., 
1942, p. 208 (Uracas, Pagan, Medinilla, Saipan, Grimes, West Fayu, Truk, Marshalls); 
Baker. Smithson, Misc. Coll., vol. 107, no. 15, 1948, p. 41 (Rota, Guam, Truk). 

Geographic range. — Throughout tropical Pacific area and south to Australia. 
In Micronesia: Mariana Islands — Asuncion, Uracas, Pagan, Medinilla, Saipan, 
Rota, Guam; Palau Islands — exact locality unknown; Caroline Islands- 
Grimes, West Fayu, Truk, Kusaie; Marshall Islands — Jaluit, Eniwetok. 

Characters. — Adult: A heavy sea bird dark brown except for white lower 
breast, belly, under tail, and auxillars; bill heavy and light bluish; face, gular 
pouch and feet greenish yellow. 

Immature: Resembles adult, but lower breast, belly and under tail mottled 
with brown; feet light yellow. 

Measurements. — Two adult males (Rota, Guam) measure: wing 386, 408; 
tail 194; exposed culmen 93, 98; tarsus 45, 49; two adult females (Rota, 
Kusaie): wing 380, 487; tail 193, 217; exposed culmen 94, 99; tarsus 45, 50. 

Weights. — The author (1948:41) records one immature female from Rota 
weighing 1042 grams. 

Specimens examined. — Total number, 6 (3 males, 3 females), as follows: Mariana Islands, 
USNM— Rota, 3 (Oct. 24); AMNH— Guam, 1 (July 23); Palau Islands, AMNH — exact 
locality not given, 1 (Dec. 1); Caroline Islands, AMNH — Kusaie, 1 (April 19). 

Nesting. — Few records have been published concerning nesting of the Brown 
Booby in Micronesia. Yamashina (1932a: 407) reports the taking of 12 eggs 
at Medinilla in the Mariana Islands on February 19, 1931. At Palau, Coultas 
(field notes) obtained reports that the bird nests at Kiangat, a small islet 
north of Babelthuap. 

Parasites.— Uchida, (1918:487, 493) obtained bird lice (Mallophaga), Meno- 
pan brevipalpe and Lipeurus potens, from the Brown Booby from the ''sea off 
Mariana Islands." 

Remarks. — The Brown Booby has not been founci abundantly by 
observers in the Micronesian area. Coultas and Kubary, who spent 
considerable time in this region, observed the bird at only a few of 
the islands. Probably the bird does not nest abundantly in Micro- 
nesia, although small colonies may be present. The NAMRU2 party 
observed a flock of twelve brown boobies on high cliffs at Taipingot 
Peninsula at Rota on October 24, 1945. Birds were seen also at 
Guam in May, July and November, 1945, and at Truk in Decem- 
ber of the same year. Coultas obtained a single specimen at Kusaie; 
the natives told him that it was not a resident of the island. The 
writer observed several Brown Boobies approximately twenty miles 
east of Eniwetok in the Marshall Islands on January 7, 1945. 
These were in the company of other sea birds. 



78 University of Kansas Publs., Mus. Nat. Hist. 

Phalacrocorax melanoleucus melanoleucus (Vieillot) 
Little Pied Cormorant 

Hydrocorax melanoleucos Vieillot, Nouv. Diet. Hist. Nat., 8, 1817, p. 88. (Type 
locality, "Australasie," restricted to New South Wales.) 

Carbo melanoleucus Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 9, 
118 (Pelew); idem, Proc. Zool. Soc. London, 1872, pp. 90, 114 (Pelew). 

Graculus melanoleucus Finsch, Journ. Mus. Godeffroy, 8, 1875, p. 48 (Pelew). 

Microcarbo melanoleucus Salvadori, Ornith. Papuasia, 3, 1882, p. 410 (Pelew); 
Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 72 
(Pelew). 

Phalacrocorax melanoleucus Ogilvie-Grant, Cat. Birds British Mus., 26, 1898, p. 
398 (Pelew); Nehrkom, Kat. Eiers., 1899, p. 235 (Palau); Takatsukasa and Kuroda, 
Tori, 1, 1915, p. 50 (Pelew); Uchida, Annot. Zool. Japon., 9, 1918, p. 486 (Palau). 

Ph [alacrocorax] melanoleucos Reichenow, Die Vogel, 1 1913, p. 127 (Palauinseln). 

Microcarbo melanoleucus Tuelanoleucus Kuroda, in Momiyama, Birds Micronesia, 
1922, p. 35 (Pelew). 

Microcarbo melanoleucus mehyillensis Mathews, Syst. Avium Australasianarum, 1, 
1927, p. 228 (Pelew); Hand-list Japanese Birds, rev., 1932, p. 186 (Babelthuap, 
Koror). 

Halietor vielanoleucos melanoleucos Peters, Check-list Birds World, 1, 1931, p. 93 
(Pelew). 

Phalacrocorax melanoleucus melanoleucus Mayr, Amer. Mus. Novit., no. 486, 1931, 
p. 5 (Pelew); Amadon, Amer. Mus. Novit., no. 1175, 1942, p. 2 (Palau); Mayr, 
Birds Southwest Pacific, 1945, pp. 50, 284 (Palau, Marianas) ; Baker, Smithson. Misc. 
Coll., vol. 107. no. 15, 1948, p. 41 (Palau). 

Phalacrocorax melanoleucos mclvillensis Hand-list Japanese Birds, 3d ed., 1942, p. 
207 (Pagan, Babelthuap, Koror, Angaur). 

Geographic range. — Tasmania, Australia, Leaser Sunda north through Mela- 
nesia to Palau Islands. In Micronesia : Palau Islands — Babelthuap, Koror, 
Garakayo, Ngabad, Peleliu, Anguar. 

Characters. — Adult: A small cormorant with upper parts black with dull 
greenish gloss; under parts white e.xcept vent and under tail-coverts which 
are sooty -black. 

Measurements. — ^The author (1948: 41) gives the following measurements of 
two adult females from Peleliu: wing, 220 and 222; tail, 153 and 157; culmen 
from notch of suture between maxilla and quadratojugal bones, 35 and 36. 

Specimens examined. — Total number, 15 (1 male, 12 females, 2 unsexed), as fol- 
lows: Palau Islands, USNM— Peleliu, 6 (Aug. 27, Sept. 7, 10, 16); AMNH— exact 
locality not given, 9 (Nov. part). 

Nesting. — Nehkorn (1899:235) recorded eggs taken at Palau. Some of 
the specimens obtained by Coultas in November, 1931, had swollen gonads. 
The author found no evidence of nesting in August and September, 1945, in 
the southern Palaus. 

Food habits. — The author (1948: 41) found small fish in the stomachs of 
birds taken in August and September. The contents of each stomach averaged 
approximately 3 cc. in volume. 

Parasites. — Uchida (1918:486) found the bird louse (Mallophaga), Lipeurus 
subsetosus, on the Little Pied Cormorant from Palau. 

Remarks. — The Palaus mark the northernmost point of range of 
the Little Pied Cormorant. It does not occur in the Philippines and 
must have reached Palau from the New Guinea region. It is un- 



Baker: The Avifauna of Micronesia 79 

known at Yap and other "high" islands in the Carolines. A sight 
record of this species at Pagan in the northern Marianas, made by 
Orii and reported in the Hand-list of Japanese Birds (Hachisuka et 
al., 1942:207), may be questioned. Amadon (1942:1) has studied 
the races of this species and points out that there is little geographic 
variation in the species; it is divisible into three subspecies. One 
of these is confined to New Zealand. Another occurs only on Ren- 
nell Island, Solomons. The six specimens taken by the NAMRU2 
party at Peleliu included only two adults, whose measurements are 
within the range of those studied by Amadon. 

The NAMRU2 party found the birds numerously in the southern 
Palaus in 1945. Birds were concentrated in the areas of mangrove 
swamp and on the tidal flats. In August and September, they were 
observed frequently in groups of 10 to 15, either sitting on the 
ground or perched on low mangroves or dead snags sunning them- 
selves. Coultas (field notes) received reports that they nested at 
a freshwater lake on the "main island" (Babelthuap?) 

Ripley (1948) reports the occurrence of "about a dozen anhingas 
(presumably Anhinga melanogaster)" at Babelthuap on 12 Novem- 
ber 1946. 

Fregata minor minor (Gmelin) 
Pacific Man-o'-War 

PcleraniLS minor Giiielin, Syst. Nat., 1, pt. 2, 1789, p. 572. (No type locality = 
Christmas Island, Indian Ocean.) 

Pelecanus aquilaf Quoy and Gaimard, Voy. "Uranie," Zool., 1824, p. 154 (Caro- 
lines). 

Pelecanus aqicilus? Lesson, Man. d'Ornith., 2, 1828, p. 354 (Carolines). 

Atagen aquilus Gray, Cat. Birds Trop. Is. Pacific Ocean, 1859, p. 61 (Ladrone or 
Marian Islands). 

Tachypetes aquila Finsch, Proc. Zool. Soc. London, 1880, p. 577 (Ruk); Hartert, 
Novit. Zool., 7, 1900, p. 11 (Ruk); Prowazek, Die deutschen Marianen, 1913, p. 100 
(Marianen). 

Tachypetes aquilus Finsch, Ibis, 1880, p. 333 (Taluit) ; idem, Joum. f. Omith., 1880, 
pp. 296, 310 (Ponape, Kuschai) ; idem. Ibis, 1881, pp. 109, 115 (Kuschai, Ponape) ; 
Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, pp. 299, 353 (Mortlock, 
Ruk). 

Fregata aquila Salvadori, Ornith. Papuasia, 3, 1882, p. 403 (Carolines, Marshalls); 
Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891), p. 71 (Ruk, 
Luganor, Ponape, Ualan, Marshalls); Ogilvie-Grant, Cat. Birds British Mus., 26, 1898, 
p. 443 (Carolines, Marshalls); Finsch, Deut. Ver. zum Schultze der Vogelwelt, 25, 1900, 
p. 452 (Ponape, Kuschai, Marshalls); Seale, Occ. Papers Bernice P. Bishop Mus., 1, 
1901, p. 24 (Guam); Safford, The Plant World, 7, 1904, p. 267 (Guam); Schnee, Zool. 
Jahrbucher, 20, 1904, p. 390 (Marschall Inseln); Safford, Contr. U. S. Nat. Herb,, 9, 
1905, p. 80 (Guam); Cox, Island of Guam, 1917, p. 22 (Guam). 

Fregata aquila palmerstoni Kuroda, in Momiyama, Birds Micronesia, 1922, p. 35 
(Carolines, Marshalls). 

Fregata minor peninsulae Mathews, Syst. Avium Australasianarum, 1, 1927, p. 233 
(Carolines, Marshalls); Peters, Check-list Birds World, 1, 1931, p. 96 (Carolines?, 
Marshalls?). 



80 University of Kansas Publs., Mus. Nat. Hist. 

Fregata minor palmerstoni Hand-list Japanese Birds, rev., 1932, p. 186 (Yap, 
Faraulep, Truk, Lukunor, Ponape, Kusaie, Namu, Likieb) ; Bryan, Guam Rec, vol. 
13, no. 2, 1936, p. 15 (Guam); Yamasiiina, Tori, 10, 1940, p. 676 (Maug, Bikar). 

Fregata minor minor Hand-list Japanese Birds, 3d ed., 1942, p. 207 (Maug, Yap, 
Faraulep, Truk, Lukunor, Ponape, Kusaie, Namu, Bikar, Likieb). 

Fregata minor Borror, Auk, 64, 1947, p. 416 (Agrihan). 

Geographic range. — Eastern Indian Ocean to western Pacific Ocean. Limits 
of range not certainly known. In Micronesia: Mariana Islands — Agrigan, 
Maug, Saipan, Guam; Caroline Islands — Yap, Faraulep, Truk, Lukunor, 
Ponape, Ku.saie; Marshall Islands — Namu, Bikar, Likieb, Kwajalein, Bikini. 

Characters. — Adult male : Large sea bird with deeply forked tail ; blackish 
but wing-coverts paler; head and back glossy purple and blue; breast lighter 
than belly. Adult female; Resembles adult male, but head blacker; chin and 
throat grayer; breast more whitish. Immature: Resembles adult, but head and 
throat whitish washed with buff; breast dark brown; belly whitish. 

Measurements. — Two adult males measure: wing, 572; tail, 354, 396; exposed 
culmen, 98. 103; two adult females; wing. 583. 604; tail, 365; exposed culmen, 
119, 127. These four specimens are from Bikini. 

Specimens examined. — Total number, 10 (3 males, 7 females), from Marshall Islands, 
USNM— Bikini (March 11, 22, 29, 30, April 13, 29, May 3, 14). 

Remarks. — The systematic position of the subspecies of Fregata 
minor in the Pacific area is not well established. I am following the 
committee who prepared the Hand-list of Japanese Birds (Hachi- 
suka et al., 1942:207) in using the name F. m. minor, although a 
thorough study may show that these birds have closer relationships 
to one of the other subspecies of the Pacific area. 

Fregata minor has been reported only occasionally in the Marianas 
and probably is not resident there. Borror (1947:416) reports the 
bird at Agrihan on August 11, 1945, and Seale (1901:24) mentions 
one taken at Guam in November, 1889. No records are known 
from the Palaus. In the Carolines the birds are probably resident, 
especially in the eastern part. In the Marshalls the species is a 
conspicuous member of the bird colonies on the coral atolls. Wal- 
lace (field notes) observed two birds at Loi Island in Kwajalein 
Atoll on May 7, 1944. Morrison obtained ten specimens at Bikini 
in the period from March through May in 1946. 

Fregata ariel arid (Gray) 
Least Man-o'-War 

Atagen (sic) Ariel Gray, Gen. Birds, 3, 184.5, col. pi. [185]. (Type locality, Raine 
Island, Queensland.) 

Pelecanus minor Lesson, Traite d'Ornith., 1831, p. 607 (Mariannes, Carolines). 

Tachypetes minor Hartlaub, Proc. Zool. Soc. London, 1867 (1868), p. 831 (Mac- 
kenzie Group); Hartlaub and Finsch, Proc. Zool. Soc. London, 1872, p. 90 (Uap) ; 
Griiffe, Joum. Mus. Godeffroy, 2, 1873, p. 123 (Yap). 

Fregata minor Salvadori, Ornith. Papuasia, 3, 1882, p. 40.5 (Mariannes, Mac- 
kenzie); Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 
(1891), p. 71 (Uap, Ngoli or Matelotas). 



Baker: The Avifauna of Micronesia 81 

Tachypetes aquila var. minor Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 
8, 1896, p. 65 (Rota, Carolines, Marshatis) ; Hartert, Novit. Zool., 5, 1898, p. 68 
(Marianne). 

Fregata ariel Ogilvie-Grant, Cat. Birds British Mus., 26, 1898, p. 447 (Marianas, 
Carolines); Seale, Occ. Pai>ers Bemice P. Bishop Mus., 1, 1901, p. 25 (Guam?); 
Safford, Osprey, 1902, p. 70 (Marianas); Bryan, Guam Rec, vol. 13, no. 2, 1936, 
p. 15 (Guam). 

Fregata ariel ariel Mathews, Birds Australia, 4, 1914-15, p. 285 (Carolines, 
Marshalls); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 36 (Yap, Ngoli, Rota); 
Hand-list Japanese Birds, rev., 1932, p. 186 (Rota, Yap, Ngulu, Uluthi) ; Hand-list 
Japanese Birds, 3d ed., 1942, p. 208 (Rota, Yap, Ngulu, Uluthi). 

Geographic range. — China coast and Philippines south to Australia and east 
to Pacific islands. In Micronesia: Mariana Islands — Guam?, Rota; Caroline 
Islands — Yap, Ngulu, Ulithi. 

Characters. — Adult male : Resembles F. m. minor, but smaller and blacker 
with upper parts lustrous greenish-blue and white patch on lower flank. 

Adult female: Resembles adult male, but browner with paler nape and 
white breast. Immature: Re.=embles adult, but with head, chin, throat, and 
belly white washed with rufous. 

Remarks. — Like F. minor, the Least Man-o'-War has not been 
observed often in Micronesia. Marclie obtaineci one female at Rota 
in June, 1888. D. H. Johnson saw a bird thought to be of this species 
at Agfayan Bay, Guam, on 4 June 1945. Records from the western 
Carolines are few. There are no reports of this bird from the Palaus 
and the Marshalls. It may breed on some of the atolls in the 
Carolines. 

The two species of man-o'-war birds may be difficult to distinguish 
in the field. The smaller size of Fregata ariel is perhaps the most 
useful character although it may be easily recognized also by the 
presence of the white flank patch, if it can be observed. 

Both of the species of Fregata discussed in this report have repre- 
sentatives in the Atlantic, Indian and Pacific oceans. Murphy 
(1936:920) has shown that the man-o'-war birds are able to cross 
the Isthmus of Panama between the Pacific and Atlantic oceans. 
This route may also be the means of dispersal for other species. The 
irregular distribution of these birds as well as of other sea birds in 
the oceanic islands of the Pacific may be caused by their remaining 
over waters which contain preferred foods and their avoidance of 
waters which lack preferred foods. 

Butorides striatus amurensis Schrenck 
Amur Green Heron 

Ardea (.Butorides) virescens var. amurensis Schrenck, Reise Amur Lande, 1, pt. 2, 
1860, p. 441. (Type locality, Amurland.) 

Butorides striatus javanicus Hand-list Japanese Birds, rev., 1932, p. 183 (Koror, 
Babelthuap). 

Butorides striatus amurensis Hand-list Japanese Birds, 3d ed., 1942, p. 204 (Babel- 
thuap, Koror); Mayr, Birds Southwest Pacific, 1945, p. 302 (Palau). 

&— 8131 



82 University of Kansas Publs., Mus. Nat. Hist. 

Geographic range. — Breeds in northeastern Asia, China, Japan, Bonins. Win- 
ters south to Philippines and Malaysia. In Micronesia: Palau Islands — Babel- 
thuap, Koror. 

Specimens examined. — Total number, 2 females, from Palau Islands, AMNH — exact locality 
not given (Nov. 13, Dec. 17-18). 

Remarks. — The Amur Green Heron has been recorded as a winter 
visitor to the Palau Islands. Two females taken by Coultas in No- 
vember and December, 1931, are imature. He comments (field notes) 
that he saw, in all, three birds in taro patch and mangrove swamp 
habitat. 

Bubulcus ibis coromandus (Boddaert) 
Cattle Egret 

Cancroma Coromanda Boddaert, Table PI. enlum., 1783, p. 54. (Type locality, 
Coromandel.) 

Ardeola ibis coromanda Hand-list Japanese Birds, rev., 1932, p. 183 (Koror). 

Bubulcus ibis coromandus Hand-list Japanese Birds, 3d ed., 1942, p. 204 (Koror, 
Babelthuap); Mayr, Birds Southwest Pacific, 1945, p. 302 (Palau). 

Geographic range. — India, Ceylon, east to China and Japan and south to 
Malaysia. In Micronesia: Palau Islands — Babelthuap, Koror. 

Remarks. — The Japanese ornithologists have recorded the Cattle 
Egret from Babelthuap and Koror in the Palau Islands. It is a win- 
ter migrant. 

Egretta intermedia intermedia (Wagler) 
Plumed Egret 

Ardea intermedia Wagler, Isis, 1829, p. 659. (Type locality, Java.) 
Egretta intermedia intermedia Hand-list Japanese Birds, rev., 1932, p. 183 (Koror); 
Hand-list Japanese Birds, 3d ed., 1942, p. 203 (Koror); Mayr, Birds Southwest Pacific, 
1945, p. 302 (Palau); Baker, Smithson. Misc. Ck)ll., vol. 107, no. 15, 1948, p. 42 
(Rota, Guam, Peleliu, Angaur, Ulithi). 

Egretta intermedia Wharton and Hardcastle, Journ. Parasitology, 32, 1946, pp. 306, 
310 (Ulithi); Baker, Ecol. Monogr., 16, 1946, p. 408 (Guam). 

Geographic range. — India and Ceylon east to Malaysia, Phihppines, China 
and Japan. In Micronesia: Mariana Islands — Saipan, Rota, Guam; Palau Is- 
lands — Koror, Peleliu, Angaur; Caroline Islands — Ulithi. 

Characters. — Adult: A large white heron with green facial skin; black legs, 
feet and toes. In breeding plumage: Head with crest; neck and back with 
ornamental plumes; bill black. Winter plumage: Without crest or plumes; 
bill yellow with blackish tip. Immature: Resembles adult in winter plumage, 
but feathers soft and downy. 

Measurements. — Five males from Saipan, Rota, Guam, and Angaur meas- 
ure: wing, 295-321 (308); tail, 112-127 (119); culmen, 85-87 (87); tarsus, 111- 
118 (114); three females from Saipan, Ulithi, Angaur: wing, 294-301 (297); 
tail, 101-116 (110); culmen, 77-83 (80); tarsus, 108-115 (107). 

Weights. — The author (1948:43) records the weights of two males from 
Guam as 445 and 463. 

Specimens examined. — Total number, 8 (5 males, 3 females), as follows: Mariana Islands. 
rSNM— Saipan, 2 (Sept. 29, Oct. 2)— Rota, 1 (Oct. 31)— Guam, 2 (June 13); Palau Islands, 
USNM— Angaur, 2 (Sept. 21); Caroline Islands, USNM— Ulithi, 1 (Aug. 15). 



Baker: The Avifauna of Micronesia 83 

Food habits. — The NAMRU2 party found grasshoppers, other insects, spiders 
and lizards in the stomachs of egrets taken at Guam, Ulithi, and Angaur. 

Parasites. — Wharton and Hardcastle (1946:306, 310) obtained the chiggers 
(Acarina), Neoschongastia egretta and N. emingi, from this egret from TTlithi. 

Remarks. — The NAMRU2 party obtained Plumed Egrets at 
Rota, Guam, Ulithi, and Angaur in 1945. Previously, the only 
known record was from Koror, as reported in the Hand-list of Japa- 
nese Birds (Hachisuka et al., 1932:183). In addition, in 1945, Joe 
T. Marshall, Jr., obtained two birds at Saipan, and Gleise (1945: 
220) reported seeing "white herons" at Tinian, which probably were 
egrets. Gleise estimated the number of these birds at Tinian to be 
fifty; he found them in swampy areas. At Rota, the NAMRU2 
party found a flock of sixteen birds in a cultivated field on October 
31. At Guam, egrets were first observed on February 25, 1945, when 
a flock of fourteen was found in a fallow rice paddy near Piti. This 
flock remained in this area and were seen occasionally until as late 
as June 13, when two were taken as specimens. A short time later 
(June 30) the entire area was cleared for military use and the birds 
were seen no more. At Agfayan Bay a flock of sixteen birds was 
found on the beach on July 24 and on August 6. These birds kept 
apart from Reef Herons which were also in the area. In June, 1946, 
M. Dale Arvey observed egrets in swamps along the Ylig River at 
Guam. At Ulithi Atoll, three egrets were seen on August 15 at 
Potangeras Island, feeding in grassy areas adjacent to the beach. 
In the southern Palaus, the NAMRU2 party found egrets in August 
and September on tidal flats and open grasslands at Peleliu and 
Angaur. At Peleliu, a flock of twenty-five birds was seen on Sep- 
tember 8 and a flock of eight birds on September 16. At Angaur 
approximately twenty birds were seen in groups of five or more on 
September 21. These birds, unlike the Reef Herons, preferred 
grasslands to beach areas for feeding and were usually seen in size- 
able flocks. 

There was no evidence of breeding; specimens examined were 
either immatures or adults in winter plumage, since they had yellow 
bills tipped with black and slight or no development of ornamental 
plumes. Birds taken at Guam in June and at Angaur in September 
had no ornamental plumes, while birds taken at Ulithi in August, at 
Saipan in September and October, and at Rota in late October show 
some development of the back plumes. Wharton and Hardcastle 
(1946:306) found the same species of chigger on Plumed Egrets 
from Ulithi and from Okinawa in the Riu Kiu Islands. The 



84 University of Kansas Publs., Mus. Nat. Hist. 

NAMRU2 party observed the birds in Micronesia from February 
until October in 1945, and although the Plumed Egret may be con- 
sidered as merely a visitor to Micronesia, it would not be surprising 
to find nests there. The fact that several new distributional records 
were obtained for Micronesia in 1945 may indicate that the birds 
have been overlooked by ornithologists in the past or that the birds 
are increasing the breadth of their winter (or breeding?) range. 

Demigretta sacra sacra (Gmelin) 
Reef Heron 

Ardea sacra Gmelin, Syst. Nat., 1, pt. 2, 1789, p. 640. (Type locality, Tahiti.) 
Ardea jugularis Kittlitz, Observ. Zool., in Lutke, Voy. "Le Seniavine," 3, 1836, pp. 
286, 299, 304 (Ualan, Lougounor, Guahan); Hartlaub, Journ. f. Omith., 1854, p. 167 
(Mariannen) ; Kittlitz, Denkw. Reise russ. Amer. Micron, und Kamchat., 2, 1858, p. 63 
(Ualan); Pelzeln, Reise "Novara," Vogel, 1865, pp. 118, 162, 120, 121 (Puynipet, 
Ualan). 

Ardea (Herodias) atra Gray, Cat. Birds Trop. Is. Pacific Ocean, 1859, p. 48 
(Ladrone or Marian Islands, Caroline Islands). 

Ardea sacra Hartlaub, Proc. Zool. Soc. London, 1867 (1868), p. 831 (Matelotas 
Islands); Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 8, 118 (Pelews); 
Finsch and Hartlaub, Journ. f. Ornith., 1870, p. 137 (Pelews, Matelotas) ; Gray, Hand- 
list Birds, 8, 1871, p. 28 (Marian, Carolines, Pelews, Matelotas); Hartlaub and Finsch, 
Proc. Zool. Soc. London, 1872, pp. 89, 104 (Pelew, Uap, Ualan); Griiffe, Journ. Mus. 
Godeffroy, 2, 1873, p. 123 (Yap); Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 5, 32 
(Palau); idem, Journ. Mus. Godeffroy, 12, 1876, pp. 18, 38 (Ponape, Ualan); idem, 
Proc. Zool. Soc. London, 1877 (1878), p. 781 (Ponape); idem, Journ. f. Ornith., 1880, 
pp. 294, 306 (Ponape, Kuschai); idem. Ibis., 1880, pp. 220, 330, 332 (Taluit) ; idem, 
Proc. Zool. Soc. London, 1880, p. 577 (Ruk); idem. Ibis, 1881, pp. 105, 106, 109, 115 
(Kushai, Ponape); Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, pp. 
299, 353 (Mortlocks, Ruk); Finsch, Mitth. Ornith. Ver. Wien, 1884, p. 51 (Jaluit, 
Kuschai); Oustalet, Le Nat., 1889, p. 261 (Mariannes); Wiglesworth, Ibis, 1893, p. 
211 (Marshalls); Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 8, 1896, p. 36 
(Guam, Marshalls, Palaos, Carolines); Schnee, Zool. Jahrbiicher, 20, 1904, p. 390 
(Marschall-Inseln). 

Demiegretta sacra Salvador!, Ornith. Papuasia, 3, 1882, p. 348 (Marshalls, Ualan, 
Ponape, Ruck, Pelew, Mariannis); Wiglesworth, Abhandl. Ber. Zool. Mus. Dresden, 
no. 6, 1890-1891 (1891), p. 67 (Marianne, Pelews, Luganor, Ruk, Ponape, Ualan, 
Taluit); Hartert, Novit. Zool., 5, 1898, p. 64 (Saipan) ; Sharpe, Cat. Birds British 
Mus., 26, 1898, p. 137 (Pelew, Carolines, Marshalls); Hartert, Novit. Zool., 7, 1900, 
p. 11 (Ruk); Seale, Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 29 (Guam); 
Safford, Osprey, 1902, p. 67 (Marianas); idem, The Plant World, 7, 1904, p. 266 
(Guam); Kuroda, Avifauna Riu Kiu, 1925, p. 129 (Micronesia); Bryan, Guam, Rec, 
vol. 13, no. 2. 1936, p. 15 (Guam); Bequaert, Occ. Papers Bernice P. Bishop Mus., 16, 
1941, p. 266 (Kusaie). 

Demigretta sacra Safford, Contr. U. S. Nat. Herb., 9, 1905, p. 79 (Guam); 
Prowazek, Die deutschen Marianen, 1913, p. 101 (Saipan, Tinian) ; Cox, Island of 
Guam, 1917, p. 21 (Guam); Bequaert, Mushi, 12, 1939, p. 81 (Kusaie); Warton, Ecol. 
Monogr., 16, 1946, p. 175 (Guam); Warton and Hardcastle, Journ. Parasitology, 32, 
1946, pp. 306, 316 (Ulithi, Guam). 

Demiegretta jugularis Takatsukasa and Kuroda, Tori, 1, 1915, p. 50 (Truk, Ponape, 
Pelew). 

Demiegretta jugulari-s grayi Uchida, Annot. Zool. Japon., 9, 1918, pp. 484, 488, 490 
(Ponape). 

Demiegretta sacra sacra Kuroda, in Momiyama, Birds Micronesia, 1922, p. 36 
(Guam, Saipan, Angaur, Luganor, Yap, Ngoli, Ruk, Ponape, Kusaie, Taluit). 

Demigretta sacra sacra Wetmore, in Townsend and Wetmore, Bull. Mus. Conip. 
Zool., 63, 1919, p. 171 (Kusaie); Mathews, Syst. Avium Australasianarum, 1, 1927, 
p. 198 (Carolines); Yaniashina, Tori, 7, 1932, p. 406 (Ponape); Hand-list Japanese 



Baker: The Avifauna of Micronesia 85 

Birds, rev., 1932, p. 183 (Saipan, Guam, Babelthuap, Peliliu, Angaur, Ngulu, Yap, 
Truk, Lukunor, Ponape, Kusaie, Jaluit, Majuro); Mayr and Amadon, Amer. Mus. 
Novit., no. 1144, 1941, p. 10 (Guam, Saipan, Palau, Ponape, Kusaie, Ruk, Tah) ; 
Hand -list Japanese Birds, 3d ed., 1942, p. 203 (Saipan, Rota, Babelthuap, Peliliu, 
Angaur, Ngulu, Yap, Ulithi, Truk, Lukunor, Ponape, Kusaie, Jaluit, Arhno, Majuro, 
Moloclab, Wotze, Likieb, Ailuk); Majt, Birds Southwest Pacific, 1945, pp. 51, 284 
(Micronesia); Downs, Trans. Kansas Acad. Sci., 49, 1946, p. 90 (Tinian) ; Strophlet, 
Auk, 68, 1946, p. 585 (Guam); Borror, Auk, 64, 1947, p. 417 (Agrihan) ; Stott, Auk, 
64, 1947, p. 524 (Saipan); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 42 
(Rota, Guam, Peleliu, Ulithi, Truk). 

Derrugretta sacra micronesiae Momiyama, Tori, 5, no. 22, 1926, p. 110 (Type lo- 
cality, Caroline Islands ; Pelew, Yap, Truk, Ponape, Kusaie). 

Geographic range. — Coasts of Asia and adjacent islands from Korea and 
Japan south to Malaysia, Australia, Melanesia, Polynesia and Micronesia. In 
Micronesia: Mariana Islands — Agrigan, Tinian, Saipan, Rota, Guam; Palau 
Islands — Babelthuap, Koror, Garakayo. Ngesebus, Peleliu, Ngabad, Anguar; 
Caroline Islands — Ulithi, Yap, Ngulu, Truk, Lukunor. Ponape, Kusaie; Mar- 
shall Island-s — Jaluit, Arhno, Majuro, Maloclab, Wotze, Likieb, Ailuk, Bikini, 
Eniwetok, Kwajalein. 

Characters. — A medium-sized heron with three color phases: in gray phase 
color of body varies from "deep blackish-slate" to light bluish-slate, particu- 
larly on the breast, with a white gular stripe ; wear and fading causes the color 
of the body to change to brownish-slate ; bluish-gray ornamental plumes may 
be present on adult ; in white phase color of body is pure white in adult stage ; 
plumage of immature may be mottled; in mottled phase there may be a 
variable amount of gray and white (for complete study of plumages of Demi- 
gretta sacra see Mayr and Amadon. 1941:4). 

Measurements. — Mayr and Amadon (1941:1) record the length of the wing 
of thirty adults from the Marianas and Carolines as 268-309 (284). Seven 
adult males obtained by the NAMRU2 party at Rota, Guam and Peleliu 
measure: wing, 287-307 (294); tail, 95-114 (101); culmen, 91-101 (96); tarsus, 
78-87 (82); .«cvcn adult females, from Rota and Guam: wing, 265-285 (275); 
tail. 87-96 (91); culmen, 86-92 (89); tarsus, 72-79 (76). 

Weights. — The author (1948:42) lists the following weights: four adult 
males from Guam (gray phase) 590-667 (614) ; two adult males from Guam 
(white phase) 600 and 662; five adult females from Guam and Rota (gray 
phase) 477-553 (506). 

Specimens examined. — Total number, 80 (38 males, 40 females, 2 unsexed) as follows: 
Mariana Islands, USNM— Rota. 3 (Oct. 18, Nov. 2, 5)— Guam, 21 (May 11, June 6, 18, 
July 6, 8, 16, 24, 27, Aug. 6, 8, 27); AMNH— Saipan, 2 (July 22)— Guam, 9 (Feb. 11, Mar. 
6, 7, April 11, Aug. 15, Sept. 14, 16, Nov. 27, Dec. 20); Palau Islands, USNM — Peleliu, 3 
(Sept. 10, 16); AMNH — exact locality not given, 5 (Nov. 8, 21, 23); Caroline Islands, 
USNM— Ulithi, 1 (Aug. 15)— Kusaie, 1 (Feb. 8); AMNH— Truk, 3 (Feb. 18, May 20, Nov. 
5)— Tah, 2 (Oct. 18) — Ponape, 2 (Nov. 21, undated) — Kusaie, 26 (Jan. 25, 26, Feb., Mar. 
10-20, 20-30, April 1-10, 18); Marshall Islands, USNM— Bikini, 2 (March 29, April 2). 

Nesting. — The Reef Heron apparently nests on most of the islands in Mi- 
cronesia. The eggs are laid in a nest of grass and twigs on or near the ground. 
Hartert (1898:64) records a nest found in grass at Saipan on July 28, 1895. 
Yamashina (1932a :406) reports on one egg taken at Ponape on July 23, 1931. 
Marshall (1949:219, fig. 37) found a breeding bird in April at Tinian. Coultas 
(field notes) learned from the natives at Ponape that the Reef Heron builds 



86 University of Kansas Publs., Mus. Nat. Hist. 

a nest of small sticks near the ground in the mangrove thickets. Two or three 
eggs are laid, and nests can be found at various times of the year. Mayr and 
Amadon (1941:4) comment on the prolonged breeding season and report six 
sets of eggs from Polynesia taken in January, March, April, September, Octo- 
ber, and November. 

Food habits. — The author (1948:42) found fish and crabs in the stomachs 
of birds taken at Guam, Ulithi and Peleliu. 

Parasites. — Vchida (1918:484, 488, 490) found the following bird lice 
(Mallophaga) on the Reef Heron at Ponape: Nirmus orarius, C olpocephalum 
importunum, and Myrsidea teraokai. Bequaert (1939:81 and 1941:266) found 
the fly (Hippoboscidae), Omithoctona plicata, on the heron at Kusaie. Whar- 
ton (1946:175) and Wharton and Hardcastle (1946:306, 316) obtained chiggers 
(Acarina), Neoschongastia egretta and A'', carveri, from the Reef Heron at 
Guam and Ulithi. 

Remarks. — The species Demigretta sacra contains two subspecies, 
the widespread D. s. sacra and a larger form, D. s. albolineata 
(Gray), known from New Caledonia and the Loyalty Islands. The 
latter subspecies is surrounded by the former, a distribution which 
closely parallels that in each of the species Phalacrocorax melano- 
leucus and Gygis alba of Oceania. Recently Delacour (in Delacour 
and Mayr, 1945b: 105) has dropped the name Demigretta placing 
all of the forms of this genus in Egretta. He says, ''We cannot ac- 
cept the genus Demigretta, which is based on the more extended 
feathering of the tibia, the different length and texture of the 
feathers of the trains, the shortness of the tarsus and the presence 
of a dark gray color phase. The latter exists in the Madagascan 
and African subspecies of Egretta garzetta." 

The Reef Heron is a conspicuous member of the bird life of Micro- 
nesia, being recorded from most of the island groups. It prefers 
the placid and shallow waters of the lagoons and tidal beaches 
where it obtains the littoral animal life as food. The birds are 
seldom seen inland and usually freciuent the beaches and rocky 
coasts. In this respect there is little opportunity for competition 
with the migratory Plumed Egret, which prefers the grassy upland 
and marsh areas and inland ponds. The Reef Heron is a quiet, 
usually solitary, and retiring bird, being exceedingly difficult to 
approach, especially when found on the open tidal flats. 

The problem of plumages and color phases in the Reef Heron 
has been treated by Mayr and Amadon (1941:4-10). Specimens 
which they examined from Micronesia were found to be 54 percent 
gray, 40 percent white, and 6 percent mottled. Of the birds obtained 
by NAMRU2 field parties, fewer than 40 percent were white. Field 
counts showed a considerable variation in the ratio of grays to 



Baker: The Avifauna of Micronesia 87 

whites: Guam— 6 grays to 4 whites; Ulithi — 4 grays, 6 whites, 1 
mottled; Palau — equal number of grays and whites; Truk— 2 
whites, 1 gray, 1 mottled. For some unknown reason, the gray 
birds were more easily approached than the white birds. Gleise 
and Genelly (1945:221) saw one white Reef Heron at Eniwetok. 
Wallace (field notes) found white herons more numerous than gray 
ones at Kwajalein in 1944 and 1945. Borror (1947:417) observed 
gray birds at Agrigan. Stott (1947:524) saw one blue heron on 
December 24, at Saipan. The 150 birds seen by him at Lake Susupe 
in December probably were Plumed Egrets. 

In discussing the variation in the color phases of the Reef Heron 
throughout its range, Mayr (1924b: 237) suggests that the reduced 
variability of small populations may not be due to accidental gene 
loss, but instead to the population having descended from a single 
pair or from one fertilized female. The descendents would naturally 
possess only those characters provided for in the genetic make-up 
of the parents. Reef Herons on New Zealand and in the Marquesas 
Islands all are gray, while at other island groups different propor- 
tions of gray and white individuals occur; such phenomena may 
result because of the genetic constitution of the "founders." 

Nycticorax nycticorax nycticorax (Linnaeus) 
Black-crowned Night Heron 

Ardes Nycticorax Linnaeus, Syst. Nat., ed. 10, 1, 1758, p. 142. (Tj-pe locality, 
Southern Europe.) 

Nycticorax griseus Hartlaub and Finsch, Proc. Zool. Soc. London, 1872, pp. 89, 105 
(Uap); Griiffe, Joum. Mus. Godeffroy, 2, 1873, p. 123 (Yap); Wiglesworth, Abhandl. 
und Bar. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 69 (Uap). 

Nycticorax nycticorax Sharpe, Cat. Birds British Mus., 26, 1898, p. 146 (Yap). 

Nycticorax nycticorax nycticorax Kuroda, in Momiyama, Birds Micronesia, 1922, 
p. 36 (Mackenzie, Yap); Hand-list Japanese Birds, rev., 1932, p. 183 (Yap, Uluthi) ; 
Hand-list Japanese Birds, 3d ed., 1942, p. 204 (Yap. Uluthi); Mayr, Birds Southwest 
Pacific, 1945, p. 302 (Marianas, Yap). 

Geographic range. — Europe and Africa east to Japan and Malaysia. In 
Micronesia: Mariana Islands — Tinian; Palau Islands — Koror; Caroline Islands 
—Yap, Ulithi, Truk. 

Specimens examined. — Total number, 2 immature females, as follows : Palau Islands, USNM 
—Koror, 1 (Nov. 27); Caroline Islands, AMNH— Truk, 1 (June 18). 

Remarks. — The Black-crowned Night Heron is a winter visitor to 
western Micronesia. Marshall (1949:221) records six of these birds 
on Tinian on April 4, 1945, and one on Koror on November 27. 

Nycticorax caledonicus pelewensis Mathews 
Rufous Night Heron 

Nycticorax caledonicus pelewensis Mathews, Bull. British Ornith. Club, 46, 1926, p. 
60. (Type locality, Pelew Islands.) 

Nycticorax caledonicus Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 
117, 118 (Pelew); Sharpe, Cat. Birds British Mus., 26, 1898, p. 158 (Pelew); Hartert, 



88 Univeksity of Kansas Publs., Mus. Nat. Hist. 

Novit. Zool., 7, 1900, p. 10 (Ruk); Keichenow, Die Vogel, 1, 1913, p. 255 (Palauin- 
seln); Takatsukasa and Kuroda, Tori, 1, 1915, p. 50 (Pelew); Uchida, Annot. Zool. 
Japon., 9, 1918, p. 486 (Palau) ; Wetmore, in Towsend and Wetmore, Bull. Mus. 
Comp. Zool., 63, 1919, p. 172 (Uala, Truk Atoll); Kuroda, in Momiyama, Birds 
Micronesia, 1922, p. 37 (Pelew, Ruk). 

Nycticorax manillensis Hartlaub and Finsch, Proc. Zool. Soc. London, 1872, pp. 89, 
105 (Pelew); Finsch, Joum. Mus. Godeffroy, 8, 1875, pp. 5, 33 (Palau); idem, Proc. 
Zool. Soc. London, 1880, p. 577 (Ruk); Schmeltz and Krause, Ethnogr. Abth. Mus. 
Godeffroy, 1881, p. 353 (Ruk); Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, 
no. 6, 1890-1891 (1891), p. 69 (Pelew, Ruk). 

Nycticorax caledonicus pelewensis Mathews, Syst. Avium Australasianarum, 1, 1927, 
p. 200 (Pelew, Carolines); Peters, Proc. Boston Soc. Nat. Hist., 39, 1930, p. 271 
(Pelew, Carolines); Peters, Check-list Birds World, 1, 1931, p. 115 (Pelew); Hand-list 
Japanese Birds, rev., 1932, p. 183 (Palau, Truk); Hand-list Japanese Birds, 3d ed., 
1942, p. 204 (Babelthuap, Koror, Coracal, Truk); Amadon, Amer. Mus. Novit., no. 
1175, 1942, p. 6 (Palau, Ruk); Mayr, Birds Southwest Pacific, 1945, p. 285 (Palau, 
Truk); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 43 (Angaur, Peleliu, 
Garakayo, Truk). 

Geographic rangrg.— Micronesia: Palau Islands — Babelthuap, Koror, Coracel, 
Garakayo, Peleliu, Ngabad, Angaur; Caroline Islands — Truk. 

Characters. — Adult : Size medium ; head and nape dark ?laty-black ; occip- 
ital plumes white with dark tips and shafts; back dark reddish-brown, lighter 
on sides of neck, wings, wing coverts, rump, and tail; under parts whitish with 
light reddish-brown on sides of neck extending to throat and upper breast; 
tibia with some brownish feathers; underwing pinkish; feet yellowish-brown; 
bill black. 

Immature: Resembles adult, but upper parts mottled black with reddish- 
brown; underparts with lighter streaks of brown and whitish on breast; feet 
yellowish; bill black above, yellowish below. 

Adult resembles A^. c. manillensis Vigors, but is duller above. 

Measureinents. — Two adult males from Peleliu measure: wing, 293, 299; 
tail, 105, 107; culmen, 82, 89; tarsus, 79, 81; seven adult females from Peleliu: 
wing, 269-286 (280); tail. 101-106 (104); culmen, 76-84 (80); tarsus, 78-83 (80); 
one adult female from Truk: wing, 280; tail, 97; culmen, 83; tarsus, 79. 

Specimens examined.- — Total number, 27 (5 males, 18 females, 4 unsexed), as follows: 
Palau Islands, USNM— Peleliu, 9 (Aug. 31, Sept. 1, 5, 6, 8, Dec. 6); AMNH— exact 
locality not given, 16 (Nov. 7, 8, 13, 23, 25, Dec. 1, undated); Caroline Islands, USNM — 
Truk, 1 (Feb. 16); AMNH— Truk, 1 (May 25). 

Nesting. — The NAMRU2 party observed a nesting colony of these night 
herons at Peleliu on August 29, 1945. Approximately eight nests were ob- 
served in a grove of saplinglike trees at the edge of a mangrove swamp. 
These nests were 15 to 20 feet above the ground; mo.st of them contained one 
or two nestling birds. Two subadults and three nestlings in postnatal molt were 
obtained; no eggs were found. Marshall (1948:219) records breeding in Au- 
gust, September and December. 

Food habits. — Baker (1948:43) reports that .stomachs of night herons ob- 
tained by the NAMRU2 party at Peleliu contained a great variety of animal 
foods, including eels, fish, lizards (skinks), crabs, shrimp, and insects. The 
stomach of one adult contained 14 large grasshoppers and four fish, totaling 
about 15 cc. in volume. The nestlings had eels, skinks, and insects in their 
stomachs.. 

Parasites. — Uchida (1918:486) found the bird louse (Mallophaga), Lipeurus 
baculus, on the night heron at Palau. 



Baker: The Avifauna of Micronesia 89 

Remarks. — Amadon (1942:4-8) has made the most recent study 
of the species Nycticorax caledonicus and recognizes eight subspecies 
from Australia and New Calendonia north to the Caroline and 
Bonin islands. This is one of the few tropical and subtropical species 
wliich has extended its range to the Bonin islands. The discontin- 
uous distributions of this species prevents an accurate estimation of 
the route by whicli it reached the Bonins. The presence of the bird 
at Palau and at Truk makes it difficult to account for its absence at 
Yap and other intervening, and seemingly suitable, islands. Popula- 
tions at Palau and Truk appear to be similar and are placed in the 
same subspecies, but when adequate material is available from Truk, 
further study may reveal that the populations on the two islands 
(Truk and Palau) are recognizably different. 

At the southern Palau Islands, night herons were found by the 
NAMRU2 party in mangrove swamps, lagoons and on beaches. 
I found them to be inactive during the daytime; the birds were 
usually perched singly in trees or at the edge of the water. The 
birds appeared to have special roosting places and were observed 
sitting in the same place on several different occasions. McElroy 
of the NAMRU2 party reported seeing three night herons at Tmk 
in December, 1945. 

Gorsachius goisagi (Temminck) 

Japanese Bittern 

Nycticorax goisagi Temminck, PI. CoL, livr. 98, 1835, pi. 582. (Type locality, 
Japan.) 

Gorsakius goisagi Hand-list Japanese Birds, rev., 1932, p. 184 (Koror) ; Hand-list 
Japanese Birds, 3d ed., 1942, p. 204 (Koror); Mayr, Birds Southwest Pacific, 1945, p. 
302 (Palau). 

Geographic range. — Eastern China, Japan, Riu Kius, Formosa, and Philip- 
pine Islands. In Micronesia : Palau Islands — Koror. 

Remarks. — Gorsachius goisagi has been recorded from Koror in 
the Palau Islands. It may be classed as a rare migrant to western 
JMicronesia. 

Gorsachius melanolophus melanolophus (Raffles) 
Malay Bittern 

Ardca melanolopha Raffles, Trans. Linn. Soc. London, 13, 1822, p. 326. (Type 
locality. Western Sumatra.) 

Nycticorax goisagi Hartlaub and Finsch, Proc. Zool. London, 186S, pp. 8, IIS 
(Pelew); idem, Proc. Zool. Soc. London, 1872, p. 89 (Pelew); Wiglesworth, Abhandl. 
und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 68 (Pelew). 

Nycticorax melanolophus Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 5, 35 (Palau). 

Gorsachius melanolophus Sharpe, Cat. Birds Brit. Mus., 26, 1898, p. 166 (Pelew). 

Gorsahius melanolophus melanolophus Hand-list Japanese Birds, rev.. 1932, p. 184 
(Pelew); Hand-list Japanese Birds, 3d ed., 1942, p. 204 (Palau); Mayr, Birds South- 
west Pacific, 1945, p. 302 (Palau). 

Gorsachius melanolophus melanolophus Mathews, Syst. Avium Australasianarum, 
1. 1927, p. 200 (Pelew). 



90 University of Kansas Publs., Mus. Nat. Hist. 

Geographic range. — India, Ceylon, southern China, Formosa, Indochina, 
Malaysia. In Micronesia: Palau Islands — exact locality unknown. 

Remarks. — Captain Tetens obtained a specimen of this bittern at 
the Palau Islands which was reported on by Hartlaub and Finsch 
(1868a: 8, 1868b: 118). It is probably a rare straggler to western 
Micronesia. The specimen has not been seen by me; it may be of the 
subspecies G. m. kutteri (Cabanis), which is known from the Philip- 
pine Islands. 

Ixobrychus sinensis (Gmelin) 

Chinese Least Bittern 

Ardea Sinensis Gmelin, Syst. Nat., 1, pt. 2, 1789, p. 642. (Type locality, China.) 

Ardea lepida Lesson, Traite d'Ornith., 1831, p. 573 (Marianne); Hartlaub, Journ. 
f. Omith., 1854, p. 167 (Mariannen). 

Ardea sinensis Kittlitz, Obser. Zool., in Lutke, Voy. "Le Seniavine," 3, 1836, p. 305 
(Guahan); Gray, Hand-list Birds, 3, 1871, p. 31 (Marian); Hartlaub and Finsch, 
Proc. Zool. Soc. London, 1872, pp. 89, 105 (Uap) ; Griiffe, Journ. Mus. Godeffroy, 2, 
1873, p. 123 (Yap); Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 5, 33 (Palau, Yap); 
idem, Proc. Zool. Soc. London, 1880, p. 577 (Ruk) ; Schmeltz and Krause, Ethnogr. 
Abth. Mus. Godeffroy, 1881, p. 353 (Ruk). 

Ardea (Ardetta) sinensis Gray, Cat. Birds Trop. Is. Pacific 0c«an, 1859, p. 49 
(Ladrone or Marian Islands). 

Ardetta Sinensis Salvador!, Ornith. Papuasia, 3, 1882, p. 364 (Pelew, Carolines, 
Mariannis); Oustalet, Le Nat., 1889, p. 261 (Mariannes); Wiglesworth, Abhandl. und 
Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 68 (Marianne, Uap, Ruk, Pelew); 
Oustalet, Nouv. Arch, Mus. Hist. Nat. Paris, (3), 8, 1896, pp. 38, 39 (Guam, Saypan, 
Ponapi, Ruk, Palaos) ; Hartert, Novit. Zool., 5, 1898, p. 65 (Guam); Sharpe, Cat. 
Birds British Mus., 26, 1898, p. 227 (Marianne, Carolines, Pelew); Hartert, Novit. 
Zool., 7, 1900, p. 11 (Ruk); Safford, Contr. U. S. Nat. Herb., 9, 1905, p. 79 (Guam); 
Prowazek, Die deutschen Marianan, 1913, p. 100 (Saipan) ; Cox, Island of Guam, 1917, 
p. 21 (Guam). 

Ardetta bryani Seale, Occ. Papers Bemice P. Bishop Mus., 1, 1901, p. 27 (Type 
locality, Guam) ; Safford, Osprey, 1902, p. 66 (Guam) ; idem. The Plant World, p. 
266 (Guam). 

Ardetta sinensis sinensis Takatsukasa and Kuroda, Tori, 1, 1915, p. 50 (Ruk, 
Pelew). 

Ixobrychus sinensis bryani Wetmore, in Townsend and Wetmore, Bull. Mus. Comp. 
Zool., 63, 1919, pp. 173, 175 (Guam) ; Kuroda, in Momoyama, Birds Micronesia, 1922, 
p. 37 (Guam, ?Yap, ?Mackenzie, ?Pelew) ; idem. Avifauna Riu Kiu. 1925, p. 134 
(Guam, ?Yap, ?Pelew); Mathews, Syst. Avium Australasianarum, 1, 1927, p. 202 
(Guam, ?Pelew); Peters, Check-list Birds World, 1, 1931, p. 121 (Guam); Hand-list 
Japanese Birds, rev., 1932, p. 184 (Saipan, Tinian, Rota, Guam) ; Oberholser, Bull. 
U. S. Nat. Mus., 159, 1932, p. 18 (Guam); Bryan, Guam Rec, vol. 13, no. 2, 1936, 
p. 15 (Guam); Hand-list Japanese Birds, 3d ed., 1942, p. 205 (Saipan, Tinian, Rota, 
Guam) ; Amadon, Bull. Bemice P. Bishop Mus., 186, 1945, p. 25 (Guam) ; Stott, Auk, 
64, 1947, p. 525 (Saipan); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 44 
(Rota, Guam). 

Ixobrychus sinensis moorei Wetmore, in Townsend and Wetmore, Bull. Mus. Comp. 
Zool., 63, 1919, p. 173 (Type locality, Uala, Truk group); Kuroda, in Momiyama, 
Birds Micronesia, 1922, p. 38 (Ruk); Kuroda, Avifauna Riu Kiu, 1925, p. 134 (Ruk); 
Mathews, Syst. Avium Australasianarum, 1, 1927, p. 202 (Middle Carolines); Peters, 
Check-list Birds World, 1, 1931, p. 121 (Truk); Hand-list Japanese Birds, rev., 1932, 
p. 184 (Palaus, Yap, Truk); Oberholser, Bull. U. S. Nat. Mus., 159, 1932, p. 17 (Caro- 
lines, ?Pelews) ; Hand-list Japanese Birds, 3d ed., 1942, p. 205 (Babelthuap, Koror, 
Yap, Truk); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 44 (Truk, 
Peleliu). 



Baker: The Avifauna of Micronesia 



91 



Ixobrychm sinensis Hartert, Vogel pal. Fauna, 10, 1920, p. 1260 (Truk, Palau, 
Guam); Mayr, Birds Southwest Pacific, 1945, p. 285 (Marianas, Palau, Yap, Truk); 
Watson, The Raven, 17, 1946, p. 41 (Guam); Downs, Trans. Kansas Acad. Sci., 49, 
1946, p. 91 (Tinian); Wharton, Ecol. Monogr., 16, 1946, p. 174 (Guam); Delacour 
and Mayr, Birds Philippines, 1946, p. 29 (Guam); Strophlet, Auk, 63, 1946, p. 536, 
(Guam); Baker, Condor, 49, 1947, p. 125 (Guam). 

Ixohrychus sinensis sinensis Hachisuka, Birds Philippines, 1, 1932, p. 365 (Guam, 
Truk); Robinson and Chasen, Birds Malay Peninsula, 3, 1936, p. 195 (Marianne). 

Ixobrychus sinensis palewensis Momiyama, Bull. Biogeogr. Soc. Japan, 2, 1932, p. 
333 (Type locality, Pelew) ; Mathews, Ibis, 1933, p. 88 (Pelew). 

Ixobrychus sinensis yapensis Momiyama, Bull. Biogeogr. Soc. Japan, 2, 1932, p. 333 
(Type locality, Yap); Mathews, Ibis, 1933, p. 89 (Yap). 

Geographic range. — Northeastern China and Japan south to Micronesia, 
Malaysia, Burma, India and Ceylon. Winter visitor to Papuan region. 
In Micronesia: Mariana Islands — Saipan, Tinian, Rota, Guam; Palau Islands 
— Babelthuap, Koror, Peleliu; Caroline Islands — Yap, Truk. 

Characters. — Adult male: A small bittern with crown and short occipital 
crest slaty-black; mantle light buffy-brown ; back and rump gray; tail black; 
wing-coverts brownish-buff; primaries and secondaries slaty-black; underparts 
yellowish buff; chin and throat whitish; sides of head and neck and a line of 
feathers across chest blackish edged with buff; bill yellowish green; feet 
greenish yellow. 

Adult female: Resembles adult male, but with upper parts mottled brown 
and golden chestnut; underparts deep buff streaked with pale brown on neck. 

Immature: Resembles adult, with upper parts heavily streaked with 
blackish-brown, and underparts streaked with chestnut and dark brown. 

Measurements. — Measurements of specimens from Micronesia are given in 
table 16. 



Table 16. Measurements of Ixohrychus sinensis From Micronesla 



Location 


Sex 


No. 


Wing 


Tail 


Full 
culmen 


Tarsus 


Yap, Truk 


males 

males 

females 


6 
11 
10 


132 
130-131 

134 
127-138 

130 
127-134 


43 

41-47 

47 
45-50 

46 
44-49 


56 
54-59 

57 
55-60 

57 
55-59 


44 


Guam 


42-47 
46 


Guam 


45-47 
45 




43-47 



Weights. — The author (1948: 44) records the weights of eight adult males 

from Guam as 82-103 (92) and eight adult females from Guam as 84-109 (95). 

Specimens examined. — Total number, 69 (34 males, 27 females, 8 unsexed), as follows: 
Mariana Islands, USNM— Saipan, 1 (Sept. 30)— Tinian, 1 (Oct. 13)— Guam, 29 (May 16, 
June 4, 6, 7, 8, 14, 18, 19, July 10, 16, 18, 24, 27, Aug. 4); AMNH— Saipan, 1 (Aug. 6) 
—Tinian, 3 (Sept. 13)— Guam, 14 (Feb. 1, Mar. 13, 29, July 11, 13, 25, Aug. 1, 7, 13. 
Sept. 4, 10, Dec. 8); Palau Islands, AMNH — exact locality not given, 6 (Nov. 19, 21, 23, 
25, Dec. 1, 18); Caroline Islands, USNM— Truk, 1 (Feb. 16); AMNH— Yap, 1 (not dated) 
—Truk, 12 (Feb. 9, Mar. 5, 17, May 7, June 13, 14, 15, Oct. 3, Nov. 1, 5, Dec. 20). 



92 University of Kansas Publs., Mus. Nat. Hist. 

Nesting.— The author (1948:44) records a nest found by the NAMRU2 
party near Achang Bay on Guam on June 6, 1945. It was found in a cane 
thicket at the edge of a fallow rice paddy, approximately four feet from the 
ground and was constructed of about three quarts of reeds and cane. Two 
eggs found in the nest are oval, white with a greenish cast and measure 33 by 
24 and 34 by 24. On February 1, 1945, the writer found two recently occupied 
nests of the Chinese Least Bittern at Oca Point, Guam. These nests were in 
dense inkberry brush approximately five feet above the ground. The area was 
not marshy, the nearest water being at the beach some 300 yards away. Nearby 
one of the nests was found a young bittern, which apparently had only re- 
cently left the nest. The pin feathers were growing. A parent bird remained 
in the vicinity with the young bird until it left the area after March 9. 

Food habits. — The Chinese Least Bittern feeds on animal foods obtained 
along waterways, marshes and beaches as well as in forests and fields. The 
NAMRU2 party observed several types of insects in the stomachs of birds 
taken at Guam. Scale (1901:27) found black crickets in stomachs of bitterns 
taken at Guam. Coultas (field notes) learned from the natives of the Palau 
Islands that the bittern feeds on land mollusks. 

Parasites. — Wharton (1946:174) obtained the chigger (Acarina), Tromhicula 
acuscutellari-s, from the Chinese Least Bittern at Guam. 

Remarks. — The Chinese Least Bittern lias been regarded by many 
workers as consisting of several geographic races; as many as eight 
have been recognized. Other workers have concluded that /. sinensis 
is made up of highly variable populations and that it lacks well- 
fined geographic variation. Hartert (1920:1260), Hachisuka (1932: 
365), and Mayr (1945a:285) have reached the latter conclusion. As 
yet this problem has not been satisfactorily solved ; a thorough study 
is needed, but may not be possible until additional material, espe- 
cially from the continental areas, can be obtained. In coloration 
there appears to be little difference between birds from the various 
localities in Micronesia. These birds may average slightly paler than 
populations from the continental areas, but on this basis I doubt 
that a person could recognize the Micronesian birds in a group of 
skins from many other localities. Birds in fresh plumage may show 
geographic differences better than slightly worn specimens. Meas- 
urements made by the author offer no clear-cut differences either. 

I. sinensis was first recorded in Micronesia by Quoy and Gaimard 
(1824:536), whose ship, the "Uranie," stopped at Guam. They 
called the bird "Petit Heron aux ailes noires." Most of the ornitho- 
logical collectors in the years following Quoy and Gaimard obtained 
this bittern in Micronesia. At Guam, its abundance and the ease 
with which it may be approached and shot is attested by the large 
series obtained by collectors: Seale (1901:27) took eight birds; 



Baker: The Avifauna of Micronesia 93 

Marche (Oustalet, 1896:36) took eighteen skins; the NAMRU2 
party took twenty-nine skins. 

The Chinese Least Bittern is found in habitats associated with 
both salt water and fresh water, as well as in upland habitat in 
Micronesia. The bird appears to be well adapted to areas of open 
forest and coconut groves. Coultas (field notes) found the birds in 
taro patches in the Palaus. Although a considerable amount of field 
observing was done in the southern Palaus, the NAMRU2 party saw 
only one bird (September 13, 1945, at Peleliu). Perhaps the birds 
prefer Babelthuap and other large islands farther north in the chain. 
McElroy found bitterns in taro patches at Truk in December, 1945. 
The NAMRU2 party did not find any birds at Rota in October and 
November, 1945. Downs (1946:91) found the birds in upland sugar 
cane and beach habitats on Tinian. 

Regarding the bittern in the Palaus, Coultas (field notes) writes, 
"Always found alone, never a pair. A bird that is not easily fright- 
ened. In the heat of the day, one finds it standing in the shade of a 
taro leaf quietly viewing the intruder and very reluctant about mov- 
ing. I have tossed pieces of earth and sticks at the bird to encourage 
him to fly so that I would not blow him to pieces when I shot, but 
my efforts at dislodgement have been rewarded by harsh scolding 
squawks. It became necessary for me to move into proper gun range. 
I have also found them perched in low trees at the edge of taro 
swamps. In flight they are atrociously awkward. They can't keep a 
course and their legs dangle every-which way. Their jerky, slow 
flight usually ends abruptly when the bird becomes entangled in 
weeds or the branches of trees. Extracting himself from his predica- 
ment he is soon in another and invariably resorts to blasphemy." 

Ixobrychus eurhythmus (Swinhoe) 
Schrenck's Least Bittern 

Ardetta eurhythma Swinhoe, Ibis, 1873, p. 74, pi. 2. (Type locality, Amoy 
Shanghai.) 

Ixobrychus eurythmus Mayr, Birds Southwest Pacific, 1945, p. 302 (Patau). 

Geographic range. — Southeastern Siberia and Japan south to India and 
Malaysia. In Micronesia: Palau Islands — exact locality unknown. 

Specimens examined. — Total number, 3 (2 males, 1 female), from Palau Islands, AMNH — 
exact locality not given (Nov. 19, 21, Dec. 3). 

Remarks. — Coultas obtained three immature specimens at Palu 
in November and December, 1931. 



94 University of Kansas Publs., Mus. Nat, Hist. 

Dupetor flavicollis flavicollis (Latham) 
Black Bittern 

Ardea flavicollis Latham, Ind. Ornith., 2, 1790, p. 701. (Type locality, India.) 
Dupetor flavicollis Seale, Occ. Papers Bemice P. Bishop Mus., 1, 1901, p. 26 
(Guam); Bryan, Guam Rec., vol. 13, no. 2, 1936, p. 15 (Guam). 

Dupetor f. flavicollis Mayr, Birds Southwest Pacific, 1945, p. 302 (Guam). 

Geographic range. — Central China south to Malaysia and India. In 
Micronesia: Mariana Islands — Guam. 

Remarks. — Seale (1901:26) records a female shot at the Agana 
River on Guam on June 11, 1900. The skin probably is in the 
Bernice P. Bishop Museum in Honolulu. 

Anas oustaleti Salvadori 
Marianas Mallard 

Anas oustaleti Salvadori, Bull. British Ornith, Club, 4, 1894, p. 1. (Type locality, 
Mariannis Islands.) 

Anas oustaleti Salvadori, Cat. Birds British Mus., 27, 1895, p. 189 (Guaham) ; 
Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 8, 1896, p. 49 (Guam); Hartert, 
Novit. Zool., 5, 1898, p. 66 (Guam, Saipan); Wheeler, Report Island of Guam, 1900, 
p. 13 (Guam); Seale, Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 25 (Guam, 
Saipan); Matschie, Journ. f. Ornith., 1901, pp. 110, 113 (Guam, Saipan); Safford, 
Osprey, 1902, p. 66 (Mariannas); idem, Amer. Anthro., 4, 1902, p. 711 (Guam); 
idem. The Plant World, 7, 1904, p. 267 (Guam) ; Dubois, Syn. Avium, 2, 1904, p. 990 
(Mariannes); Safford, Contr. U. S. Nat. Herb., 9, 1905, pp. 80, 126 (Guam); 
Prowazek, Die deutschen Marianen, 1913, pp. 47, 100 (Marianen); Cox, Island of 
Guam, 1917, p. 22 (Guam); Phillips, Nat. Hist. Ducks, 2, 1923, p. 53 (Guam, 
Saipan); Mathewsi, Syst. Avium Australasianarum, 1, 1927, p. 214 (Guam, Saipan); 
Berlioz, Bull. Mus. Hist. Nat. Paris, 2d ser., 1, 1929, p. 67 (Guam); Peters, Check- 
list Birds World, 1, 1931, p. 159 (Guam, Tinian, Saipan); Hand-list Japanese Birds, 
rev., 1932, p. 184 (Guam, Tinian, Saipan); Bryan, Guam Rec, vol. 13, no. 2, 1936, 
p. 15 (Guam); Kuroda, Tori, 11, 1941-42, pp. 99, 443 (Marianas); Hand-list Jap- 
anese Birds, 3d ed., 1942, p. 205 (Guam, Tinian, Saipan); Amadon, Amer. Mus. Novit., 
no. 1237, 1943, p. 1 (Marianne); Mayr, Birds Southwest Pacific, 1945, p. 285 
(Marianas); idem, Audubon Mag., 47, 1945, p. 282 (Marianas); Baker, Trans, 11th 
N. Amex. Wildlife Conf., 1946, p. 208 (Guam); Stott, Auk. 64, 1947, p. 525 
(Saipan); Baker, Smithson, Misc. Coll., vol. 107, no. 15, 1948, p. 45 (Saipan, 
Tinian); Momiyama, Pacific Science, 2, 1948, p. 121 (Saipan, Tinian, Guam). 

Polionetta oustaleti Kuroda, in Momiyama, Birds Micronesia, 1922, p. 39 (Guam, 
Saipan). 

Anas superciliosa oustaleti Hartert, Novit. Zool., 36, 1930, p. 112 (Guam, Saipan). 

Anas platyrhynchos oustaleti Delacour and Mayr, Wilson Bull., 57, 1945, pp. 21, 
39 (Marianas). 

Geographic range. — Micronesia: Mariana Islands — Guam, Tinian, Saipan. 

Characters. — From study of a large series of specimens of Anas oustaleti, 
Yamashina (1948) described two types of plumages: one type resembles that 
of A. platyrhynchos and another type resembles that of A. poecilorhyncha. 
He based his conclusions on both a study of prepared skins and observations 
of the molt of living specimens as reported by Kuroda (1941-1942). The fol- 
lowing descriptions are quoted from Yamashina (1948:122). 

Adult male in nuptial plumage of A. platyrhynchos type: "Whole head 
is dark green, except at the sides where buff feathers are plentifully inter- 
mingled, a dark brown streak through the eye, and faint white ring on the 
lower neck. Feathers on scapulars and sides of body are as those of Anas 



Baker: The Avifauna of Micronesia 95 

poecilorhyjicha. Sides of body are vermiculated but some brown feathers are 
found even in the full nuptial plumage. Upper breast is dark reddish chestnut 
with dusky spots. Upper and under tail-coverts are as in Anas platyrhynchos. 
Speculum is as that of Ar^as platyrhynchos, but the tips of the greater coverts 
are buff instead of white. Central tail feathers are more or less curled upward. 
Base of bill is black, tip is olive color. Iris is dark brown. Feet, reddish- 
orange, webs darker." Eclipse plumage of adult male resembles that of A. 
platyrhynchos. 

Adult male in nuptial plumage of A. poecilorhyncha type: "Resembles Anas 
poecilorhyncha pelewensis from the Palau Islands and Truk Island, but sides 
of head are browner, superciliary stripes and ground color of cheeks are more 
buffy. Feathers on upper breast and sides of body are more broadly edged 
with brown. Speculum is usually violet-purple as in the platyrhynchos type, 
but in two specimens from Saipan and Tinian, respectively, it is dark green 
as in Anas poecilorhyncha pelewensis. Tips of the secondaries are usually 
white, but sometimes very faint as in Anas poecilorhyncha pelewensis, and in 
one specimen from Saipan they are buffy. Bill is olive color with a black 
spot in the center of the upper mandible. Iris, dark brown. Feet, dark 
orange, darker in joints and webs." Eclipse plumage of adult male resembles 
the nuptial plumage. 

Measurements. — Measurements of nine ducks from Guam and Saipan are: 
wing, 238-266 (252); tail, 75-84 (81); exposed culmen, 49^53 (51); tarsus, 41-43 
(42). 

Specimens examined. — Total number, 9 (5 males, 2 females, 2 unsexed), as follows: Mari- 
ana Islands, USNM — Saipan, 2 (Oct. 2, 3) — Guam, 1 (June 6); AMNH— Saipan, 2 (Aug. 7, 
ID— Guam, 4 (Jan. 10, April 6, Dec. 11, 16). 

Nesting. — At Guam, Seale (1901:25) found nests of the Marianas Mallard 
"among the reedy swamps and streams of the island." He obtained two downy 
young in June. Kuroda (1941-1942) reports nesting at Lake Challankanoa, 
Saipan, in July. He writes that nests contained 7 to 12 eggs. Ducklings and 
incubated eggs were obtained in June and July, but he is of the opinion that 
the breeding season may be longer. He notes that adults exhibit both nuptial 
plumage and eclipse plumage at the same time, suggesting that breeding may 
occur at various times in the year. A nest with seven eggs taken on July 4, 
1941, at Hagoi Lake, Tinian, is described by Kuroda as having been found 
among rushes and constructed of dead leaves, stems, and roots and lined with 
down. He describes the eggs as being grayish-white with a pale greenish tinge, 
and measuring 61.6 by 38.9. Marshall (1949:202) saw a family of ducklings 
in April. 

Remarks. — The Marianas Mallard is rare; probably it never has 
been very abundant in the small chain of islands to which it is re- 
stricted, because fresh water marshes and swamps are not extensive. 
The bird was first recorded by Bonaparte as Anas boschas a. Frey- 
cineti in 1865. This name was a nomen nudum and later the same 
specimen in the Paris Museum was named by Salvadori (1894) as 
Anas oustaleti. In 1888, Marche obtained six specimens at Guam; 
these were reported on by Oustalet (1896:49). Later collecting 



96 University of Kansas Publs., Mus. Nat. Hist. 

showed that the duck inhabited also the islands of Saipan and 
Tinian. There have been no records of this duck in the more north- 
ern islands of the Marianas. According to Yamashina (1948:121) 
in the period from 1931 to 1940, the Japanese obtained 38 specimens 
of the Marianas Mallard at Tinian and Saipan. In 1940, four birds 
from Tinian were shipped alive to Japan and kept in an aviary by 
Kuroda. At Tinian in 1940, one of the collectors observed two 
flocks of A. oustaleti, each containing 50 or 60 individuals. The 
Japanese took specimens at a lagoon area and at fresh water lakes. 
Yamashina describes one of the localities, Lake Hagoi on Tinian, as 
"a small body of fresh water surrounded by about 40 acres of 
marsh." During the war, sei-vicemen reported the presence of the 
Marianas Mallard at both Saipan and Tinian. Moran (1946:261) 
counted twelve ducks at Saipan. Stott (1947:525) saw seven birds 
at Lake Susupe on Saipan in December, 1945. He writes that the 
birds were gentle and easily approached and that they preferred 
winding channels in reed beds to open water. Marshall obtained 
two ducks at Lake Susupe in early October, 1945. These specimens 
are in the United States National Museum. He (1949:202) found 
ducks at both Saipan and Tinian; twelve was the greatest number 
seen at any one time. Gleise (1945:220) estimated that there were 
twelve birds on Tinian in 1945, remarking that their habitat was 
swamp area. 

At Guam and Rota, the NAMRU2 party failed to obtain any 
specimens but received reports of the presence of ducks on both 
islands. At Guam, reports were obtained of ducks of unknown spe- 
cies at a fallow rice paddy in August, 1944, and in a marsh near Agat 
on June 13, 1945. The presence of Japanese soldiers in the interior 
of Guam made it inadvisable to investigate marshes and swamps of 
the interior and the upper courses of streams. H. G. Hornbostel, as 
quoted by Phillips (1923:54), reported that ducks were found at 
Guam only in the Tolofofo River Valley. The NAMRU2 field par- 
ties investigated the lower reaches of this valley and found no evi- 
dence of the ducks. The upper part of this valley was used as an 
artillery range in 1945. Probably the firing of field guns was a dis- 
turbing influence to any birds that might have been there. If the 
ducks were on Guam at that time, they must have been secretive and 
restricted in their movements. At Rota, two ducks which might have 
been A. oustaleti were seen by the NAMRU2 party on October 20, 
1945, in a cultivated field. 

These recent reports indicate that the Marianas Mallard is secure 
for the present on the islands of Saipan and Tinian, but thoughtful 



Baker: The Avifauna of Micronesia 97 

conservation practices need to be placed in operation to insure its 
survival in the future. 

Evolutionary history of Anas oustaleti. — In the past, most of the 
studies have pointed to a northern ancestry for A. oustaleti. Bryan 
(1941:187) has noted a relationship between A. oustaleti and the 
Laysan Duck [A. laysanensis Rothschild) and the Hawaiian Duck 
{A. wyvilliana Sclater). Amadon (1943:1) suggests that these three 
species of ducks are rather recent derivatives of the Common Mal- 
lard {A. platyrhynchos) and postulates the evolution of A. ivyvil- 
liana from migrants from North America. He further states that A. 
laysanensis and A. oustaleti may have been derived from A. wyvil- 
liana or may represent independent colonizations. Delacour and 
Mayr (1945:21) go a step further and make these forms subspecies 
of A. platyrhynchos, saying that they are "dull-colored editions" of 
the Common Mallard, that because of isolation they have become 
reduced in size and have lost many of the characteristics of their 
ancestors. Recently, however, Yamashina (1948) has concluded that 
the Marianas Mallard has evolved as the result of hybridization 
between the two species, A. platyrhynchos and ^4. poecilorhyncha. 
His conclusions are based on a study of a large number of specimens, 
both museum skins and captive birds, in which he has been able to 
detect plumages of the A. platyrhynchos type and of the A. poecilo- 
rhyncha type (see Characters) . He has noted specimens which have 
ninety percent of the characteristics of A. platyrhynchos and ten 
percent of the A. poecilorhyncha type. These percentages are re- 
versed in specimens favoring the A. poecilorhyncha type. In his 
series of skins he finds the A. poeciolrhyncha type of plumage most 
frequently, in forty-four specimens out of fifty examined, while only 
six specimens have the A. platyrhynchos type of plumage. Yama- 
shina cites also as evidence favoring his conclusion that hybridiza- 
tion has taken place the results obtained from the crossing of captive 
A. platyrhynchos and A. poecilorhyncha. It is his assumption that 
there has been a resident form of A. poecilorhyncha in the Marianas, 
apparently resembling closely that which occurs in the Palaus and at 
Truk {A. p. pelewensis) , and that stragglers of A. platyrhynchos 
from the north occasionally reach the Marianas where hybridization 
between the two species occurs. Yamashina remarks (1948:123): 
"The opportunity for hybridization should occur more rarely in the 
south, and thus more frequent back-crossing of the hybrid with the 
indigenous Anas poecilorhyncha on Tinian and Guam explains the 
superabundance there of the poecilorhyncha type. As the hybridiza- 
7—8131 



98 University of Kansas Publs., Mus. Nat. Hist. 

tion should have taken place more frequently to the north in Saipan, 
the ratio of the occurrence of the platyrhynchos type is logically 
higher there." The Common Mallard (^4. p. platyrhynchos) has not 
been recorded in Micronesia, but according to Yamashina (1948: 
123) "winters frequently just north of the Marianas in the Bonin 
and Volcano Islands." 

This remarkable explanation for the development of the Marianas 
Mallard is not questioned by this author, who feels that hybridiza- 
tion may be found to be the cause for other unusual forms of life in 
island habitats whose ancestry has not been explained. As Yama- 
shina comments, the special environments of islands together with 
small and restricted populations of animals are factors which could 
favor such development. 

Anas poecilorhyncha pelewensis Hartlaub and Finsch 
Australian Gray Duck 

Anas superciliosa var. pelewensis Hartlaub and Finsch, Proc. Zool. Soc. London, 
1872, p. 108. (Type locality, Pelew Islands.) 

Anas superciliosa Hartlaub and Finsch, Proc. Zool. London, 1868, pp. 8, 118 
(Pelew); Sclater, Proc. Zool. Soc. London, 1869, p. 659 (Pelew); Gray, Hand-list 
Birds, 3, 1871, p. 82 (Pelew); Salvadori, Ornith. Papua&ia, 3, 1882, p. 395 (Pelew); 
Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 70 
(Pelew); Salvadori, Cat. Birds British Mus., 27, 1895, p. 206 (Pelew); Oustalet, 
Nouv. Arch. Mus. Hist. Nat. Paris, (3), 8, 1896, p. 50 (Palaos). 

Anas superciliosa pelewensis Dubois, Syn. Avium, 2, 1904, p. 990 (Pelew); 
Mathews, Birds Australia, 4, 1915, p. 90 (Pelew); Phillips, Nat. Hist. Ducks, 2, 
1923, p. 113 (Pelew); Mathews, Syst. Avium Australasianarum, 1, 1927, p. 215 
(Pelew); Hartert, Novit. Zool., 36, 1930, p. 112 (Pelew); Peters, Check-list Birds 
World, 1, 1931, p. 160 (Pelew); Hand-list Japanese Birds, rev., 1932, p. 184 (Palaus, 
Truk) ; Hand-list Japanese Birds, 3d ed., 1942, p. 205 (Babelthuap, Peliliu) ; Amadon, 
Amer. Mus. Novit., no. 1237, 1943, p. 3 (Palau) ; Mayr, Birds Southwest Pacific, 
1945, p. 286 (Palaus, Truk); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 
45 (Peleliu, Truk). 

Anas pelewensis Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 5, 40 (Palau); 
Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, p. 407 (Palau); Bolau, 
Mitteil. Naturhist. Mus. Hamburg, 1898, p. 71 (Palau). 

Polionetta superciliosa pelewensis Kuroda, in Momiyama, Birds Micronesia, 1922, 
p. 38 (Pelew). 

Anas superciliosa rukensis Kuroda, "Gan to Kamo" (Geese and Ducks), 1939, page 
not numbered, description between pis. 52 and 53 (Type locality, Ruk) ; Hand-list 
Japanese Birds, 3d ed., 1942, p. 206 (Truk). 

Anas poecilorhyncha superciliosa Delacour and Mayr, Wilson Bull., 57, 1945, pp. 21, 
39 (no locality given); Yamashina, Pacific Science, 2, 1948, p. 122 (Palau, Truk). 

Geographic range. — Islands of Micronesia, Polynesia, and Melanesia. In 
Micronesia: Palau Islands — Babelthuap, Peleliu; Caroline Islands — Truk. 

Characters. — Adult: A medium-sized duck with upper parts dark brown, 
feathers edged with buff; top of head blackish merging into gray on hind neck 
with narrow buff line below; eye-stripe broad and blackish; lower parts uni- 
formly dark brown to gray brown, feathers edged with buff; face, chin and 
throat light buff with some dark streakings; under wing white; speculum 
green; bill plumbeous with nail black; legs yellow-brown to yellowish, webs 



Baker: The Avifauna of Micronesia 99 

dusky. A. p. pelewensis resembles A. p. rogersi Mathews, but is smaller with 
a wing length averaging as much as 20 mm. shorter. 

Measurements. — As given by Amadon (1943:4) seven unsexed skins from 
the Palaus, studied by Finsch (1875:40), have wing lengths of 207, 212, 212, 
214, 223, 235, 230. For an adult male taken by Coultas at Palau, the exposed 
culmen measures 45 and the tarsus 37. 

Specimens examined. — Total number, 3 males from Palau Islands, AMNH — exact locality 
not given (Oct. 26, Nov. 25). 

Remarks. — A. p. yelewensis is apparently rare in the Palau Is- 
lands. Coultas, who visited the Palaus in October to December, 
1931, writes (field notes) that he received reports that the birds 
were present and nested in numbers on fresh water lakes. He took 
specimens in taro patches and comments that the ducks probably 
feed at night and have retiring habits during the day. At Peleliu in 
1945, the NAMRU2 party received several reports of ducks but 
failed to find the birds. At Truk, in December, 1945, McElroy of 
the NAMRU2 party found ducks to be fairly numerous in rice 
paddies, marshes, and swamps. He observed that the birds roosted 
at Moen Island at night but that they apparently flew to outlying 
islands to spend the day. Richards observed ducks on Moen Island 
on August 28 and 29, 1947, and again in the period from January 
19 to FebiTiary 10, 1948. He saw several flocks of ducks including 
one containing "about a dozen ducks" at ponds along a roadway 
and at an airstrip. Kuroda named the population at Truk as distinct 
in 1939. I have not been able to examine his description and no 
specimens are available for study, but if the birds at Truk represent 
an independent colonization (different from that of the birds at 
Palau) they might exhibit recognizable variation. Amadon (1943: 
5) has already pointed out that the shortness of the wing of speci- 
mens in the Palaus may merit subspecific status for the population. 
Delacour and Mayr (1945:21) propose that the Palau Gray Duck 
is a subspecies of A. poecilorhyncha; this treatment is followed in 
the present work. 

Evolutionary history. — A. p. pelewensis, as Amadon (1943:1) 
has stated, represents a population of mallards which became sepa- 
rated from the ancestral stock in the Australian or Malayan area 
and when once differentiated, invaded New Zealand and other parts 
of Polynesia, Melanesia, and southwestern Micronesia. Amadon 
points out that its range in the Pacific islands is more or less com- 
plimentary to that of A. oustaleti in the Marianas and the Philippine 
Mallard (.4. poecilorhyncha luzonica Fraser)^ as well as to the 
Hawaiian forms (^4. wyvilUana Sclater and .4. laysanensis Roths- 



100 University of Kansas Publs., Mus. Nat. Hist. 

child). The range of A. p. pelewensis gives one the impression that 
its present distribution may be only a stage in a gradual spreading 
of the species, for it certainly has not yet occupied all habitats suit- 
able for it in southern Micronesia nor elsewhere in Oceania. As in 
the case of A. oustaleti, A. p. pelewensis appears to prefer areas of 
fresh, and possibly brackish, water on the larger islands. 

A. p. luzonica is a near relative of A. p. pelewensis but has rufous- 
brown instead of buffy-brown coloring on the chin, throat, sides of 
head, and superciliary region. The underparts of the Philippine 
Mallard are much less mottled. The specula are similar. Both of 
these forms were probably derived from a mallard of the A. p. 
poecilorhyncha type. 

Anas querquedula Linnaeus 
Garganey Teal 

Anas Querquedula Linnaeus, Syst. Nat., ed. 10, 1, 1758, p. 12G. (Type locality, 
Europe, restricted to Sweden.) 

Anas querquedula Marshall, Condor, 51, 1949, p. 221 (Tinian). 

Geographic range. — Breeds in Europe and Asia. Winters from northern 
Africa to New Guinea. In Micronesia: Mariana Islands — Tinian. 

Remarks. — Marshall (1949:221) obtained one of a pair of these 
ducks which he observed "daily in April on Lake Hagoi" at Tinian. 

Anas crecca crecca Linnaeus 
European Teal 

Anas Crecca Linnaeus, Syst. Nat., ed. 10, 1, 1758, p. 126. (Type locality, Europe, 
restricted to Sweden.) 

Querquedula crecca crecca Hand-list Japanese Birds, rev., 1932, p. 185 (Pagan). 
Anas crecca crecca Hand-list Japanese Birds, 3d ed., 1942, p. 206 (Pagan). 
Anas crecca Mayr, Birds Southwest Pacific, 1945, p. 302 (Micronesia). 

Geographic range. — Breeds in Iceland, northern Europe, Asia, and Aleutians. 
Winters south to northern Africa, Asia and Philippines. In Micronesia: Mari- 
ana Islands — Pagan. 

Remarks. — The European Teal has been recorded by the Japanese 
at Pagan in the northern Marianas. It appears to be an uncommon 
winter visitor to Micronesia. 

Anas crecca carolinensis Gmelin 
Green-winged Teal 

Anas carolinensis Gmelin, Syst. Nat., 1, pt. 2, 1789, p. 533. (Type locality, Carolina 
to Hudson Bay.) 

Anas carolinensis Reichenow, Omith. Monatsber., 1901, p. 17 (Jaluit); Schnee, 
Ornith. Monatsber., 1901, p. 131 (Marshalls) ; idem, Zool. Jahrbiicher, 20, 1904, p. 390 
(Marschall Inseln) ; Phillips, Nat. Hist. Ducks, 2, 1923, p. 235 (Marshall Islands). 

Querquedula crecca carolinensis Hand-list Japanese Birds, rev., 1932, p. 185 (Mar- 
shall Islands). 



Baker: The Avifauna of Micronesia 101 

Anas crecca carolinensis Hand-list Japanese Birds, 3d ed., 1942, p. 206 (Marshall 
Islands). 

Geographic range. — Breeds in northwestern and northcentral North America. 
Winters to West Indies, Central America and Mexico. In Micronesia: Mar- 
shall Islands — Jaluit. 

Remarks. — Reichenow (1901:17) and Schnee (1901:131) record 
the Green-wing Teal in the Marshall Islands. It is the only record 
known for Micronesia. Bryan and Greenway (1944:104) record the 
teal as a migrant to the Hawaiian Islands. 

Anas acuta acuta Linnaeus 
Pintail 

Anaa acuta Linnaeus, Syst. Nat., ed. 10, 1, 1758, p. 126. (Type locality, Europe, 
restricted to Sweden.) 

Dafila acuta acuta Hand-list Japanese Birds, rev., 1932, p. 185 (Pagan). 
Anas acuta acuta Hand-list Japanese Birds, 3d ed., 1942, p. 206 (Pagan). 
Anas acuta Mayr, Birds Southwest Pacific, 1945, p. 302 (Micronesia). 

Geographic range. — Breeds in Iceland, northern Europe and Asia. Winters 
south to northern Africa, Asia and Philippines. In Micronesia: Mariana Islands 
— Pagan, Guam; Palau Islands — exact locality unknown. 

Remarks. — The Pintail has been recorded from Pagan and Guam 
in the northern Marianas and from the Palau Islands and is thought 
to be an uncommon visitor to Micronesia. At Guam, Flavin (fieiu 
notes) recorded one female on October 27, 1945, and three females 
and two drakes on January 19, 1946. Marshall (1949:221) saw a 
flock of fifteen Pintails at Saipan on February 7, 1945. 

Anas acuta tzitzihoa Vieillot 
Pintail 

Anas tzitzihoa Vieillot, Nouv. Diet. Hist. Nat., 5, 1816, p. 163. (Type locality, 
Mexico, ex Hernandez.) 

Anas acuta americana Reichenew, Ornith. Monatsber., 1901, p. 17 (Jaluit); Schnee, 
Zool. Jahrbucher, 20, 1904, p. 390 (Marschall Inseln). 

Anas acuta Schnee, Ornith, Monatsber., 1901, p. 131 (Marshalls); Phillips, Nat. 
Hist. Ducks, 2, 1923, p. 316 (Jaluit). 

Anas acuta tzitzihoa Hand-list Japanese Birds, 3d ed., 1942, p. 206 (Marshall 
Islands). 

Geographic range. — Breeds in northwestern and northcentral North America. 
Winters south to West Indies, Panama, and west to Hawaiian Islands. In 
Micronesia: Marshall Islands — Jaluit. 

i^emarA's.— Reichenow (1901:17) and Schnee (1901:131) re- 
ported that flocks of ducks belonging to this and other American 
species were observed in the Marshall Islands in October, 1899, and 
May, 1900. This species may winter in the Hawaiian Islands, ac- 
cording to Peters (1931:167). If so it is not surprising that occa- 
sional visitors reach eastern Micronesia. 



102 University of Kansas Publs., Mus. Nat. Hist. 

Anas penelope Linnaeus 
Widgeon 

Anas penelope Linnaeus, Syst. Nat., ed. 10, 1, 1758, p. 126. (Type locality, 
Europe, restricted to Sweden.) 

Anas penelope Finsch, Ibis, 1880, pp. 332, 333 (Taluit) ; Schnee, Zool. Jahrbiicher, 
20, 1904, p. 390 (Marschall Insein) ; Phillips, Nat. Hist. Ducks, 2, 1923, p. 175 
(Taluit); Hand-list Japanese Birds, 3d ed., 1942, p. 206 (Tinian, Yap, Jaluit) ; Mayr, 
Birds Southwest Pacific, 1945, p. 302 (Micronesia). 

Mareca penelope Finsch, Mitth. Ornith. Ver. Wien, 1884, p. 56 (Jaluit); Wigles- 
worth, Abhandl. und Ber Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 71 (Taluit); 
Finsch, Deut. Ver. zum Schultze der Vogelwelt, 18, 1893, p. 458 (Marshalls) ; Kuroda, 
in Momoyama, Birds Micronesia, 1922, p. 38 (Taluit) ; Hand-list Japanese Birds, 
rev., 1932, p. 185 (Tinian, Yap, Jaluit). 

Geographic range. — Breeds in Iceland, northern Europe and Asia. Winters 
south to Africa, southern Asia and Philippines; casual to eastern North 
America. In Micronesia: Mariana Islands — Tinian; Caroline Islands — Yap; 
Marshall Islands — Jaluit. 

Remarks. — The Widgeon may be an occasional winter visitor to 
Micronesia. The recorci at Jaluit in the Marshall Islands may be 
questioned. 

Anas clypeata Linnaeus 
Shoveller 

Anas clypeata Linnaeus, Syst. Nat., ed. 10, 1, 1758, p. 124. (Type locality. 
Coasts of Europe, restricted to southern Sweden.) 

Spatula clypeata Hand-list Japanese Birds, rev., 1932, p. 185 (Pagan); Yamashina, 
Tori, 10, 1940, p. 676 (Pingelap) ; Hand-list Japanese Birds, 3d ed., 1942, p. 206 
(Pagan, Pingelap). 

Anas clypeata Mayr, Birds Southwest Pacific, 1945, p. 302 (Micronesia). 

Geographic range. — Breeds in northern Europe, Asia, North America and 
adjacent islands. Winters to northern Africa, southern Asia, Philippines, 
Hawaiians, southern United States to Central America. In Micronesia: Mari- 
ana Islands — Pagan, Tinian; Caroline Islands — Ponape, Pingelap. 

Specimens examined. — One female from Mariana Islands, USNM — Tinian (Oct. 12). 

Remarks. — The Shoveller is known from localities in the Mari- 
anas and in the Carolines. In the collections of the American Mu- 
seum of Natural History there is a female taken by Rollo Beck at 
Kauehi, Tuamotu Archipelago, on March 6, 1923. A specimen ex- 
amined from Tinian was taken there by Joe T. Marshall, Jr., at 
Lake Hogoya on October 12, 1945. Richards obtained two Shovel- 
lers (one immature male and one immature female) at Ponape on 
December 21, 1947, and January 6, 1948, respectively. He found 
them in a pond in a bomb crater. This duck appears to be a casual 
winter visitor to Micronesia and other parts of Oceania. 



Baker: The Avifauna of Micronesia 103 

Aythya fuligula (Linnaeus) 
Tufted Duck 

Anas fuligula Linnaeus, Syst. Nat., ed. 10, 1, 1758. p. 128. (Type locality, Europe, 
restricted to Sweden.) 

Fuligula cristata Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 9, 118 
(Pelew); idem, Proc. Zool. Soc. London, 1872, p. 90 (Pelew) ; Finsch. Journ. Mus. 
Godeffroy, 8, 1875, pp. 5. 40 (Palau) ; Wiglesworth, Abhandl. und Ber. Zool. Mus. 
Dresden, no. 6, 1890-1891 (1891), p. 71 (Pelew); Oustalet, Nouv. Arch. Mus. Hist. 
Nat. Paris, (3), 8, 1896, p. 50 (Mariannes, Palaos). 

Fuligula fuligula Salvadori, Cat. Birds British Mus., 27, 1895, p. 363 (Pelew); 
Hartert, Novit. Zool., 5, 1898. p. 68 (Marianne); Seale, Oce. Papers Bernice P. 
Bishop Mus., 1. 1901, p. 26 (Micronesia); Safford, Osprey, 1902, p. 70 (Mariannes); 
idem. The Plant World. 7, 1904, p. 268 (Guam) ; Kuroda, in Moniiyama, Birds Micro- 
nesia, 1922, p. 38 (Mariane, Pelew, Yap); idem, Av-ifauna Riu Kiu, 1925, p. 143 
(Pelew, Marianne). 

Marila fuligula McGregor, Man. Philippine Birds, 1909, p. 199 (Marianne, Pelew). 

Nyroca fuligula Phillips. Nat. Hist. Ducks, 3, 1925, p. 234 (Guam); Hand-list 
Japanese Birds, rev.. 1932. p. 185 (Pagan, Saipan, Palau. Yap); Bryan, Guam Rec, 
vol. 13, no. 2, 1936, p. 15 (Guam); Mayr, Birds Southwest Pacific, 1945, p. 302 
(Micronesia). 

Aytha fuligula Hand-list Japanese Birds, 3d ed., 1942. p. 207 (Pagan. Saipan, 
Tinian, Guam, Yap, Palau). 

Geographic range. — Breeds in Iceland, Europe, northern Asia. Winters in 
Europe, Africa, Asia, Malaysia, and parts of Oceana. In Micronesia : Mariana 
Islands— Pagan, Saipan, Tinian, Guam ; Palau — exact locality unknown ; Caro- 
line Islands — Yap. 

Remarks. — The Tufte(i Duck is a winter migrant to western Mi- 
cronesia. It has been recorded only a few times and may be an 
irregular visitor. Flavin observed a duck, which he thought to be 
of this species, at Guam on January 19, 1946. Marshall (1949: 221) 
reports that two Tufted Ducks were seen at Lake Hagoi in April 

1945. 

Aythya valisineria (Wilson) 

Canvasback 

Anas valisineria Wilson. Amer. Ornith.. 8, 1814. p. 103. pi. 70, f. 5. (Type locality. 
Eastern United States.) 

Nyroca vallisneria Reichenow, Ornith. Monatsber., 1901, p. 17 (Jaluit) ; Schnee. 
Ornith. Monatsber., 1901, p. 131 (Marshalls); idem, Zool. Jahrbiicher, 20, 1904, p. 
390 (Marschall Inseln). 

Nyroca valisineria Phillips, Nat. Hist. Ducks, 3, 1923, p. 124 (Marshall Islands). 

Aythya valisineria Hand-list Japanese Birds, 3d ed., 1942, p. 207 (Marshall Islands). 

Geographic range. — Breeds in northwestern and northcentral North America. 
Winters south to Gulf States, Florida and Mexico. In Micronesia: Marshall 
Islands — Jaluit. 

i2emar/cs.— Reichenow (1901:17) and Schnee (1901:131) re- 
ported three species of American ducks {Aythya valisineria, Anas 
acuta tzitzihoa and Anas crecca carolinensis) in the Marshalls in 
October, 1899, and May, 1900. These species may be stragglers to 
eastern Micronesia. 



104 University of Kansas Publs., Mus. Nat. Hist. 

Accipiter soloensis (Horsfield) 
Chinese Goshawk 

Falco Soloensis Horsfield, Trans. Linn. Soc. London, 13, 1821, p. 137. (Type lo- 
cality, Java.) 

Accipiter soloensis Hand-list Japanese Birds, rev., 1932, p. 182 (Yap); Hand-list 
Japanese Birds, 3d ed., 1942, p. 203 (Yap, Rota); Mayr, Birds Southwest Pacific, 
1945, p. 302 (Yap). 

Geographic range. — Breeds in northern China south to Kwangtung. Winters 
to Malaysia. In Microne.sia : Mariana Islands — Rota ; Carohne Islands — Yap. 

Remarks. — The Chinese Goshawk is a winter visitor to Micro- 
nesia and has been recorded at Rota and Yap. The NAMRU2 party 
saw several unidentified hawks in Micronesia in 1945. At Mt. Tenjo, 
Guam, Muennink saw a small hawk, resembling an accipiter, darting 
at swiftlets on June 8, 1945. At Angaur, the writer saw a small hawk 
flying through heavy vegetation along the rugged coast line on Sep- 
tember 21, 1945. A hawk "Butio ( ?) " was reported at Saipan in 1945 
by Moran (1946:262) ; this hawk may have been Butastur indicus 
(Gmelin). Marshall (1949:221) reports seeing "three kinds of 
hawks" on Palau in November, 1945. Obviously, further observa- 
tions and collecting will increase our knowledge of the known num- 
ber of kinds of hawks which visit Micronesia. 

Accipiter virgatus gularis (Temminck and Schlegel) 
Asiatic Sparrow Hawk 

Astur (Nisus) gularis Temminck and Schlegel, in Siebold, Fauna Japon., Aves, 1845, 
p. 5, pi. 2. (Type locality, Japan.) 

Accipiter nisoides Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, p. 166 
(Guam); Hartert, Novit. Zool., 5, 1898, p. 51 (Marianne); Scale, Occ. Papers Bernice 
P. Bishop Mus., 1, 1901, p. 44 (Guam); Safford, Osprey, 1902, p. 70 (Marianas). 

Accipiter gularis Kuroda, in Momiyama, Birds Micronesia, 1922, p. 39 (Guam). 

Accipiter virgatus gularis Hand-list Japanese Birds, rev., 1932, p. 182 (Guam); 
Hand-list Japanese Birds, 3d ed., 1942, p. 203 (Guam); Mayr, Birds Southwest Pacific, 
1945, p. 302 (Micronesia). 

Accipiter virgatus nisoides Bryan, Guam. Rec, vol. 13, no. 2, 1936, p. 15 (Guam). 

Geographic range. — Breeds in Japan and northern China. Winters south to 
Philippines and Malaysia. In Micronesia: Mariana Islands — Guam. 

Remarks. — Oustalet (1895:166) records a male bird shot by 
Marche at Guam in October, 1887. Scale (1901:44) records a speci- 
men taken at Guam by Owston's Japanese collectors. These are the 
only records found for Micronesia, and the hawk may be classed as 
a casual winter visitor. Strophlet (1946:535) observed "a small 
light-throated" falcon at Guam on November 7, 1945, which may 
have been of this species. 



Baker: The Avifauna of Micronesia 105 

Pandion haliaetus melvillensis Mathews 

Osprey 

Pandion haliaetus melvillensis Mathews, Australian Avium Rec, 1, 1912, p. 34. 
(Type locality, Melville Island.) 

Pandion leucocephalus Finsch, Joum. Mus. Godeffroy, 8, 1875, p. 49 (Palau). 

Pandion haliaetus leucocephalus Wiglesworth, Abhandl. und Ber. Zool. Mus. Dres- 
den, no. 6, 1890-1891 (1891), p. 1 (Pelew). 

Pandion haliaetus cristatus Kuroda, in Momiyama, Birds Micronesia, 1922, p. 40 
(Pelew); Hand-list Japanese Birds, rev., 1932, p. 182 (Pelew); Hand-list Japanese 
Birds, 3d ed., 1942, p. 203 (Palau). 

Pandion haliaetus melvillensis Mayr, Birds Southwest Pacific, 1945, pp. 55, 286 
(Palau); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 46 (Guam, Palau). 

Geographic range. — Malaysia, northern Australia, Melanesia. In Micronesia: 
Mariana Islands — Guam; Palau Islands — Peleliu. 

Remarks. — The Osprey was first recorcied at Palau by Finsch 
(1875:49). The author (1948:46) cites recorcis obtained by C. 
K. Dorsey at Peleliu in 1944 and 1945. Dorsey saw the Osprey on 
several occasions; the NAMRU2 party did not find the bird while 
on their stay there in August and September, 1945. B. V. Travis of 
NAMRU2 saw an Osprey at Agana Bay, Guam, in December, 1945. 
He observed the bird to be carrying a fish in its talons. Flavin ob- 
served the Osprey at Guam on January 28, 1945, and on December 
23, 1945. Mayr (1945a: 286) says that the Osprey apparently 
breeds at Palau. The bird seen in the Marianas may have been P. 
h. haliaetus (Linnaeus) , a visitor from Asia, which is known to 
winter in the Philippines and adjacent areas. 

The Osprey is the only resident member of the order Falconifor- 
mes, and it is principally a fish eater. The few records of mammal 
and bird eating hawks in Micronesia indicate that predation on 
insular vertebrate populations from this source is at a minimum. 
The absence of this predation may have a pronounced effect on the 
resident land birds, particularly from the standpoint of the per- 
petuation of nonadaptive mutations, which might be "weeded out" 
under what might be considered as normal predatory pressure in 
continental bird populations. 

Faico peregrinus japonensis Gnielin 
Peregrine Falcon 

Falco japonensis Gmelin, Syst, Nat., 1, pt. 1, 1788, p. 267. (Type locality. Off the 
coast of Japan.) 

Falco peregrinus Hartlaub and Finsch, Proc. Zool. Soc. London, 1872, pp. 89, 90 
(Mackenzie); Griiffe, Journ. Mus. Godeffroy, 2, 1873, p. 122 (Yap); Finsch, Joum. 
Mus. Godeffroy, 8, 1875, pp. 4, 8 (Palau); Schmeltz and Krause, Ethnogr. Abth. Mus. 
Godeffroy, 1881, p. 391 (Yap) ; Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, 
no. 6, 1890-1891 (1891), p. 1 (Yap, Pelew); Hand-list Japanese Birds, rev. 1932, p. 



106 University of Kansas Publs., Mus. Nat. Hist, 

182 (Yap, Palau); Hand-list Japanese Birds, 3d ed., 1942, p. 202 (Yap, Palau) ; 
Mayr, Birds Southwest Pacific, 1945, p. 302 (Yap, Palau); Baker, Smithson. Misc. 
Coll., vol. 107, no. 15, 1948, p. 46 (Guam). 

tFalco peregrinus calidus Kuroda, in Momiyama, Birds Micronesia, 1922, p. 40 
(Yap, Pelew). 

Geographic range. — Breeds in northern Asia. Winters to southern Asia, 
Malaysia and Melanesia. In Micronesia: Mariana Islands — Guam; Palau 
Islands — exact locality unknown ; Caroline Islands — Yap. 

Remarks. — The Peregrine Falcon may be classed as a casual 
winter visitor to Micronesia. It has been recorded by Hartlaub and 
Finsch at Yap and Palau. A specimen from Yap was taken by 
Kubary in November, 1870. On November 2, 1945, at Guam as 
previously recorded (Baker, 1948:46) Irvin O. Buss saw a falcon 
alight on the superstructure of his ship. He watched it catch and 
eat a Common Noddy [Anous stolidus) . As the ship approached 
the island, the bird flew to the rugged cliffs near Facpi Point. 
Strophlet (1946:535) saw a large falcon, ''presumed to be a Duck 
Hawk," at Guam on November 16, 1945. Possibly these two ob- 
servers saw the same bird. In July, 1945, Flavin observed a Pere- 
grine Falcon at Guam. F. p. fruitii Momiyama, which is known 
from the Volcano Islands, may occur in Micronesia. 

Megapodius laperouse senex Hartlaub 
Micronesian Megapode 

Megapodius senex Hartlaub, Proc. Zool. Soc. London, 1867 (1868), p. 820. (Type 
locality, Pelew Islands.) 

Megapodius senex Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 7, 118 
(Pelew); Gray, Hand-list Birds, 2, 1870, p. 256 (Pelew); Hartlaub and Finsch, Proc. 
Zool. Soc. London, 1872, pp. 89, 103 (Pelew); Finsch, Joum. Mus. Godeffroy, 8, 1875, 
pp. 5, 29, pi. 5, fig. 2, 3 (Palau); Giebel, Thes. Ornith., 2, 1875, p. 547 (Pelew); 
Sehmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, p. 407 (Palau); Oustalet, 
Ann. Sci. Nat., (6), art. 2, 1881, pp. 63, 140, 145, 171, 175 (Pelew); Tristram, Cat. 
Birds, 1889, p. 30 (Pelew); Wiglesworth, Abhandl. und Bar. Zool. Mus. Dresden, no. 
6, 1890-1891 (1891), p. 58 (Pelew); Oustalet, Ann. Sci. Nat., Zool., 11, 1891, p. 196 
(Peleu); idem, Nouv. Arch Mus. Hist. Nat. Paris, (3), 8, 1896, p. 30 (Palaos) ; 
Ogilvie-Grant, Hand-book Game-birds, 2, 1897, p. 182 (Pelew); Hartert, Novit. Zool., 
5, 1898, p. 62 (Pelew); Bolau, Mitteil, Naturhist. Mus. Hamburg, 1898, p. 69 (Palau); 
Finsch, Sammlung wissensch. Vortriige, 14 ser., 1900, p. 659 (Palau); Matschie, Joum. 
f. Omith., 1901, p. 113 (Palau); Lister, Proc. Zool. .Soc. London, 1911, p. 757 (Pelew). 

Megapodius laperousii Ogilvie-Grant (part). Cat. Birds British Mus., 22, 1893, p. 
460 (Pelew) ; Takastukasa and Kuroda, Tori, 1, 1915, p. 51 (Pelew) ; Kuroda, Dobutsu. 
Zasshi, 27, 1915, p. 390 (Pelew); idem, Dobutsu. Zasshi, 28, 1916, p. 69 (Pelew). 

Megapodius laperousi Seale (part), Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 
39 (Pelew); Safford (part). The Plant World, 7, 1904, p. 265 (Pelew); Uchida, Annot. 
Zool. Japon., 9, 1918, pp. 486, 487 (Palau). 

Megapodius laperousii var. senex Dubois, Syn. Avium, 2, 1904, p. 787 (Pelew). 

M[egapodius] lapeyrousei Reichenow (part), Die Vogel, 1, 1913, p. 273 (Palauin- 
seln). 

Megapodius laperov^ei senex Kuroda, in Momiyama, Birds Micronesia, 1922, p. 40 
(Pelew). 

Megapodius laperouse senex Mathews, Syst. Avium Australasianarum, 1, 1927, p. 14 
(Pelew); Takstsukasa, Birds Nippon, vol. 1, pt. 1, 1932, p. 13, pi. 4, 5 (Pelew); 
Yamashina, Tori, 7, 1932, p. 412 (Ngesebus, Auror, Peliliu) ; Hand-list Japanese Birds, 



Baker: The Avifauna of Micronesia 107 

rev., 1932, p. 198 (Palau) ; Peters, Check-list Birds World, 2, 1934, p. 6 (Palau) ; 
Yamashina, Tori, 10, 1940, p. 679 (Gayangas, Arumidin) ; Amadon, Amer. Mus. Novit., 
no. 1175, 1942, p. 9 (Palau); Mayr, Birds Southwest Pacific, 1945, p. 286 (Palau); 
Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 46 (Garakayo, Peleliu, Ngabad). 

Megapodius la peroitse senex Hand-list Japanese Birds, 3d ed., 1942, p. 223 (Babel- 
thuap, Koror, Auror, Ngesebus, Peliliu, Gayangas, Arumidin). 

Megapodius laperouse Wharton and Hardcastle, Joum. Parasitology", 32, 1946, p. 
294 (Garakayo). 

Geographic range. — Micronesia: Palau Islands — Babelthuap, Koror, Auror, 
Kayangel, Garakayo, Ngesebus, Peleliu, Ngabad, Gayangas, Arumidin. 

Characters. — Adult: A small megapode with top of head near "mouse gray"; 
forehead, sides of face and neck, chin, and throat thinly covered with feathers 
of the same color; mantle and upper breast grayish-black becoming dark olive- 
brown on wings; lower back, rump and upper tail-coverts dark brown; tail 
blackish-brown; underparts grayi.sh-brown, lighter on midline of belly; under 
wings dark brown; exposed skin of head reddish to yellowish-red; bill yellow- 
ish, basally blackish; legs yellowish; feet and claws black; iris tan. 

Measurements. — Measurements of three adult males: wing, 178, 182, 188; 
tail, 55, 63; culmen, 22.7, 23.3; tarsus, 55, 56, 57; of seven adult females: wing, 
171-189 (182); tail, 46-68 (58); culmen, 25-30 (27); tarsus, 45-60 (55). Taka- 
tsukasa (1932:14) lists the following measurements: males — wing, 176-181; 
tail, 59-67; culmen, 25.5-26.0; tarsus, 58-61; females— wing, 177-187; tail, 62-68; 
culmen, 24.0-26.0; tarsus, 55-58. 

Specimens examined. — Total number, 23 (11 males, 8 females, 4 unsexed), as follows: 
Palau Islands, USNM— Koror, 1 (Nov. 28)— Garakayo, 5 (Sept. 17, 18, 19)— Peleliu, 2 (Aug. 
31, Sept. 1)— Ngabad, 1 (Sept. 11); AMNH— Palau, 16 (Nov., Dec., not dated). 

Nesting. — The megapodes do not incubate their eggs, but the female de- 
posits them in a moundlike structure of sand, volcanic ash, and forest litter 
or some other type of soil in which there is warmth sufficient to hatch the 
eggs after an extended period (perhaps 40 days or more) without further at- 
tention from the parent bird. The young dig out and lead an independent 
existence. Several megapodes may utilize one nest site, which ordinarily is 
at a low elevation near a beach or lagoon. 

The NAMRU2 party obtained two downy chicks at Gayakayo Island on 
September 18 and 19, 1945. A female taken on September 1 at Peleliu con- 
tained large eggs. Coultas obtained two chicks (one in postnatal molt) in 
November and December, 1931. Kubarj', as quoted by Takatsukasa (1932: 
15), says that eggs may be found in the mounds throughout the year at 
Palau but are found most numerously in the south-east monsoon (April to 
November). Yamashina (1932a :412) reports on eggs taken in 1932 as fol- 
fows: eight eggs from Auror Island on Januarj' 15; one egg from Ngesebus 
Island on January 16; and four eggs from Peleliu Island on January 16. 
Takatsukasa (1932:15) states that eggs are most numerous in the mounds 
in the months of May and June. The chicks obtained by NAMRU2 in Sep- 
tember were of such a size as to suggest that they too had been laid in 
June. 

Takatsukasa (1932:15) comments, "Whilst Dr. Yaichiro Okada was in the 
Pelew Group, he found two nests on Kajangel Island, which is an uninhabited 
island about twelve sea-miles southeast of the island of Malacal. He says 
that he found two nests, one of which was obsolete and the other was in use. 



108 University of Kansas Publs., Mus. Nat. Hist. 

The first one was oval in shape; the diameter of the largest part was twenty- 
four feet, and the smallest part was twenty feet, and it had a height of four 
feet. The second one was fan-shaped, as an obstacle existed at one side of 
the nest, and its diameter was twelve feet and the height was a little more 
than four feet, and the native whom he asked to dig out the eggs got three. 
One of the eggs contained a well-advanced embryo and the others were not 
so advanced as the first one. This distance from the top of the mound to 
the spot where the eggs were laid was about two and a half feet, and the 
natives made a great deal of effort to get these eggs. These nests were 
found in the bush by the natives." The NAMRU2 party observed a mound 
on Ngabad Island, a small islet near Peleliu, on September 11. It was much 
like those described by Takatsukasa, being approximately six feet high and 
some twelve or fifteen feet across. It was not excavated. 

Molt. — Birds taken in August, September and November were molting body 
feathers. Birds taken in December were molting wing feathers. 

Food habits. — Takatsukasa (1932:16) comments, "My collector reports to 
me that this bird diets on insects and tender shoots which it gets from under 
the soil by scratching with its large and powerful feet." According to Captain 
Tetens, as noted by Takatsukasa, the food of the bird consists of insects and 
berries. Birds taken by the NAMRU2 party had the following food items 
in their stomachs: adult female — 2 cc. seeds, grit; adult female — 3 cc. crab 
parts, grit; adult female — 2 cc seeds, sand; male chick — 1 cc. ground food, grit; 
female chick — 1 cc. ground food, grit, in crop 3 cc. small wood roaches 
(Blattidae). 

Parasites. — Wharton and Hardcastle (1946:294) obtained the chigger 
(Acarina), Neoschongastia yeomansi, from the megapode at Palau. Uchida 
(1918:486, 487) found the bird lice (Mallophaga), Goniocotes minor and 
Ldpeurtis sinuatus, on megapodes from the Palaus. 

Remarks. — The NAMRU2 party arrived at the Palau Islands on 
August 23, 1945, with little notion that the megapode would be found 
on the war-torn island of Peleliu. As reported by the author (1946b: 
209 and 1948:46) we found birds in small numbers in the relatively 
undisturbed areas of rough coral covered by jungle and a few birds 
in the heavy matting of viny and brushy vegetation which was rap- 
idly covering the battlefields. The finding of a higher population on 
the more isolated and relatively undisturbed offshore islets (Ngabad, 
Garakayo) by the NAMRU2 party was an observation similar to 
those of Takatsukasa (1932:15, 16) and Coultas (field notes). 
Takatsukasa (1932:16) remarks, "Dr. Finsch said that this Mega- 
pode frequents nearly all the islands of the Pelew Group . . . 
but it is very noticeable that this bird has either disapj^eared, or only 
very rarely exists in the following islands: Koror, Ngarekobasanga, 
and especially the main island of Babelthuap." He quotes Otto 
Finsch as remarking that, "It seems that the bird occasionally moves 
from one island to another, as the bird is a good flier." Takatsukasa 



Baker: The Avifauna of Micronesia 109 

continues, "According to Tetens, this Megapode runs very swiftly 
among the bushes, and when it is startled it takes to the nearest tree. 
. . . Captain Wilson says nothing about the Megapode, but Dr. 
Finsch wrote that Captain Wilson is probably referring to the egg of 
this bird under 'Wild Fowls,' when he said that the natives of the 
Palaus do not eat the flesh of the birds, but they go to the woods and 
bring back the eggs; they do not appreciate the newly laid eggs, but 
they consider it as a delicacy to swallow the well advanced embryo." 
The NAMRU2 party found the birds to prefer rough, coral jungle 
where there was considerable heavy undergrowth and ground litter. 
The birds were located by their loud screeches and cackles but were 
difficult to stalk. It was best to remain quiet and let them approach 
within shooting distance. Young chicks were extremely active and 
wild. One of the two chicks taken at Garakayo was obtained by a 
fortunate shot as the bird was flying rapidly through the brush. The 
natives use them as food, and I learned of one serviceman who had 
worked out a technique for trapping the birds. He traded the live 
birds to the natives for island souvenirs. As Wilson and Takatsukasa 
note, the natives apparently prefer the eggs to the adults as food, and 
in normal times of food abundance they probably do not molest the 
adults but hunt for their eggs. This seems logical, since if a deter- 
mined trapping program were in operation by the natives, it should 
not take many decades to eliminate completely the entire population. 
On four islands visited by the NAMRU2 party in August and Sep- 
tember, 1945, I estimated the following populations: Garakayo — 20 
to 30; Ngabad— 5 to 10; Peleliu— 10 to 20; Angaur— less than 10. 

Megapodius laperouse laperouse Gaimard 
Micronesian Megapode 

Megapodius La Perouse Gaimard, Bull. Gen. Univ. Annon. Nouv. Sci., 2, 1823, 451. 
(Type locality, Tinian, Archipel des Mariannes.) 

Megapodius La Perouse Quoy and Gaimard, Voy. "Uranie," Zool., 1824, pp. 127, 
693, Atlas, pi. 33 (Tinian); idem, Ann. Sci. Nat. Paris, 6, 1825, p. 149 (Tinian) 

Megapodius La Pcrousii Quoy and Gaimard, Voy. "Uranie," Zool., 1824, p. 127, pi. 
33 (Tinian); Wagler, Isis, 1829, p. 735 (Tinian, Guam, Rota); Gray, Cat. Birds Trop. 
Is. Pacific Oceon, 1859, p. 46 (Tinian); Oustalet, Ann. Sci. Nat., (6), art. 2, 1881, pp. 
63, 138, 140, 143, 171, 175, 176, 177 (Tinian); idem, Le Nat., 1889, p. 261 (Mariannes); 
idem, Ann. Sci. Nat., Zool., 11, 1891, p. 196 (Tinian, Seypan, Pagon). 

Megapodius La Peyrouse Lesson, Man. d'Ornith., 2, 1828, p. 221 (Tinian) ; idem, 
Compl. de Buffon, 2d ed., 2, Ois., 1838, p. 255 and accompanying plate (Tinian). 

Megapodius laperousii Lesson, Traite d'Ornith., 1831, p. 478 (Mariannes); Gray, 
Hand-list Birds, 2, 1870, p. 256 (Marian); Ogilvie-Grant, Hand-book Game-birds, 2, 
1897, p. 183 (Marianne); Dubois, Syn. Avium, 2, 1904, p. 787 (Mariannes); Lister, 
Proc. Zool. Soc. London, 1911, p. 757 (Marianne). 

Megapodius Lapeyrousii Hartlaub, Journ. f. Ornith., 1854, p. 167 (Mariannen). 

Megapodius La Peyrousii Reichenbach, Tauben, 1861, p. 5 (Marianen). 

Megapodius la-perousi Gray, Proc. Zool. Soc. London, 1864, p. 43 (Guam, Botta, 
Tinian). 



110 University of Kansas Publs., Mrs. Nat. Hist. 

Megapodius laperousi Giebel, Thes. Omith., 2, 1875, p. 547 (Marianae); Seale, Occ. 
Papers Bernice P. Bishop Mus., 1, 1901, p. 39 (Marianas); Safford ; Osprey, 1902, p. 
68 (Tinian); idem. The Plant World, 7, 1904, p. 265 (Tinian); idem, Contr. U. S. Nat. 
Herb., 9, 1905, p. 78 (Rota, Saipan, Pagan, Agrigan) ; Schnee, Zeitschr, f. Naturwisch., 
82, 1912, p. 467 (Marianen); Prowazek, Die deutschen Marianen, 1913, pp. 47, 101 
(Marianen); Linsley, Guam, Rec, vol. 12, no. 8, 1935, p. 249 (Rota, Saipan, Pagan, 
Agrigan). 

Megapodius perousei Finsch, Journ. Mus. Godeffroy, 8, 1875, p. 30 (Marianen); 
Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 58 
(Guam, Botta, Tinian, Pagon). 

Megapodius laperousii Ogilvie-Grant (part). Cat. Birds British Mus., 22, 1893, p. 
460 (Marianne). 

Megapodius la perousei Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 8, 1896, 
p. 26 (Saypan, Pagan, Rota, Agrigan, Tinian). 

Megapodius laperouse Hartert, Novit. Zool., 5, 1898, p. 61 (Saipan, Tinian, Rota, 
Guam). 

Megapodius laperousei Finsch, Sammlung wissensch. Vortrage, 14 ser., 1900, p. 600 
(Marianen); Prowazek, Die deutschen Marianen, 1913, p. 87 (Marianen). 

Megapodius lapeyrouse Matschie, Journ. f. Ornith., 1901, p. 113 (Guam, Saipan). 

M[egapodius] lapeyro^isei Reichenow (part). Die Vogel, 1, 1913, p. 273 (Mariannen). 

Megapodius laperousei laperousei Kuroda, in Momiyama, Birds Micronesia, 1922, p. 
40 (Guam, Saipan, Rota, Tinian, Pagan, Agrigan). 

Megapodius laperouse laperouse Mathews, Syst. Avium Australasianaruni, 1, 1927, 
p. 16 (Marianas); Takatsukasa, Birds Nippon, vol. 1, pt. 1, 1932, p. 6, pi. 4, 5 
(Marianne); Yamashina, Tori, 7, 1932, p. 411 (Pagan Agrigan); Hand-list Japanese 
Birds, rev., 1932, p. 198 (Marianas); Peters, Check-list Birds World, 2, 1934, p. 7 
(Marianne Islands); Yamashina, Tori, 10, 1940, p. 679 (Assongsong) ; Aniadon, Amer. 
Mus. Novit., no. 1175, 1942, p. 9 (Asuncion, Saipan, Guam); Mayr, Birds Southwest 
Pacific, 1945, p. 286 (Marianas). 

Megapodius laperousi laperousi Bryan. Guam Rec, vol. 13, no. 2, 1936, p. 15 
(Guam). 

Megapodius la perouse la perouse Hand-list Japanese Birds, 3d ed., 1942, p. 223 
(Assongsong, Agrigan, Pagan, Alniagan, Saipan, Tinian, Agiguan, Rota, Guam). 

Geographic range. — Micronesia: Mariana Islands — Asuncion, Agrihan, 
Pagan, Almagan, Saipan, Tinian, Agiguan, Rota, Guam. Probably extinct on 
Saipan, Tinian, Rota, Guam. 

Characters. — Adult: Resembles M. I. senex, but crown slightly darker gray; 
back, wing-coverts and scapulars more heavily washed with olivaceous-brown; 
mantle less slate; underparts paler and more brownish, especiallj' belly. (Note 
— The specimens examined from the Marianas are old and rather worn in 
appearance.) 

Measurements. — Two males measure: wing 180?, 182?; tail 62, 63; tarsus 
55, 55; three females: wing 181?, 181?; tail 55, 59, 62; tarsus 54, 54, 56. 
Takatsukasa (1932: 10) lists the following measurements: males — wing, 155- 
169; tail, 54-62; culmen, 22.5-24; tarsus, 51-54; females— wing, 158-170; tail, 
56-65; culmen, 23-25; tarsus, 50-55. 

Specimens examined. — Total number, 10 (3 males. 4 females, 3 unsexed), as follows; 
Mariana Islands, AMNH — Guam, 1 (June 6) — Saipan, 6 (1895) — Asuncion, 3 (1904). 

Nesting. — Concerning the nest of the Micronesian Megapode in the Mari- 
anas, Takatsukasa (1932:10) writes: "The nest is a large mound of sand mixed 
with grass and is made in the wooded land along the seashore. The mound 
is over one hundred feet in circumference and a few yards in height, and is 
built by the united efforts of the male and female, by scratching sand and 
grass with their large feet. The eggs are laid in this mound and they are 
hatched by the heat of the sun and that produced by the fermentation of the 



Baker: The Avifauna of Micronesia 111 

grass, and they are never hatched by the parent birds. The egg is of a pale 
brown, but always stained by nesting materials." 

Takatsukasa (1932:11) quotes Oustalet as follows: "Specimens collected 
by Mr. Marche have proved that the breeding season of La Perouse's Mega- 
pode is rather long, like the other species of the same family, it begins to 
breed in January or February and ends in June. Accordingly, in this period 
the eggs just laid, the chicks, the young and adult can be seen at one place, 
but Mr. Marche did not obtain any egg." Hartert (1898:61) records a chick 
taken on July 17. Yamashina (1932a: 411) records eggs taken in 1931 as 
follows: two eggs from Pagan, February 17; three eggs from Pagan, May 15; 
four eggs from Agrihan, June 24. The breeding season for both of the incuba- 
tor birds, M. I. senex and M. I. laperov^e, is apparently from about January 
to August. 

Remarks. — The Micronesian Megapode was first taken in the 
Marianas by the expedition of the Uranie. Berard, a member of 
the expedition, obtained the bird at Tinian in December, 1820. 
Quoy and Gaimard (1824:27), who studied the birds of this ex- 
pedition, reported that according to native tradition the species was 
in former times widely distributed in the Marianas and domesti- 
cated by the ancient people of the islands, but that in 1819 and 
1820 the birds were not numerous on Tinian and not found on Guam 
and Rota. IMarche (in Oustalet, 1896:27) obtained twenty-three 
birds at Saipan, one from Rota, two from Agrihan, and five from 
Pagan in 1887, 1888, and 1889; it is apparent that Quoy and Gai- 
mard missed the bird at Rota. Marche was of the opinion that the 
megapodes were never domesticated and that they would probably 
not last much longer at Saipan and Rota owing to the incessant 
hunting for them by the natives. As in the Palaus, the natives ap- 
parently prefer the eggs to the adults. The latest collections of 
these birds in the Marianas were made by the Japanese. Yama- 
shina (1932:411) obtained eggs in 1931 at Pagan and Agrihan, and 
again in 1940. He (1940:679) reported birds at Assongsong 
(Asuncion). Takatsukasa (1932:12) says, "A collector, working 
for Marquis Yamashina and myself, lately procured many speci- 
mens in Saipan and Pagan." Linsley (1935:249, 250) in search- 
ing for the megapode at Guam found little evidence of the birds. 
He interviewed people between the ages of forty-five and eighty and 
only two or three remembered seeing the bird. He said he saw one 
or two cross the road; but I suspect that they might have been rails 
{Rallus owstoni). Service personnel stationed at various islands in 
the Marianas during the late war have not reported the birds. The 
NAMRU2 party found no trace of the bird at Guam or Rota. Joe 
T. Marshall, Jr. (1949:203), did not find the bird at Saipan, Tin- 



112 University of Kansas Publs., Mus. Nat. Hist. 

ian, or Guam in 1945. Its status on Agiguan is unknown; isolated 
Japanese troops present on this small island from the time of the 
American invasion (1944) until the armistice (1945) may have used 
the birds for food and depleted the population seriously. At present 
the birds apparently still occur on islands in the northern Marianas. 
It seems that if these birds are to survive, they must be given some 
protection. 

Evolutionary history. — The genus Megapodius consists, according 
to Peters (1934:1-7), of nine species which are distributed through 
the islands from the Philippines and Borneo to Australia and Mela- 
nesia. These have been redesignated under three specific names by 
Mayr (1938). Outlying forms occur in the Nicobar Group to the 
west and in Tonga (Niuafou Island) in the east and in the Palaus 
and Marianas to the extreme northeast. Lister (1911:757) is of 
the opinion that the megapodes may have reached these outlying 
islands by having been transported by the natives, by whom the 
eggs were highly valued as food. This idea is also maintained by 
Rutland (1896:29-30) and Christian (1926:260). Possibility and 
not factual evidence support this hypothesis. From their seeming 
ancestral stocks, M. pritchardii Gray of Niuafou Island and M. 
laperouse of Micronesia are remarkably distinct which may indicate 
their early arrival at these islands and subsequent change from their 
ancestral stocks. 

Like M. pritchardii, the Micronesian species is smaller than its 
relatives to the southwest and has short, rather rounded wings, al- 
though its feet are heavily built whereas those of M. pritchardii are 
lightly constructed. In comparing these birds with the species of 
megapode found in the Philippines, Celebes and Melanesia, it seems 
that both M. prichardii and M. Laperouse are closely related to the 
widespread species, M. freycinet, which may have been ancestral to 
both. The differences between M. prichardii and M. laperouse indi- 
cate that they represent independent invasions. Nevertheless these 
megapodes may have had a wider range in Oceania in former times; 
man may have eliminated the birds from some islands by using their 
eggs. The eggs are laid in conspicuous mounds which are easily found 
by man. 

M. laperouse differs from il/. jreycinet of New Guinea and other 
parts of Melanesia and the Philippines; its small size, short wing and 
pearl gray head are distinctive characters. It shows greatest resem- 
blance to the subspecies in Celebes iM. f. gilberti) in size and to the 
subspecies in the Moluccas (M. /. freycinet) in coloring; possibly M. 



Baker: The Avifauna of Micronesia 113 

laperouse represents stock from one of these regions. Apparently the 
group as a whole evolved from a center of dispersal in the New 
Guinea area. Mayr (1942b:167) regarded all the species of Megapo- 
dius as belonging to one polytypic species, except M. laperouse and 
M. pritchardii, which are allopatric species. 

Coturnix chinensis lineata (Scopoli) 
Painted Quail 

Oriolus lineatus Scopoli, Del. Flor. et Faun. Insubr., fasc. 2, 1786, p. 87. (Type 
locality, Luzon, ex Sonnerat.) 

Excalfactoria sinensis Hartert, Novit. Zool., 5, 1898, p. 61 (Guam); Seale, Occ. 
Papers Bernice P. Bishop Mus., 1, 1901, p. 37 (Guam); Safford, Osprey, 1902, p. 68 
(Guam); idem, Amer. Anthro., 4, 1902, p. 711 (Guam); idem. The Plant World, 7, 
1904, p. 265 (Guam); idem, Contr. U. S. Nat. Herb., 9, 1905, p. 78 (Guam); Cox, 
Island of Guam, 1917, p. 21 (Guam); Nelson, Proc. 1st Pan-Pacific Sci. Conf., 1921, 
p. 273 (Guam). 

Excalfactoria chinensis lineata Wetmore, in Townsend and Wetmore, Bull. Mus. 
Comp. Zool., 63, 1919, p. 176 (Guam); Kuroda, in Momiyama, Birds Micronesia, 1922, 
p. 41 (Guam); Mathews, Syst. Avium Australasianarum, 1, 1927, p. 20 (Marianne); 
Hand-list Japanese Birds, rev., 1932, p. 198 (Guam); Peters, Check-list Birds World, 
2, 1934, p. 96 (Guam); Brj'an, Guam. Rec, vol. 13, no. 2, 1936, p. 15 (Guam); Hand- 
list Japanese Birds, 3d ed., 1942, p. 223 (Guam); Mayr, Birds Southwest Pacific, 1945, 
p. 287 (Guam). 

Excalfactoria chinensis Strophlet, Auk, 1946, p. 536 (Guam). 

Coturnix chinensis lineata Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 47 
(Guam). 

Geographic range. — Philippinos and parts of Malaysia. In Micronesia: Mari- 
ana Islands — Guam (introduced). 

Characters. — Adult: A small quail with upper parts brown splotched with 
black and streaked with buff; males with face and throat black surrounded by 
white line, upper breast blue gray, lower breast, belly and under tail-coverts and 
tail near "burnt sienna"; females lighter than males, underparts pale brown, 
mottled with blackish on breast and sides of body; bill dark lead colored, feet 
yellow. 

Measurements. — Three adult males from Guam measure: wing, 66, 67, 67; 
culmen, 9.2, 10.0, 10.3; tarsus, 18.1, 18.7, 22.6. 

Weights. — Two adult males taken by NAMRU2 at Guam weigh 34.5 and' 
35.5 grams. 

Specimens examined. — Total number, 3 males from Mariana Islands, USNM — Guam (Feb. 
24, June 13, 28). 

Remarks. — Seale (1901:37) writes that the Painted Quail was 
introduced to Guam from Manila, or the island of Luzon in the Phil- 
ippine Islands, by Captain Pedro Duarty of the Spanish Army in 
1894. It was a successful introduction; the bird is well adapted to the 
grasslands, open hillsides, and fallow rice paddies. The bird appears 
to offer no serious competition to native species, because there are 
few native birds which depend largely on this habitat. The 
NAMRU2 party obtained specimens at Mt. Santa Rosa and near 
Agat ; others were seen as singles or pairs near Umatac and on Mount 

8—8131 



114 University of Kansas Publs., Mus. Nat. Hist. 

Tenjo. Strophlet (1946:536) observed the birds in the southern part 
of Guam in 1945. He found them as singles or pairs in the months of 
September, October and November. Wilfred Crabb reported a covey 
of seven birds in June, 1945. Two males taken in June had enlarged 
testes. Seale (1901:37) obtained a nest of seven eggs. 

Gallus gallus (Linnaeus) 
Red Jungle Fowl 

Phasianus Gallus Linnaeus, Syst. Nat., ed. 10, 1, 1758, p. 158. (Type locality, 
"India orientali, Pouli condor etc.," restricted to Pulo Condor, off noouths of the 
Mekong.) 

Phasianus Gallus Kittlitz, Obser. Zool., in Lutke, Voy. "Le Seniavine," 3, 1836, 
p. 284 (Ualan = Kusaie). 

Gallus bankiva Hartlaub and Finsch, Proc. Zool. Soc. London, 1872, pp. 89, 103 
(Pelew); Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 5, 29 (Palau) ; Schmeltz and 
Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, pp. 281, 298, 353 (Ponape, Mortlock, 
Ruk); Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), 
p. 59 (Pelew, Caroline, Marianne, Marshall); Oustalet, Nouv. Arch. Mus. Hist. Nat., 
Paris, (3), 8, 1896, p. 25 (Saypan, Palaos, Marshall); Hartert, Novit. Zool., 5, 1898, 
p. 61 (Saipan); Seale, Occ. Papers Bernice P. Bishop Mus., 1. 1901, p. 38 (Marianas); 
Safford, Osprey, 1902, p. 70 (Marianas). 

Gallus ferrugineus Finsch, Proc. Zool. Soc. London, 1877 (1878), p. 780 (Ponape); 
idem. Ibis, 1881, p. 114 (Ponap6, Kushai). 

Gallus gallus bankiva Kuroda, in Moniyania, Birds Micronesia, 1922, p. 41 (Saipan, 
Pelew, Ponape, Marshall). 

Gallus gallus Mathews, Syst. Avium Australasianarum, 1, 1927, p. 21 (Micronesia); 
Cram, Bull. U. S. Nat. Mus., 140, 1927, pp. 238, 328 (Guam); Bequaert, Mushi, 12, 
1939, p. 81 (Kusaie) ; idem, Occ. Papers Bemice P. Bishop Mus., 16, 1941, p. 266 
(Kusaie); Mayr, Birds Southwest Pacific, 1945, pp. 57, 286 (Marianas, Carolines, 
Palaus); Wharton and Hardcastle, Journ. Parasitology, 32, 1946, pp. 294, 310 (Ulithi, 
Garakayo); Stott, Auk, 1947, p. 525 (Saipan). 

Gallus gallus domesticus Hand-list Japanese Birds, rev., 1932, p. 198 (Marianas, 
Palaus, Carolines, Marshalls). 

Gallus gallus micronesiae Hachisuka, Tori, 10, 1939 (1940), p. 600 (Type locality, 
Truk, also from Pelew, Rota, Yap, Ponape) ; Hand-list Japanese Birds, 3d ed., 1942, p. 
222 (Saipan, Rota, Babelthuap, Koror, Yap, Truk, Lukunor, Ponape, Kusaie, Mar- 
shalls). 

Gallus gallus gallus Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 47 
(Peleliu, Ngabad, Garakayo, Ulithi, Truk). 

Geographic range. — Southeastern Asia and Malaysia; introduced into many 
islands of Oceana. In Micronesia: Mariana Islands — Saipan, Rota; Palau 
Islands — Kayangel, Babelthuap, Koror, Garakayo, Peleliu, Ngabad, Angaur; 
Caroline Islands — Ulithi, Yap, Truk, Lukunor, Ponape, Kusaie; Marshall Is- 
lands — exact locality not known. 

Specimens examined. — Total number, 3 (1 male, 2 females) as follows : Palau Islands, 
USNM— Garakayo, 1 (Sept. 19) — Peleliu, 1 (Sept. 13)— Ngabad, 1 (Sept. 11). 

Parasites.— Cram (1927:238, 328) found the round worms (Nematoda), 
Dispharnyx nas-uta and Oxyspirura rnansoni in birds from Guam. Bequaert 
(1939:81 and 1941:266) found the fly (Hippoboscidae) Ornithoctona plicata, 
Dn fowl from Kusaie. Wharton and Hardcastle (1946:294, 310) obtained the 
chiggers (Acarina), Neoschongastia yeomansi and N. exvingi from fowl at 
Ulithi and Garakayo. 

Remarks. — The Red Jungle Fowl has been introduced in Micro- 



Baker: The Avifauna of Micronesia 115 

nesia, as it has been in other parts of Oceania. It is found on many 
of the islands of Micronesia, including the volcanic islands and the 
atolls. The NAMRU2 party did not find feral fowl at Guam but 
found the wary birds at Ulithi and in the Palaus. The birds at 
Ulithi were small and of a mixed breed. At Palau some fine ex- 
amples of typical jungle fowl were observed. Coultas obtained simi- 
lar specimens at Ponape and Kusaie. The natives have apparently 
allowed these birds to go wild, but catch them for food. These wild 
stocks may represent the earlier "liberations" while domestic fowl 
kept by natives at present appear to include several other breeds 
probably obtained from Europeans. 

The committee that prepared the Hand-list of Japanese Birds 
(Hachisuka et al., 1942:222) points out that although many orni- 
thologists believe the Red Jungle Fowl to be introduced in Micro- 
nesia and other parts of Oceania, it is their opinion (based on a series 
of more than 100 skins before them) that the population in Micro- 
nesia is racially distinct. They further comment, as did Hachisuka 
(1939b: 600), that one may find hybrids between these birds and the 
domestic fowl belonging to the native peoples; this is commonly seen 
on the more populated islands such as Koror and Saipan. I have no 
doubt that these skins show distinct features; nevertheless, I am 
reluctant to recognize these by subspecific name, since the birds may 
be a mixture of domestic strains introduced by man at different times 
after the jungle fowl was first brought by the early Micronesians. It 
seems that the production of hybrids between the feral and domestic 
fowl, which we find there today, may have been going on ever since 
the European colonists arrived with their fancy breeds of chickens. 

Phasianus colchicus Linnaeus 
Ring-necked Pheasant 

Phasianus colchicus Linnaeus, Syst. Nat., ed. 10, 1, 1758, p. 158. (Type locality, 
Africa, Asia = Rion.) 

Phasianus torquatus Baker, Sniithson. Misc. Coll., vol. 107, no. 15, 1948, p. 47 
(Guam). 

Geographic range. — Eastern China and northeastern Tonkin. Widely intro- 
duced into North America, Europe, and New Zealand. In Micronesia: Mari- 
ana Islands — Guam (introduced). 

Remarks. — On July 4, 1945, fifty-seven Ring-necked Pheasants 
(sixteen cocks and forty-one hens) were liberated at Guam by per- 
sonnel of the U. S. Navy. The birds were eleven weeks old when 
released, having been brought by plane from the hatcheries of the 
State Division of Game and Fish in California. Twenty-four birds 
were liberated at the site of CincPoa headquarters near Mt. Tenjo. 



116 Univeesity of Kansas Publs., Mrs. Nat. Hist. 

Thirty-three were placed near the FEA dairy farm, approximately 
one and one-fourth miles west of Price School. One month after 
release the birds were present at the liberation sites, although there 
were reports that some had drifted as far away as a mile or more. 
The birds were not banded. This liberation has been reported on by 
Quinn (1946:32-33) and by the author (1946b :211 and 1948:47). 
In using the name P. colchicus, I am following Delacour (in McAtee, 
1945: 8) and the twenty-third supplement to the American Ornithol- 
ogists' Union check-list of North American birds (Auk, 65, 1948: 
440). 

Rallus philippensis pelewensis (Mayr) 

Banded Rail 

Hypotaenidia -philippensis pelewensis Mayr, Amer. Mus. Novit., no. 609, 1933, p. 3. 
(Type locality, Palau Islands.) 

Rallus philippensis Hartlaub, Proc. Zool. Soc. London, 1867 (1868), p. 831 (Pelew) ; 
Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 5, 37 (Palau); idem, Proc. Zool. Soc. 
London, 1877, p. 587 (Palau); Wiglesworth (part), Abhandl. und Ber. Zool. Mus. 
Dresden, no. 6, 1890-1891 (1891), p. 59 (Pelew); Finsch, Deut. Ver. zum Schulze der 
Vogelwelt, 18, 1893, p. 459, Palau). 

Rallus pectoralis Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 8, 117, 
118 (Pelew); idem, Proc. Zool. Soc. London, 1872, pp. 89, 107 (Pelew). 

Eulabeornis forsteri Gray (part). Hand-list Birds, 3, 1871, p. 57 (Pelew). 

Hypotaenidia philippensis Salvador! (part), Omith. Papuasia, 3, 1882, p. 261 
(Pelew); Sharps (part), Cat. Birds British Mus., 23, 1894, p. 39 (Pelew); Kuroda, 
in Momiyama, Birds Micronesia, 1922, p. 42 (Pelew). 

Eulabeornis philippensis? Mathews, Birds Australia, 1, 1910-1911, p. 199 (Pelew). 

Hypotaenidia philippinensis philippensis Hand-list Japanese Birds, rev., 1932, p. 
196 (Palau). 

Rallus philippensis pelewensis Hand-list Japanese Birds, 3d ed., 1942, p. 220 
(Babelthuap, Koror) ; Mayr, Birds Southwest Pacific, 1945, p. 287 (Palau); Baker, 
Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 48 (Peleliu, Garakayo). 

Geographic range. — Micronesia: Palau Islands — Babelthuap, Koror, Araka- 
besan, Garakayo, Peleliu, Angaur. 

Characters. — Adult: A large, slender rail with black crown streaked with 
brown; .superciliary stripe ashy-gray, lighter toward bill; eye stripe brown 
becoming more rufous behind eye and on nape; chin a.shy-gray; throat near 
"mouse gray" tinged with olive especially toward breast; breast, belly and 
sides barred with black and white, with a broad "tawny" band on breast; 
posterior part of belly and vent buffy with some barring; under tail-coverts 
barred with black, white, and buff; mantle black with feathers subtcrminally 
barred with white; back, scapulars, inner wing-coverts, and rump black with 
white spotting and feathers edged with olive brown; outer wing-coverts, sec- 
ondaries, and primaries barred with black and rufous with some buffy-white 
on outer webs; under wing barred black and white with some brownish mark- 
ings; tail with both bars and blotches of black, white, and buffy -rufous; max- 
illa horn-colored; mandible yellowi.?h; feet light brown. 

R. p. pelewensis resembles R. p. philippensis Linnaeus of the Philippines, 
but is darker with nape more rufous-brown ; upper parts marked with nar- 
rower and darker edgings to feathers and with pronounced whitish spotting. 



Baker: The Avifauna of Micronesia 117 

Resembles R. p. chandleri (Mathews) of Celebes, but with wing shorter; more 
pronounced band on breast; bird darker above and below; rump and upper 
tail-coverts less spotted. 

Measurements. — Specimens in the collection of the United States National 
Museum measure as follows: four adult males — wing, 130-134 (132); tail, 59- 
63 (61) ; full culmen, 30-37 (34) ; tarsus, 38-45 (43) ; four adult females— wing, 
125-130 (127); tail, 54-61 (58); full culmen, 29-35 (32); tarsus, 38-42 (40). 
Mayr (1933c :4) lists the following measurements: twelve adult males — 127- 
143 (134.6); tail, 54-65 (60); exposed bill, 25-28 (27.7); tarsus, 41-46 (43.5); 
three adult females— wing, 129, 136, 136; tail, 56, 57, 58; exposed bill, 23, 24, 
25; tarsus, 40, 41, 42. 

Specimens examined. — Total number, 27 (18 males, 9 females), as follows: Palau Islands, 
USNM— Garakayo, 4 (Sept. 18, 19, 20)— Peleliu, 4 (Aug. 27, 28, Sept. 16)— Arakabesan, 1 
(Nov. 26); AMNH — exact locality not given, 18 (Oct., Nov., Dec). 

Nesting. — The condition of the gonads in specimens obtained indicates that 
the breeding season is principally in the fall and winter. Of adult rails taken 
by Coultas in October, November and December, 1931, 6 of 12 males and 3 
or 4 females had enlarged gonads. In September, 1945, the NAMRU2 party 
obtained two adult males with swollen testes. Marshall (1949:219) recorded 
breeding in September and November. 

Food habits. — Stomachs of rails obtained by the NAMRU2 party contained 
insects, seeds and small mollusks. Coultas (field notes) notes that the birds 
eat snails, roots and other vegetable matter. 

Remarks. — Ralhis philippensis is geographically widespread, being 
found from Tasmania and Australia north to Malaysia and the Phil- 
ippines west to Cocos Keeling Island east to ^Melanesia and western 
Polynesia and north to the Palau Islands. The species is divisible 
into several subspecies. The one in the Palaus, although distinctive, 
does not appear to have undergone a higher degree of differentiation 
(even though isolated as a small population) than any of the sub- 
species in Malaysia or Melanesia. Perhaps the form on Palau as 
well as the relatively undifferentiated Poliolimnas cinereus are rather 
recent invaders of Micronesia, as compared with Ralln^ owstoni and 
Ayhanolimnas monasa. 

The Banded Rail is less secretive in habits than Rallus owstoni of 
Guam, and neither was seen to fly. At Angaur, Peleliu and Gara- 
kayo, the NAMRU2 party found the rail in areas of swamp and 
marsh as well as in the rocky uplands; it probably prefers the former 
habitats. Several rails were observed and shot in open places, but 
they probably prefer to remain in dense cover. Coultas found the 
birds at taro patches and swamps. I watched a rail feeding along an 
open trail on Garakayo. The bird was eating small mollusks and 
other items which were in the open area. Being a true skulker, the 
bird would make a quick dash to the feeding place, remain only a 



118 University of Kansas Publs., Mus. Nat. Hist. 

few moments, hurriedly return to the protective cover, and then 
repeat the process. The best means that I found of obtaining these 
birds was using traps baited with peanut butter and oatmeal. The 
trape had to be visited frequently or the ants made short work of the 
captured birds. 

Rallus owstoni (Rothschild) 

Guam Rail 

Hypotaenidia owstoni Rothschild, Novit. Zool., 2, 1895, p. 481. (Type locality, 
Guam.) 

f Rallus philippinus Gray, Cat. Birds Trop. Is. Pacific Ocean, 1859, p. 51 (Marian or 
Ladrone Is.). 

Rallus pectoralis Finsch and Hartlaub, Fauna Centralpolynesiens, 1867, p. 157 
(Guam). 

Eulabeornis forsteri Gray (part), Hand-list Birds, 3, 1871, p. 57 (Marian). 

Hypotaenidia philippensis Pelzeln, Ibis, 1873, p. 41 (Marianne Isl.); Salvadori (part), 
Ornith. Papuasia, 3, 1882, p. 261 (Marianas); Sharpe (part). Cat. Birds British Mus., 
23, 1894, p. 39 (Guam). 

Rallus philippinus Wiglesworth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 
1890-1891 (1891), p. 59 (Guam). 

Hypotaenidia owstoni Hartert, Novit. Zool., 5, 1898, p. 62 (Guam); Safford, Osprey, 
1902, pp. 41, 67 (Guam); idem, The Plant World, 7, 1904, p. 265 (Guam); Dubois, 
Syn. Avium, 2, 1904, p. 961 (Mariannes); Safford, Contr. U. S. Nat. Herb., 9, 1905, 
p. 79 (Guam) ; Cox, Island of Guam, 1917, p. 21 (Guam) ; Kuroda, in Momiyama, 
Birds Micronesia, 1922, p. 42 (Guam); Hartert, Novit. Zool., 34, 1927, p. 22 (Guam); 
Mathews, Syst. Avium Australasianarum, 1, 1927, p. 84 (Guam); Hand-list Japanese 
Birds, rev., 1932, p. 196 (Guam). 

Hypotaenidia marchei Oustalet, Nouv. Arch. Mus. Hist. Nat., Paris, (3), 8, 1896, 
p. 32 (Type locality, Guam). 

Hypotaenidia oustini Seale, Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 30 
(Guam). 

Rallus owstoni Peters, Check-list Birds Worid, 2, 1934, p. 166 (Guam); Bryan, 
Guam Rec, vol. 13, no. 2, 1936, p. 15 (Guam); Hand-list Japanese Birds, 3d ed., 
1942, p. 220 (Guam); Mayr, Birds Southwest Pacific, 1945, p. 287 (Guam); idem, 
Audubon Mag., 47, 1945, p. 279 (Guam); Watson, Raven, 17, 1946, p. 41 (Guam); 
Strophlet, Auk, 1940, p. 536 (Guam); Baker, Smithson. Misc. Coll., vol. 107, no 15, 
1948, p. 48 (Guam). 

Geographic range. — Micronesia: Mariana Islands — Guam. 

Characters. — Adult: A large rail with head, neck, and eye stripe near 
"mummy brown" with feathers on sides of neck tipped with "russet"; super- 
ciliary stripe to back of neck, throat and upper breast near "mouse gray"; 
mantle, back, scapulars, and some upper wing-coverts dark olive-brown be- 
coming browner on rump and upper tail-coverts; wings dark with brownish 
spots and barred with white ; lower breast, abdomen, under tail-coverts, and 
tail blackish with white barrings; bill lead colored; feet dark brown; tibia 
brown; iris red. 

Measurements. — Four adult males measure: wing, 120-123 (121); tail, 46- 
54 (50) ; full culmen, 37-43 (41) ; tarsus, 47-51 (50) ; si.x adult females measure: 
wing, 108-118 (112) ; tail, 38-46 (42) ; full culmen, 36-39 (37) ; tarsus, 43-47 (45). 

Weights. — The NAMRU2 party obtained specimens with the following 
weights: two adult males 256, 257; four females 147, 153, 210, 252 grams. 

Specimens examined. — Total number, 13 (5 males, 6 females, 2 unsexed), from Mariana 
Islands, USNM — Guam (Jan. 29, May 8, June 19, 20, 23, 28, 30, July 14, 19, 23, Sept. 8). 



Baker: The Avifauna of Micronesia 119 

Nesting. — A nest was found by McElroy of the NAMRU2 party at Guam 
on October 24, 1945, in dense grass on a hillside near Mount Santa Rosa. The 
nest contained three eggs, which the author (1948:48) describes as "white 
with a pinkish cast and a scattering of small spots of colors near 'russet' and 
near 'pear blue' which are concentrated at the large ends. They measure 37.5 
by 29.1, 39.1 by 28.0, and 40.7 by 29.0." Downey, black chicks were found on 
April 1, May 16, and May 26. M. Dale Arvey found a chick on August 2, 
1946, near Tumon Bay. A parent bird with young ones was seen near Merizo 
on October 2. A male taken on January 26 had enlarged gonads. Seale (1901: 
30) obtained a black chick in June or July. On the basis of the above obser- 
vations it seems that the nesting season extends from spring to fall, although 
Marshall (1949:219) assumes that this rail breeds the year around. 

Remarks. — The Guam Rail was first reported by Quoy and Gai- 
mard who called it "Rale tiklin," but was not described as new until 
1895 by Rothschild. It appears to be equally at home in upland 
grassy areas and in jungle areas. The species was not seen fre- 
quently by the NAMRU2 party, although birds were occasionally 
observed crossing the roads. Few birds were shot; most of the 
specimens were taken in rat traps, which may be the most satisfac- 
tory method of obtaining them. Coultas took his specimens with 
the aid of a dog. On June 19, 1945, a small patch of woodland was 
being removed by a bulldozer. Four rails, which were hiding in this 
thicket, were surrounded and three were captured by hand. These 
birds tried to escape over the cleared ground by running with wings 
flapping but made no effort to fly. I am inclined to believe, as the 
natives do,that these birds are virtually incapable of actual flight. 

The Guam Rail usually appeared to be a quiet bird, but at Tarague 
Point on July 12, 1945, 1 heard its loud penetrating cry ; it was a series 
of rapid screeches. At the same time rapid movement made consid- 
erable noise in the undercover. The bird making the call suddenly 
appeared, either rapidly chasing, or being chased by, another rail. 
The birds had abandoned their usual skulking habits and had little 
concern for the observer. I took this to be breeding behavior, com- 
parable to that of some of the North American rails during the 
mating period. 

The Guam Rail is probably not in serious danger of extermination. 
It is utilized by the natives as food; they capture the bird, using dogs 
and trail snares. Its skulking habits and ability to inhabit most 
types of cover on the island should insure its existence for a long time 
to come. 

Evolutionary history. — Rallus owstoni is endemic to the island of 
Guam with no closelv related forms nearby. It is one of the several 



120 



University of Kansas Publs., Mus. Nat. Hist. 



rails found in the Pacific which live on isolated islands. In com- 
parison with other species in the region, it has some resemblance to 
both R. torquatus and R. philippensis. In general, the underparts of 
R. owstoni resemble those of the R. philippensis group, although the 
upper parts resemble somewhat those of R. torquatus. Certain speci- 
mens of R. owstoni have a slight indication of a pale pectoral band. 




c wo looo 8000 Miles 



Po '■zonula 



O' PennulO 



Nesophylax 



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



The bill is shorter and heavier than that of R. torquatus, possibly- 
more like that of R. philippensis. The short rounded wing is a dis- 
tinctive character. The bird came from an ancestral stock possibly 
resembling R. philippensis and probably originated in the Philippine 
or Papuan areas. It may have invaded Micronesia at an early date 
and may have had a wider distribution in the islands in former times. 
Perhaps this same invasion resulted in the establishment of R. 
wakensis (Rothschild) at Wake. The supposed route of colonization 
is shown in figure 9. 

Rallina fasciata (Raffles) 
Malay Banded Crake 

Rallus fasciatus Raffles, Trans. Linn. Soc. London, 13, pt. 2, 1822, p. 328. (Typo 
locality, Benkulen, western Sumatra.) 

Rallina fasciata Hartlaub, Proc. Zool. Soc. London, 1867 (1868), p. 831 (Pelew) ; 
Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 7, 118 (Pelevv) ; idem, Proc. 
Zool. Soc. London, 1872, pp. 89, 106 (Pelew); Finsch, Journ. Mus. Godeffroy, 8, 1875, 
pp. 5, 37 (Palau); Salvador!, Oniith. Papuasia, 3, 1882, p. 264 (Pelew); Wiglesworth, 
Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 60 (Pelew); Sharps, 
Cat. Birds British Mus., 23, 1894, p. 75 (Pelew); Finsch, Deut. Ver. zum Sehulze der 



Baker: The Avifauna of Micronesia 121 

Vogelwelt, 18, 1893, p. 459 (Palau); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 
41 (Pelew); Mathews, Syst. Avium Australasianarum, 1, 1927, p. 88 (Pelew) ; Hand- 
list Japanese Birds, rev., 1932, p. 196 (Palau); Peters, Check-list Birds World, 2, 1934, 
p. 171 (Pelew); Hand-list Japanese Birds, 3d ed., 1942, p. 221 (Palau); Mayr, Birds 
Southwest Pacific, 1945, p. 287 (Palau); Delacour, Birds Malaysia, 1947, p. 77 (Palau). 

Geographic range. — Burma east and south to Malaysia and the Philippines. 
In Micronesia: Palau — exact locality unknown. 

Remarks. — The Malay Ban(ie(i Crake is known in the Palau Is- 
lands from birds taken by captains Tetens, Heinsohn, and Peters 
and by Kubary according to Finsch (1875: 37). It has not been 
taken by later collectors. Two unsexed and undated skins are in 
the collection of the American Museum of Natural History; these 
are from the Kubary collection. 

Rallina eurizonoides eurizonoides (Lafresna3'e) 
Philippine Banded Crake 

Gallinula eurizonoides Lafresnaye, Rev. Zool., 1845, p. 368. (No locality; the 
type agrees with specimens from the Philippine Islands.) 

Rallina eurizonoides eurizonoides Hand-list Japanese Birds, rev. 1932, p. 196 
(Koror); Hand-list Japanese Birds, 3d ed., 1942, p. 221 (Koror). 

Rallina eurizonoides subsp. Mayr, Birds Southwest Pacific, 1945, p. 302 (Palau). 

Geographic range. — Philippine Islands. In Micronesia: Palau Islands — 
Koror. 

Remarks. — This crake is apparently a straggler to western Micro- 
nesia from the Philippine area. 

Aphanolimnas monasa (Kittlitz) 

Kusaie Black Rail 

Rallus monasa Kittlitz, Denks. Riese russ. Amer. Micron, und Kamchat., 2, 1858, 
p. 30. (T>'pe locality, Kushai.) 

Rallus tabuennis? Kittlitz, Obser. Zool., in Lutk^, Voy. "Le Seniavine," 3, 1836, p. 
286 (Ualan). 

Ortygometra tabuensis Finsch, Joum. f. Ornith., 1880, pp. 297, 307 (Kuschai) ; 
idem. Ibis, 1881, pp. 106, 109 (Kushai); Wiglrsworth, Abhandl. und Ber. Zool. Mus. 
Dresden, no. 6, 1890-1891 (1891), p. 60 (Ualan). 

Kittlitzia monasa Hartlaub, Abhandl. nat. Ver. Bremen, 12, 1892, p. 391 (Ku- 
schai); Finsch, Mitth. Ornith. Ver. Wien, 17, 1893, p. 1 (Kuschai). 

Aphanolimnas monasa Sharpe, Bull. British Ornith. Club, 1892, p. 20 (Kuschai); 
Finsch, Deut. Ver. zum Schulze der Vogelwelt, 18, 1893, p. 457, pi. 4 (Ualan); Wigles- 
worth, Ibis, 1S93, p. 214 (Kushai); Sharpe, Cat Birds British Museum, 23, 1894, p. 
115 (Kushai); Matschie, Joum. f. Ornith., 1901, pp. 110, 113 (Ualan); Mathews, 
Syst. Avium Australasianarum, 1, 1927, p. 93 (Caroline Islands); Hand-list Japanese 
Birds, rev., 1932, p. 197 (Kusaie); Peters, Check-list Birds World, 2, 1934, p. 189 
(Kusaie); Hand-list Japanese Birds, 3d ed., 1942, p. 221 (Kusaie); Mayr, Birds 
Southwest Pacific, 1945, p. 288 (Kusaie); idem, Audubon Mag., 47, 1945, p. 280 
(Kusaie). 

Porzana tabuensis Sharpe, Cat. Birds British Mus., 23, 1894, p. Ill (Kushai). 

Pennula monasa Dubois, Syn. Avium, 2, 1904, p. 969 (Kuschai). 

Porzana tabuensis tabuensis Kuroda, in Momiyama, Birds Micronesia, 1922, p. 42 
(Kusaie). 

Geographic range. — Micronesia: Caroline Islands — Kusaie (probably ex- 
tinct). 



122 University of Kansas Publs., Mus. Nat. Hist. 

Characters. — Sharpe (1894:115) gives the following description: "Adult. 
Black with a bluish-grey reflexion; quills and tail somewhat browner; inner 
wing-coverts brownish with white spotting, outer edge of first primary dull 
brownish, chin and middle of the throat somewhat paler; bill blackish (Hart- 
laub.)." 

Remarks. — Two specimens of this rail are known. The two were 
taken by Kittlitz on his visit to Kusaie in December and January 
of 1827-'28. Coultas made a search for the bird in 1931 and failed 
to obtain it; he suggested that the high population of introduced 
rodents may have been a factor contributing to its extinction. The 
bird is considered to be extinct by the authors of the Hand-list of 
Japanese Birds (Hachisuka et al., 1942:221). 

The two known specimens are in Leningrad, and Mayr sent ex- 
amples of Porzana tabuensis there for comparison. The following 
is a translation of the letter received by Mayr from Boris Stegmann 
dated at Leningrad, December 7, 1937. 

"I have compared the two specimens of Porzana tabuensis with 
our specimens of Aphanolimnas monasa. The difference is in my 
opinion of generic value. Aphanolimnas is distinctly larger and 
more robust. The bill is not only absolutely but also relatively 
longer. Its length (measured from the forehead) reaches to the end 
of the second phalanx of the middle toe while it not nearly reaches 
it in tabuensis. The proportions of feet and toes are the same in 
both, but the feet are distinctly heavier in Aphanolimnas. The 
wings are relatively shorter in Aphanolimnas and the wing feathers 
are very soft. The wing is also much more rounded, the first pri- 
mary is about 21 mm. shorter than the wing tip. The tail consists 
of very soft loose feathers which resemble only distantly true tail 
feathers. It is therefore questionable whether this bird was at all 
able to fly. 

"The coloration is in general dull black, brownish black on head 
and wings, chin and upper throat are dark slate colored lighter in 
the middle. The under wing and tail-coverts are marked with 
scattered white spots (querflecken). The first primary has an 
irregular whitish brown margin on the outer web. The bill is dark 
and the feet yellowish." 

Possibly this rail represents an ancient colonization of Kusaie from 
an ancestral stock of Porzana in Polynesia. Mayr. (1941b: 203) is 
also of this opinion, and if this is true there is no close relationship 
between Aphanolimnas and the rails at Guam and Wake, Rallus 
owstoni and R. wakensis, which are probably colonizers from the 
Philippines or the Papuan area. Mayr (1943:46) remarks further 



Bakee: The Avifauna of Micronesia 123 

that the Hawaiian flightless rail [Peuula) is of doubtful taxonomic 
position, but may be related to the "Ap/iawohmnas-Porzanoidea- 
Nesophylax stock," although there is no evidence that Pennula is not 
related to Rallus. Supposed colonization routes are shown in figure 9. 

Poliolimnas cinereus micronesiae Hachisuka 
White-browed Rail 

Poliolimnas cinereus micronesiae Hachisuka, Bull. British Ornith. Club, 59, 1939, 
p. 151. (Type locality. Yap.) 

Ortygometra quadristrigata Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 
8, 118 (Pelew); idem, Proc. Zool. Soc. London, 1872, pp. 90, 107 (Pelew, Uap). 

Ortygometra cinerea Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 5, 38 (Palau, 
Yap); idem, Proc. Zool. Soc. London, 1880, p. 577 (Ruk); Salvadori, Ornith. Papuasia, 
3, 1882, p. 273 (Yap, Pelew); Wiglesworth, Abhandl. and Ber. Zool. Mus. Dresden, 
no. 6, 1890-1891 (1891), p. 61 (Pelew, Yap, Ruk); Finsch, Deut. Ver. zum Schulze 
der Vogelwelt, 18, 1893, p. 459 (Palau). 

Ortygometra cinerea = quadristrigata Schmeltz and Krause, Ethnogr. Abth. Mus. 
Godeffroy, 1881, p. 353 (Ruk). 

Poliolimnas cinereus Sharpe, Cat. Birds British Mus., 23, 1894, p. 130 (Pelew, Yap, 
Ruk); Hartert, Novit. Zool., 5, 1898, p. 64 (Guam); idem, Novit. Zool., 7, 1900, p. 9 
(Ruk); Scale, Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 30 (Guam); Safford, 
Osprey, 1902, p. 67 (Mariannes); idem. The Plant World, 7, 1904, p. 265 (Guam); 
idem, Contr. U. S. Nat. Herb., 9, 1905, p. 79 (Guam) ; Cox, Island of Guam, 1917, p. 
21 (Guam); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 42 (Guam, Pelew, Yap, 
Ruk). 

Porzana cinerea Stresemann, Novit. Zool., 21, 1914, p. 54 (Guam, Truk). 

Porzana cinerea ocularis Hartert, Novit. Zool., 31, 1924, p. 264 (Ruk, Guam). 

Poliolimnas cinereus collingwoodi Mathews, Syst. Avium Australasianarum, 1, 1927, 
p. 95 (Pelew, Marianne, Carolines); Hand-list Japanese Birds, rev., 1932, p. 197 (Guam, 
Koror, Yap, Truk); Hachisuka, Birds Philippine Islands, 1, 1932, p. 236 (Marianne, 
Pelew, Caroline); Peters, Check-list Birds World, 2, 1934, p. 198 (Marianne, Caroline, 
Pelew); Bryan, Guam Rev., vol. 13, no. 2, 1936, p. 15 (Guam); Mayr, Birds Southwest 
Pacific, 1945, p. 288 (Guam, Palau, Yap, Truk, Bikini) ; Delacour and Mayr, Birds 
Philippines, 1946, p. 64 (Micronesia); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 
1948, p. 48 (Ulithi?, Truk). 

Porzana cinerea collingwoodi Rensch, Mitt. Zool., 1931, p. 468 (Marianne, Karolinen, 
Palau). 

Poliolimnas cinereus micronesiae Yamashina, Tori, 10, 1940, p. 679 (Bikini); Hand- 
list Japanese Birds, 3d ed., 1942, p. 221 (Guam, Babelthuap, Koror, Yap, Truk, Bikini). 

Geographic range. — Micronesia: Mariana Islands — Guam; Palau Islands — 
Koror, Babelthuap; Caroline Islands — Yap, Ulithi?, Truk; Marshall lislands — 
Bikini. 

Characters. — Adult: A slightly built, long-legged rail with forehead and 
anterior crown light gray with darker, slate-colored feather shafts; color more 
olive-brown on occiput and nape; eyestripe dark slate extending to occiput; 
superciliary from bill to eye, and stripe below eye, white; chin and throat ashy- 
white; sides of head, neck and breast ashy-gray, lighter on breast and whitish 
on abdomen; sides of abdomen ashy-brown becoming more buffy on tibia and 
under tail-coverts; mantle olive-colored becoming lighter and more brownish 
on back, rump, and scapulars; wing-coverts similar in color but feathers with 
broad dark brown shaft-marks; wings brown, first primary with whitish outer 
web; under wing gray with some lighter streaks; tail dark brown, lighter on 
edges; bill horn colored, tan below; feet brown; iris vermillion. 

Immature : Resembels adult, but head more rufous, upper parts marked with 
buffy rufous; eye stripe light rufous-brown; underparts tinged with rufous. 



124 



University of Kansas Publs., Mus. Nat. Hist. 



P. c. micronesiae differs from P. c. collingwoodi Mathews of the PhiHppines 
by having more pale gray and less olivaceous-brown on the nape and shoulder; 
darker on the under tail-coverts; and having a shorter culmen. P. c. brevipes 
(Ingram) of the Volcano Islands differs from P. c. micronesiae by being paler 
on upper parts, particularly back and wing-coverts and more washed with buff 
below; by having a shorter, thicker culmen; and by having a shorter tarsus. 

Measurements. — Measurements are shown in table 17. 

Table 17. Measurements of Three Subspecies of Poliolimnas cinereus 



Locality 


No. 


Wing 


Tail 


Culmen 


Tarsus 


Poliolimnas cinereus 
collingwoodi 
Philippines, Talaut. Celebes 

Poliolimnaff cinereus 
micronesiae 
Guam 


13 

10 

10 

5 

8 


98 
92 108 

95 
91-102 

93 
89-95 

95 
94-97 

96 
94-97 


51 
50-53 

f)l 
51-53 

51 
51-53 


22.5 
21.0-24.0 

21.0 
20.0-22.5 

21.0 
20.0-23. 

21.0 
20.5-22. 

19.0 
17.0 20.0 


38.0 
35.0-41.0 

37.0 


Palau 


34.5-39.0 
37.0 


Truk 


34.0-38.0 
36.0 


Poliolimnas cinereus 
brevipes 
S. Dionisio Island 


35.0-37.0 

30.0 
29.0-32.0 



Specimens examined. — Total number, 25 (11 males, 13 females, 1 unsexed), as follows: 
Mariana Islands, AMNH — Guam, 10 (July 13, Aug. 1, 5, 7, 13, 19, 23, 31); Palau Islands, 
AMNH — exact locality not given, 10 (Nov. 11, 13, 15, 23, 25); Caroline Islands, AMNH — 
Truk, 5 (June 3, 8, 16, 17, 18). 

Nesting. — Hartert (1900:9) describes two nests found on swampy ground. 
One contained three eggs, the other four eggs. He writes, "The eggs are pale 
buff, or cream-colour, speckled all over with brownish rufous, more frequently 
near the broad end. In some eggs, these spots are larger, in others minute, and 
there are often some, underlying pale purplish gray spot£." 

Remarks. — Superficially, the White-browned Rail of Micronesia 
is distinct from its near relative, P. c. collingwoodi, but the differ- 
ences are not so well marked as they are between insular populations 
of other species of rails. It is probably a comparatively recent addi- 
tion to the Micronesian avifauna, and its pattern of distribution may 
represent an early stage in the development of endemism in contrast 
to the pattern of later stages in the development of insular forms 
shown by the isolated rails, Rallus owstoni and Aphanolimnas mo- 



Bakek: The Avifauna of Micronesia 125 

nasa. The fact that Poliolimnas cinereiis is found only on widely 
separated islands in Micronesia does not necessarily mean that it has 
become "extinct" on the intervening islands, but that it may be par- 
tial to the larger, "high" islands, or that it is actually present but 
remains to be discovered on these intervening islands when more 
intensive field investigations are made. Hachisuka (1939a: 151), in 
naming the Micronesian form, comments that it has a shorter bill 
than P. c. colUngivoodi of the Philippines and Celebes, and that it is 
intermediate between this subspecies and P. c. brevipes of the Vol- 
cano Island to the north. Within these three subspecies there are 
trends toward a shorter culmen and shorter tarsus and, less markedly, 
toward a shorter wing. From the evidence at hand, it can be con- 
cluded that Poliolimnas first colonized Micronesia probably from the 
Philippine area (or Papuan area) through the Palaus and Carolines, 
to the Marianas and north to the Volcano Islands. Further, this has 
probably been a relatively recent invasion, although the subspecies 
in the Volcano Islands shows marked change in length of tarsus and 
culmen. This extension of range to the islands north of the Marianas 
is unusual and resembles somewhat the distribution of Nycticorax 
caledonicus in the same general area. 

The Micronesian White-browed Rail is a shy bird with the typical 
skulking habits of most rails. The NAMRU2 party did not find the 
bird at Guam, although reports were obtained that it was present in 
the marsh and swamp areas. Coultas (field notes) tells of observing 
the rail at Palau at a fresh water lake on Babelthuap, where it was 
difficult to obtain and apparently rare. Seale (1901:30) obtained a 
female specimen at Guam from native boys who snared it in a sweet 
potato patch near the Agafia River. This bird, taken in June or 
July, had eggs ready for laying. McElroy of the NAMRU2 party 
observed rails at Truk in brackish swamps, where he found them to 
be fairly common. A male which was taken in December had en- 
larged gonads. At Asor in the Ulithi Atoll, the NAMRU2 party 
learned that a small rail (possibly of this species) was found at taro 
patches in the early days of occupation, but that it was apparently 
eliminated by clearing operations. The taking of a bird at Bikini, 
as reported by Yamashina (1940:679), is further evidence that these 
birds may subsist on coral atolls as well as on the high volcanic 
islands; possibly the bird of the Marshalls may have been derived 
from the south rather than from the west. Unlike Rallus owstoni, 
this bird is apparently restricted to swampy areas, and may be elimi- 
nated from its habitat by drainage or clearing by man. It may 



126 University of Kansas Publs., Mus. Nat. Hist 



-•) 



always persist, however, in the taro patches maintained by the 
natives. 

Gallinula chloropus subsp. near orientalis Horsfield 

Gallinule 

Gallinula orientalis Horsfield, Trans. Linn. Soc. London, 13, 1821, p. 195. (Type 
locality, Java.) 

Gallinula chloropus indicus Hand-list Japanese Birds, rev., 1932, p. 197 (Babel- 
thuap); Takatsukasa and Yamashina, Dobutsu. Zasshi, 44, 1932, p. 266 (Pelew, 
Coror). 

Gallinula chloropus indica Hand-list Japanese Birds, 3d ed., 1942, p. 221 (Babel- 
thuap). 

Gallinula chloropus subsp. Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, 
p. 49 (Peleliu, Angaur). 

Geographic range. — Malaysia from southern Malay Peninsula to Celebes. 
In Micronesia: Palau Islands — Babelthuap, Koror, Peleliu, Angaur. 

Characters. — Adult: Resembles G. c. indica Blyth, G. c. lozanoi Lletget and 
G. c. guami Hartert, but smaller and paler; upper wing-coverts less olivaceous- 
brown and more slate-colored; back, rump, and scapulars less richly washed 
with olivaceous-brown. Resembles G. c. orientalis from Java in size, but much 
paler. 

Measurements. — An unsexed adult bird from Angaur measures: wing, 150; 

bill from rictes, 27.1; bill from nostril, 13.4; tarsus, 46. 

Specimens examined. — Total number, 3 (2 males, 1 unsexed) from Palau Islands, USNM 
— Angaur (Sept. 21). 

Remarks. — Owing to the lack of sufficient material, I am unable 
to determine the exact status of the resident gallinule in the Palau 
Islands. On the basis of a single, unsexed adult and two immatures 
there is not very much that can be said. The adult is smaller and 
paler than G. c. indica, G. c. lozonoi, and G. c. guami. It resembles 
specimens of the subspecies G. c. orientalis in size but is also paler 
than the skins of this race which I have examined. It seems closest 
to this latter subspecies to which I tentatively refer it. If it is 
closest to this subspecies, it probably reached Palau from the Cele- 
bean region, rather than from the Philippines or some other route. 
Whether specimens taken by the Japanese at Babelthuap and Koror 
are G. c. indica is questionable, unless the skins were from migrants 
which may visit Palau from the west or northwest. The Hand-list 
of Japanese Birds (Hachisuka et al., 1942:177) records G. c. indica 
from the Bonin Islands. 

The three Gallinules were taken by the NAMRU2 party at fresh 
and brackish water swamps at Angaur on September 21, 1945. Sev- 
eral Gallinules were seen in the area and several were observed also 
at Peleliu Island. One of the immatures was just growing its wing 
feathers, indicating that the birds must breed in the Palau Islands. 



Baker: The Avifauna of Micronesia 127 



Gallinula chloropus guami Hartert 
Gallinule 

Gallinula chloropus guami Hartert, Novit. ZooL, 24, 1917, p. 268. Type locality, 

Guam). 

Fulica chloropus Quoy and Gaimard, Voy. "Uranie," Zool., 1824, p. 703 (Guam); 
Kittlitz, Obser. Zool., in Lutke, Voy. "Le Seniavine," 8, 1836, p. 305 (Guahan). 

Gallinula galeata var. sandwichensis Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, 
(3), 8, 1896, p. 34 (Saypan, Tinian, Guam). 

Gallinula chloropus Hartert, Novit. Zool., 5, 1898, p. 62 (Guam); Scale, Occ. 
Papers Bernice P. Bishop Mus., 1, 1901, p. 31 (Guam); Safford, Osprey, 1902, p. 67 
(Marianas); idem, Amer. Anthro., 4, 1902, p. 711 (Guam); idem, The Plant World, 
7, 1904, p. 265 (Guam); idem, Contr. U. S. Nat. Herb., 9, 1905, p. 79 (Guam); 
Prowazek, Die deutschen Marianen, 1913, p. 101 (Marianen) ; Cox, Island of Guam, 
1917, p. 21 (Guam); Wetmore, in Townsend and Wetmore, Bull. Mus. Comp. Zool., 
63, 1919, p. 177 (Guam); Strophlet, Auk, 63, 1946, p. 536 (Guam). 

Gallinula chloropus guami Hartert, Vogel pal. Fauna, 15, 1921, p. 1843 (Guam); 
Kuroda, Avifauna Riu Kiu, 1925, p. 199 (Guam); Kuroda, in Momiyama, Birds Micro- 
nesia, 1922, p. 43 (Guam, Tinian, Saipan); Mathews, Syst. Avium Australasianarum, 1, 
1927, p. 99 (Mariana Islands); Takatsukasa and Yamashina, Dobutsu. Zasshi, 44, 1932, 
p. 226 (Pagan); Hand-list Japanese Birds, rev., 1932, p. 197 (Guam, Tinian, Saipan, 
Pagan); Hachisuka, Birds Philippine Islands, 1, 1932, p. 241 (Guam); Peters, Check- 
list Birds World, 2, 1934, p. 204 (Marianne Islands); Bryan, Guam. Rec, vol. 13, no. 2, 
1936, p. 15 (Guam); Hand-list Japanese Birds, 3d ed., 1942, p. 222 (Guam, Tinian, 
Saipan, Pagan); Mayr, Birds Southwest Pacific, 1945, p. 288 (Marianas); Downs, 
Trans. Kansas Acad. Sci., 49, 1946, p. 92 (Tinian); Stott, Auk, 1947, p. 525 (Saipan); 
Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 49 (Guam, Tinian, Saipan). 

Geographic range. — Micronesia: Mariana Islands — Pagan, Saipan, Tinian, 
Guam. 

Characters. — Adult: Head and neck sooty black; upper back dark, bluish 
slate-gray; lower back and wing-coverts brownish; tail blackish-brown; wings 
dark brown, outer edge of first primary white; breast and upper abdomen dark 
slate-gray, feathers on sides of breast with longitudinal white streak; under 
wing dark with white edges; lower abdomen grayish with white-tipped feathers; 
vent black; under tail-coverts white; bill and frontal shield red, tip of bill 
yellowish; legs and feet olive-green. 

Adult female : Resembles adult male but usually with smaller frontal shield. 

Immature: Resembles adult, but forehead mottled white and brown, with 
sides of head less distinctly speckled with brown ; crown, neck and upper back 
dusky brown; back, scapulars and upper tail-coverts olivaceous-brown; chin 
and throat whitish; breast feathers pearly-gray tipped with white; abdomen 
white; sides gray, washed with buff. Older birds are darker above and more 
brownish-gray below; frontal shield small. 

G. c. guami resembles G. c. indica, but upper wing-coverts darker and near 
"olivaceous black"; back, rump and scapulars darker and less olivaceous brown, 
although not so dark as in G. c. orientalis. From G. c. lozanoi, G. c. guami 
differs in: slightly darker upper wing-coverts; richer olivaceous-brown on back, 
scapulars and rump; thinner culmen with possibly less yellow coloring on tip. 
G. c. guami resembles G. c. sandvicensis Streets of the Hawaiian Islands, but 
has less olive wash on the feathers and a smaller frontal shield. 

Measurements. — Measurements of Gallinula chloropus are presented in table 
18. In general, females are smaller than males. 



128 University of Kansas Publs., Mus. Nat. Hist. 

Table 18. Mr-^surements of Gallinula chloropus 



Subspecies 



G. c. indica. 



G. c. orientalis. 



G. c. lozanoi. 



G. c. guami. 



G^ c. sandvicensis . 



No. 


Wing 


Bill 

from 

rictus 


Bill 

from 

nostril 


Tarsus 


15 


164 
158-173 


27 
24-29 


14.4 
13.1-18.1 


48 
44-50 


3 


152 
146-152 


27 
26-29 


13.8 
13.1-14.4 


45 
44-46 


11 


164 
153-170 


27 
24-29 


14.5 
13.1-15.2 


50 
45-57 


11 


164 
156-171 


27 

24-28 


14.7 
13.1-16.2 


49 
47-56 


2 


150-158 


27 


13.4 


52-56 



Weights. — From Guam an adult male weighed 291 grams and an adult female 
256 (Baker, 1948:49). 

Specimens examined. — Total number, 42 (16 males, 22 females, 4 unse.xed), as follows: 
Mariana Islands, USNM— Guam, 5 (Feb. 24, May, June 5, 7, 18— Tinian, 3 (Oct. 12, 18)— 
Saipan, 3 (Sept. 28, 30); AMNH— Guam, 25 (Feb. 21, April 6, July 13, 28, 30, Aug. 1, 3, 6, 7, 
13, 19, 23, 30, 31, Sept. 3, 17, Dec. 11— Tinian, 5 (June 11, Sept. 12, 13, 14). 

Nesting. — Hartert (1898:63) reports nests of the Gallinule at Guam in grass 
and on swampy ground in December and March. A male with enlarged gonads 
was taken by the NAMRU2 party at Guam on June 7. Marshall (1949:219) is 
of the opinion that this bird breeds all year. 

Food habits. — Seale (1901:31) found grass, insects, and larvae in stomachs 
obtained at Guam. 

Remarks. — The subspecies G. c. indica, G. c. lozanoi, G. c. guami, 
and G. c. sandvicensis bear a close resemblance to one another in size 
and color. G. c. guami and G. c. lozanoi resemble each other so 
closely that it would be difficult to separate unlabeled specimens of 
the two subspecies. G. c. orientalis differs from all of the gallinules 
in smaller size and darker color. Study of these forms indicates that 
the Gallinule has colonized the Marianas from Asia probably by way 
of Japan and the Bonin and Volcano islands. The Hawaiian sub- 
species is probably of American origin, as pointed out by Mayr 
(1943:46), and is not a close relative of the Mariana subspecies. 
The fact that these insular subspecies have not undergone much dif- 
ferentiation does not necessarily mean that they are recent arrivals, 
but probably is a reflection of the lack of plasticity of the species; as 
a whole the species does not exhibit anywhere a great amount of geo- 
graphic variaton. A thorough study of all insular populations of this 
species (including specimens from the Azores, Seychelles, Reunion, 



Baker: The Avifauna of Micronesia 129 

Mauritus, and the Greater and Lesser Antilles) might reveal the 
effect of isolation on the species in general. Its ability to become 
established on isolated islands is apparent, although it is indeed 
peculiar that the species has not reached the Caroline Islands. 

The Gallinule in the Marianas is restricted to fresh water lakes, 
marshes and swamps on the islands of Guam, Tinian, Saipan and 
Pagan. Coultas (field notes), on visiting the island of Tinian in 
1931, comments that the bird is rare and found at only one lake on 
the island. Downs (1946:92) noted the species in 1945, and Joe T. 
Marshall Jr. obtained three specimens at Lake Hagoya in October of 
the same year. Gleise (1945:220) estimated the population of 
Gallinules on Tinian in 1945 at 70 individuals. Stott (1947:525) 
reports that the birds were abundant at Lake Susupe, Saipan, in 
1945. Scale (1901:31) found the Gallinule to be abundant at Guam 
in marshes and taro patches. In 1945, the NAMRU2 party found 
fairly large populations of the Gallinule in fresh water marshes and 
fallow rice paddies at Guam. The greatest concentration of birds 
appeared to be in the Agaiia Swamp and along the Ylig River. They 
seldom ventured out into open water but preferred weedy edges into 
which they could suddenly dart when disturbed. It was interesting 
to note such wary behavior, for an observer would think that after 
the bird had been in an environment virtually devoid of birds of prey 
(except for an occasional migrant) for a number of generations, it 
would have lost such behaviorisms as a result of the absence of the 
selective processes involved in predation. 

Porphyrio porphyrio pelewensis Hartlaub and Finsch 
Purple Swamphen 

Porphyrio melanotus Temm. var. pelewensis Hartlaub and Finsch, Proc. Zool. Soc. 
London, 1872, p. 107. (Type locality, Pelew Islands.) 

Porphyrio melanotus Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 8, 
117, 118 (Pelew); Gray, Hand-list Birds, 3, 1871, p. 64 (Pelew). 

Porphyrio melanotus pelewensis Wiglesworth, Abhandl. und Ber. Zool. Mus. Dres- 
den, no. 6, 1890-1891 (1891), p. 61 (Pelew); Bolau, Mitteil. Naturhist. Mus. Ham- 
burg, 1898, p. 70 (Palau); Dubois, Syn. Avium, 2, 1904, p. 976 (Pelew); Mathews, 
Birds Australia, 1, 1911, p. 241 (Pelew); Kuroda, in Momiyama, Birds Micronesia, 
1922, p. 43 (Pelew); Mathews, Syst. Avium ,\ustralasianarum, 1, 1927, p. 100 
(Pelew) ; Hand-list Japanese Birds, rev., 1932, p. 197 (Palau) ; Hachisuka, Birds 
Philippines, 1, 1932, p. 245 (Pelew). 

Porphyrio pelewensis Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 5, 39 (Palau); 
Salvadori, Atti Accad. Sci. Torino, 14, 1879, p. 1169 (Pelew); Schmeltz and Krause, 
Ethnogr. Abth. Mus. Godeffroy, 1881, p. 407 (Palau); Finsch, Deut. Ver. zum 
Schulze der Vogelwelt, 18, 1893, p. 459 (Palau); Sharpe, Cat. Birds British Mus., 23, 
1894, p. 206 (Pelew); Nehrkorn, Nat. Eiers., 1899, p. 205 (Palau-lnseln) ; Matschie, 
Journ. f. Ornith., 1901, p. 113 (Palau); Reichenow, Die Vogel, 1, 1913, p. 216 (Palau- 
inseln); Takatsukasa and Kuroda, Tori, 1, 1915, p. 51 (Pelew). 

Porphyrio cyanocephalus Elliot, Stray Feathers, 7 1878, pp. 10, 13 (Palau). 

9—8131 



130 University of Kansas Publs., Mus. Nat. Hist. 

Porphyria poliocephaltis pelewensis Peters, Check-list Birds World, 2, 1934, p. 208 
(Pelew); Hand-list Japanese Birds, 3d ed., 1942, p. 222 (Koror). 

Porphyria porphyria pelewensis Mayr, Birds Southwest Pacific, 1945, p. 288 
(Palau); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 49 (Angaur). 

Geographic range. — Micronesia: Palau Islands — Koror, Angaur. 

Characters. — Adult: A large, purplish-blue, marsh bird with crown and sides 
of head dusky-black; wing-coverts purplish-blue; rest of upper parts dark, 
washed with olivaceous-brown ; outer webs of primaries and secondaries tinged 
with purplish-blue; chin, axillaries and under wing-coverts dusky; under tail- 
coverts whitish; rest of underparts purplish-blue, blacker on abdomen. 

Porphyrio p. pelewensis resembles P. p. palliatiis Bruggemann of Celebes 
and P. p. melanopterus Bonaparte of the Moluccas and New Guinea but upper 
parts paler and slightly less glossy; lesser and primary wing-coverts more 
purplsh-blue and less greenish-blue; outer webs of primaries and secondaries 
lighter purpHsh-blue ; underparts less blue with patch on throat and breast 
paler blue with less green (patch present on only one specimen from the 
Palaus). 

Measurements. — Measurements of one male: wing, 227; tail, 81; culmen 
and shield, 62.5; tarsus, 77; of three females: wing, 212, 218, 227; tail, 77, 81, 
86; culmen and shield, 57, 61, 64; tarsus, 75, 75, 77. 

Specimens examined. — Total number, 6 (1 male, 3 females, 2 unsexed), as follows: Palau 
Islands, USNM — Angaur, 1 chick (Sept. 21) AMNH — exact locality not given, 5 (Nov. 13, 
19, Dec. 17-19, undated). 

Nesting. — A black, downy chick was captured on September 21, 1&45, at 
the edge of a fresh-water lake on Angaur by Davidson of the NAMRU2 party 
(Baker, 1948:49). Two females taken by Coultas in December had enlarged 
gonads. 

Remarks. — The Purple Swamphen in the Palaus stands out as one 
of the more distinctive subspecies of P. porphyrio. It also marks 
the most northeastern extension of the range of this species. The 
subspecies in the Palaus shows affinities to that found to the south 
and southwest and probably reached Micronesia via the Papuan 
area, Celebes or the Moluccas rather than from the Philippines. 
It is interesting that this bird, as well as several other species, has 
been able to establish itself at the Palau Islands, but has not ex- 
tended its range farther into other islands of Micronesia. Perhaps, 
the bird is now in an early stage in its island occupation. 

The Purple Swamphen is probably not abundant in the Palaus, 
It is a large and conspicuous bird, and its restriction to swamps and 
areas around lakes may allow native hunters to obtain it rather 
easily, particularly by snares or by organized drives. Coultas (field 
notes) obtained specimens in taro swamps; he saw only 4 individ- 
uals and remarks that the birds utter harsh cries at night. The 
NAMRU2 party flushed an adult from lake side vegetation at An- 



Baker: The Avifauna of Micronesia 131 

gaur on September 21, 1945. This bird was not taken, but a downy- 
young was obtained in the area the same day. 

Fulica atra atra Linnaeus 
Common Coot 

Fulica atra Linnaeus, Syst. Nat., ed. 10, 1, 1758, p. 152. (Type locality, Europe, 
restricted to Sweden.) 

Fulica atra Hartert, Novit. Zool., 5, 1898, pp. 64, 69 (Guam); Seale, Ouc. Papers 
Bernice P. Bishop Mus., 1, 1901, p. 32 (Guam); Safford, Osprey, 1902, p. 70 (Mari- 
anas); idem. The Plant World, 7, 1904, p. 268 (Guam); Kuroda, in Momiyama, Birds 
Micronesia, 1922, p. 43 (Guam) ; Bryan, Guam Rec, vol. 13, no. 2, 1936, p. 15 
(Guam). 

Fulica atra atra Hartert, Vogel pal. Fauna, 15, 1921, p. 1852 (Guam); Hand-list 
Japanese Birds, rev., 1932, p. 197 (Tinian, Guam); Hand-list Japanese Birds, 3d ed., 
1942, p. 222, (Tinian, Guam); Mayr, Birds Southwest Pacific, 1945, p. 302 (Micro- 
nesia). 

Geographic range. — Breeds in Europe, northern Africa, and Asia. Winters 
south to Africa, Malaysia, southern Asia. In Micronesia: Mariana Islands — 
Tinian, Guam. 

Remarks. — The Common Coot is a straggler to Micronesia in win- 
ter. It has been recorded from Guam and Tinian. An unsexed speci- 
men in the collections of the American Museum of Natural History 
was taken at Guam in the fall of 1896 by one of Owston's collectors. 

Squatarola squatarola (Linnaeus) 
Black-bellied Plover 

Tringa Squatarola Linnaeus, Syst. Nat., ed. 10, 1, 1758, p. 149. (Type locality, 
Europe, restricted to Sweden.) 

Charadrius squatarola Hartert, Novit. Zool., 5, 1898, p. 66 (Saipan); Seale, Occ. 
Papers Bernice P. Bishop Mus., 1, 1901, p. 35 (Micronesia); Safford, Osprey, 1902, p. 
67 (Marianas). 

Squatarola squatarola Hartert, Novit. Zool., 7, 1900, p. 9 (Ruk); Safford, The Plant 
World, 7, 1904, p. 266 (Guam); idem, Contr. U. S. Nat. Herb., 9, 1905, p. 80 (Guam); 
Cox, Island of Guam, 1917, p. 22 (Guam); Ridg^vay, Bull. U. S. Nat. Mus., 50, pt. 8, 
1919, p. 72 (Ruk); Bryan, Guam Rec, vol. 13, no. 2, 1936, p. 15 (Guam); Hand-list 
Japanese Birds, 3d ed., 1942, p. 216 (Saipan, Truk) ; Mayr, Birds Southwest Pacific, 
1945, p. 36 (Truk); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 50 (Guam). 

Squatarola helvetica Takatsukasa and Kuroda, Tori, 1, 1915, p. 61 (Marianas, Ruk). 

Squatarola squatarola hypomelaena Kuroda, in Momiyama, Birds Micronesia, 1922, 
p. 43 (Ruk, Saipan); Hand-list Japanese Birds, rev., 1932, p. 193 (Saf^Jan, Truk). 

Geographic range. — Breeds in arctic regions of Holarctica. Winters in South- 
ern Hemisphere. In Micronesia: Mariana Islands — Guam, Saipan; Caroline 
Islands — Truk; Marshall Islands — Eniwetok. 

Specimens examined. — One female from Mariana Islands. USNM — Guam (Aug. 27). 

Remarks. — The Black-bellied Plover is an uncommon visitor to 
Micronesia. One bird was obtained by Markley of the NAMRU2 
party at Guam on August 27, 1945; Flavin recorded five of these 
birds from November, 1944, to January, 1946. Bryan and Green- 
way (1944:109) record this species as an occasional visitor to the 



132 University of Kansas Publs., Mus. Nat. Hist. 

Hawaiian Islands. Gleise and Genelly (1945:221) observed the 
Black-bellied Plover at Eniwetok in 1945. 

Pluvialis dominica fulva (Gmelin) 
Pacific Golden Plover 

Charadrius fulmis Gmelin, Syst. Nat., 1, pt. 2, 1789. p. 687. (Type locality, 
Tahiti.) 

Charadrius pluvialis Kittlitz, Obser. ZooL, in Lutke., Voy. "Le Seniavine," 3, 1836, 
pp. 287, 299, 304 (Ualan, Longounor, Guahan) ; idem, Denkw. Reise russ. Anier. 
Micron, und Kamchat., 2, 1858, pp. 32, 55 (Ualan). 

Charadrius virginianus Hartlaub, Joum. f. Ornith., 1854, p. 167 (Mariannen, Caro- 
linen). 

Charadrius longipes? Gray, Cat. Birds Trop. Is. Pacific Ocean, 1859, p. 47 (Ladrone 
or Marian Islands, Oualan). 

Pluvialis fulvus Schlegel, Mus. Pays-Bas, 6, no. 29, 1865, p. 52 (Micronesie). 

Charadrius fulvus Finsch and Hartlaub, Fauna Central -polynesiens, 1867, p. 196 
(Marianen, Ualan); Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 8, 117, 
118 (Pelews); Finsch and Hartlaub, Joum. f. Ornith., 1870, p. 139 (Pelew) ; Finsch, 
Journ. f. Ornith., 1872, p. 52 (Pelew, Carolinen) ; Hartlaub and Finsch, Proc. Zool. 
Soc, London, 1872, pp. 89, 104 (Pelew, Mackenzie, Uap) ; Graffe, Joum. Mus. Godef- 
froy, 2, 1873, p. 123 (Yap); Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 5, 31 (Pa- 
lau); idem, Journ. Mus. Godeffroy, 12, 1876, pp. 18, 38 (Ponape) ; idem. Proc. Zool. 
Soc. London, 1877 (1878), p. 781 (Ponape); idem., Proc. Zool. Soc. London, 1880, p. 
576 (Ruk); idem., Journ. f. Ornith., 1880, pp. 293, 305 (Ponape, Kuschai) ; idem. Ibis, 
1880, pp. 220, 331, 332 (Taluit) ; Schmeltz and Krause, Ethnogr. Abth. Mus. Godef- 
froy, 1881, pp. 281, 353 (Ponape, Ruk); Finsch, Ibis, 1881, pp. 105, 106, 109, 113, 
115 (Kushai, Ponape); Salvadori, Ornith. Papuasia, 3, 1882, p. 395 (Carolines, Pe- 
lews, Marianas) ; Finsch, Mitth. Ornith. Ver. Wien, 1884, p. 55 (Jaluit, Milli, Ku- 
schai) ; Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), 
p. 63 (Marshall Islands, Ualan, Luganor, Ponape, Ruk, Uap, Pelew, Marianne) ; 
Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 8, 1896, p. 46 (Guam, Hogoleu, 
Marshalls, Palaos) ; Hartert, Novit. Zool. 5, 1898, p. 66 (Guam); idem, Novit. Zoo!., 
7, 1900, p. 9 (Ruk); Scale, Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 36 (Mi- 
cronesia); Schnee, Ornith. Monatsber., 1901, p. 132 (Marshalls); Safford, Osprey, 
1902, p. 68 (Marianas); idem. The Plant World, 7, 1904, p. 266 (Guam); Schnee, 
Zool. Jahrbiicher, 20, 1904, p. 389 (Marsehall-Insein) ; Takatsukasa and Kuroda, Tori, 
1, 1915, p. 51 (Ponape). 

Charadrius doviinicus fulvus Safford, Contr. U. S. Nat. Herb., 9, 1905, p. 80 
(Guam); Cox, Island of Guam, 1917, p. 22 (Guam). 

Charadrius dominicus Sharpe, Cat. Birds British Mus., 24, 1896, p. 195 (Micro- 
nesia). 

Pluvialis dominicus fulvus Ridgway, Bull. U. S. Nat. Mus., 50, pt. 8, 1919, p. 89 
(Kuschai, Pelew, Ruk, Marianas, Mackenzie, Ponape) ; Wetmore, in Townsend and 
Wetmore, Bull. Mus. Comp. Zool., 63, 1919, p. 177 (Uala, Arhno, Rongelab); Kuroda, 
in Momiyama, Birds Micronesia, 1922, p. 44 (Guam, Angaur, Ualan, Luganor, Ponape, 
Ruk, Yap, Arhno) ; Hand-list Japanese Birds, 3d ed., 1942, p. 216 (Saipan, Tinian, 
Guam, Babelthuap, Koror, Peliliu, Angaur, Yap, Ulithi, Truk, Lukunor, Ponape, Ku- 
saie, Mille, Arhno, Majuro, Likieb). 

Pluvialis apricarius fulvus Hand-list Japanese Birds, rev., 1932, p. 193 (Saipan, 
Tinian, BabeJthuap, Koror, Pelilieu, Angaur, Yap, Uluthi, Truk, Lukunor, Ponape, 
Kusaie, Mille, Arhno, Majuro, Likieb). 

Pluvialis dominica fulva Peters, Check-list Birds World, 2, 1934, p. 244 (Oceania); 
Bryan, Guam, Rec, vol. 13, no. 2, 1936, p. 24 (Guam); Stickney, Amer. Mus. Novit., 
no. 1248, 1943, p. 3 (Saipan, Guam, Palau, Ponape, Kusaie, Ruk, Tarawa) ; Mayr, 
Birds Southwest Pacific, 1945, p. 39 (Oceania); Downs, Trans. Kansas Acad. Sci., 49, 
1946, p. 93 (Tinian); Strophlet, Auk, 1946, p. 536 (Guam); Borror, Auk, 1947, p. 
417 (Agrihan); Stott, Auk, 1947, p. 525 (Saipan); Baker, Smithson. Misc. Coll., vol. 
107, no. 15, 1948, p. 50 (Guam, Rota, Peleliu, Garakayo, Ulithi, Truk). 



Baker: The Avifauna of Micronesia 133 

Pluvialis dominica Wharton and Hardcastle, Journ. Parasitology, 32, 1946, pp. 306, 
310, 313, 316, 318 (Ulithi, Guam); Wharton, Ecol. Monogr., 16, 1946, pp. 174, 175 
(Guam). 

Geographic range. — Breeds from Siberia to western Alaska. Winters from 
India east to Oceania; stragglers occur west to Africa and east to Pacific coast 
of North America. In Micronesia: Mariana Islands — Guam, Rota, Tinian, 
Saipan, Agrihan, Asuncion; Palau Islands — Angaur, Peleliu, Ngabad, Gara- 
kayo, Koror, Babelthaup; Caroline Islands — Yap, Ulithi, Truk, Lukunor, Pon- 
ape, Kusaie; Marshall Islands — Mille, Arhno, Rongelab, Majuro, Likieb, 
Bikini. 

Specimens examined. — Total number, 69 (39 males, 26 females, 4 unsexed), as follows: 
Mariana Islands, USNM— Guam, 17 (July 8, 19, 24, Aug. 31, Sept. 4, 17, 19, 26, Oct. 5, 8, 
23, 24)— Rota, 5 (Oct. 20, 25); AMNH— Guam, 6 (Mar. 7, 8, 27, Aug. 15)— Saipan, 1 (Sept. 
8)— Asuncion, 2 (Feb. 16); Palau Islands, USNM— Peleliu, 9 (Sept. 6-20)— Garakayo, 1 
(Sept. 20); AMNH — exact locality not given, 7 (Oct. 13, Nov. 13, 15; Caroline Islands, 
USNM— Ulithi, 4 (Aug. 16, 21); AMNH— Kusaie, 9 (Mar. 10-30)— Ponape, 2 (Dec. 15)— 
Truk, 3 (Feb. 6); Marshall Islands, USNM— Bikini, 3 (Mar. 4, 7, May 3). 

Parasites. — Wharton (1946:174, 175) records the following chiggers (Aca- 
rina) from Pluvialis taken by the NAMRU2 party at Guam: Acariscus pluvius, 
A. anoiis, Neoschongastia carveri, and N. namrui; and at Ulithi: N. pauensis 

and N. ewingi. 

Weights. — Birds taken at Guam and Rota weighed as follows: seven males, 
107-125 (117); four females, 109-120 (114). 

Remarks. — The Pacific Golden Plover is one of the most abundant 
migratory shore birds to visit IMicronesia. So characteristic of 
Micronesia is this species that ahnost all ornithologists who have 
made observations in the area have recorded it. Finsch observed the 
plover in the Carolines and Marshalls. Coultas made notes on, and 
collected specimens of, it in the Marianas, Carolines, and Palaus. 
The Hand-list of Japanese Birds (Hachisuka et al., 1942:216) lists 
Pluvialis from 17 islands in Micronesia. 

Stickney (1943:3, 4) discusses the migrations of the Pacific 
Golden Plover through Oceania, using as a basis for her remarks the 
data from the extensive collections made by the Whitney South Sea 
Expedition. She states that the northward migration begins in 
March from the southern islands (New Zealand and southern Aus- 
tralia) . At Guam in 1945, the writer observed flocks of plover begin- 
ning on February 11. Birds were seen in small groups in March and 
April. In the latter month most of the birds seen were in nuptial 
plumage. For the year 1945, the latest spring record at Guam was 
April 28. In the same year, Gleise (1945:220) observed his last 
spring record at Tinian "between April 26 and 27." In 1946, Mor- 
rison obtained plover in nuptial plumage at Bikini on May 3. 

In an effort to obtain dates when shore birds appeared at Guam, 
field parties of NA]\IRU2 made observations at several beaches in 



134 University of Kansas Publs., Mus. Nat. Hist. 

late spring, summer, and early fall, as is shown in table 8. Pacific 
Golden Plovers in post-nuptial molt were first observed and collected 
on July 8. Following this date, small flocks and later large flocks 
were more numerous; by September 29, plover were abundant. Sim- 
ilar findings were obtained at Ulithi (see table 9) and in the Palau 
Islands (see table 10) in August and September. The birds collected 
by the NAMRU2 party at Guam, Ulithi, Peleliu, and Garakayo in 
July, August, September, and early October were in postnuptial molt. 
Birds taken at Rota on October 20 and 26 were in winter plumage. 
Downs (1946:93) observed plover in small flocks at Tinian in 1945, 
beginning after September 5. Borror (1947:417) saw two birds at 
Agrihan on August 10, 1945. 

The flocks of plover seen by the NAMRU2 party varied in size 
from three to 30 birds, the average being less than ten. Coultas 
(field notes) noted "large flocks" at the Palaus from October to 
December, 1931. Although plover was often found on the same 
beach as other birds, the NAMRU2 observers rarely saw plover 
together with other shore birds. However, on air strips Pluvialis 
occasionally occurred with small numbers of Arenaria, Heteroscelus 
spp., and Numenius phaeopus. Pluvialis and N. phaeopus were the 
only shore birds found to use open grassy flats and other inland areas 
at Guam and Peleliu in 1945. 

Stickney (1943) records Pluvialis in late spring and summer from 
Polynesia, indicating these to be birds remaining in the winter range 
during the breeding season. The NAMRU2 party observed no 
Pacific Golden Plovers at Guam which might be regarded as non- 
migrants, but other species of shore birds were found which might be 
considered as such. The lingering of individuals in the winter range 
is not unusual among migratory birds, and as Stickney points out, 
most of the non-migrants retain their winter dress or assume an 
incomplete breeding plumage. 

Charadrius hiaticula semipalmatus Bonaparte 
Semipalmated Plover 

Charadrius semipalmatus Bonaparte, Journ. Acad. Nat. Sci. Phila., 5, 1825, p. 98. 
New name for Tringa hiaticula Ord. not Charadrius hiaticula Linnaeus, in Wilson's 
Amer. Ornith., Ord. repr., 7, 1824, p. 65. (Type locality, Coast of New Jersey.) 

Charadrius hiaticula Finsch, Ibis, 1880, p. 331 (Taluit) ; Wiglesworth, Abhandl. und 
Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 64 (Taluit or Bonham); Schnee, 
Zool. Jahrbiicher, 20, 1904, p. 389 (Marschall-Inseln) ; Kuroda, in Momiyama, Birds 
Micronesia, 1922, p. 45 (Taluit). 

Geographic range. — Breeds from Arctic America south to coastal Canada. 
Winters from southern United States to South America. In Micronesia: 
Marshall Islands — Jaluit. 



Baker: The Avifauna of Micronesia 135 

Remarks.— F'msch (1880d:331) reported this bird (sight record) 
at Jaluit in the Marshall Islands. Other than this observation, there 
is no history of the species in Micronesia. 

Chardrius dubius curonicus Gmelin 
Ring-necked Plover 

Charadrius curonicus Gmelin, Syst. Nat., 1, pt. 2, 1789, p. 692. (Type locality, 
Kurland.) 

Charadrius dubius curonicus Hand-list Japanese Birds, rev., 1932, p. 194 (Yap); 
Hand-list Japanese Birds, 3d ed., 1942, p. 217 (Yap); Mayr, Birds Southwest Pacific, 
1945, p. 37 (Micronesia). 

Geographic range. — Breeds in northern Europe and Asia. Winters from 
Africa east to Malaysia and Melanesia. In Micronesia: Caroline Islands — Yap. 

Remarks. — The Ring-necked Plover has been recorded at Yap by 
the Japanese collectors. Mayr (1945a: 37) remarks that the bird 
is an occasional migrant through IMicronesia. Gleise and Genelly 
(1945:221) observed four "Papuan" Ring-necked Plovers at Eni- 
wetok in 1945. Apparently no specimen was obtained. 

Charadrius alexandrinus nihonensis Deignan 
Kentish Plov^er 

Charadrius alexandrinus nihonensis Deignan, Joum. Washington Acad. Sci., vol. 31, 
1941, p. 106. (Type locality, Aomori, Hondo.) 

Charadrius cantianus Hartlaub and Finsch, Proc. Zool. Soo. London, 1868, pp. 117, 
118 (Pelew); idem, Proc. Zool. Soc. London, p. 89 (Pelew); Finsch, Journ. Mus. Godef- 
froy, 8, 1875, p. 31 (Palau). 

Aegialitis cantianus Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 
1890-1891 (1891), p. 64 (Pelew); Takatsukasa and Kuroda, Tori, 1, 1915, p. 62 (Pelew). 

Aegialitis alexandrinus dealbatua Kuroda, in Momiyama, Birds Micronesia, 1922, 
p. 45 (Pelew). 

Charadrius alexandrinus dealbatus Hand-list Japanese Birds, rev., 1932, p. 194 
(Palau); Hand-list Japanese Birds, 3d ed., 1942, p. 217 (Palau); Mayr, Birds South- 
west Pacific, 1945, p. 37 (Palau). 

Geographic range. — Breeds in Japan and possibly on adjacent parts of the 
Asiatic mainland. Winters south to Malaya. In Micronesia : Palau Islands — 
exact locality unknown. 

Remarks. — The Kentish Plover is known from a single record ob- 
tained by Semper in the Palau Islands. It is tentatively assigned to 
C. a. nihonensis, which breeds directly north of the Palau Islands on 
Japan. C. a. dealbatus (Swinhoe) breeds more to the west on the 
Asiatic mainland and adjacent islands south of Japan. Additional 
specimens are needed before the subspecific status of migrants to 
Micronesia can be accurately determined. 

Chardrius mongolus stegmanni Stresemann 
Mongolian Dotterel 

Charadrius mongolus stegmanni Stresemann, Omith. Monatsb., 48, 1940, p. 55. 
New name for Charadrius mongolus littoralis Stegmann, 1937, preoccupied. (Type lo- 
cality, Berhing Island.) 

Charadrius sanguineus Lesson, Man. d'Omith., 2, 1828, p. 330 (No type locality 
= Mariana Islands, ex Oustalet, Nouv. Arch. Mua. Hist. Nat. Paris, (3), 8, 1896, p. 



136 University of Kansas Publs., Mrs. Nat. Hist. 

48); idem, Traite d'Omith., 1831, p. 544 (no locality =z Mariana Islands); Hartlaub, 
Journ. f. Ornith., 1854, p. 167 (Mariannen). 

Charadrius monoglicus Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 8, 1896, p. 
48 (Guam, Jaluit, Palaos, Carolines); Hartert, Novit. Zool., 5, 1898, p. 66 (Guam); 
Seale, Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 36 (Guam); Safford, Osprey, 
1902, p. 68 (Guam). 

Aegialitis mongolus Hartert, Novit. Zool., 7, 1900, p. 9 (Ruk). 

Aegialis mongola Safford, Contr. U. S. Nat. Herb., 9, 1905, p. 80 (Guam). 

Aegialites mongola Cox, Island of Guam, 1917, p. 22 (Guam). 

Ochthodromus mongoUcus Takatsukasa and Kuroda, Tori, 1, 1915, p. 62 (Marianas, 
Ruk). 

Charadrius mongolus Ridgway, Bull. U. S. Nat. Mus., 50, pt. 8, 1919, p. 132 
(Ruk); Mayr, Birds Southwest Pacific, 1945, p. 38 (Micronesia). 

Charadrius mongolus mongolus Hartert, Vogel pal. Fauna, 11-12, 1920, p. 1543 
(Marianen, Karolinen) ; Hand-list Japanese Birds, rev.. 1932, p. 194 (Guam, Truk, 
luripik, Kusaie, Jaluit, Majuro); Peters, Check-list Birds World, 2, 1934, p. 253 
(Carolines, Marianas); Bryan, Guam Rec, vol. 13, no. 2, 1936, p. 24 (Guam); Baker, 
Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 50 (Guam, Peleliu, Uhthi). 

Cirrepidcsmus mongolus mongolus Kuroda, in Momiyama, Birds Micronesia, 1922, 
p. 44 (Guam, Ruk). 

Charadrius mongolus stegmanni Hand-list Japanese Birds, 3d ed., 1942, p. 217 
(Guam, Peliliu, Truk, luripik, Kusaie, Jaluit, Majuro). 

Geographic range. — Breeds in northeastern Siberia and Bering Sea area. 
Winters south to eastern Malaysia, Melanesia, and Australia. In Micronesia: 
Mariana Islands — Guam ; Palau Island.s — Angaur, Peleliu ; Caroline Islands — 
Ulithi, Truk, luripik, Ku.saie; Marshall Islands — Jaluit, Majuro. 

Specimens examined. — Total number, 10 (4 males, 5 females, 1 unsexed), as follows: 
Mariana Islands, USNM — Guam, 2 (June 7, Sept. 1); AMNH — Guam, 3 (Aug. 15, 18, Nov. 
30); Palau Islands, USNM— Peleliu, 3 (Sept. 7-12); Caroline Islands, USNM— Ulithi, 1 
(Aug. 22); AMNH— Truk, 1 (Feb. 8). 

Remarks. — According to Oustalet (1896:48), Lesson used two 
specimens of this species, which were collected in the Marianas by 
the expedition in the "Uranie," as types for his Charadrius sangui- 
neus. 

The Mongolian Dotterel is a regular visitor to western Micro- 
nesia. It is recorded also from the Marshall Islands, which it prob- 
ably reaches from the westward by way of the Carolines, since the 
species has not been recorded in the Hawaiian Islands. 

A bird taken by the writer at Guam on June 7, 1945, was in 
winter plumage and probably nonmigratory. The species was re- 
corded also at Guam in September. At Peleliu in September, 1945, 
the Mongolian Dotterel was seen frequently on tidal fiats by the 
NAMRU2 party. On September 8 there was a flock of approxi- 
mately fifty birds, in company with Charadrius leschenaultii, at 
Akarakoro Point. In August at Ulithi, birds were on the beaches in 
company with Crocethia alba. At Angaur on September 21, 1945, 
the species was with other shore birds in small groups at fresh water 
ponds. 

I am tentatively referring all specimens examined to C. m. steg- 



Baker: The Avifauna of Micronesia 137 

manni although at this writing (1948) I am inclined to the opinion 
that a critical reexamination of the referred specimens might reveal 
one or a few individuals of the subspecies C. m. mongolus Pallas. 

Charadrius leschenaultii Lesson 
Large Sand Dotterel 

Charadrius Leschenaultii Lesson, Diet. Sci. Nat., ed. Levrault, 42, 1826, p. 36. 
(Type locality, Pondichery, India.) 

Charadrius griseus Lesson, Traite d'Omith., 1831,, p. 544 (Oulan). 

Charadrius geoffroyi Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 117, 
118 (Pelew); idem, Proc. Zool. Soc. London, 1872, p. 89 (Pelew); Finsch, Journ. Mus. 
Godeffroy, 8, 1875, pp. 5, 31 (Palau). 

Aegialitis geoffroyi Salvadori, Omith. Papuasia, 3, 1882, p. 299 (Ualan, Pelew); 
Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 64 
(Pelew, Ualan). 

Ochthodromus geoffroyi Sharpe, Cat. Birds British Mus., 24, 1896, p. 217 (Pelew, 
Ualan); Takatsukasa and Kuroda, Tori, 1, 1915, p. 62 (Pelew). 

Pagoa leschenaultii Kuroda, in Momiyama, Birds Micronesia, 1922, p. 44 (Pelew, 
Kusiae, Yap). 

Charadrius leschenaultii leschenaultii Hand-list Japanese Birds, rev., 1932, p. 193 
(Yap, Kusaie, Palau); Hand-list Japanese Birds, 3d ed., 1942, p. 216 (Yap, Kusaie, 
Palau). 

Charadrius leschenaultii Mayr, Birds Southwest Pacific, 1945, p. 38 (Micronesia); 
Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 51 (Peleliu). 

Geographic range. — Breeds in Asia south to Persia. Winters from Malaysia 

east to Australia and Melanesia. In Micronesia: Palau Islands — Peleliu; 

Caroline Islands — Yap, Kusaie. 

Specimens examined.- — Total number, 9 (2 males and 7 females), as follows: Palau Islands, 
USNM— Peleliu, 7 (Sept. 6-12); AMNH— exact locality not given, 2 (Nov. 21, 25). 

Remarks. — The Large Sand Dotterel is a regular visitor to the 
Palau Islands. It has been recorded also at Yap and Kusaie in the 
Carolines, where it may be considered as an uncommon visitor. 

At Peleliu, the species was seen on several occasions in September, 
1945, by the NAMRU2 party. The birds were found on tidal flats 
in company with Charadrius mongolus stegmanni in flocks of 10 to 
30 individuals. 

Numenius phaeopus variegatus (Scopoli) 
Whimbrel 

Tantalus variegatus Scopoli, Del. Flor. et Faun. Insubr., fasc. 2, 1786, p. 92. (Type 
locality, Luzon, ex. Sonnerat.) 

Scolopax phaeopus Lesson, Traite d'Ornith., 1831, p. 566 (Marianas). 

Numenitis phaeopus Kittlitz, Obser. Zool., in Lutke, Voy. "Le Seniavine," 3, 1836, 
pp. 287, 304 (Ualan, Guahan), Hartlaub, Joum. f. Omith., 1854, p. 167 (Mariannen); 
Kittlitz, Denkw. Reise russ. Amer. Micron, und Kamchat., 2, 1858, p. 129 (Ualan); 
Hartlaub, Proc. Zool. Soc. London, 1867 (1868), p. 831 (Pelew, Matelotas) ; Hartlaub 
and Finsch, Proc. Zool. Soc. London, 1868, pp. 8, 118 (Pelew); idem, Proc. Zool. Soc. 
London, 1872, pp. 89, 106 (Uap, Pelews) ; Graffe, Journ. Mus. Godeffroy, 2, 1873, p. 
123 (Yap); Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 5, 35 (Palau); idem, Joum. f 
Ornith., 1880, pp. 294, 307 (Ponape, Kuschai) ; idem, Proc. Zool. Soc. London, 1880, 
p. 576 (Ruk); idem. Ibis, 1881, pp. 107, 109, 115 (Kushai, Ponape); Schmeltz and 
Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, pp. 281, 299, 353 (Ponape, Mortlock, 



138 University of Kansas Publs., Mus. Nat. Hist. 

Ruk); Wharton and Hardcastle, Joum. Parasitology, 32, 1946, pp. 308, 316, 318, 320 
(Ulithi, Guam); Wharton, Ecol. Monogr., 16, 1946, pp. 174, 175 (Guam). 

Numenius tenuirostris Kittlitz, Denkw. Reise russ. Amer. Micron, and Kamchat., 2, 
1858, p. 55 (Marianas, Ualan). 

Numenius uropygialis Gray, Hand-list Birds, 3, 1871, p. 43 (Pelew). 

Numenius variegatui Salvadori, Ornith. Papuasia, 3, 1882, p. 332 (Pelew, Ponape) ; 
Wiglesworth, Abhandl. und Bar. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 66 
(Marianne, Pelew, Matalotas, Luganor, Ruk, Ponape, Ualan); Sharpe, Cat. Birds 
British Mus., 24, 1896, p. 361 (Micronesia); Safford, The Plant World, 7, 1904, p. 266 
(Guam). 

Numenius phaeopus variegatus Oustalet, Nouv. Arch. Mus. Hist. Nat Paris, (3), 8, 
1896, p. 39 (Mariannes, Palaos, Carolines, Jaluit) ; Hartert, Novit. Zool., 5, 1898, 
p. 65 (Guam); idem, Novit. Zool., 7, 1900, p. 8 (Ruk); Seale, Occ. Papers Bernice P. 
Bishop Mus., 1, 1901, p. 34 (Guam); Safford, Osprey, 1902, p. 67 (Marianas); idem, 
Contr. U. S. Nat. Herb., 9, 1905, p. 80 (Guam); Takatsukasa and Kuroda, Tori, 1, 
1915, p. 62 (Marianas, Carolines, Pelews) ; Cox, Island of Guam, 1917, p. 21 (Guam); 
Hartert, Vogel pal. Fauna, 13-14, 1921, p. 1649 (Guam); Hand-list Japanese Birds, 
rev., 1932, p. 192 (Marianas, Carolines, Palaus, Marshalls) ; Peters, Check-list Birds 
World, 2, 1934, p. 261 (Caroline, Marianne, Pelew); Bryan, Guam Rec, vol. 13, no. 2, 
1936, p. 24 (Guam); Hand-list Japanese Birds, 3d ed., 1942, p. 215 (Guam, Koror, 
Babelthuap, Ngulu, Yap, Uluthi, luripik, Truk, Lukunor, Ponape, Kusaie, Jaluit, 
Wotze); Mayr, Birds Southwest Pacific, 1945, p. 39 (Micronesia); Strophlet, Auk, 1946, 
p. 537 (Guam); Stott, Auk, 1947, p. 525 (Saipan); Baker, Smithson. Misc. Coll., vol. 
107, no. 15, 1948, p. 51 (Guam, Angaur, Peleliu, Ulithi). 

Phaeopus phaeopus variegatus Wetmore, in Townsend and Wetmore, Bull. Mus. 
Comp. Zool., 63, 1919, p. 178 (Guam); Kuroda, in Momiyama, Birds Micronesia, 1922, 
p. 45 (Palaus, Carolines, Marians). 

Geographic range. — Breeds in northeastern Asia. Winters from Malaysia 
east to Oceania. In Micronesia: Mariana Islands — Guam; Palau Islands — 
Angaur, Peleliu, Koror, Babelthuap; Caroline Islands — Ngulu, Yap, Ulithi, 
Truk, Lukunor, luripik, Ponape, Kusaie; Marshall Islands — Jaluit, Wotze. 

Specimens examined. — Total number, 26 (9 males, 17 females), as follows: Mariana Islands, 
USNM— Guam, 16 (June 4, 6, July 24, 26, 27, Sept. 1, 19, 25, Oct. 8); Palau Islands, USNM 
—Peleliu, 5 (Sept. 8, 12, 14)— Angaur, 4 (Sept. 21); Caroline Islands, USNM— Ulithi, 1 
(Aug. 17). 

Weights. — At Guam, the NAMRU2 party obtained the weights of two males, 
373 and 435, and of six females, 295-426 (384). 

Parasites. — Wharton (1946:174, 175) lists the following species of chiggers 
(Acarina) taken from the Whimbrel at Guam: Acariscus pluvins, A. anous, 
Neoschongastia strongi, and A'^. carveri; and at Ulithi: N. namrui and A''. 
atollensis. 

Remarks. — The Whimbrel is an abundant visitor to western Mi- 
cronesia. It was first taken by Quoy and Gaimard, who found it 
in the Marianas. It is recorded in the Marshall Islands (Jaluit and 
Wotze), but apparently reaches these islands from the west, since 
the species is unknown in the Hawaiin Islands. 

As shown in table 8, the NAMRU2 party observed the Whimbrel 
at Guam on spring migration in March, 1945, the last record being 
on March 21. In June and July, single birds or small groups were 
occasionally seen on the tidal flats. Some of these birds may have 
been nonmigratory. Beginning on July 24, more birds were re- 
corded as they began to migrate south after their nesting season. 



Baker: The Avifauna of Micronesia 139 

Whimbrels were numerous from August until the conclusion of the 
observations in October. Birds were abundant at the Palaus in 
September; only a few were noted at Ulithi in late August. The 
Whitney South Sea Expedition of the American Museum of Natu- 
ral History made collections of this species at several islands in 
Micronesia. At Ponape, Coultas (field notes) writes that in No- 
vember and December, 1930, a few birds were seen on the reefs and 
at the edges of mangrove swamps. At Peleliu in October to Decem- 
ber, 1931, he found Whimbrels concentrated on a small islet between 
Koror and Babelthuap. At both Ponape and Palau Coultas re- 
ceived reports that the birds remain at the islands throughout the 

year. 

Numenius tahitiensis (Gmelin) 

Bristle-thighed Curlew 

Scolopax tahitiensis Gmelin, Syst. Nat., 1, pt. 2, 1789, p. 656. (Type locality, 
Tahiti, Society Islands, based on the Otaheiti Curlew of Latham, Gen. Syn., 3, pt. 1, 
1785, p. 122, no. 4.) 

Numenius femoralis Finsch, Ibis, 1880, pp. 220, 331, 332 (Jaluit, Arno). 

Nu7neniiis tahitiensis Seebohm, Geogr. Dist. Charadriidae, 1887, p. 332 (Marshalls) ; 
Wiglesworth, Abhandl. und Ber Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 66 
(Marianne?, Marshalls); Sharpe, Cat. Birds British Mus., 24, 1896, p. 367 (Marianas, 
Marshalls); Schnee, Zool. Jahrbiicher, 20, 1904, p. 390 (Marschall-Inseln) ; Takatsu- 
kasa and Kuroda, Tori, 1, 1915, p. 62 (Marianas, Pelews) ; Kuroda, in Momiyama, 
Birds Micronesia, 1922, p. 49 (Marianas, Marshalls); Bent, Bull. U. S. Nat. Mus., 
146, 1929, p. 143 (Jaluit); Hand-list Japanese Birds, rev., 1932, p. 192 (Saipan, 
Marshalls); Peters, Check-list Birds World, 2, 1934, p. 261 (Marshalls); Yamashina, 
Tori, 10, 1940, p. 677 (Jarchi) ; Hand-list Japanese Birds, 3d ed., 1942, p. 215 (Sai- 
pan, Jaluit, Arhno, Maloelab, Wotze, Ailuk, Ringelab, Larchi) ; Stickney, Amer. Mus. 
Novit., no. 1248, 1943, p. 4 (Ponape, Marshalls); Mayr, Birds Southwest Pacific, 
1945, p. 39 (Marshalls, straggler to Carolines and Marianas). 

Phaeopiis tahitiensis Wetmore, in Townsend and Wetmore, Bull. Mus. Comp. Zool., 
63, 1919, p. 179 (Rongelab); Ridg^vay, Bull. U. S. Nat. Mus., 50, pt. 8, 1919, p. 
407 (Marianas, Marshalls). 

Geographic range. — Breeds in western Alaska. Winters in eastern and 
central Polynesia. In Micronesia: Mariana Islands — Saipan; Caroline Islands 
— Ponape; Marshall Islands — Jaluit, Arhno, Moloelab, Wotze, Ailuk, Ronge- 
lab, Larchi, Bikini. 

Specimens examined. — Total number, 6 (3 males, 3 females), as follows: Caroline Islands, 
AMNH— Ponape, 2 (Dec. 15); Marshall Islands, USNM— Bikini, 4 (Mar. 10, 14, April 2, 
30). 

Remarks. — The Bristle-thighed Curlew is a regular migrant 
through the Marshall Islands of eastern Micronesia. It is recorded 
as a straggler to the Caroline and Mariana islands. Stickney (1943: 
4, fig. 1) shows a map and discusses the breeding and wintering 
ranges of this curlew. As can be observed from her map, the prin- 
cipal wintering areas are east and south of Micronesia. She records 
the species from the Bonin Islands, which is the westernmost record. 

It is difficult to offer plausible reasons for the present migratory 



140 University of Kansas Publs., Mus. Nat. Hist. 

habits of the Bristle-thighed Curlew. It is related to both the Asia- 
tic form, N. phaeopus, and to the American species, N. hudsonicus, 
but its origin is not understood. The characteristics of its route of 
migration resemble that of some continental migrants and might 
have come about by a slow adjustment of the species to its environ- 
ment, probably through an expansion of range from the west, 

Numenius madagascariensis (Linnaeus) 
Long-billed Curlew 

Scolopax madagascariensis Linnaeus, Syst. Nat., ed. 12, 1, 1766, p. 242. (Type 
locality, Madagascar, error = Manila, Philippine Islands, fide Stresemann.) 

Numenius cyanopus Hartert, Novit. Zool., 5, 1898, p. 65 (Guam); Seale, Occ. 
Papers Bernice P. Bishop Mus., 1, 1901, p. 35 (Micronesia); Safford, Osprey, 1902, 
p. 67 (Marianas); idem. The Plant World, 7, 1904, p. 266 (Guam); idevi, Contr. 
U. S. Nat. Herb., 9, 1905, p. 80 (Guam); Cox, Island of Guam, 1917, p. 21 (Guam); 
Hartert, Vogel pal. Fauna, 13-14, 1921, p. 1645 (Guam); Kuroda, in Momiyama, 
Birds Micronesia, 1922, p. 45 (Guam); Hand-list Japanese Birds, rev., 1932, p. 192 
(Guam). 

Numenius madagascariensis Hand-list Japanese Birds, 3d ed., 1942, p. 214 (Guam); 
Mayr, Birds Southwest Pacific, 1945, p. 40 (Micronesia); Baker, Smithson. Misc. Coll., 
vol. 107, no. 15, 1948, p. 51 (Guam, Ngesebus). 

Geographic range. — Breeds in eastern Siberia. Winters from Malayia east 
to Australia and Melanesia. In Micronesia: Mariana Islands — Guam; Palau 
Islands — Peleliu, Ngesebus. 

Remarks. — The Long-billed Curlew is a regular visitor to western 
Micronesia, especially to the Palau Islands. It is apparently a less 
common migrant in the Marianas, although it has been recorded 
from Guam. At Guam, the NAMRU2 party observed a single bird 
on June 6 and two on October 3 at tidal beaches. At Peleliu these 
large curlews were seen on several occasions between September 9 
and 16, 1945. They were found usually as singles feeding on tidal 
flats in company with other shorebirds. 

Limosa lapponica baueri Naumann 
Pacific Godwit 

Limosa Baueri Naumann, Naturg. Vog. Deutschl., 8, 1836, p. 429. (Type locality, 
New Holland = Victoria, apud Mathews; Novit. Zool., 18. 1912, p. 220.) 

Limosa uropygialis Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, p. 
299 (Mortlock). 

Limosa novae-sealandiae Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 
6, 1890-1891 (1891), p. 66 (Luganor). 

Limosa lapponica baueri Hartert. Novit. Zool., 5, 1898, p. 65 (Guam); Seale, Occ. 
Papers Bernice P. Bishop Mus., 1, 1901, p. 34 (Guam); Safford, Osprey, 1902, p. 67 
(Marianas); idem. The Plant World, 7, 1904, p. 266 (Guam); idem, Contr. U. S. Nat. 
Herb., 9, 1905, p. 80 (Guam); Prowazek, Die deutschen Marianen, 1913, p. 101 (Mari- 
anen); Cox, Island of Guam, 1917, p. 21 (Guam); Hartert, Vogel pal. Fauna, 13-14, 
1921, p. 1641, (Guam); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 46 (Caro- 
lines, Marianas); Hand-list Japanese Birds, rev., 1932, p. 191 (Marianas. Carolines); 
Bryan, Guam Rec, vol. 13, no. 2, 1936, p. 24 (Guam); Stickney, Amer. Mus. Novit., 
no. 1248, 1943, p. 5 (Guam, Palau); Mayr, Birds Southwest Pacific, 1945, p. 41 



Baker: The Avifauna of Micronesia 141 

(Oceania); Strophlet, Auk, 1946, p. 537 (Guam); Baker, Smithson. Misc. Coll., vol. 
107, no. 15, 1948, p. 52 (Guam, Peleliu). 

Limosa lapponica novazealandiae Hartert, Novit. Zool., 7, 1900, p. 8 (Ruk); Hand- 
list Japanese Birds, 3d ed., 1942, p. 214 (Guam, Truk). 

Limosa rufa uropygialis Takatsukasa and Kuroda, Tori, 1, 1915, p. 62 (Marianas, 
Ruk). 

Geographic range. — Breeds in northeastern Asia and northwestern North 
America. Winters from Malaysia east to Oceania. In Micronesia: Mariana 
Islands — Guam; Palau Islands — PeleHu; Caroline Islands — Truk. 

Specimens examined. — Total number, 5 (2 males, 3 females), as follows: Mariana Islands, 
AMNH— Guam, 2 (Sept. 26); Palau Islands, USNM— Peleliu, 1 (Sept. 7); AMNH— exact 
locality not given, 2 (Nov. 21, 23). 

Remarks. — The principal wintering grounds of the Pacific Gociwit 
are probably in Australia and New Zealand according to Stickney 
(1943:5). The bird reaches these areas from Arctic breeding 
grounds by migrating to a great extent along the edge of the Asiatic 
Continent. It may also be considered as a regular migrant in west- 
ern Micronesia, and probably reaches eastern Micronesia as an un- 
common visitor, since it is occasionally recorded in the Hawaiian 
Islands. 

At Guam in 1945, the NAMRU2 party found the Pacific Godwit 
at tidal beaches on April 26 and October 15. Strophlet (1946:537) 
recorded one bird from Guam on October 20, 1945. At Peleliu, the 
NAMRU2 party found birds at beaches on September 7 and 16. 
Coultas (field notes) reported that "a few" were seen at Peleliu 
from October to December, 1931. McElroy did not find any of 
these birds at Truk in December, 1945. 

Tringa nebularia (Gunnerus) 
Greenshank 

Scolopax nebularis Gunnerus, in Leem, Beskr. Finm. Lapper, 1767, p. 251. (Type 
locality, District of Trondhjem, Norway.) 

Glottis nebularius Kuroda, in Momiyama, Birds Micronesia, 1922, p. 47 (Yap) ; 
Takatsukasa and Yamashina, Dobutsu. Zas.shi, 44, 1932, p. 225 (Truk); Hand-list 
Japanese Birds, rev., 1932, p. 191 (Yap, Truk). 

Tringa nebularis Hand-list Japanese Birds, 3d ed., 1942, p. 214 (Yap, Truk); Mayr, 
Birds Southwest Pacific, 1945, p. 41 (Yap, Truk) ; Baker, Smithson. Misc. Coll., vol. 
107, no. 15, 1948, p. 52 (Peleliu). 

Geographic range. — Breeds in northern Eurasia. Winters in Mediterranean 
area, Africa, southern Asia, Malaysia, Australia and Melanesia. In Micronesia: 
Palau Islands— Peleliu ; Caroline Islands — Yap, Truk. 

Specimens examined. — Total number, 4 (1 male, 3 females) from Palau Islands, USNM — 
Peleliu (Aug. 28, Sept. 14, 15). 

Remarks. — The Greenshank has been recorded at the Palau 
Islands and at Yap and Truk in the Caroline Islands. It is appar- 
ently a regular visitor to western Micronesia. It probably reaches 
the western Carolines as an occasional visitor from the region of the 



142 University of Kansas Publs., Mus. Nat. Hist. 

Palaus to the westward, rather than from the northward, since the 
bird has not been observed in the Marianas. 

The NAMRU2 party observed two small flocks of these birds at 
Peleliu in August and September, 1945. One group of six birds was 
found wading in the shallow water of a mangrove swamp on August 
28. Another group of three birds was seen on a tidal beach on Sep- 
tember 14 and 15, where they were observed feeding apart from 
other species of shore birds. 

Tringa melanoleuca (Gmelin) 

Greater Yellow-legs 

Scolopax melanoleuca Gmelin, Syst. Nat., 1, pt. 2, 1789, p. 659. (Type locality, 
Sandy shores of abrador := Chateau Bay, Labrador.) 

Tringa melanoleuca Kuroda, Dobutsu. Zasshi, 46, 1934, p. 313 (Jaluit) ; Hand-list 
Japanese Birds, 3d ed., 1942, p. 214 (Jaluit). 

Geographic range. — Breeds in Alaska and Canada. Winters from California 
east to the Gulf States and the West Indies and south to South America. In 
Micronesia : Marshall Islands — Jaluit. 

Remarks. — Kuroda records one specimen of the Greater Yellow- 
legs from Jaluit Atoll in the Marshall Islands. It is a straggler to 
Oceania and has not been recorded in the Hawaiian Islands. 

Tringa glareola Linnaeus 
Wood Sandpiper 

Tringa glareola Linnaeus, Syst. Nat., ed. 10, 1, 1758, p. 149. (Europe, restricted 
type locality, Sweden.) 

Totanus glareola Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 8, 1896, p. 43 
(Guam); Hartert, Novit. Zool., 5, 1898, pp. 65, 69 (Guam); Seale, Occ. Papers Bemice 
P. Bishop Mus., 1, 1901, p. 34 (Guam); Safford, Osprey, 1902, p. 70 (Guam); idem. 
The Plant World, 7, 1904, p. 268 (Guam). 

Rhyacophilus glareola Kuroda, in Momiyama, Birds Micronesia, 1922, p. 48 (Guam, 
Angaur). 

Tringa glareola Hand-list Japanese Birds, rev., 1932, p. 191 (Guam, Angaur, 
Koror) ; Hand-list Japanese Birds, 3d ed., 1942, p. 213 (Guam, Anguar, Koror) ; 
Mayr, Birds Southwest Pacific, 1945, p. 41 (Guam, Palau) ; Baker, Smithson. Misc. 
Coll., vol. 107, no. 15, 1948, p. 52 (Anguar). 

Tringa glariola Bryan, Guam Rec, vol. 13, no. 2, 1936, p. 24 (Guam). 

Geographic range. — Breeds in northern Eurasia from Norway and Germany 
east to Siberia, Sakhalin, and Kamchatka. Winters from Africa east to south- 
ern Asia, Malaysia, and Australia. In Micronesia: Mariana Islands — Guam; 
Palau Islands — Anguar, Koror. 

Specimens examined. — Total number, 2 (1 male, 1 female), as follows: Palau Islands, 
USNM — Angaur, 1 (Sept. 21); AMNH — e.xact locality not given, 1 (October 26). 

Remarks. — Marche, in 1877, first recorded the Wood Sandpiper 
in Micronesia (at Guam). In the Marianas it is apparently an un- 
common migrant but it is considered to be a regular visitor in the 
Palau Islands. At the Palaus in September, 1945, the writer found 



Baker: The Avifauna of Micronesia 143 

the bird at a fresh water pond on Angaur. It was not observed on 
the tidal beaches at Peleliu. 

Actitis hypoleucos Linnaeus 
Common Sandpiper 

Tringa Hypoleucos Linnaeus, Syst. Nat., ed. 10, 1> 1858, p. 149 (Europe, restricted 
type locality, Sweden.) 

Totanus hypoleucos Lesson, Traite d'Ornith., 1831, p. 552 (Marianas). 

Totanus (Tringoides) hypoleucus Gray, Birds Trop. Is. Pacific Ocean, 1859, p. 51 
(Marianas). 

Actitis hypoleuca Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, p. 8 (Pelew). 

Actitis hypoleucus Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, p. 118 
(Pelew); idem, Proc. Zool. Soc. London, 1872, pp. 89, 106 (Pelew); Finsch, Journ. 
Mus. Godeffroy, 8, 1875, p. 36 (Pelew); Schmeltz and Krause, Ethnogr. Abth. Mus. 
Godeffroy, 1881, pp. 299, 353 (Ruk, Mortlock) ; Wiglesworth, Abhandl. und Ber. Zool. 
Mus. Dresden, no. 6, 1890-1891 (1891), p. 64 (Luganor, Marianne, Pelew); Oustalet, 
Nouv. Arch. Mus. Hist. Nat. Paris (3), 8, 1896, p. 43 (Guam, Palaos, Luganor). 

Tringoides hypoleucos Gray, Hand-list Birds, 3, 1871, p. 46 (Pelew, Ladrone) ; 
Salvadori, Ornith. Papuasia, 3, 1882, p. 318 (Pelew). 

Tringoides hypoleucus Sharpe, Cat. Birds British Mus., 24, 1896, p. 456 (Micro- 
nesia); Takatsukasa and Kuroda, Tori, 1, 1915, pp. 51, 62 (Pelews, Marianas). 

Totanus hypoleucus Hartert, Novit. Zool., 5, 1898, p. 65 (Saipan) ; Scale, Occ. 
Papers Bernice P. Bishop Mus., 1, 1901, p. 34 (Guam); Safford, Osprey, 1902, p. 70 
(Mariannes); idem. The Plant World, 7, 1904, p. 268 (Guam). 

Actitis hypoleucos Ridgway, Bull. U. S. Nat. Mus., 50, pt. 8, 1919, p. 372 (Micro- 
nesia); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 47 (Marianas, Carolines, 
Pelews) ; Peters, Check-list Birds World, 2, 1934, p. 269 (Micronesia) ; Bryan, Guam 
Rec, vol. 13, no. 1, 1936, p. 24 (Guam); Mayr, Birds Southwest Pacific, 1945, p. 42 
(Micronesia); Strophlet, Auk, 1946, p. 537 (Guam); Baker, Smithson. Misc. Coll., 
vol. 107, no. 15, 1948, p. 52 (Guam, Peleliu, Ulithi). 

Tringa hypoleucos Hand-list Japanese Birds, rev., 1932, p. 191 (Marianas, Carolines, 
Pelews); Hand-list Japanese Birds, 3d ed, 1942, p. 214 (Saipan, Babelthuap, Koror, 
Peleliu, Angaur, Ulithi, Truk). 

Geographic range. — Breeds in Europe and Asia. Winters from Africa east to 
Polynesia. In Micronesia: Mariana Islands — Guam, Saipan; Palau Islands — 
Angaur, Peleliu, Koror, Babelthuap; Caroline Islands — Ulithi, Truk, Lukunor. 

Specimens examined. — Total number, 12 (4 males, 7 females, 1 unsexed), as follows: 
Mariana Islands, USNM— Guam, 4 (July 16, Sept. 20); AMNH— Saipan, 1 (July 27); Palau 
Islands, USNM— Peleliu, 3 (Sept. 9, 14),— Koror, 1 (Nov. 7); AMNH— exact locality not 
given, 2 (Nov. 11, 19); Caroline Islands, USNM— Ulithi, 1 (Aug. 22). 

Weights. — The present author (1948:52) recorded the weight of one male 
taken at Guam as 67 grams, and of two females as 57 and 63 grams. These 
were fall migrants taken by the NAMRU2 party. 

Remarks. — The Common Sandpiper has been known from Micro- 
nesia since the time of Lesson. Tetens, Peters and Kiibary obtained 
specimens in the Palaus; the latter collector obtained the bird at 
Lukunor and probably also at Truk. In recent years several col- 
lectors have taken the birds in western Micronesia, where the species 
appears to be a regular visitor. Field observations by the NAMRU2 
party indicate that the birds are usually found as singles and remain 
apart from other species of migratory shorebirds which visit the 
islands. The margins of inland ponds and beaches consisting of 



144 University of Kansas Publs., Mus. Nat. Hist. 

rocks and pebbles appear to be preferred over the sandy, tidal flats. 
At Peleliu on September 9, 1945, two birds were taken at a bare bank 
of coral at an inland pond. These were the only two Common Sand- 
pipers seen at the island. A specimen taken by the NAMRU2 party 
at Ulithi on August 22 at a beach, piled with debris from ships, has 
its entire and underparts stained by fuel oil. 

Heteroscelus brevipes (Vieillot) 
Gray-tailed Tattler 

Totanus brevipes Vieillot, Nouv. Diet. Hist. Nat., 6, 1816, p. 410. (No locality 
given, the type is from Timor.) 

Totanus pedestris Lesson, Traite d'Ornith., 1831, p. 552 (Marianne, Ualan). 

Totanus brevipes Kittlitz, Obser. Zool., in Lutke, Voy. "he Seniavine," 3, 1836, pp. 
287, 299, 304 (Ualan, Lougounor, Guahan) ; Gray, Cat. Birds Trop. Is. Pacific Ocean, 
1859, p. 51 (Ladrone or Marian Is.); Pelzeln, Raise "Novara," Vogel, 1865, p. 129, 
162 (Puynipet, Ualan). 

Totanus incanus Finsch and Hartlaub (part). Fauna Centralpolynesians, 1867, p. 187 
(Mariannen, Ualan, Puynipet); Salvadori (part), Ornith. Papuasia, 3, 1882, p. 322 
(Micronesia); Wiglesworth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890- 
1891 (1891), p. 65 (Mulgrave, Taluit, Ualan, Ponape, Ruk, Luganor, Uap, Pelew, 
Marianas); Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 8, 1896, p. 41 
(Saypan, Guam, Jaluit, Carolines, Palaos). 

Heteractitis brevipes Seale, Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 35 
(Marianas); Safford, Osprey, 1902, p. 67 (Marianas); idem, Contr. U. S. Nat. Herb., 
9, 1905, p. 80 (Guam); Cox, Island of Guam, 1917, p. 21 (Guam); Wetmore, in 
Townsend and Wetmore, Bull. Mus. Comp. Zool., 63, 1919, p. 180 (Uala = Truk); 
Mathews, Syst. Avium Australasianarum, 1, 1927, p. 170 (Carolines). 

Heteractitis brevis Prowazek, Die deutschen Marianen, 1913, pp. 47, 101 (Marianen). 

Heteroscelus brevipes Ridgway, Bull. U. S. Nat. Mus., 50, pt. 8, 1919, p. 367 
(Western Pacific); Peters, Check-list Birds World, 2, 1934, p. 270 (Carolines). 

Tringa incana brevipes Hartert, Vogel pal. Fauna, 13-14, 1921, p. 1623 (Guam, 
Truk); Hand-list Japanese Birds, rev., 1932, p. 191 (Palaus, Carolines); Hand-list 
Japanese Birds, 3d ed., 1942, p. 213 (Babelthuap, Koror, Angaur, Yap, luripik, 
Faraulep, Truk, Ponape). 

Heteroscelus incaiius brevipes Kuroda, in Momiyama, Birds Micronesia, 1922, p. 47 
(Pelew, Yap, Ruk); Kuroda, Avifauna Riu Kiu, 1925, p. 177 (Micronesia); Stickney, 
Amer. Mus. Novit., no. 1248, 1943, p. 5 (Saipan, Guam, Palau, Ruk, Kusaie) ; Mayr, 
Birds Southwest Pacific, 1945, p. 43 (Micronesia) ; Baker, Smithson. Misc. Coll., vol. 
107, no. 15, 1948, p. 52 (Guam, Peleliu, Truk). 

Heteroscelus incanus Wharton and Hardcastle (part), Journ. Parasitologj', 32, 1946, 
pp. 296, 316, 318 (Guam, Peleliu). 

Geographic range. — Breeds in eastern Siberia and adjacent areas. Winters 
south to Malaysia and east to Australia and Oceania. In Micronesia: Mari- 
ana Islands — Guam, Saipan; Palau Islands — Angaur, Peleliu, Koror, Babel- 
thuap; Caroline Islands — Yap, Truk, luripik, Faraulep, Ponape, Kusaie. 

Specimens examined. — Total number, 39 (11 males, 27 females, 1 unsexed), as follows: 
Mariana Islands, USNM— Guam, 16 (June 4, 6, July 16, 24, Aug. 6, 27, Sept. 4, 6, 27, 
Oct. 23); AMNH— Saipan, 1 Sept. 8),— Guam, 5 (Feb. 11, Mar. 4, 13, Sept. 14, Dec. 5); 
Palau Islands, USNM— Peleliu, 7 (Sept. 6-8, 16); AMNH — exact locality not given, 4 (Nov. 
8); Caroline Islands, USNM— Truk, 1 (Dec. 13); AMNH— Truk, 3 (Feb. 6, 26, Oct. 14),— 
Kusaie, 2 (Mar., April). 

Weights. — Weights of birds obtained by the NAMRU2 party were as fol- 
lows: three males from Guam, 90-104 (95); six females from Guam, 99-116 
(104). 



Baker: The Avifauna of Micronesia 145 

Remarks. — It is not clear whether some of the accounts cited 
above refer to this species or to the species, Heteroscehis incanus. 
Owing to the fact that specimens used in some of these early re- 
ports have not been examined by me, the identifications of the birds 
concerned cannot be verified and consequently it is impossible to 
be certain to which species some of the references pertain. In list- 
ing these accounts in the literature, I am following Sharpe (1896: 
455) whenever possible. 

Tattlers were among the first birds observed and taken in Micro- 
nesia. Quoy and Gaimard found them in the Marianas, and Kitt- 
litz and Kubary recorded the species in the Carolines. Kubary also 
reported the birds at the Palaus. 

The Gray-tailed Tattler apparently does not reach the Marshall 
Islands but visits only the western part of Micronesia. Stickney 
(1943:2) shows a map of the known geographic range of this species 
in Micronesia. The separation of H. brevipes and H. incanus in the 
field is not always possible. For identification, the NAMRU2 party 
depended primarily on specimens collected. At Guam, specimens of 
H. brevipes, thought to be nonmigratory, were taken in early June. 
These were in winter plumage. Beginning in mid-July there was an 
increase in the number of tattlers seen; apparently fall migration 
had begun. At Peleliu in September, 1945, the NAMRU2 party 
found tattlers to be numerous. Apparently all were of this species ; 
no H. incanus were taken there. On September 8, approximately 75 
individuals in small and large flocks were counted at Akarakoro 
Point on the tidal flats. The birds remained apart from the other 
shorebirds which were feeding at the same locality. 

Heteroscelus incanus (Gmelin) 
American Wandering Tattler 

Scolopax incana Gmelin, Syst. Nat., 1, pt. 2, 1789, p. 6.58, (Type locality, Eimeo 
= Moorea, Society Islands and Palinerton Islands.) 

Totanus oceanicus Lesson, Mamm. et Ois., 2, 1847, p. 244 (Kusaie) ; Hartlaub, 
Archiv f. Naturgesch., 1852, p. 135 (Carolinen) ; idem, Journ. f. Ornith., 1854, pp. 
167, 168 (Carolinen, Mariannen). 

Tryanga glareola Kittlitz, Denkw. Reise russ. Amer. Micron, und Kamchat., 1, 1858, 
p. 365, 2, pp. 55, 86 (Ualan). 

Totanus incanus Schlegel, Mus. Pays-Bas, 5, no. 27, 1864, p. 74 (Micronesie) ; 
Salvadori (part), Ornith. Papuasia, 3, 1882, p. 322 (Ualan, Puynipet, Marshalls, 
Mariannis); Wiglesworth (part), Abhandl. und. Ber. Zool. Mus. Dresden, no. 6, 1890- 
1891 (1891), p. 65 (Mulgrave, Taluit, Ualan, Ponape, Ruk, Luganor, Uap, Marianne, 
Pelew) ; Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 8, 1896, p. 41 (Say- 
pan, Guam, Jaluit, Carolines, Palaos) ; Hartert, Novit. Zool., 5, 1898, p. 64 (Guam); 
idem, Novit. Zool. 7, 1900, p. 8 (Ruk); Schnee, Zool. Jahrbucher, 20, 1904, p. 389 
(Marschall-Inseln). 

Actitis incanus Finsch and Hartlaub (part). Fauna Centralpolynesions, 1867, p. 187 
(Mariannen, Ualan, Puynipet); Hartlaub and Finsch, Proc. Zool. Soc. London, 1872, 

10—8131 



146 University of Kansas Publs., Mus. Nat. Hist. 

pp. 89, 106 (Uap, Ualan); GriifTe, Joum. Mus. Godeffroy, 2, 1873, p. 123 (Yap); 
Finsch, Joum. Mus. Godeffroy, 8, 1875, pp. 5, 36 (Palau); idem, Joum. Mus. Godeffroy, 
12, 1876, pp. 18, 38 (Ponape) ; idem, Joum. f. Omith., 1880, pp. 294, 306 (Ponape, 
Kuschai, Marshalls) ; idem. Ibis, 1881, pp. 105, 109, 115 (Kushai, Ponape); Schmeltz 
and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, p. 299 (Mortlock); Finsch, Mitth. 
Ornith. Ver. Wien, 1884, p. 55 (Jaluit, Arno. Kuschai). 

Actitis incana Finsch, Proc. Zool. Soc. London, 1877 (1878), p. 781 (Ponape); idem, 
Proc. Zool. Soc. London, 1880, p. 576 (Ruk) ; idem, Ibis, 1880, pp. 219, 220, 330, 332 
Milli or Mulgrave, Taluit). 

Heteractitis incanus Sharpe, Cat. Birds British Mus., 24, 1906, p. 455 (Oceania); 
Safford, The Plant World, 7, 1904, p. 268 (Guam); Takatsukasa and Kuroda, Tori, 1, 
1915, p. 62 (Yap, Ruk, Ponape, Kusaie) ; Wetmore, in Townsend and Wetmore, Bull. 
Mus. Comp. Zool., 63, 1919, p. 179 (Kusaie); Mathews, Syst. Avium Australasianarum, 
1, 1927, p. 70 (westcentral Pacific). 

Heteroscelus incanus Ridgway, Bull. U. S. Nat. Mus., 50, pt. 8, 1919, p. 367 (Caro- 
lines, Marianas); Peters, Check-list Birds World, 2, 1934, p. 270 (Micronesia); Bryan, 
Guam Rec, vol. 13, no. 2, 1936, p. 24 (Guam): Watson, The Raven, 17, 1946, p. 42 
(Guam); Wharton and Hardcastle (part), Joum. Parasitology, 32, 1946, pp. 296, 316, 
318 (Guam, Peleliu); Downs, Trans. Kansas Acad. Sci., 49, 1946, p. 93 (Tinian) ; 
Strophlet, Auk, 1946, p. 537 (Guam); Wharton, Ecol. Monogr., 16, 1946, pp. 174, 175 
(Guam); Borror, Auk, 1947, p. 417 (Agrihan). 

Tringa incana incana Hartert, Vogel pal. Fauna, 13-14, 1921, p. 1623 (Guam); 
Hand-list Japanese Birds, rev., 1932, p. 191 (Marianas, Carolines, Marshalls, Palaus) ; 
Hand-list Japanese Birds, 3d ed., 1942, p. 214 (Saipan, Guam, Koror, Angaur, Yap, 
Faraulep, Lamatrek, Truk, Ponape, Kusaie, Jaluit, Mille, Arhno, Majuro, Maloelab, 
Wotze, Likieb, Ailuk). 

Heteroscelus incanus incanus Kuroda, in Momiyama, Birds Micronesia, 1922, p. 46 
(Kusaie, Ruk, Ponape, Yap, Marianas, Mulgrave, Taluit, Pelew); Stickney, Amer. Mus. 
Novit., no. 1248, 1943, p. 7 (Guam, Palau, Ponape, Ruk, Kusaie); Mayr, Birds South- 
west Pacific, 1945, p. 42 (Palau, Marianas); Baker, Smithson. Misc. Coll., vol. 107, 
no. 15, 1948, p. 53 (Guam, Rota Ulithi). 

Geographic range. — Breeds in Alaska south to Prince William Sound. Win- 
ters in North and South America and west in Oceania to Melanesia. In Micro- 
nesia: Mariana Islands — Guam, Rota, Saipan. Agrihan; Palau Islands — Angaur, 
Koror; Caroline Islands — Yap, Ulithi, Truk, Faraulep, Lamatrek, Ponape, 
Kusaie; Marshall Islands — Jaluit, Mille, Arhno, Majuro, Maloelab, Wotze, 
Likieb, Ailuk, Bikini. 

Specimens examined. — Total number, 47 (23 males, 20 females, 4 unsexed) as follows: 
Mariana Islands, USNM— Guam, 13 (May 21-29, Sept. 19-27, Oct. 10, 23),— Rota, 2 (Oct. 
23, 25); AMNH— Guam, 4 (April 23, Aug. 16); Palau Islands, AMNH— exact locality not 
given, 1 (no date); Caroline Islands, USNM— Ulithi, 3 (Aug. 20, 22); AMNH— Tmk, 1 
(June 25), — Ponap6, 1 (Dec. 15), — Kusaie, 19 (Feb., Mar., April 1-10); Marshall Islands, 
USNM— Bikini, 3 (Feb. 26, 28, April 28). 

Weights.— In 1948 (1948:53) I listed weights of two males from Guam as 
175 (May) and 109 (September); weights of two females from Guam were 
175 and 192 (both in May). These data were obtained by the NAMRU2 field 
party. 

Parasites.— Wharton and Hardcastle (1946:296, 316, 318) list the following 
chiggers (Acarina) from tattlers taken by NAMRU2 collectors at Guam and 
Peleliu: Neoschongastia bougainvillensis, N. evoingi, N. carveri, and N. namrui. 
Wharton (1946:174, 175) records the chiggers, Acari.scus pluvius and A. anous, 
from tattlers from Guam. It is not certain from which species of Heteroscelus 
these chiggers were obtained. 

Remarks. — Records indicate that the American Wandering Tat- 
tler is a regular visitor to eastern Micronesia, and that it only occas- 
ionally reaches the Palau Islands in western Micronesia. 



Baker: The Avifauna of Micronesia 147 

The NAMRU2 field parties found H. brevipes as singles or in 
small groups of five or less. They remained apart from other species 
and appeared to prefer rocky beaches and coral-reef rocks to the 
sandy beaches. At Guam in 1945, the latest spring migrants were 
taken on May 29. These birds were in nuptial plumage. Birds 
taken at Bikini by Morrison on February 26 and April 28, 1946, 
were in worn, winter plumage. At Guam, the NAMRU2 observers 
obtained the first fall migrants on September 19. These observa- 
tions in 1945, showed that H. incanus arrived at Guam on its south- 
bound flight fully one month after the first individuals of H. brevi- 
pes began to appear (mid-July). This difference may partly result 
from the fact that the distance to the Asiatic breeding grounds of 
H. brevipes is not so great as that to the American breeding grounds 
of H. incanus. 

Whether the two tattlers, H. brevipes and H. incanus, are distinct 
species (allopatric species insofar as breeding ranges are concerned), 
or whether they are mere subspecies (geographic races) is open to 
question. I failed to find evidences of intergradation in the few 
specimens which I examined critically; however, the final answer to 
the problem might be obtained by collecting series of birds from 
breeding grounds where ranges closely approach each other or over- 
lap (if they do). Stickney (1943:6, 7) lists the distinctive differ- 
ences in these two birds, particularly the character of the nasal 
groove, and does not mention having found any evidence of inter- 
gradation. Wetmore (in Townsend and Wetmore, 1919:180) gives 
evidence that they belong to two separate species. 

Arenaria interpres interpres (Linnaeus) 
Turnstone 

Tringa Interpres Linnaeus, Syst. Nat., ed. 10, 1, 1758, p. 148, (Type locality, 
Europe and North America, restricted to Gotland, Sweden.) 

Tringa interpres Quoy and Gaimard, Voy. "Uranie," Zool., 1824, p. 708 (Guam). 

Strepsila coUaris Kittlitz, Obser. Zool., in Lutke, Voy. "Le SeniavLne," 3, 1836, pp. 
287, 299, 304 (Ualan, Lougounor, Guahan) ; idem, Denkw. Raise russ. Amer. Micron, 
und Kamchat., 2, 1858, p. 32 (Ualan). 

StrepsUas interpres Kittlitz, Denk. Reise russ. Amer. Micron, und Kamchat., 2, 
1858, pp. 32, 55, 86 (Ualan); Pelzeln, Reise "Novara," Vogel, 1865, p. 117 (Mari- 
annen) ; Finsch and Hartlaub, Fauna Ornith. Centralpolynesian, 1867, p. 200 (Mari- 
annen) ; Hartlaub, Proc. Zool. Soc. London, 1867 (1868), p. 831 (Pelew) ; Hartlaub 
and Finsch, Proc. Zool. Soc. London, 1868, pp. 8, 118 (Pelew); idem, Proc. Zool. Soc. 
London, 1872, pp. 89, 104 (Pelew, Uap, Mackenzie) ; Griiffe, Joum. Mus. Godeffroy, 
2, 1873, p. 123 (Yap); Finsch, Joum. Mus. Godeffroy, 8, 1875, pp. 5, 32 (Palau); 
idem, Proc. Zool. Soc. London, 1877 (1878), p. 781 (Ponape) ; idem. Ibis, 1880, pp. 
220, 330, 332 (Taluit) ; idem, Joum. f. Omith., 1880, pp. 294, 306 (Ponape, Kuschai); 
idem, Proc. Zool. Soc. London, 1880, p. 576 (Ruk); idem. Ibis, 1881, pp. 105, 109, 
115 (Kushai, Ponape); Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, 
pp. 281, 330, 353 (Ponape, Nukuor, Ruk); Salvadori, Ornith. Papuasia, 3, 1882, p. 
289 (Pelew, Mariannis) ; Finsch, Mitth. Ornith. Ver. Wien, 1884, p. 56 (Jaluit, Ku- 
schai); Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891); 



148 University of Kansas Publs., Mus. Nat. Hist. 

p. 63 (Ualan, Ponape, Luganor, Nukucr, Ruk, Mackenzie, Pelew, Marianne) ; Oustalet, 
Nouv. Arch. Mus. Hist. Nat. Paris, (3), 8, 1896, p. 45 (Guam, Saypan, Hogoleu, Mar- 
shalls, Mackensie, Palaos) ; Hartert, Novit. Zool., 5, 1898, p. 66 (Guam) ; idem, Novit. 
Zool., 7, 1900, p. 9 (Ruk); Takatsukasa and Kuroda, Tori, 1, 1915, p. 51 (Ponape); 
Uchida, Annot. Zool. Japon., 9, 1918, p. 489 (Ponape). 

Cinclus interpres Gray, Cat. Birds Trop. Is. Pacific Ocean, 1859, p. 48 (Ladrones). 

Arenaria interpres Sharpe, Cat. Birds British Mus., 24, 1896, p. 92 (Micronesia); 
Seale, Occ. Papers Bemice P. Bishop Mus., 1, 1901, p. 37 (Guam); Safford, Osprey, 
1902, p. 68 (Marianas); idem. The Plant World, 7, 1904, p. 266 (Guam); Schnee, 
Zool. Jahrbucher, 20, 1904, p. 38'9 (Marshall Islands) ; Safford, Contr. U. S. Nat. Herb., 
9, 1905, p. 80 (Guam), Cox, Island of Guam, 1917, p. 22 (Guam); Wharton and 
Hardcastle, Journ. Parasitology, 32, 1946, pp. 316, 320 (Guam, Peleliu); Downs, Trans. 
Kansas Acad. Sci., 49, 194o, p. 105 (Tinian) ; Strophlet, Auk, 1946, p. 537 (Guam); 
Wharton, Ecol. Monogr., 16, 1946, pp. 174, 175 (Guam); Borror, Auk, 1947, p. 417 
(Agrihan); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 53 (Guam, Rota, 
Peleliu, Truk). 

Are7iaria interprex oahuensis Wetmore, in Townsend and Wetmore, Bull. Mus. Comp. 
Zool., 66, 1919, p. 177 (Jaluit, Rongelab, Uaia); Kuroda, in Momiyama, Birds Micro- 
nesia, 1922, p. 45 (Guam, Saipan, Pelew, Angaur, Kusaie, Ponape, Luganor, Nukuor, 
Ruk, Yap, Mackenzie, Taluit, Rongelab). 

Arenaria interpres interpres Ridgway, Bull. U. S. Nat. Mus., 50, pt. 8, 1919, p. 45 
(Micronesia); Hand-list Japanese Birds, rev., 1932, p. 194 (Guam,, Saipan, Anguar, 
Kusaie, Ponape, Luganor, Nukuor, Ruk, Yap, Mackenzie, Taluit, Rongelab, Mille, 
Majuro, Wotze, Likieb); Bryan, Guam Rec, vol. 13, no. 2, 1936, p. 24 (Guam); Hand- 
list Japanese Birds, 3d ed., 1942, p. 217 (Guam, Saipan, Angaur, Kusaie, Ponape, 
Luganor, Ruk, Yap, Mackenzie, Taluit, Rongelab, Mille, Majuro, Wotze, Likieb); 
Stickney, Amer. Mus. Novit., no. 1248, 1943, p. 8 (Guam, Palau, Ponape, Kusaie). 

Geographic range. — Breeds in northern parts of the Northern Hemisphere. 
Winters to Southern Hemisphere. In Micronesia: Mariana Islands — Guam, 
Rota, Saipan; Palau Islands — Angaur, Peleliu, Caroline Islands — Yap, Ulithi, 
Truk, Lugunor, Nukuor, Ponape, Kuasaie; Marshall Islands — Jaluit, Rongelab, 
Mille, Majuro, Wotze, Likieb, Bikini. 

Specimens examined. — Total number, 36 (17 males, 16 females, 3 unsexed), as follows: 
Mariana Islands. USNM— Guam, 7 (Oct. 10-26)— Rota, 2 (Oct. 20, Nov. 2); AMNH— Guam, 
4 (Mar. 22, 27, Aug. 18); Palau Islands, USNM— Peleliu, 1 (Sept. 8); AMNH— e.xact locality 
not given, 3 (Dec. 8); Caroline Islands, USNM — Truk, 1 (Dec. 22); AMNH— Ponape, 4 
(Dec. 16)— Truk, 4 (Feb. 5, 7, July 14) — Kusaie, 7 (Mar. 10-30); Marshall Islands, USNM— 
Bikini, 8 (Feb. 26, Mar. 4). 

Weights — The NAMRU2 party obtained the weights of four males taken at 
Guam and Rota as 77-99 (92) and one female from Guam as 90. These birda 
were obtained in October and November. 

Parables. — Wharton and Hardcastle (1946:316, 320) list the following chig- 
gers (Acarina) from the Turnstone from Guam and Peleliu: Neoschongastia 
carveri and A^. sirongi. Wharton (1946:174) records also Acariscus anous from 
the Turnstone at Guam. Uchida (1918:489) records the bird louse (Malloph- 
aga), Colpocephalum pediculoides, from this bird at Ponape. 

Remarks. — The Turnstone is a regular visitor to Micronesia and 
to most other parts of Oceania. As pointed out by Stickney (1943:8), 
the material obtained by the Whitney South Sea Expedition yields 
evidence that the population which winters in Oceania is as wide- 
spread as that of Pluvialis dominica fulva but less abundant. The 
writer's observations at Guam, Ulithi and the Palaus are in agree- 
ment with this evidence. Stickney suggests that the reason the 



Baker: The Avifauna of Micronesia 149 

Turnstone was not recorded by the Whitney South Sea Expedition in 
eastern Polynesia was because of "a tendency of the turnstone to hug 
the continental coasts more closely, avoiding extensive overseas mi- 
grations." 

At Guam in 1945, the NAMRU2 party recorded the Turnstone on 
its northward migration as late as March 19 ; on its southward migra- 
tion it was first seen at Guam on July 24. On its southward migra- 
tion the bird was not numerous until September. Our observations 
indicated that in 1945, the principal waves of migration of the Turn- 
stone appeared approximately two weeks after those of the Pacific 
Golden Plover and the Whimbrel. Stickney remarks that the spring 
migratory season in Oceania is completed in May and that the fall 
migratory season begins in August. Borror (1947:417) found small 
flocks on the beaches at Agrihan on August 10 and 11, 1945. 

Bryan and Greenway (1944:112) indicate that the subspecies, 
Arenaria interpres morinella, which breeds in North America, east of 
Point Barrow, Alaska, may reach the Hawaiians. Careful examina- 
tion of specimens from eastern Micronesia might reveal its presence 
there also. The name Areneria interpres oahuensis (Bloxham) may 
apply to specimens from eastern Micronesia but Peters (1934:271) 
considers oahuensis to be inseparable from Arenaria interpres inter- 
pres (Linnaeus). 

Gallinago megala Swinhoe 
Marsh Snipe 

Gallinago ynegala Swinhoe, Ibis, 1861, p. 343. (Type locality. Between Takoo and 
Pekin, China. 

Gallinago heteroeaca Finsch, Joum. Mus. Godeffroy, 8, 1875, pp. 5, 36 (Palau). 

Gallinago megala Salvadori, Ornith. Papuasia, 3, 1882, p. 337 (Pelew) ; Wiglesworth, 
Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 67 (Pelew); 
Sharpe, Cat. Birds British Mus., 24, 1896, p. 624 (Pelew); Hartert, Novit. Zool., 5, 
1898, p. 65 (Guam); Seale, Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 33 (Mari- 
annas); Safford, Osprey, 1902, p. 67 (Mariannas); idem. The Plant World, 7, 1904, p. 
266 (Guam); idem, Contr. U. S. Nat. Herb., 9, 1905, p. 80 (Guam); Cox, Island of 
Guam, 1917, p. 21 (Guam); Hartert, Vogel pal. Fauna, 13-14, 1921, p. 1665 (Palau, 
Guam) ; Mayr, Birds Southwest Pacific, 1945, p. 44 (Guam, Palau) ; Strophlet, Auk, 
63, 1946, p. 537 (Guam); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 54 
(Angaur). 

Subspilura megala Kuroda, in Momiyama, Birds Micronesia, 1922, p. 49 (Guam, 
Pelew). 

Capclla megala Hand-list Japanese Birds, rev., 1932, p. 193 (Guam, Koror) ; Bryan, 
Guam Rec., vol. 13, no. 2, 1936, p. 24 (Guam) ; Robinson and Chasen, Birds Malay 
Peninsula, 3, 1936, p. 170 (Pelew, Marianne); Hand-list Japanese Birds, 3d ed., 1942, 
p. 316 (Guam, Koror). 

Geographic range. — Breeds in east-central Asia. Winters south to Malaysia, 
Australia, and parts of Melanesia. In Micronesia: Mariana Islands — Guam; 
Palau Islands — Koror, Angaur. 

Specimens examined. — One female from Palau Islands, USNM — Angaur (Sept. 21). 



150 University of Kansas Publs., Mus. Nat. Hist. 

Remarks. — The Marsh Snipe is a regular visitor to western Micro- 
nesia, being recorded from the Mariana and Palau islands. At An- 
gaur on September 21, 1945, the NAMRU2 party observed four birds 
at the edge of a brackish water swamp, which was margined with 
reeds and other vegetation. Birds were not seen on tidal beaches at 
Peleliu. Strophlet (1946:537) records the Marsh Snipe at Guam on 
October 21 and December 3, 1945. 

Gallinago gallinago gallinago (Linnaeus) 
Common Snipe 

Scolopax Gallinago Linnaeus, Syst. Nat., ed. 10, 1, 1758, p. 147. (Europe, restricted 
type locality, Sweden.) 

Capella gallinago roddei Takatsukasa and Yamashina, Dobutsu. Zasshi, 44, 1932, 
p. 224 (Saipan). 

Capella gallinago gallinago Hand-list Japanese Birds, rev., 1932, p. 193 (Saipan); 
Hand-list Japanese Birds, 3d ed., 1942, p. 216 (Saipan). 

Gallinago gallinago Mayr, Birds Southwest Pacific, 1945, p. 44 (Saipan). 

Geographic range. — Breeds in northern Eurasia. Winters in southern part of 
breeding range and south to Africa and east to Malaysia. In Micronesia: 
Mariana Islands — Saipan. 

Remarks. — From Micronesia there is a single record of the taking 
of this bird at Saipan, apparently by Japanese collectors. It is prob- 
ably an occasional straggler to the area, but owing to its similarity 
to Gallinago megala it may not often be recognized in the field. 

Crocethia alba (Pallas) 
Sanderling 

Trynga alba Pallas, in Vroeg's Cat., 1764, Adunibr., p. 7. (Type locality. Coast of 
the North Sea.) 

Calidris arenaria Finsch, Ibis, 1880, pp. 331, 332 (Taluit) ; idem, Mitth. Ornith. 
Ver. Wien, 1884, p. 56 (Jaluit) ; Seale, Occ. Papers Bernice P. Bishop Mus., 1, 1901, 
p. 33 (Guam); Safford, Osprey, 1902, p. 70 (Mariannes); idem. The Plant World, 7, 
1904, p. 268 (Guam); Schnee, Zool. Jahrbucher, 20, 1904, p. 390 (Marschall-Inseln). 

Tringa arenaria Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1390- 
1891 (1891), p. 64 (Taluit); Hartert, Novit. Zool., 5, 1898, pp. 65, 69 (Guam). 

Calidris alba Ridgway, Bull. U. S. Nat. Mus., 50, pt. 8, 1919, p. 308 (Marshall 
Islands). 

Crocethia alba Kuroda, in Momiyama, Birds Micronesia, 1922, p. 48 (Taluit, 
Guam); Hand-list Japanese Birds, rev., 1932, p. 193 (Taluit, Guam); Br>'an, Guam 
Rec, vol. 13, no. 2, 1936, p. 24 (Guam) ; Hand-list Japanese Birds, 3d ed., 1942, p. 
215 (Jaluit, Guam); Stickney, Amer. Mus. Novit., no. 1248, 1943, p. 9 (Guam, Ja- 
luit); Mayr, Birds Southwest Pacific, 1945, p. 44 (Marianas, Marshalls) ; Baker, 
Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 54 (Ulithi). 

Geographic range. — Breeds in Arctic regions of the Northern Hemisphere. 
Winters to Southern Hemisphere. In Micronesia: Mariana Islands — Guam; 
Caroline Islands — Ulithi; Marshall Islands — Jaluit. 

Specimens examined. — Total number, 5 (2 males, 3 females), as follows: Mariana Islands, 
AMNH— Guam, 4 (Dec. 2-4); Caroline Islands, USNM, 1 (Aug. 21). 

Remarks. — Stickney (1943:8, 9) summarizes the available infor- 
mation concerning the Sanderling in Oceania. The bird may be 



Baker: The Avifauna of Micronesia 151 

classed as a regular visitor in eastern Micronesia; the most western 
record is from Ulithi in the western Carolines. It has been recorded 
also at Guam and Jaluit. 

The NAMRU2 party secured one Sanderling from a flock of ap- 
proximately thirty birds containing this species and Charadrius 
mongohis stegmanni at Pau Island, Ulithi Atoll, on August 21, 1945. 

Calidris tenuirostris (Horsfield) 
Asiatic Knot 

Totanus tenuirostris Horsfield, Trans. Linn. Soc. London, 13, pt. 1, 1821, p. 192. 
(Type locality, Java.) 

Calidris tenuirostris Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 54 
(Peleliu). 

Geographic range. — Breeds in northeastern Siberia. Winters from India 
east to Malaysia and Australia. In Micronesia: Palau Islands — Peleliu. 
Specimens examined. — Four males from Palau Islands, USNM — Peleliu (Sept. 16). 

Remarks. — The Asiatic Knot was observed and obtained by the 
NAMRU2 party at Peleliu in September, 1945. Flocks containing 
fifteen to twenty birds were noted at the tidal flats of Akarakoro 
Point on September 8 and 16. The birds appeared to remain apart 
from other shore birds in this area. 

Erolia minuta ruficollis (Pallas) 
Little Stint 

Trynga ruficollis Pallas, Reise versch. Prov. Russ. Reicha, 3, l"7fi, p. 700. (Type 
locality, "Circa lacus salsos Dauriae campestris" =: Kulussutai, southern Transbaikalia.) 

Tringa miuuta Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 8, 118 (Pe- 
lew); Gray, Hand-list Birds, pt. 3, 1871, p. 50 (Pelew) ; Hartlaub and Finsch, Proc. 
Zool. Soc. London, 1872, pp. 89, 106 (Pelew) ; Finsch, Journ. Mus. Godeffroy, 8, 1875, 
pp. 5, 36 (Palau). 

Tringa albescens Salvadori, Ornith. Papuasia, 3, 1882, p. 316 (Pelew) ; Wiglesworth, 
Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 64 (Pelew). 

Limonites minuta Takatsukasa and Kudora, Tori, 1, 1915, p. 62 (Pelew). 

Pisobia ruficollis Ridgway, Bull. U. S. Nat. Mus., 50, pt. 8, 1919, p. 290 (Pelew). 

Pisobia minuta ruficollis Kuroda, in Momiyama, Birds Micronesia, 1922, p. 48 
(Palau, Ulithi); Hand-list Japanese Birds, rev., 1932, p. 192 (Palau, Ulithi). 

Calidris ruficollis ruficollis Hand-list Japanese Birds, 3d ed., 1942, p. 215 (Palau, 
Ulithi). 

Calidris minuta ruficollis Mayr, Birds Sauthwest Pacific, 1945, p. 45 (Micronesia); 
Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 54 (Rota, Peleliu). 

Geographic range. — Breeds from northeastern Siberia to northwestern Alaska. 

Winters south from the Malay area to Australia. In Micronesia: Mariana 

Islands — Rota; Palau Islands — Angaur, Peleliu; Caroline Islands — Ulithi. 

Specimens examined. — Total number, 16 (4 males, 12 females), as follows: Mariana Islands, 
USNM— Rota, 1 (Oct. 20); Palau Islands, USNM— Peleliu, 14 (Sept. 6-14)— Angaur, 1 
(Sept. 21). 

Remarks. — The Little Stint is apparently a regular visitor to the 
Palau Islands and a less common visitor to the Mariana Islands. At 
Peleliu and Angaur the NAMRU2 party found these birds in small 



152 University of Kansas Publs., Mus. Nat. Hist. 

flocks of 10 to 15 at tidal flats and at inland ponds. On tidal flats the 
species appeared to remain apart from other kinds of shore birds, but 
at inland ponds the Little Stint was found in company with other 
species. On shooting into a mixed flock of shore birds at an island 
pond at Angaur, the writer secured specimens of this species and also 
of Erolia acuminata. 

Erolia subminuta (Middendorff) 
Least Sandpiper 

Tringa subminuta Middendorff, Reise Nord. und Ost. Siberien, 2, Th. 2, 1853, p. 
222, pi. 19, fig. 6. (Type locality, Western slopes of the Stanovoi Mountains and mouth 
of the Uda.) 

Pisobia minutilla subminuta Hand -list Japanese Birds, rev., 1932, p. 192 (Koror). 

Calidris minutilla subminuta Hand-list Japanese Birds, 3d ed., 1942, p. 215 (Koror); 
Mayr, Birds Southwest Pacific, 1945, p. 45 (Palau). 

Geographic range. — Breeds in northeastern Asia. Winters south to India and 
east to Malaysia. In Micronesia : Palau Islands — Koror. 

Remarks. — The Least Sandpiper has been recorded in the Palau 
Islands by the Japanese investigators. It is probably an uncommon 
visitor to this area. 

Erolia melanotos (Vieillot) 
Pectoral Sandpiper 

Tringa melanotos Vieillot, Nouv. Diet. Hist. Nat., 34, 1819, p. 462. (Type locality, 
Paraguay.) 

Pisobia melanota Hand-list Japanese Birds, rev., 1932, p. 192 (Ponape). 
Calidris melanotos Hand-list Japanese Birds, 3d ed., 1942, p. 215 (Ponape). 
Calidris melanota Mayr, Birds Southwest Pacific, 1945, p. 45 (Ponape). 

Geographic range. — Breeds on the Arctic coast of northeastern Asia and 
eastward into Arctic America. Winters to South America. In Micronesia: 
Caroline Islands — Ponape. 

Remarks. — The Pectoral Sandpiper has been recorded from Po- 
nape. Brj'an and Greenway (1944:114) list the species as an "acci- 
dental" visitor to the Hawaiian Islands from North America. 

Erolia acuminata (Horsfield) 
Sharp-tailed Sandpiper 

Totanus acum-inatus Horsfield, Trans. Linn. Soo. London, 13, pt. 1, 1821, p. 192. 
(Type locality, Java.) 

Tringa acuminata Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 8, 118 
(Pelew); idem, Proc. Zool. Soc. London, 1872, pp. 89, 106 (Pelew); Finsch, Journ. 
Mus. Godeffroy, 8, 1875, pp. 5, 35 (Palau); Salvadori, Ornith. Papuasia, 3, 1882, p. 
314 (Pelew); Wigleswbrth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 
(1891), p. 64 (Pelew); Hartert, Novit. Zool., 5, 1898, p. 65 (Marianne); Seale, Occ. 
Papers Bernice P. Bishop Mus., 1, 1901, p. 33 (Guam); Safford, Osprey, 1902, p. 70 
(Marianas); idem. The Plant World, 7, 1904, p. 268 (Guam). 

Hrteropygia acuminata Sharpe, Cat. Birds British Mus., 24, 1896, p. 566 (Pelew); 
Hartert, Novit. Zool., 7, 1900, p. 8 (Ruk) ; Takatsukasa and Kuroda, Tori, 1, 1915, 



Baker: The Avifauna of Micronesia 153 

p. 62 (Marianas, Ruk, Pelew); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 48 
(Pagan, Pelew, Ruk). 

Tringa maculata var. acwninata Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3) 
8, 1896, p. 44 (Pagan, Palaos). 

Pisobia acuminata Ridgway, Bull. U. S. Nat. Mus., 50, pt. 8, 1919, p. 276 (Caro- 
line Islands). 

Erolia acuminata Hartert, Vogel pal. Fauna, 11-12, 1920, p. 1586 (Palau, Karo- 
linen); Brjan, Guam Rec, vol. 13, no. 2, 1936, p. 24 (Guam). 

Pisobia acuminatus Hand-list Japanese Birds, rev., 1932, p. 192 (Ponape, Truk, 
Pagan, Jaluit, Koror). 

Calidris acum-inata Hand-list Japanese Birds, 3d ed., 1942, p. 215 (Pagan, Jaluit, 
Koror, Truk, Ponape); Mayr, Birds Southwest Pacific, 1945, p. 45 (Micronesia); 
Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 54 (Guam, Angaur). 

Geographic range. — Breeds in northeastern Siberia. Winters from the Ma- 
lay Archipelago and Australia to the Southwest Pacific. In Micronesia: Mari- 
ana Islands — Guam, Pagan; Palau Islands — Angaur; Caroline Islands — Truk, 
Ponape; Marshall Islands — Jaluit. 

Specimens examined. — Total number, 4 (2 males, 2 females), as follows: Mariana Islands, 
USNM— Guam, 1 (Sept. 17); Palau Islands, USNM— Angaur, 3 (Sept. 21). 

Remarks. — The Sharp-tailed San(ipiper is a regular visitor to 
western Micronesia and an uncommon visitor to eastern Micronesia. 
It was first recorded from the Palau Islands in 1868, where the bird 
was taken by Tetens, Heinsohn, and Kubary. In 1896 and 1898, 
records of this bird in the Mariana and Caroline islands were pub- 
lished by Oustalet and Hartert. 

The NAMRU2 party obtained one specimen at Guam on Septem- 
ber 17 and three at Angaur on September 21. At Angaur several 
birds of this species were seen at fresh water ponds in company with 
Erolia minuta ruficollis, Limicola falcinellus sihirica, Tringa glare- 
ola, and other shore birds. 

Erolia ferruginea (Pontoppidan) 
Curlew Sandpiper 

Tringa Jcrrugineus Pontoppidan, Danske Atlas, 1, 17C3, p. 624. (No type locality 
^ Denmark.) 

Calidris ferruginea Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 55 
(Peleliu). 

Geographic range. — Breeds in northern Asia. Winters from Africa east to 
Australia. In Micronesia: Palau Islands — Peleliu. 

Specimens examined. — One female from Palau Islands, USNM — Peleliu (Sept. 6). 

Remarks. — The NAMRU2 party obtained one female on Sep- 
tember 6 at a tidal flat on Peleliu. The Curlew Sandpiper is seem- 
ingly a rare visitor to the Palau Islands from Asia. In using this 
specific name, I am following Mayr (in Delacour and Mayr, 1945: 
107). 



154 University of Kansas Publs., Mus. Nat. Hist. 

Limicola falcinellus sibirica Dresser 
Broad-billed Sandpiper 

Limicola sibirica Dresser, Proc. Zool. Soc. London, 1876, p. 674. (Type locality, 
Siberia and China.) 

Limicola falcinellus sibirica Baker, Sniithson. Misc. Coll., vol. 107, no. 15, 1948, 
p. 55 (Angaur). 

Geographic range .—Breeds in northeastern Asia. Winters from India east 
to Australia. In Micronesia: Palau Islands — Angaur. 

Specimens examined. — One male from Palau Islands, USNM — Angaur (Sept. 21). 

Remarks. — A single male bird was taken by the NAMRU2 party 
at a fresh water pon(i on Angaur Island on September 21, 1945. 
This is the only known record for this bird from Micronesia. 

Phalaropus lobatus (Linnaeus) 
Northern Phalarope 

Tringa lobata Linnaeus, Syst. Nat., ed. 10, 1, 1758, p. 148, in Emendanda, p. 824. 
(Type locality, Hudson Bay.) 

Geographic range. — Breeds throughout Arctic region. Winters at sea in 
tropical and subtropical waters. 

Remarks. — The Northern Phalarope has not been found in Micro- 
nesia. Mayr (1945a :46) records it in the pelagic areas north of the 
New Guinea region. The occurrence there suggests that migration 
is through the Microesian area. 

Larus argentatus vegae Palmen 
Herring Gull 

Larus argentatus Briinn. var. Vegae Palmen, in Nordenskiold, Vega-Exped. Vetensk. 
lakttag., 5, 1887, p. 370. (Type locality, Pidlin, northeastern Siberia.) 

Larus vegae Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 8, 1896, p. 56 
(Agrigan) ; Hartert, Novit. Zool., 5, 1898, p. 68 (Marianne); Scale, Occ. Papers Bemice 
P. Bishop Mus., 1, 1901, p. 20 (Marianas); Safford, Osprey, 1902, p. 70 (Marianas); 
idem. The Plant World, 7, 1904, p. 268 (Guam?). 

Larus vegae Takatsukasa and Kuroda, Tori, 1, 1915, p. 62 (Marianas). 

Larus argentatus vegae Kuroda, in Momiyama, Birds Micronesia, 1922, p. 49 
(Agrigan); Hand-list Japanese Birds, rev., 1932, p. 196 (Agrigan); Hand-list Japanese 
Birds, 3d ed., 1942, p. 220 (Agrigan). 

Geographic range. — Breeds in northern Siberia. Ranges east to Alaska and 
south to the Philippines and the China coast. In Micronesia : Mariana Islands 
— Agrihan. 

Remarks. — The Herring Gull is ascribed to Micronesia on the 
basis of one bird obtained by Marche in January, 1889, at Agrihan 
in the nothern Marianas and reported on by Oustalet (1896:56). 
The gull is considered a straggler to the northern Marianas from the 
northward. Stott (1947:525) observed a gull, which was thought to 
be this species or Larus ridibundus, at Lake Susupe, Saipan, in 1945. 



Baker: The Avifauna of Micronesia 155 

Chlidonias leucopterus (Temminck) 
White-winged Black Tern 

Sterna leucoptera Temminck, Man. d'Ornith., 1815, p. 483. (Type locality. Coasts 
of the Mediterranean.) 

Hydrochelidon leucoptera Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 8, 1896, 
p. 57 (Guam); Hartert, Novit. Zool., 5, 1898, p. 67 (Guam); Seale, Occ. Papers Bernice 
P. Bishop Mus., 1, 1901, p. 20 (Guam); Safford, Osprey, 1902, p. 70 (Marianas); 
idem. The Plant World, 7, 1904, p. 268 (Guam); Hartert, Vogel pal. Fauna, 13-14, 
1921, p. 1686 (Guam); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 51 (Guam). 

Chlidonias leucoptera Hand-list Japanese Birds, rev., 1932, p. 194 (Guam); Hand- 
list Japanese Birds, 3d ed., 1942, p. 217 (Guam); Baker, Smithson. Misc. Coll., vol. 
107, no. 15, 1948, p. 55 (Angaur). 

Geographic range. — Breeds in central and southern Eurasia. Winters from 
Africa east to Australia. In Micronesia: Mariana Islands — Guam; Palau 
Islands — Angaur. 

Measurements. — One adult male has the following measurements: wing, 211; 
tail, 72; exposed culmen, 27; tarsus, 20; one adult female: wing, 210; exposed 
culmen, 25.5. These specimens were taken at the Palau Islands. 

Specimens examined. — Total number, 6 (3 males, 3 females), as follows: Palau Islands, 
USNM— Angaur, 1 (Sept. 21); AMNH— exact locality not given, 5 (Oct. 13). 

Remarks. — The White-winged Black Tern was first collected at 
Guam in October, 1887, by Marche and reported on by Oustalet 
(1896:57). It was later taken at the Palau Islands by Coultas in 
1931, and by the NAMRU2 party at Angaur in 1945. The bird is 
seemingly an uncommon winter visitor to Micronesia. 

At Angaur, the NAMRU2 party obtained one of four terns seen 
at a small fresh water lake. Coultas took five birds at the Palau 
Islands. He writes (field notes) that a flock of 14 of the terns 
appeared at the island following a heavy typhoon. All birds ex- 
amined are in winter plumage (September and October). 

Sterna hirundo longipennis Nordmann 

Black-billed Common Tern 

Sterna longipennis Nordmann, in Erman's Verz. Thier. Pflanz., 1835, p. 17. (Type 
locality. Mouth of the Kutchui River, Sea of Okhotsk.) 

Sterna longipennis Hartlaub and Finsch, Proc. Zool. Soc. London, 1872, pp. 90, 112 
(Pelew) ; Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 5, 41 (Palau) ; Salvadori, Ornith. 
Papuasia, 3, 1882, p. 440 (Pelew); Wiglesworth, Abhandl, und Ber. Zool. Mus. Dres- 
den, no. 6, 1890-1891 (1891), p. 74 (Pelew); Saunders, Cat. Birds British Mus., 25, 
1896, p. 67 (Pelew) ; Takatsukasa and Kuroda, Tori, 1, 1915, p. 62 (Pelew) ; Kuroda, 
in Momiyama, Birds Micronesia, 1922, p. 21 (Pelew). 

Sterna hirundo longipennis Hand-list Japanese Birds, rev., 1932, p. 195 (Palau); 
Hand-list Japanese Birds, 3d ed., 1942, p. 218, (Palau); Mayr, Birds Southwest Pa- 
cific, 1945, p. 25 (Palau). 

Geographic range. — Breeds in northeastern Asia. Winters south to Mela- 
nesia. In Micronesia: Palau Islands — exact locality unknown. 

Remarks. — Finsch (1875:41) states that Heinsohn and Kubary 
obtained specimens of this tern from the Palau Islands for the 



156 University of Kansas Publs., Mus. Nat. Hist. 

Godeffroy Museum. These are the only records for the occurrence 
of the Black-billed Common Tern in Micronesia. 

Sterna sumatrana sumatrana Raffles 
Black-naped Tern 

SterTia Sumatrana Raffles, Trans. Linn. Soc. London, 13, pt. 2, 1822, p. 329. (Type 
locality, Sumatra.) 

Sterna melanauchen Kittlitz, Obser. Zool.. irk Lutke, Voy. "Le Seniavine," 3, 1836, 
pp. 306, 308 (Guahan, Ouleai); Hartlaub and Finsch, Proc. Zool. Soc. London, 1872, 
pp. 90, 113 (Pelew, Uap); Graffe, Journ. Mus. Godeffroy, 2, 1873, p. 123 (Yap); 
Finsch. Journ. Mus. Godeffroy, 8, 1875, pp. 5, 41 (Palau); idem. Ibis, 1880, pp. 220, 
330, 332 (Taluit) ; idem, Journ. f. Ornith., 1880, p. 295 (Ponape) ; idem, Proc. Zool. 
Soc. London, 1880, p. 577 (Ruk); idem, Ibis, 1881, pp. 113, 115 (Ponape); Srlimeltz 
and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, pp. 281, 299, 330, 353 (Ponape, 
Mortlock, Nukuor, Ruk); Salvadori, Ornith. Papuasia, 3, 1882, p. 444 (Pelew, Mac- 
kenzie, Ruk, Ponape, Marshalls) ; Wiglesworth, Abhandl. und Ber. Zool. Mus. Dres- 
den, no. 6, 1890-1891 (1891), p. 74 (Pelew, Uap, Ruk, Luganor, Nukuor, Ponape, 
Taluit); Sanders, Cat. Birds British Mus., 25, 1896, p. 126 (Carolines, Pelews, Mar- 
shalls); Nehrkorn, Kat. Eiers., 1899, p. 222 (Palau); Hartert, Novit. Zool., 7, 1900, 
p. 10 (Ruk); Schnee, Zool. Jahrbiicher, 20, 1904, p. 390 (Marschall-Inseln) ; Taka- 
tsukasa and Kuroda, Tori, 1, 1915, p. 52 (Ruk, Ponape); Uchida, Annot. Zool. Japon., 
9, 1918, pp, 483, 488 (Ponape). 

Sterna sumatrana Wetmore, in Townsend and Wefniore, Bull. Mus. Conip. Zool., 
63, 1919, p. 186 (Arhno). 

Gygisterna sumatrana Kuroda, in Momiyama, Birds Micronesia, 1922, p. 52 (Pelew, 
Mackenzie, Yap, Ruk, Luganor, Nukuor, Ponape, Taluit, Arhno). 

Gygisterna smnatrana sumatrana Kuroda, Avifauna Riu Kiu, 1925, p. 192 (Caro- 
lines, Pelews). 

Sterna sumatrana sumatrana Yamashina, Tori, 7, 1932, p. 410 (Aruuo) ; Hachisuka, 
Birds Philippines, 2, 1932, p. 335 (Caroline, Pelew) ; Hand-list Japanese Birds, rev., 
1932, p. 195 (Palau, Guam, Saipan, Yap, Truk, Lukunor, Nukuoro, Ponape, Jaluit, 
Namu, Arhno, Majuro, Aurh); Peters, Check-list Birds World, 2, 1934, p. 336 (Caro- 
line Islands); Mayr, List New Guinea Birds, 1941, p. 36 (Micronesia); Hand-list Japa- 
nese Birds, 3d ed., 1942, p. 218 (Babelthuap, Koror, Yap, Truk, Lukunor, Nukuoro, 
Ponape, Jaluit, Namu, Arhno, Majuro, Aurh); Mayr, Birds Southwest Pacific, 1945, 
p. 24 (Micronesia); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 55 (Peleliu, 
Ulithi). 

Geographic range. — Micronesia, central Polynesia, northern Australia. Malay- 
sia, west to India, and north to the Riu Kiu Islands. In Micronesia: Palau 
Islands — Babelthuap. Koror, Peleliu; Caroline Islands — Yap, Ulithi, Truk, 
Lukunor, Nukunro, Ponape; Marshall Islands — Jaluit, Namu, Majuro, Aurh, 
Bikini. 

Characters. — Adult: A small tern with a long, forked tail and white plumage 
often with pinkish cast except for mantle, back, rump, tail, wing-coverts, and 
scapulars which are pale pearl-gray ; band across nape, spot in front of eye, and 
outer web of outer primary black; bill and feet black. 

Immature: Resembles adult, but black and white mottling on upper parts. 

Measurements. — Measurements are listed in table 19. 

Specimens examined. — Total number, 15 (8 males, 6 females, 1 female?), as follows: Palau 
Islands, AMNH- — exact locality not given, 4 (Oct. -Dec.); Caroline Islands, USNM — Ulithi 
Atoll, 6 (Aug. 15, 16, 20, 22); AMNH— Truk, 1 (Feb. 10); Marshall Islands, USNM— Bikini, 
4 (March 26, April 30). 

Nesting. — Nehrkorn (1899:222) recorded eggs taken at the Palau Islands. 
Yamashina (1932a :410) listed the finding of three nests containing one egg 



Baker: The Avifauna of Micronesia 157 

each on September 26, 1931, at Arhno in the Marshall Islands. The NAMRU2 
party obtained no evidence of nesting at Ulithi or Palau in August and Sep- 
tember. 1945. Coultas (field notes) obtained reports of the finding of two eggs 
at the Palau Islands in the period October to December, 1931. 

Parasites.— [J chida (1918:483, 488) records the following Mallophaga taken 
at Ponape from this tern: Docophorus albemarlensis, C olpocephalum miUeri, 
and C olpocephalum imperlunum. 

Remarks. — -There are no records for the Black-naped Tern from 
the Mariana Islands, although the species is known from the Palau, 
CaroUne and Marshall Islands. At Ulithi Atoll, the NAMRU2 party 
observed these terns at the islands of Potangeras, Mangejang, Pau, 
and Losiep in August, 1945. They were found in groups of 4 to 15, 
either sitting on sandy beaches or rocky exposures or flying over the 
reefs. Unlike the Crested Tern, tiiese birds appeared quite un- 
afraid of man and would hover over a freshly killed or wounded 
individual of their own kind, making of themselves easy targets. 
The writer saw only one Black-naped Tern at the Palau Islands 
(Peleliu, on September 16, 1945). The birds seem to prefer the 
"low" atolls to the "high" volcanic islands of Micronesia. 

Two subspecies of Sterna swtnatrana are recognized by Peters 
(1934:336) : Sterna sumatrana matheivsi known from islands of the 
western Indian Ocean and Sterna s. sumatrana from islands of Oce- 
ania, Australia, Malaysia, and China coast. There is a considerable 
area separating these subspecies. For populations in the Pacific 
area, other names which have been proposed are Sterna sumatrana 
kempi Mathews for birds from Torres Straits and Gygis decorata 
Hartlaub for birds from the Fiji Islands. A study of 201 specimens 
of this species from various parts of its range (in the collections of 
the American Museum of Natural History and the United States 
National Museum) shows that there is little color variation within 
the species. This observation is the same as that of Mathews (1912: 
372). 

As listed in table 19, measurements of the length of the wing show 
little variation. The length of the tail of birds from localities more 
remote from the continent of Asia (Micronesia, Phoenix, Union, Fiji, 
Samoa, Tonga, and the islands of the Indian Ocean: Aldabra and 
Providence) is, on the average, shorter than the length of the tail of 
birds from islands nearer the Asiatic mainland. This shortness is 
reflected also in the measurement of the difference between the short- 
est and longest tail feather. 



158 



University of Kansas Publs., Mus. Nat. Hist. 



Table 19. Measurements of Specimens of Sterna sumatrana 











Differ- 
ence: 






Locality 


No. 


Wing 


Tail 


Longest 

and 
shortest 

tail 
feather 


Exposed 
culmen 


Tarsus 


S. s. sumatrana 

Micronesia 


13 


221 
211-225 


127 
117-138 


65 
54-79 


37 
35-39 


20.5 
20.0-21.0 


Phoenix and Union 


5 


228 


113 


66 


37 

36-38 


19.5 
18.5-20.0 


Fiji, Samoa, Tonga 


29 


221 
218-229 


131 
122-142 


63 
51-74 


38 
36-41 


20.0 
18.0-21.0 


New Caledonia, 
Loyalty, New 
Hebrides 


8 


224 
221-230 


141 
135-148 


72 
68-81 


39 
37-41 


19.5 
18.5-20.0 


Queensland, Torres 
Straits 


4 


229 


142 
139-148 


78 
71-83 


38 
36-40 


19.5 
18.5-20.0 


Solomons 


52 


227 
220-232 


144 
129-162 


77 
66-95 


36 
34.0-38.5 


19.0 




18.5-20.5 


New Guinea, 
Bismarcks 


10 


224 
219-231 


143 
135-146 


76 
67-81 


34 
32.0 36.5 


19.5 
18.5-20.0 


Malay area 


49 


228 
220-234 


141 
125-153 


74 
63-84 


34 
32.0-37.0 


20.0 
19.0-20.5 


China coast, 
Riu Kiu 


21 


223 
212-234 


144 
130-151 


77 
67-85 


35 
31.5-38,0 


19.5 
19.0-*20.0 


S. S. mathewsi 
Indian Ocean: 
Aldabra, Provi- 


10 


220 


125 


71 


38 
35.0-40.0 


19.0 
18.0-20.0 


dence 











The differences in the length of the exposed culmen of these terns 
shows that birds from islands more remotely oceanic possess longer 
bills than do those from islands closer to the Asiatic continent. Mur- 
phy (1938:538) has written that this phenomenon is characteristic 
among some species which have both continental and insular popula- 
tions (or subspecies). Figure 10 shows the southeastern part of the 
range of the subspecies, Sterna s. sumatrana, and gives the average 
measurements of the exposed culmen of birds from several localities. 



Baker: The Avifauna of Micronesia 



159 



120 



140 




SO 



40 



160 

I 



180 



160 



. C (371 • • . .• -^ '• .» ~. t ^ 



900 1000 
I 




•* 






f. ®-:. 7 



o ^.. ^ 



20 



20 



2000 MILES •,_? 



^^ 



-40 



120 



140 



160 



180 



160 



_^UL 



Fig. 10. Geographic variation in the average length of the exposed culmen 
of Sterna sumatrana sumatrana. 

These localities are given in table 19. Terns with longer bills (37-39) 
were taken in Micronesia, in the Polynesian islands, and in northern 
Australia. Terns with shorter bills (34-36) were taken in Melanesia, 
Malaysia, and the coastal region of China, but there appears to be 
no abrupt line of demarkation between them. Further evidence of 
this tendency may be obtained from the literature. Kuroda (1925: 
191) gives the measurements of the exposed culmen of seven males 
and five females from the Riu Kius as averaging 35 mm. (range 31- 
40.5). It is also of interest to note that the length of the exposed 
culmen of the males averages one to two mm. longer than that of the 
females. The status of Sterna sumatrana mathewsi may be ques- 
tioned. I find no characters separating my series of mostly poor 
specimens. The systematic position of this subspecies from the 
Indian Ocean (and likewise the status of subspecies of other sea birds 
which range into the Indian Ocean) may not be known with cer- 
tainty until additional material is obtained. 



160 University of Kansas Publs., Mus. Nat. Hist. 

Sterna lunata Peale 
Spectacled Tern 

Sterna lunata Peale, U. S. Expl. Exped., 8, 1848, p. 277. (Type locality, Vincennea 
Island, Paumotu Group.) 

Sterna lunata Hartlaub, Proc. Zool. See. London, 1867 (1868), p. 831 (Pelew) ; 
Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 9, 118 (Pelew); idejn, Proc. 
Zool. Soc. London, 1872, pp. 90, 113 (Pelew); Finsch, Joum. Mus. Godeffroy, 8, 1875, 
pp. 5, 41 (Palau); Saunders, Cat. Birds British Mus., 25, 1896, p. 100 (Pelew); Takat- 
sukasa and Kuroda, 1, 1915, p. 62 (Ruk, Pelew); Hand-list Japanese Birds, rev., 1932, 
p. 195 (Palau); Hand-list Japanese Birds, 3d ed., 1942, p. 218 (Palau); Mayr, Birds 
Southwest Pacific, 1945, p. 26 (Micronesia). 

Onychoprion lunatus Salvadori, Ornith. Papuasia, 3, 1882, p. 451 (Pelew) ; Wigles- 
worth, Abhandl. und Bar. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 76 (Pelew). 

Melanostema lunata Kuroda, in Momiyama, Birds Micronesia, 1922, p. 52 (Pelew). 

Geographic range. — Breeds in Oceania from the Hawaiian Group south to 
Fiji and the Tuamotus and west to the Moluccas. In Micronesia : Palau Islands 
— exact locality not known. 

Remarks. — Finsch (1875:41) recorded specimens taken by Tetens, 
Peters and Kubary at the Palau Islands. Coultas obtained one im- 
mature male at sea south of the eastern Caroline Islands a't 1° 25' N 
and 159° E on October 19, 1930. The Spectacled Tern ranges 
throughout the tropical Pacific, spending considerable time at sea, 
and probably reaches most parts of Micronesia in its travels. 

Sterna anaetheta anaetheta Scopoli 
Bridled Tern 

Sterna Anaethetus Scopoli, Del. Flor. et Faun, Insubr., fasc. 2, 1786, p. 92. (Type 
locality, "In Guinea" = Panay, Philippine Islands, ex. Sonnerat.) 

Sterna anaestheta Takatsukasa and Kuroda, Tori, 1, 1915, p. 62 (Pelew). 

Melanostema anaestheta anaestheta Kuroda, in Momiyama, Birds Micronesia, 1922, 
p. 52 (Pelew). 

Sterna anaethetus anaethetus Hand-list Japanese Birds, rev., 1932, p. 195 (Palau); 
Yamashina, Tori, 10, 1940, p. 678 (Bikar) ; Hand-list Japanese Birds, 3d ed., 1942, p. 
218 (Palau. Bikar). 

Sterna anaetheta anaetheta Mayr, Birds Southwest Pacific, 1945, p. 26 (Palau). 

Geographic range. — Breeds from Malaysia to Australia and Oceania and 
north to Formosa. Ranges west to Ceylon and north to Japan. In Micro- 
nesia : Palau Islands — exact locality not known ; Marshall Islands — Bikar. 

Measurements. — Four adult males from the Palau Islands have the follow- 
ing measurements: wing 246-254, longest tail feather 147-177, shortest tail 
feather 71-72, exposed culmen 40-44, tarsus 21-23; one adult female: wing 266, 
exposed culmen 40.5, tarsus 22.5. 

Specimens examined. — Total number, 7 (4 males, 3 females) from Palau Islands, AMNH 
— exact locality not given (Dec. 20). 

Remarks. — The Bridled Tern is known from the Palau Islands 
and from Bikar in the Marshall Islands. In Micronesia, the species 
apparently reaches the northeastern extent of its range. In the 
Palaus, Coultas found the terns on small outlying islands. He ob- 



Baker: The Avifauna of Micronesia 161 

served them to fly to sea early in the day and to return to the 
islands in the evening. Of the seven specimens obtained by him, 
two males and one female had enlarged gonads (Dec. 20). 

Sterna fuscata oahuensis Bloxham 
Sooty Tern 

Sterna Oahuensis Bloxham, Voy. "Blonde," 1826, p. 251. (Type locality, Oahu, 
Hawaiian Islands.) 

Sterna fuliginosa Finsch, Journ. Mus. Godeffroy, 12, 1876, pp. 18, 39 (Ponape) ; 
idem, Proc. Zool. Soc. London, 1877 (1878), p. 781 (Ponape); idem. Joum. f. Omith., 
1880, p. 295 (Ponape) ; Takatsukasa and Kuroda, Tori, 1, 1915, p. 62 (Ponape). 

Onychoprion fuscata infuscata Kuroda, in Momiyama, Birds Micronesia, 1922, p. 
51 (Ponape). 

Sterna fuscata nibilosa Hand-list Japanese Birds, rev., 1932, p. 195 (Ponape) ; 
Yamashina, Tori, 10, 1940, p. 677 (Helen Reef) ; Hand-list Japanese Birds, 3d ed., 
1942, p. 218 (Ponape, Helen Reef). 

Sterna fuscata oahuensis Mayr, Birds Southwest Pacific, 1945, p. 25 (Micronesia). 

Geographic range. — Breeds from the Hawaiian, Marcus, and Benin islands 
south to the Phoenix Islands and Micronesia. In Micronesia: Mariana Is- 
lands — Asuncion; Palau Islands — Helen Reef; Caroline Islands — Ponape. 

Specimens examined. — Total number, 1 unsexed from Mariana Islands, AMNH — .Asuncion 
(Jan. 18). 

Remarks. — The systematic position of the Sooty Tern in Micro- 
nesia is uncertain; in using this name I am following Peters (1934: 
338), who comments that the species "is badly in need of revision." 
Coultas obtained one immature female at 0° 9' S and 159° 50' E, a 
position south of the eastern Caroline Islands. The bird is tenta- 
tively placed in the subspecies S. /. oahuensis. The Sooty Tern 
probably does not breed in large numbers in Micronesia, unless it be 
in the northern Marianas. Bryan (1903:97) reports that this spe- 
cies is very abundant at Marcus Island, which is north and east of 
the Marianas. 

Sterna albifrons sinensis Gmelin 
Least Tern 

Sterna sine7isis Gmelin, Syst. Nat., 1, pt. 2, 1789, p. 608. (Type locality, China, 
ex Latham.) 

Sterna albifrons Marshall, Condor, 51, 1949, p. 221 (Saipan). 

Geographic range. — Found on coastal areas from Korea and China south to 
New Guinea. In Micronesia: Mariana Islands — Saipan. 

Specimens examined. — One female from Mariana Islands, USNM — Saipan (Sept. 26). 

Remarks. — Marshall (1949:221) took one of two Least Terns at 
Lake Susupe on Saipan on September 26, 1945. The specimen 
taken, a female, is in post juvenal molt. 

11—8131 



162 University of Kansas Publs., Mus. Nat. Hist. 

Thalasseus bergii pelecanoides (King) 
Crested Tern 

Sterna pelecanoides King, Surv. Inteitrop. and Western Coasts Australia, 2, 1827, 
p. 422. (Type locality, Torres Strait, northern Queensland.) 

Sterna bergii Finsch, Journ. Mus. Godeffroy, 8, 1875, p. 50 (Palau) ; idem, Proc. 
Zool. Soc. London, 1877 (1878), p. 781 (Ponape) ; idem. Ibis, 1880, pp. 330, 332 (Ratak 
Chain); idem, Journ. f. Ornith., 1880, p. 295 (Ponape); idem, Proc. Zool. Soc. London, 
1880, p. 577 (Ruk); idem. Ibis, 1881, pp. 113, 115 (Ponape); Schmeltz and Krause, 
Ethnogr. Abth. Mus. Godeffroy, 1881, pp. 281, 299, 330, 353 (Ponape, Mortlock, 
Nukuor, Ruk); Salvador!, Ornith. Papuasia, 3, 1882, p. 434 (Ruk, Ponape, Marshalls) ; 
Finsch, Mitth. Ornith. Ver. Wien, 1884, p. 51 (Jaluit); Wiglesworth, Abhandl. und 
Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 74 (Pelew, Luganor, Nukuor, Ruk, 
Ponape, Marshall Islands); Hartert, Novit. Zool., 7, 1900, p. 10 (Ruk); Saunders, Cat. 
Birds British Mus., 25, 1890, p. 89 (Ponape, Mar.shalls) ; Takatsukasa and Kuroda, 
Tori, 1, 1915, p. 52 (Ponape); Uchida, Annot. Zool. Japon., 9, 1918, pp. 483, 488 
(Ponape). 

Sterna bergeri Schnee, Zool. Jahrbiicher, 20, 1904, p. 390 (Marschall-Inseln). 

Sterna bergii cristata Stresemann, Novit. Zool., 21, 1914, p. 58 (Truk). 

Thalasseus bergii pelecanoides Oberholser, Pros. U. S. Nat. Mus., 49, 1915, p. 523 
(Marshall Islands); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 51 (Luganor, 
Nukuor, Ruk, Ponape, Marshall Islands); Kuroda, Avifauna Riu Kiu, 1925, p. 188 
(Marshall Islands); Hand-list Japanese Birds, rev., 1932, p. 194 (Palau, Faraulep, Truk, 
Lukunor, Mukuoro, Ponape, Jaluit, Mille, Aurh, Maloelab, Ailuk) ; Yamashina, Tori, 
10, 1940, p. 677 (Helen Reef, Babelthuap); Hand-list Japanese Birds, 3d ed., 1942, 
p. 218, (Babelthuap, Helen Reef, Faraulep, Truk, Lukunor, Nukuoro, Ponape, Jaluit, 
Mille, Aurh, Maloelab, Ailuk). 

Thalasseus bergii cristatus Peters, Check-list Birds World, 2, 1934, p. 342 (Caro- 
lines, Marshalls); Mayr, Birds Southwest Pacific, 1945, p. 26 (Micronesia); Baker, 
Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 55 (Peleliu, Ngajangel [Kayangel], 
Truk). 

Geographic range. — Malaysia and east coast of Australia south to Tasmania, 
east to Melanesia and Polynesia, north to Phoenix Islands and Microne.sia (see 
figure 11). In Micronesia: Palau Islands — Helen Reef, Peleliu, Babelthuap; 
Caroline Islands — Ulithi, Truk, Faraulep, Lukunor, Nukuoro. Ponape; Marshall 
Islands— Jaluit, Mille, Aurh, Moloelab, Ailuk, Bikini. 

Characters. — Adult: A large, white tern with back, rump, tail, wing-coverts, 
wing, and axillaries pearl gray; outer edges of primaries pearly grayish-black; 
crown black with crest; bill greenish-yellow with blackish base; feet black. 
Crown black, mottled with white and mantle paler in postnuptial plumage. 

Immature: Resembles adult, but crown and back dark, mottled with white 
and crest small. 

Measurements. — Measurements of Crested Terns of the Pacific area are 
listed in table 20. 

Specimens examined. — Total number, 10 (6 males, 4 females), as follows: Caroline Islands, 
USNM— Ulithi, 1 (Aug. 21); AMNH— Truk, 2 (May 7, Dec. 5)--Ponape, 3 (Nov. 1, 7); 
Marshall Islands, USNM— Bikini, 4 (March 4, 11, 12). 

Parasites. — Uchida (1918:483, 488) obtained the following species of bird 
lice (Mallophaga) from the Crested Tern at Ponape: Docojphorus alhemarlen- 
sis and Colpocephalum importunum. 

Remarks. — Oberholser (1915:520-526, pi. 66) lists five subspecies 
{T. b. cristatus, T. b. halodramus, T. b. 'pelecanoides, T. b. rectiros- 
tris, anci T. b. polioceraus) in the region including the coast of 



Baker: The Avifauna of Micronesia 



163 



China, the Riu Kiu Islands, Malaysia, Melanesia, eastern Australia, 
Polynesia, and Micronesia. Only one subspecies, T. b. cristatus, is 
recognized in this area by Stresemann (1914:58), Hartert (1921: 
1695-1696), and Peters (1934:341-342), who mention that there is 
much variation in size and coloring. Measurements, as shown in 
table 20, indicate a wide range of sizes but, in most series, the aver- 

Table 20. Measurements of Thalasseiis bergii in the Pacific Area 



Location 



No. 



Wing 



Longest 

tail 
feather 



Shortest 

tail 
feather 



Exposed 
culmen 



Tarsus 



Thalasseus bergii pelecanoides 



Palaus, Carolines, 
Marshalls 

Christmas, 
Phoenix, 
Tuamotus, 
Society, Fiji, 
Loyalty, 
New Hebrides 

Eastern Australia. . 



New Guinefi, 
Bismarck 
Archipelago, 
Moluccas 

Totals. . . . 



6 



48 



14 



18 



86 



Thalasseus bergii cristatus 
Philippines, 

China, 18 

Formosa, 

Riu Kius 



343 
334-352 



344 
329-362 



34.5 
338-349 



342 
332-361 



344 
329-362 



332 
324-342 



Thalasseus bergii givendolenae 



Western Australia . 



14 



354 
339-369 



168 
153-184 



170 
145-198 



165 
152-174 



168 
144-194 



169 

144-198 



162 
149-182 



171 
162-182 



82 
80-85 



83 
77-92 



88 
84-92 



81 
75-87 



83 
75-92 



81 

78-87 



86 
81-91 



60 
58-65 



58 
54-64 



58 
55-63 



59 
53-64 



58 
53-65 



58 
55-64 



58 
53-65 



27 
25-29 



27 
26-29 



27 
26-28 



27 
25-29 



28 
26-30 



27 
25-29 



ages are nearly the same. Nevertheless, it is evident that birds from 
the coast of China, the Riu Kius, Formosa, and the Philippines have 
a distinctly shorter wing than birds from the Moluccas, Melanesia, 
eastern Australia, Polynesia, and Micronesia. Further evidence of 
this is presented by Kuroda (1925:186) who hsts the measure- 
ments of the wing of eight Crested Terns from the Riu Kiu Islands 
as 322 to 340 (average 330). The occurrence of populations with 



164 



University of Kansas Publs., Mus. Nat. Hist. 



shorter wings has already been pointed out in the work of Oberhol- 
ser (1915:520-526), who divided the short-winged birds into two 
subspecies. It seems advisable to recognize but one subspecies, T. b. 





160 



160 



120 







X 



A 



120 






,-40 

20 



20 

40 



160 



160 



120 



Fig. 11. Geographic distribution of Thalasseus bergii. (1) T. b. bergii; 
(2) T. b. thalassinnics ; (3) T. b. velox; (4) T. b. cristatus; (5) T. b. gwen- 
dolenae; (6) T. b. pelecanoides. 

cristatus, for the birds with short wings and another subspecies, T. 
b. pelecanoides, to include the birds with the longer wings (see fig- 
ure 11). The average measurements of the length of wings of these 
two subspecies, 332, and 344, differ significantly, although there is 
some overlap in measurements. A few specimens at hand from the 
western part of Malaysia are in poor condition and not measurable. 

Most specimens of T. b. cristatus and T. b. pelecanoides have 
lighter-colored upper parts than specimens of T. b. velox, but not 
so light-colored as specimens of T. b. gwendolenae. Size probably 
is a better character than color to use in separating these groups. 

In Micronesia, the NAMRU2 party observed Crested Terns at 
Ulithi, Peleliu and Truk, in August, September, and December, 1945, 
respectively. Birds were seen as singles or in small groups flying 
over the reefs. The birds were wary and difiicult to approach, but 
they were conspicuous and easily identified. 

Procelsterna cerulea saxatilis W. K. Fisher 
Blue-gray Tern 

Procelsterna saxatilis W. K. Fisher, Proc. U. S. Nat. Mus., 26, 1903, p. 5.59. (Type 
locality, Necker Island, Hawaiian Islands.) 

Procelsterna cerulea saxatilis Yaniashina, Tori, 10, 1940, p. 678 (Bikar) ; Hand-list 
Japanese Birds, 3d ed., 1942, p. 219 (Bikar); Mayr, Birds Southwest Pacific, 1945, p. 
27 (Micronesia). 

Geographic range. — Known from Marcus Island and the western Hawaiian 
Islands. In Micronesia: Marshall Islands — Bikar. 



Baker: The Avifauna of Micronesia 165 

Remarks. — Yamashina (1940:678) recorded the taking of eight 
of these terns (5 adult males, 3 adult females) on July 10, 1932, at 
Bikar in the Marshall Islands. He gives the following measure- 
ments: wing, 180.5-188; tail, 104-113.5; exposed culmen, 24-26.5. 
This is the only known record for the species in Micronesia. 

Anous stolidus pileatus (Scopoli) 
Common Noddy 

Sterna pileata Scopoli, Del. Flor. et Faun. Insubr., fasc. 2, 1786, p. 92. (No type 
locality r= Philippines, ex. Sonnerat.) 

Sterna stolida Chamisso, in Kotzebue's Voy. "Rurick," 3, 1821, pp. 150, 157 (Mar- 
shall Islands); Kittlitz, Kupfertaf. Naturgesch. Vogel, 3, 1833, p. 27, pi. 36, fig. 1 
(Mordloks-Inseln) ; idem, Obser. Zool., in Lutke, Voy. "Le Seniavine," 3, 1836, pp. 286, 
299, 308, 309 (Ualan, Lougounor, Ouleai) ; idem, Denkw. Reise russ. Amer. Micron, 
und Kamchat., 1, 1858, p. 364, 2, pp. 77, 86 (Ualan); Wiglesworth, Ibis, 1893, p. 212 
(Marshalls). 

Anous stolidus Hartlaub, Archiv f. Naturgesch., 18, 1852, p. 137 (Mortlock) ; idem, 
Journ. f. Omith., 1854, p. 168 (Carolinen) ; Gray, Cat. Birds Trop. Is. Pacific Ocean, 
1859, p. 59 (Carolines) ; Finsch and Hartlaub, Fauna Centralpolynesiens, 1867, p. 236 
(Mordlocks, Puynipet = Ponape) ; Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, 
pp. 9, 118 (Pelew); idem., Proc. Zool. Soc. London, 1872, pp. 90, 112 (Pelew); Finsch, 
Journ. Mus. Godeffroy, 8, 1875, pp. 6, 42 (Palau); idem, Journ. Mus. Godeffroy, 12, 
1876, pp. 18, 40 (Ponape); idem, Proc. Zool. Soc. London, 1877 (1878), p. 781 (Po- 
nape); idem, Journ. f. Ornith., 1880, pp. 295, 307 (Ponape, Ruck, Kuschai); idem, 
Proc. Zool. Soc. London, 1880, p. 577 (Ruk, Ponape, Kuschai); idem. Ibis, 1881, pp. 
105, 109, 115, 246, 247 (Kuschai, Ponape); Schmeltz and Krause, Ethnogr. Abth. Mus. 
Godeffroy, 1881, pp. 299, 330, 353 (Mortlock, Nukuor, Ruk); Salvadori, Ornith. Papu- 
asia, 3, 1882, p. 455 (Pelews, Carolines, Marshalls); Finsch, Mitth. Omith. Ver. Wien, 
1884, p. 51 (Jaluit, Ponape); Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 
1890-1891 (1891), p. 76 (Pelew, Mortlock, Ruk, Nukuor, Ponape, Ualan, Marshalls); 
Saunders, Cat. Birds British Museum, 25, 1896, p. 136 (Pelew, Carolines, Marshalls); 
Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 8, 1896, p. 59 (Saypan, Guam, Rota, 
Agrigan, Hogoleu =: Truk, Kushai, Ponapi, Marshalls); Hartert, Novit. Zool., 5, 1898, 
p. 68 (Guam); Seale, Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 21 (Guam); 
Safford, Osprey, 1902, p. 66 (Mariannas) ; Bryan, Occ. Papers Bernice P. Bishop Mus., 
2, 1903, p. 101 (Guam); Schnee, Zool. Jahrbucher, 20, 1904, p. 390 (Marshall-Inseln) ; 
Safford, The Plant World, 7, 1904, p. 267 (Guam); idem, Contr. U. S. Nat. Herb., 9, 
1905, p. 80 (Guam); Prowazek, Die deutschen Marianen, 1913, p. 100 (Marianen); 
Takastukasa and Kuroda, Tori, 1, 1915, p. 51 (Ponape, Ruk); Cox, Island of Guam, 
1917, p. 22 (Guam); Uchida, Annot. Zool. Japon., 9, 1918, pp. 484, 488 (Palau, 
Ponape); Wharton, Ecol. Monogr., 16, 1946, p. 174 (Guam); Wharton and Hardcastle, 
Journ. Parasitology-, 32, 1946, pp. 292, 296, 306 (Guam, Ulithi). 

Anous pileatus Pelzeln, Reise "Novara," Vogel, 1865, pp. 155, 162 (Puynipetrr 
Ponape). 

Anous stolidus pileatus Hartert, Novit. Zool., 7, 1900, p. 9 (Ruk); Wetmore, in 
Townsend and Wetmore, Bull. Mus. Comp. Zool., 63, 1919, p. 183 (Kusaie); Kuroda, 
in Momiyama, Birds Micronesia, 1922, p. 49 (Guam, Saipan, Pelew, Mortlock, Ruk, 
Wolea, Nukuoro, Ponape, Kusaie, Marshalls); Hand-list Japanese Birds, rev., 1932, 
p. 195 (Koror, Urukthapel, Angaur, Saipan, Guam, Wolea, Truk, Mortlock, Lukunor, 
Nukuoro, Ponape, Kusaie, Jaluit, Mille, Aurh, Wotze) ; Bryan, Guam Rec, vol. 13, no. 
2, 1936, p. 24 (Guam); Yamashina, Tori, 10, 1940, p. 678 (Assongsong, Babelthuap) ; 
Hand-list Japanese Birds, 3d ed., 1942, p. 219 (Saipan, Assongsong, Guam, Babelthuap, 
Koror, Urukthapel, Peliliu, Angaur, Wolea, Truk, Mortlock, Lukunor, Nukuoro, Ponape, 
Kusaie, Taluit, Mille, Aurh, Wotze); Borror, Auk, 1947, p. 417 (Agrihan); Baker, 
Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 56 (Rota, Guam, Peleliu, Ngabad, 
Ulithi, Truk). 

Anous stolidus unicolorf Ridgway, Bull. U. S. Nat. Mus., 50, pt. 8, 1919, p. 547 
(Guam). 



166 University of Kansas Publs., Mus. Nat. Hist. 

Table 21. Me.asurements of Anoils stolidus of the Paofic Area 



Location 


No. 


Wing 


Tail 


Exposed 
culmen 


Anoiis stolidus ridgwayi 

Isabella, Cocos, Clipperton Islands. . . . 


18 


278 
260-295 


158 
147-166 


41 
38-42 


Anoils stolidus galapagensis 

Galapagos Islands 


11 


277 
2 4-282 


151 
142-160 


40 




38-42 


Anoiis tolidus pileatus 

Hawaiian Islands: Nihoa to Mi way... 


35 


281 
268-299 


162 
149-176 


42 
38-40 


Wake Islands 


8 


278 
273-285 


159 
152-170 


41 




39-43 


Mariana Islands: Guam, Rota 


12 


280 

275-288 


167 
159-187 


41 
39-43 


Palau Islands 


9 


278 
268-283 


161 
155-166 


41 




39-42 


Caroline Islands 


41 


282 
270-291 


164 
: 50-173 


42 




39-45 


Marshall Islands 


3 


282 
270-289 


164 
154-174 


42 




41-43 


Ellice, Phoenix, Danger, Suvarov Islands 


27 


284 
265-295 


162 
152-174 


41 
39-44 


Christmas Island 


13 


287 
280-292 


162 
152-174 


43 




40-46 


Marquesas Islands 


19 


282 
275-291 


163 
155-170 


42 




40-43 


Tuamotu Archipelago 


38 


287 
277-299 


165 
154-173 


42 




39-46 


Society, Austral, Cook, Rapa Islands . . 


16 


290 
2S -301 


164 
155-173 


43 

40-45 


Oeno, Henderson, Ducie, Easter Islands 


6 


293 

285-298 


164 
154-175 


44 
41-45 


Samoa, Fiji, Tonga 


19 


285 
277-295 


164 
153-173 


42 




39 44 


Kermadecs, Norfolk 


23 


276 
269-289 


158 
148-173 


41 




38-43 


New Hebrides, Solomons, New Guinea 
area 


31 


278 
265-287 


158 
150-172 


41 




3 -44 



Baker: The Avifauna of Micronesia 



167 



Table 21. — Concluded 



LocATioisr 



Northwest Australia . 



South China Sea area, Strait of Malacca 



Riu Kius, Japan. 



Indian Ocean area: Seychelles. Aid- 
bra, Providence, Somaliland 



No. 



9 

4 
5 

20 



Wing 



263 

258-267 

271 
262-278 

268 
259-275 



276 
270-286 



Tail 



145 
138-152 

153 
148-257 

148 
143-155 



154 
146-164 



Exposed 
culmen 



40 
38-42 

39 
37-40 

39 
37-40 



41 
39-42 



Geographic range. — Islands in the Indian Ocean east to tropical parts of 
western and central Pacific. In Micronesia: Mariana Islands — Agrihan, Asun- 
cion, Saipan, Rota, Guam; Palau Islands — Kayangel, Babelthuap, Koror, 
Urukthapel, Ngabad, Peleliu, Angaur; Caroline Islands — UHthi, Truk, Wolea, 
Mortlock, Lukunor, Nukuoro, Ponape, Kusaie; Marshall Islands — Jaluit, 
Mille, Aurh, Wotze, Bikini, Kwajalein. 

Characters. — Adult: A large, dark-brown tern witli grayish crown and 
whitish forehead; line above eye white; crescent of white on lower eyelid; 
lores blackish; bill black; feet brownish, iris dark. 

Immature: Resembles adult, but lighter and browner and top of head 
grayish-brown. 

A. s. pileatus resembles A. s. ridgwayi, but darker and less browni.sh, al- 
though not so dark as A. s. galapagensis ; forehead and crown usually duller; 
length of wing and tail average larger (282 and 161) than in A. s. ridgwayi 
(278 and 158) and A. s. galapagensis (277 and 151). 

Measurements. — Measurements of the Common Noddy of the Pacific area 
are listed in table 21. 

Weights.— In 1948 (1948:56) I listed the weights of specimens from Guam 
and Rota as follows: four adult males 187-204 (197); three adult females 177- 
203 (189). 

Specimens examined.— Total number, 92 (43 males, 39 females, 10 unsexed), as follows: 
Mariana Islands, USNM— Guam, 7 (May 24, June 15, July 6, 21)— Rota, 3 (Oct. 18, 24); 
AMNH— Guam, 4 (April 21, 27, Aug. 18)— Asuncion, 1 (Jan. 18); Palau Islands, USNM— 
Peleliu, 2 (Sept. 1)— Ngabad, 1 (Sept. 11); AMNH — exact locality not given, 6 (Nov. 3, 8); 
Caroline Islands, USNM— Ulithi, 3 (Aug. 15)— Kusaie, 1 (Feb. 8); AMNH— Truk, 15 (Feb. 
1, S, 25, March 10, May 6, June 12, 13, Nov. 25, Dec. 25)— Ponape, 20 (Dec. 3. 5, 8, 12, 
15)— Kusaie, 24 (Jan., March 10-30, April 1-10); Marshall Islands, USNM— Bikini, 5 (Feb. 
28, March 2, 19). 

Nesting.— Murphy (1936:1152) writes that the Atlantic subspecies, A. s. 
stolidus, breeds in tropical localities every month of the year, although there 
may be a part of the resident population away at sea at any given time. In 



168 University of Kansas Publs., Mus. Nat. Hist. 

the Pacific area, Kirby (1925:187) found nests "on platforms of sticks built 
on tufts of grass" at Christmas Island in August. In Micronesia, Coultas ob- 
tained young birds at Kusaie in January and April and commented (field 
notes) that they probably nest "spasmodically at all times of the year." At 
Ponape, Coultas observed nests in high trees in December, and birds obtained 
by him in that month had enlarged gonads. At Bikini, Morrison obtained 
eggs on March 2 and 19, and young on March 19. At Palau, Coultas took one 
female tern in postnatal molt on November 8. Adults obtained by him in 
that month had enlarged gonads. At Ulithi, the NAMRU2 party recorded 
one nest containing a single egg on August 21. At the same atoll the 
NAMRU2 party received reports of a large colony of nesting noddys in May 
to July, 1945. In the following August few noddies were seen by the 
NAMRU2 party. McElroy found nests on cliffs and in coconut trees at Truk 
in December, 1945. Hartert (1900:10) reports on eggs taken at Truk in the 
period from March to July 1. The NAMRU2 party observed birds carrying 
nest materials at Peleliu on August 28 but failed to find the nests. At Guam, 
the writer found terns in numbers varying from 4 to 75 in May to July, 1945, 
along the rocky cliffs but no evidence of nesting activity was obtained. 
Strophlet (1946:537) reports that nests may have been present on Orote 
Peninsula at Guam on December 13, 1945. Coultas (field notes) is of the 
opinion that the birds do not nest at Guam but do nest farther north in the 
Marianas. Borror (1947:417) found two colonies at Agrihan on August 10, 
1945. Thus, there are records of nesting in nine months of the year in Micro- 
nesia; although I suspect that the larger flocks of terns have more regular 
breeding habits correlated with their pelagic feeding activities. "Stragglers" 
probably nest irregularly. 

Food habits. — The author (1948:56) records small fish and crustaceans in 
stomachs of terns taken at Ulithi and Peleliu. At Ypao Point, Guam, birds 
were seen to fly back and forth in the day from their roosts on the sea-cliffs. 
On one occasion I saw these birds feeding approximately a half mile from 
shore. 

Paro^^es.— Wharton (1946:174) and Wharton and Hardcastle (1946:292, 
296, 306) list the following species of chiggers (Acarina) from the Common 
Noddy from Guam and Ulithi : Neoschongastia bougainvillensis, N. americana 
solomonis, N. egretta, Acariscus pluvius, and A. anoiis. Uchida (1918:484, 
488) found the bird louse (Mallophaga), Nirmus separatus, on terms at Palau 
and at Ponape he found Colpocephalum milleri on the bird. Bequaert {in 
litt.) has identified a fly (Hippoboscidae) as Oljersia aenescens from a tern 
from Rota. 

Remarks. — Of the Common Noddy Tern of the Pacific area, three 
subspecies are recognized by Peters (1934:346-347). Anoiis stolidus 
ridgwayi is known from islands off the western coast of Mexico and 
Central America; A. s. galapagensis is recorded from the Galapagos 
Archipelago; and A. s. pileatus is found on tropical islands through- 
out the Pacific and west to Madagascar and the African coast in the 
Indian Ocean. These subspecies differ from one another principally 



Baker: The Avifauna of Micronesia 169 

in color, as noted by Ridgway (1919:545) ; A. s. galapagensis is the 
darkest form, A. s. ridgwayi is less blackish and more brownish in 
color of body, and A. s. pileatus is between the two in coloring. A. s. 
pileatus averages larger in length of wing and tail, but these meas- 
urements do not appear to be significant from a taxonomic stand- 
point. 

As shown in table 21, measurements of length of wing for speci- 
mens from throughout most of the Pacific area are almost the same. 
Length of tail is correspondingly uniform. There is a gradual in- 
crease in size of birds in the Tuamotus and Societies and east to 
Easter Island. In this region the average measurement for length of 
wing is 293 millimeters. The lengths of wing and tails are shorter in 
specimens from the Kermadecs and Norfolk Island, which may indi- 
cate relationships with the smaller birds of the Australian area, 
Western Melanesia and possibly Malaysia and the Riu Kiu Islands. 
I am unable to determine the subspecific status of the birds from the 
Kermadecs and Norfolk Island, because of the lack of sufficient ma- 
terial from the Australian region and Malaysia. Possibly Mathews' 
name, A. s. gilberti, is valid for the noddys of Australia and also for 
the birds at Norfolk and the Kermadecs. The small-sized birds of 
the Riu Kiu Islands have been designated as yl. s. pullus by Bangs. 
When specimens from the type locality of ^. s. pileatus in the Phil- 
ippine Islands are available, the true relationships of the populations 
from Micronesia and the other areas in the Pacific can be ascer- 
tained. 

The tern found in the Hawaiians has the palest body and the most 
chalky-white forehead of any of the birds of the Pacfiic. Bryan 
(1903:101) found terns from Marcus Island to agree with specimens 
from Guam and to be "slightly darker" than birds from Midway and 
Laysan in the Hawaiian chain. The birds from the Riu Kius are 
darker and thus similar to the few specimens seen from Malaysia. 
Birds from Polynesia and Melanesia possess the most sooty under- 
parts while those from Micronesia are only slightly less pale. This 
condition also seems to be true for the birds in the Australian area 
and for specimens seen from islands in the Indian Ocean. With fad- 
ing, or wear, or both, there is a change from dusky black to dusky 
brown in the plumage; effort was made by me to compare specimens 
with reltaively similar conditions of plumage. In summary, the sys- 
tematic position of the Common Noddy Terns of the Pacific seem- 
ingly depends on the characteristics of specimens from the type lo- 
cality in the Philippines. When topotypes are available for study, 



170 University of Kansas Publs., Mus. Nat. Hist. 

they may be found to be nearer the darker forms of Malaysia or may 
tend toward the paler, oceanic forms. The Hawaiian population 
probably is distinct. 

In Micronesia the Common Noddy Tern is not a conspicuous bird 
except during its breeding period. Probably it spends most of its life 
at sea, being unlike Gygis alba in this respect. Large flocks seem 
less wary of man than are small groups and singles, which are often 
easily disturbed. Birds of this species appear to prefer the low atolls 
and offshore islets where both tall vegetation and bare ground are 
utilized for nesting or roosting. At Ponape, Coultas (field notes) 
observed the birds to fly to sea at daybreak and to begin to return to 
their roosts by 4:00 pm. Wallace (field notes) observed similar 
activities at Kwajalein in May, 1944, w^here he saw approximately 
forty individuals in a flock with Gygis alba. 

Anoiis stolidus is divided naturally into an Atlantic subspecies, 
which is distinguished by its browner color, and into several sub- 
species which are distinguished by their blacker color in the Pacific 
and Indian oceans. Whether the genus and species evolved in the 
Atlantic or in the Pacific region is not known. If it were the Pacific 
region, the center of differentiation may very well have been the 
islands of Oceania. There, relatively little variation is observable 
within populations covering a large area. To the eastward, birds 
along the American coast are darker or lighter, to the northward, the 
birds of Hawaii are paler, to the southward and southwestward, the 
birds are smaller and to the westward, the birds are smaller and 
darker. The virtual absence of ground-living, predatory animals 
which might prey on nesting colonies has probably been a reason for 
the lack of discrimination by this tern in selecting breeding sites. 
This is probably true of other birds which nest in colonies. 

Anous tenuirostris marcusi (Bryan) 
White-capped Noddy 

Micranous marcusi Bryan, Occ. Papers Bernice P. Bishop Mus., 2, 1903, p. 101. 
(Type locality, Marcus Island.) 

Sterna tenuirostris Kittlitz, Obser. Zool., in Lutke, Voy. "Le Seniavine," 3, 1836, 
pp. 286, 308 (Ualan, Ouleai) ; idem, Denkw. Reise russ. Amer. Micron, und Kamchat., 
2, 1858, p. 64 (Ualan). 

Anoiis tenuirostris Hartlaiib and Finsch, Proc. Zool. Soc. London, 1872, pp. 90, 113 
(Pelew, Carolines); Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 6, 42 (Palau) ; 
Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, pp. 299, 330 (Mortlock, 
Nukuor); Stott, Auk, 64, 1947, p. 526 (Saipan). 

Anous melanogenys Finsch, Proc. Zool. Soc. London, 1877 (1878), p. 781 (Palau); 
idem, Journ. f. Ornith., 1880, pp. 295, 308 (Ponape, Kuschai) ; idem, Ibis, 1880, pp. 
219, 220, 332 (Taluit, Arno) ; idem, Proc. Zool. Soc. London, 1880, p. 577 (Ruk); 
idem. Ibis, 1881, pp. 107, 109, 115 (Kuschai, Ponape); Salvador!, Omith. Papuasia, 3, 
1882, p. 456 (Pelew, Ponape, Marshalls); Finsch, Mitth. Ornith. Ver. Wien, 1884, p. 
52 (Jaluit, Arno, Kuschai) ; Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 



Baker: The Avifauna of Micronesia 171 

6, 1890-1901 (1891), p. 77 (Pelew, Ualan, Ponape, Nukuor, Luganor, Ruk); Hartert, 
Katalog Vogelsamml. Senckenb., 1891, p. 238 (Ualan); Takatsukasa and Kuroda, Tori. 
1, 1915, p. 62 (Ruk); Kuroda, in Moniiyama, Birds Micronesia, 1922, p. 50 (Pelew, 
Ruk, Wolea, Luganor, Nukuor, Ponape, Kusaie, Marshalls). 

Anous leucocapillus Finsch, Proc. Zool. Soc. London, 1877 (1878), p. 781 (Ponape) ; 
Nehrkorn, Journ. f. Ornith., 1879, p. 410 (Ponape) ;Schmeltz and Krause, Ethnogr. 
Abth. Mus. Godeoffry, 1881, p. 281 (Ponape); Finsch, Mitth. Ornith. Ver. Wien, 1884, 
p. 52 (Jaluit); Tristram, Cat. Coll. Birds, 1889, p. 10 (Pelew); Salvador!, Ornith. 
Papuasia, 3, 1882, p. 457 (Pelew); Wiglesworth, Abhandl. und Ber Zool. Mus. Dres- 
den, no. 6, 1890-1891 (1891), p. 77 (Pelew); Oustalet, Nouv. Arch. Mus. Hist. Nat. 
Paris, (3), 8, 1896, p. 60 (Saypan, Palaos, Ruk, Luganor, Nukuor, Ponapi, Kuschai, 
Bonhani) ; Hartert, Novit. Zool., 5, 1898, p. 68 (Marianne); Seale, Occ. Papers Bernice 
P. Bishop Mus., 1, 1901, p. 20 (Saipan?); Safford, Osprey, 1902, p. 66 (Marianas); 
idem, The Plant World, 7, 1904, p. 267 (Guam); Schnee, Zool. Jahrbucher, 20, 1904, 
p. 390 (Marschall-Inseln) ; Safford, Contr. U. S. Nat. Herb., 9, 1905, p. 80 (Guam); 
Ck)x, Island of Guam, 1917, p. 22 (Guam). 

Micranous leucocapillus Saunders, Cat. Birds British Mus., 25, 1896, p. 145 (Pelew, 
Caroline Islands); Nehrkorn, Kat. Eiers., 1899, p. 222 (Kusai) ; Hartert, Novit. Zool., 

7, 1900, p. 9 (Ruk); Takatsukasa and Kuroda, Tori, 1, 1915, p. 51 (Pelew). 

Megalopterus minutus marcusi Mathews, Birds Au.stralia, 2, 1912, p. 423 (Mari- 
anas?); Ridgway, Bull. U. S. Nat. Mus., 50, pt. 8, 1919, p. 553 (Mariannes?); Mat- 
hews, Syst. Avium Australasianarum, 1, 1927, p. 146 (Mariannes); Hachisuka, Birds 
Philippines, 2, 1932, p. 343 (Mariannes). 

Megalopterus tenuirostris leucocapillus Kuroda, in Momiyama, Birds Micronesia, 
1922, p. 50 (Saipan, Pelew, Ruk, Ponape, Kusaie). 

Megalopterus minutus minutus Fisher and Wetmore, Proc. U. S. Nat. Mus., 79, 
1931, p. 45 (Caroline Islands). 

Anous minutus worcesteri Yamashina, Tori, 7, 1932, p. 409 (Coror, Namo, 
Iringlab); Hand-list Japanese Birds, rev., 1932, p. 195 (Saipan, Babelthuap, Koror, 
Truk, Ponape, Kusaie, Ebon, Namorik, Jaluit, Elmore, Mille, Aurh, Wotze, Ailuk) ; 
Yamashina, Tori, 10, 1940, p. 678 (Assongsong, Saipan); Hand-list Japanese Birds, 3d 
ed., 1942, p. 219 (Assongsong, Saipan, Babelthuap, Koror, Peliliu, Truk, Ponape, 
Kusaie, Ebon, Namorik, Jaluit, Elmore, Mille, Aurh, Wotze, Ailuk). 

Anous minutus marcusi Peters, Check-list Birds World, 2, 1934, p. 347 (Caroline 
Islands). 

Anous m.inutus Bequaert, Mushi, 12, 1939, p. 82 (Ponape); idem, Occ. Papers Bernice 
P. Bishop Mus., 16, 1941, p. 253 (Ponape, Palau). 

Anous tenuirostris marcusi Mayr, Birds Southwest Pacific, 1945, p. 27 (Micronesia); 
Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 56 (Peleliu, Ulithi, Truk). 

Geographic range. — Marcus, Wake, and Micronesia. In Micronesia: Mari- 
ana Islands — Asuncion, Saipan, Guam?; Palau Islands — Babelthuap, Koror, 
Peleliu; Caroline Islands — Ulithi, Truk, Ponape, Luganor, Nukuor, Wolea; 
Marshall Islands — Ebon, Namorik, Jaluit, Elmore, Mille, Aurh, Wotze, Ailuk. 

Characters. — Adult: A small tern with sooty-black plumage, grayer on rump 
and tail; forehead and crown white becoming grayer on nape to merge with 
blackish on shoulder; narrow, black superciliary stripe; lores black, lower eye- 
lid with white streak, upper eyelid with white spot. Resembles A. t. melano- 
genys but wing and tail longer and superciliary stripe narrower. Resembles 
A. t. minutus but with narrower,, black superciliary stripe. 

Immature: Resembles adult, but crown more whitish, this coloration ending 
abruptly at nape, with mottling in some birds; plumage of body with brownish 
wash. - ■ - 

Measurements — Measurements are listed in table 22. 

Specimens examined. — Total number, 51 (27 males, 22 females, 2 unsexed), as follows: 
Mariana Islands, AMNH — Asuncion, 1 (Jan. 18); Palau Islands, USNM — Peleliu, 2 (Sept. 9, 
12) ; AMNH— exact locality not given, 2 (Nov. 3) ; Caroline Islands, USNM — Ulithi, 4 (Aug. 
20); AMNH— Truk, 5 (Nov. 16, 21, 22)— Ponape, 15 (Dec. 15)— Kusaie, 17 (Jan. 10, March 
10-30, April 1-10); Marshall Islands, USNM— Bikmi, 4 (May 2, 14); AMNH— no locality 
given, 1 (Sept. 3). 



172 University of Kansas Publs., Mus. Nat. Hist. 

Table 22. Measurements of Anoils tenuirostris of the Pacific Area 



Location 


No. 


Wing 


Tail 


Exposed 
culmen 


Anoils tenuirostris melanogenys 

Hawaiian Islands 


29 


222 
210-229 


113 
105-120 


4^ 




41-48 


Anoiis tenuirostris marcusi 

Wake Islands 


8 


227 
218-231 


118 
112-124 


45 




44^8 


Mariana I lands 


1 
3 


223 

228 
227-228 


117 

122 
117-126 


44 


Palau Islands 


43 




41-45 


Caroline Islands 


32 


?29 
220-240 


120 
113-127 


4 




40-47 


Marshall Islands 


5 


224 
222-229 


118 
114-123 


44 




41-46 


Anoiis tenuirostris minutus 

Christmas Island 


13 


227 
220-234 


120 
108-128 


44 




41-46 


Phoenix, Howland, Union, Danger, 
Suvarov Islands 


9 


229 
226-233 


119 
113-124 


46 




42-48 


Marquesas Islands 


10 


226 
220-23 5 


117 
115-124 


45 




42-48 


Tuamotu Archipelago 


17 


229 
222-234 


118 
112-126 


45 




42-47 


Society, Cook, Austral Islands 


12 


230 
223-238 


118 
114-120 


46 
43-47 


Samoa, Fiji, Tonga Islands 


6 


228 
224-231 


118 
115-121 


44 




42-47 


Kermadec, Norfolk Isl'ds, New Zealand 


15 


226 
21d-235 


116 
112-121 


44 

42-47 


New Hebrides, Solomon, Bismarck, 
Admiralty Islands, New Guinea 


34 


229 
222-237 


117 
109-130 


43 
40-46 


Anoiis tenuirostris diamesus 

Clipperton, Cocos Islands 


14 


230 
224-237 


120 
114-127 


44 




41-47 



Baker: The Avifauna of Micronesia 173 

Nesting. — Few reports have been obtained concerning the nesting of the 
White-capped Noddy in Micronesia. Finsch (1881b: 107) recorded nests, and 
Nehrkom (1899:222) reported on eggs taken at Kusaie. Yamashina (1932a: 
409) recorded the taking of eggs at Koror in the Palau Islands on Januar>' 19 
and November 10 and in the Marshalls at Namo on October 19, and at Iring- 
lab on October 21. No evidence of nestings was obtained by the NAMRU2 
party in 1945, although a number of birds were seen at Ulithi in August. 
Coultas (field notes) writes that a colony of approximately 20 birds began 
nesting about Christmas time on a small offshore island near Ponape. Nests 
were placed in the crotches of limbs of mangroves, 8 to 15 feet above the 
ground. 

Food habits. — The NAMRU2 party found small fish in the stomachs of 
terns taken at Ulithi and Peleliu. 

Parasites. — Bequaert (1939:82 and 1941:253) records the fly (Hippobosci- 
dae), Alfersia aenescens, from the White-capped Noddy taken at Ponape and 
Palau. I 

Remarks. — The subspecies of Anoiis tenuirostris are well differen- 
tiated by color and to a lesser extent by measurements. Table 22 
lists measurements which show that the Hawaiian subspecies, A. t. 
melanogenys, has the shortest wing and the shortest tail whereas the 
subspecies from Cocos and Clipperton islands, A. t. diamesus, has 
the longest wing and the longest tail. The exposed culmen varies in 
length but little among the four subspecies. The systematic position 
of A. t. worcesteri from Cavilli Island in the Sula Sea has not been 
determined because of lack of material. In the third edition of the 
Hand-list of Japanese Birds (Hachisuka et al., 1942:219) the birds 
from Micronesia are referred to A. t. worcesteri as they are also in 
other recent publications by the Japanese. Specimens from the 
Philippines are needed for examination to determine satisfactorily 
the subspecies status of the birds under consideration. 

Field observations indicate that the White-capped Noddy is not 
abundant in the Mariana Islands. According to Oustalet (1896:60) , 
Marche obtained a female at Saipan in June, 1888, and Yamashina 
(1940:678) records five adults from Assongsong (Asuncion). Ows- 
ton's collectors obtained a specimen at Asuncion on January 18, 
1904. In the Palaus, Carolines, and Marshalls birds of this species 
are numerous and have been observed or collected at many of the 
islands. Coultas with the Whitney South Sea Expedition obtained 
specimens at Kusaie, Ponape and Palau. He found them along the 
shores of the large islands and, especially, on the smaller offshore 
islets. At Ulithi Atoll in August, 1945, the NAMRU2 party ob- 
served small flocks of four to ten individuals flying offshore and 
feeding inside the reef. They were frequently observed in company 



174 University of Kansas Publs., Mus. Nat. Hist. 

with Sterna siimatrana. Fewer birds were seen in September, 1945, 
at the Palau Islands by the NAMRU2 party. 

Gygis alba Candida (Gmelin) 
White Tern 

Sterna Candida Gmelin, Syst. Nat., 1, pt. 2, 1789, p. 607. (Type locality, Christ- 
mas Island.) 

Gygis Candida Finsch, Ibis, 1880, p. 220 (Taluit); Saunders (part). Cat. Birds 
British Mus., 25, 1896, p. 149 (Marshalls) ; Schnee, Zool. Jahrbucher, 20, 1904, p. 390 
(Marschall-Inseln). 

Gygis alba Finsch, Ibis, 1880, pp. 330, 332 (Taluit); Wiglesworth (part), Abhandl. 
und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 78 (Marshalls); Oustalet 
(part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 8, 1896, p. 58 (Saypan, Pagan, Agri- 
gan, Marshalls); Safford, Guam, 1912, p. 19 (Guam); Strophlet, Auk, 63, 1946, p. 
537 (Guam); Baker, Condor, 49, 1947, p. 125 (Guam); Stott, Auk, 64, 1947, p. 525 
(Saipan); Baker (part), Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 57 (Guam, 
Rota, Saipan). 

Gygis alba kittlitzi Hartert, Novit. Zool., 5, 1898, p. 67 (Saipan, Guam) ; Seale, 
Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 21 (Guam); Safford, Osprey, 1902, 
66 (Marianas); idem, The Plant World, 7, 1904, p. 267 (Guam); idem, Contr. U. S. 
Nat. Herb., 9, 1905, p. 80 (Guam); Mathews (part). Birds Australia, 2, 1912, p. 443 
(Marianas); Prowazek, Die deutschen Marianen, 1913, p. 100 (Marianaii) ; Cox, Island 
of Guam, 1917, p. 22 (Guam); Ridgway (part), Bull. U. S. Nat. Mus., 50, pt. 8, 
1919, p. 559 (Mariannes)); Kuroda, Avifauna Riu Kiu, 1925, p. 193 (?Mariannes); 
Yamashina, Tori, 7, 1931, p. 410 (Saipan^ ; Yamashina, Tori, 7, 1932, p. 409 (Iringlab, 
Namo, Aruno) ; Hand-list Japanese Birds (part), rev., 1932, p. 196 (Guam, Tinian, 
Saipan, Pagan, Agrigan, Jaluit, Mille, Aurh, Wotze, Likieb, Mejit) ; Yamashina (part). 
Tori, 10, 1940, p. 678 (Assongsong). 

Gygys alba Wheeler, Report Island of Guam, 1900, p. 13 (Guam). 

Gygis albus kittlitzi Kuroda (part), in Momiyama, Birds Micronesia, 1922, p. 50 
(Guam, Saipan, Pagan, Agrigan, Marshalls). 

Leucanous albus kittlitzi Mathews (part), Syst. .\vium Australasianarum, 1, 1927, 
p. 143 (Marianne). 

Gygis alba microrhyncha La Touche (part), Handbook Birds Eastern China, 2, 1933, 
p. 335 (Marianne). 

Gygis alba Candida Bryan, Guam Rec, vol. 13, no. 2, 1936, p. 24 (Guam); Hand- 
list Japanese Birds (part), 3d ed., 1942, p. 219 (Guam, Tinian, Saipan, Pagan, Agrigan, 
Assongsong, Jaluit, Mille, Aurh, Wotze, Likieb, Mejit) ; Downs, Trans. Kansas Acad. 
Sci., 49, 1946, p. 94 (Tinian); Borror, Auk, 64, 1947, p. 417 (Agrihan). 

Geographic range. Northern Pacific from Bonins and Marianas east to 

Wake and Hawaiian Chain, south to Marshall, Phoenix, Christmas and Fan- 
ning islands (see figure 12). In Micronesia: Mariana Islands — Guam, Rota, 
Tinian, Saipan, Pagan, Agrihan; Marshall Islands — Jaluit, Mille, Aurh, Wotze, 
Likieb, Mejit, Eniwetok, Bikini, Kwajalein. 

Characters. — Adult: A small tern with ivory-white plumage except for black, 
narrow, orbital ring; shafts of primary quills dark brown; shafts of tail feathers 
blackish; bill black with bluish base; tarsus dark bluish with yellowish webs; 
iris and skin black. 

Immature : Resembles adult, but with light brown mottlings on upper parts, 
especially on the mantle; feathers softer, bill shorter. 

Measurements. — Measurements are listed on table 23. 

Weights.— The NAMRU2 party obtained weights of 11 adult males from 
Guam and Rota as 110 (97-124); weights of 6 adult females from Guam as 108 
(100-116). These specimens were taken from May to October, 1945. 



Baker: The Avifauna of Micronesia 175 

Specimens examined. — Total number, 4l (23 males, 14 females, 4 unsexed), as follows: 
Mariana Islands, USNM— Guam, 20 (May 24, 29, June 6, 8, 14, 15, 16, 18, 23, July 10, 19, 
20)— Rota, 2 (Oct. 19, 27)— Saipan, 1 (Sept. 26); AMNH— Guam, 4 (March 7, 9, 20)— 
Tinian, 1 (Sept. 8) — Asuncion, 4 (Jan. 1, 18, 25); MCZ — Saipan, 3 (Jan. 7, March 20, April 
17); Marshall Islands, USNM— Bikini, 6 (Feb. 27, March 2, 16, 19). 

Nesting. — Gygis alba does not construct a nest but places its single egg rather 
precariously in the crotch of a branch in a tree (or on rock). In Micronesia 
nesting activities have been observed at various times of the year. Yamashina 
(1932a :409, 410) reported on eggs taken in the Marianas at Saipan on February 
2 and in the Marshalls at Arhno on September 26, at Iringlab on October 21 and 
at Namo on October 19. At Guam a pair of White Terns was seen in a large 
tree on March 27, 1945, by the NAMRU2 observers. Because of their behavior, 
it was suspected that they had an egg or young in the tree. Further inspection 
revealed, on March 31, a downy young sitting in the tree. The young bird was 
attended by the parents until it began to fly on April 17. Hartert (1898:68) 
reports that eggs of the White Tern were taken at Saipan on July 28 and August 
11. Morrison obtained a male nestling on March 16 and eggs on March 22 at 
Bikini in 1946. 

Remarks. — The White Tern is usually restricted to the remote 
islands in the Pacific, Indian and South Atlantic oceans; there, ac- 
cording to the latest treatment, which is that of Peters (1934:348, 
349), six subspecies are recognized. In studying the geographical 
variation of the species, the writer has examined 595 adult specimens, 
including previously unstudied material collected by the Whitney 
South Sea Expedition, which is deposited in the American Museum 
of Natural History. 

This ivory-white species presents an unsual problem in that there 
are few characters available to distinguish the subspecies. IMeasure- 
ments of taxonomic value include those of the wnng, tail, exposed 
culmen, and depth and the shape of the culmen. There appears to be 
no significant secondary sexual difference between males and fe- 
males, and measurements of the two sexes are combined. The chief 
problem within this species seems to hinge on how to classify isolated, 
but relatively similar, populations. The examination of the large 
series of specimens from the Whitney collections has yielded more 
complete information to assist in the solution of this problem. 

Gygis alba alba (Sparrman) of the South Atlantic Ocean (Fer- 
nando de Noronha, South Trinidad, Ascension, and St. Helena 
islands) and G. a. monte Mathews of the Indian Ocean (Seychelles, 
Aldabra, Mascarene and Chagos islands) are isolated populations. 
Specimens examined are those which have previously been studied 
by other workers; measurements are shown in table 23. 

With the exception of G. a. microrhyncha, G. a. monte has the 
smallest average length of wing of all of the subspecies of G. alba 



176 



University of Kansas Publs., Mus. Nat. Hist. 



In G. a. alba the length of wing as well as most of the other measure- 
ments differ but slightly from those of some of the populations in the 
Pacific area although the slender bill of the Atlantic bird is a distinc- 
tive character, as pointed out by Murphy (1936:1166). 

Table 23. Measurements of SuBSPEaES of Gygis alba From the Atlantic and 

Indian Ocean Area 



Subspecies 


No. 


Wing 


Longest 

tail 
feather 


Shortest 

tail 
feather 


Exposed 
culmen 


Depth 
culmen 


Tarsus 


Gygis alba alba . . . 
Gygis alba monte 


24 
35 


246 
239-256 

232 

224-244 


99 
93-111 

106 
98-116 


71 

68-77 

71 
64-81 


40 
35-44 

39 
37-44 


8.0 
7.5-9.0 

8.5 
8.0-8.5 


14.5 
13.0-16.5 

13.5 
12.5-14.0 



The taxonomic position of the White Terns of the Pacific area has 
been one of uncertainty for a long time; as Peters (1934:349) puts it, 
"It is obvious that the last word on the Pacific races of Gygis has not 
yet been said." A principal feature of the problem in this region is 
the presence in the Marquesas of a well-marked subspecies, G. a. 
microrhyncha, virtually surrounded by a wide-ranging and rela- 
tively undifferentiated form, G. a. pacifica (Lesson) (see figure 12). 
The small cormorant {Phalacrocorax melanoleucus brevicauda 
Mayr) from Rennell Island, Solomons, is another example of a 
distinct form surrounded by a widely distributed subspecies. 

In all, 55 adult specimens of G. a. microrhyncha have been exam- 
ined from the following islands in the Marquesas Group: Mukahiva, 
Eiau, Motane, Hivaoa, Uapu, Tahuata, Uahuka, Fatuhiva. The 
measurements are listed in table 24, and show that the White Tern in 
the Marquesas is a much smaller bird than the other subspecies and 
has a shorter bill, wing, and tail. The tail possesses a shallow fork 
as compared with the deeper fork of the tail of other subspecies. In 
addition, the depth of the culmen averages two millimeters less in the 
subspecies in the Marquesas. The presence of a wider, black eye-ring 
is also a distinguishing character in this subspecies. 

Gygis a. microryhncha was for a long time treated as a species 
distinct from G. alba but has recently been considered as a sub- 
species G. alba by Peters and others. On the islands of Hatutu and 
Motane in the Marquesas, the Whitney South Sea Expedition ob- 
tained some birds which appear to be intergrades between the two 



Baker: The Avifauna of Micronesia 



177 



subspecies of White Terns in the area. The measurments of nine 
birds which show intergradation between G. a. microrhyncha and 
G. a. pacifica are listed in table 24. Probably the Marquesas popu- 
lation is tending toward complete reproductive isolation. 

Peters (1934:348, 349) recognizes three other subspecies from 
the Pacific area: G. a. rothschildi Hartert from Laysan, Lisiansky, 
and Krusenstem islands; G. a. Candida (Gmelin) from "the Caro- 
lines east to Christmas Island and south to the Tonga and Society 
Islands"; and G. a. royana Mathews from Norfolk and the Ker- 
madec Islands. Birds from Revilla Gigedo, Cocos and Clipperton 
islands, although geographically isolated, are placed in G. a. Can- 
dida. On the basis of a critical study of specimens at hand, the 
populations in the Pacific fit into three groups. Small birds, G. a. 
Candida, are found in the North Pacific from the Bonins and Mari- 
anas east to Wake and the Hawaiian Chain and south to the Mar- 
shall, Phoenix, Christmas and the Fanning islands (see figure 12). 




Fig. 12. Geographic distribution of Gygis alba in the Pacific area. (1) 
G. a. Candida; (2) G. a. pacifica; (3) G. a. microrhyncha ; (4) G. a. royana. 

Larger birds, G. a. pacifica, are found in the Central Pacific and 
South Pacific from the Carolines in the west southeastward through 
Melanesia and eastward through Samoa, to the Tuamotus and 
Easter to Cocos, Clipperton, and Revilla Gigedo islands. In the 
Southwest Pacific, at Norfolk and the Kermadec Islands, a longer- 
winged populations occurs; it is separable as G. a. royana. The 
measurements of these birds are given in table 24. 

12—8131 



178 University of Kansas Publs., Mus. Nat. Hist. 

Table 24. Measurements op Gygis alba From the Pacific Area 



Location 


No. 


Wing 


Longest 

tail 
feather 


Shortest 

tail 
feather 


Exposed 
culmen 


Depth 
culmen 


Tarsus 


Gygis alba Candida (Gmelin) 
Japan, Bonins 


4 
35 
10 
36 

4 

8 
19 


2.38 


109 


65 


36 
34-38 

38 
36-41 

38 
37-41 

37 
33-40 

39 
38-40 

39 
37-41 

38 
37-42 












Mariana Islands 


237 

227-246 

236 
232-243 

235 
220-246 

234 
231-238 

238 
237-240 

238 
227-242 


111 
98-120 

109 
101-118 

109 
102-118 

111 
107-115 

107 
101-116 

107 
97-119 


69 
61-75 

69 
64-77 

68 
64-74 

71 
70-73 

70 
64-76 

68 
65-72 


9.0 

8.5 
8.0-9.0 


13 


Wake Islands 


12.0-14.0 
13 


Hawaiian Islands 


13.0-14.0 
13 


Marshall Islands 


12.0-14.0 








Phoenix, Howland, Hull, 
Canton Islands 


8.5 


14 






Fanning, Washington, 

Christmas Islands 


8.0 
7.5-9.0 


13 5 




12.0-15.0 


Totals 


116 

33 

12 

20 
13 
29 
37 
118 

54 
10 


236 
220-246 


109 
107-120 


69 
61-77 


38 
33-42 


8.5 
7.5-9.0 


13 




12.0-15.0 


Gygis alba pacifica (Lesson) 
Caroline, Palau Islands 

Bismarck Arch., Solomon 

Islands 


245 
236-253 

247 
242-256 

247 
239-254 

245 
238-252 

247 
241-255 

249 
241-257 

245 
236-252 

247 
240-255 

245 
240-253 


116 
112-125 

116 
105-129 

115 
110-127 

115 
107-118 

114 
104-124 

113 
107-126 

114 
107-127 

113 
106-126 

115 
110-120 


73 

67-76 

74 
68-78 

71 
67-78 

73 
69-78 

73 
65-78 

71 
62-76 

72 
62-82 

73 
63-84 

72 
71-73 


42 
38-44 

42 
39-45 

42 
39-44 

41 
39-42 

42 
40-45 

42 
40-45 

42 
38-46 

41 
40-45 

40 
38-43 


8.5 


13.5 
13.0-13.5 








Samoa, Wallis, Fiji, Tonga, 
Niue Islands 












Line, Danger Islands 












Cook, Austral Islands 












Society Islands 


8.5 
8.0-9.0 


13.5 


Tuamotu Arch 


12.0-14.0 








Rapa, Bass Rocks, Oeno, 
Henderson, Ducie, Pitcairn, 












Clipperton, Cocos Islands 


8.5 
8.5-9.5 


13.5 
13.0-14.0 


Totals 


326 


246 
236-257 


114 
104-129 


72 
62-84 


42 
38-46 


8.5 
8.0-9.5 


13.5 
12.0-14.0 



Baker: The Avifauna of Micronesia 



179 





Table 24.- 


—Concluded 








Location 


No. 

9 
55 

16 
12 
28 


Wing 


Longest 

tail 
feather 


Shortest 

tail 
feather 


Exposed 
culmen 


Depth 
culmen 


Tarsus 


Intergrades between G. a. mi- 
crorhyncha and G. a. pacifica. . 

Gygis alba microrhyncha 

Gygis alba rnyana Mathews 


237 
230-247 

218 
211-235 

250 
242-257 

251 
244-255 


105 
93-122 

78 
72-96 

113 
105-124 

115 
110-121 


74 
67-89 

64 
60-75 

73 

68-79 

75 
71-81 


38 
36-41 

36 
32-39 

42 
41-44 

43 
40-46 


7.5 
7.0-8.0 

6.5 
6.0-8.0 


13.0 
12.0-14.0 

12.0 
11.0-12.5 


























Totals 


250 
242-257 


114 
105-124 


74 
68-81 


42 
40-46 



















The measurements indicate that there is a gradient in size from 
small in the north to large in the south ; however, there is a definite 
separation in average measurements — ten millimeters in length of 
wing and four millimeters in length of exposed culmen — between the 
two populations which are designated as G. a. Candida and G. a. 
pacifica. In studying material from Micronesia and the Hawaiian 
Islands, I (1948:57) pointed out the similarities between birds of 
the Marianas and the Hawaiians and separated these from terns 
found in the Caroline Islands. The systematic position of the White 
Tern in the Gilbert and Ellice islands will remain in doubt until 
specimens are available for examination. 

G. a. royana is provisionally retained as the name for the Fairy 
Tern of the Kermadecs and Norfolk Island; there is considerable 
overlap in measurements between G. a. royana and G. a. pacifica. 
Measurements have given evidence of the degrees of structural re- 
semblance of the White Terns of the different islands, but it is not 
certain that the groupings made on this basis are natural; more 
data is needed on ecology and life history. Of particular importance 
is to learn whether these birds fly regularly from island to island. 
On the basis of eleven months of rather continuous observation in 
Micronesia, I suspect that the White Tern has little tendency to 
make inter-island migrations. This might account for the differ- 
ences in size in the populations at Guam in the Marianas {G. a. 
Candida) and at Ulithi in the Carolines (G. a. pacifica) where only 
approximately 400 miles of open water separate the tw^o islands. 
The occurrence of the distinct G. a. microrhyncha in the Marquesas 
may be accounted for by such nonmigratory behavior, Mayr 



180 University of Kansas Publs., Mus. Nat. Hist. 

(1945a:27), however, is of the opinion that White Terns found in 
the Bismarck Archipelago, the Solomons, Santa Cruz and New Heb- 
rides islands may not breed there, which is another way of saying 
that they are migrants. Swarth (1934:221) and Murphy (1936: 
1268) record the wandering of the White Tern to the Galapagos Is- 
lands, probably from breeding grounds at Cocos Island. Swarth 
suggests that the tern is not established at the Galapagos because 
of the presence of colder water in the area. Murphy (1936:1166) 
is of the opinion that the South Atlantic White Terns are sedentary, 
but reports evidence of pelagic migration in the Pacific at the Ker- 
madecs. The fact that G. alba is restricted in its distribution to 
widely separated groups of islands in tropical and subtropical areas 
of the South Atlantic, Indian and Pacific oceans may indicate that 
the birds at one time had a more extensive range than at present, 
probably including even coastal regions of the continents and large 
continental islands. 

Gygis alba pacifica (Lesson) 

White Tern 

Sterna pacifica Lesson, Ann. Sci. Nat., 4, 1825, p. 101. (Type locality, Society 
Islands, Paumotu Islands, and Bora Bora.) 

Sterna alba Kittlitz, Kupfertaf. Naturgesch. Vogel, 3, 1833, p. 28 (Carolinen); 
idem, Obser. Zool., in Lutke, Voy. "Le Seniavine," 3, 1836, pp. 286, 299, 308 (Ualan, 
Lougounor, Ouleai). 

Gygis Candida Hartlaub, Archiv f. Naturgesch., 18, 1852, p. 137 (Carolinen); 
Hartlaub, Journ. f. Omith., 1854, p. 168 (Carolinen); Kittlitz, Denkw. Reise russ. 
Amer. Micron, und Kamchat., 1, 1858, p. 382, 2, 1858, pp. 39, 60 (Ualan); Gray, Cat. 
Birds Trop. Is. Pacific Ocean, 1859, p. 59 (Caroline Islands); Saunders (part), Cat. 
Birds British Mus., 25, 1896, p. 149 (Pelew, Carolines); Takatsukasa and Kuroda, Tori, 
1, 1915, p. 51 (Ruk, Pelew). 

Gygis alba Finsch and Hartlaub, Fauna Centralpolynesiens, 1867, p. 233 (Carolinen); 
Hartlaub, Proc. Zool. Soc. London, 1867 (1868), p. 832 (Pelew); Hartlaub and Finsch, 
Proc. Zool. Soc. London, 1868, pp. 9, 118 (Pelew); Finsch and Hartlaub, Journ. f. 
Ornith., 1870, p. 140 (Pelew); Hartlaub and Finsch, Proc. Zool. Soc. London, 1872, pp. 
90, 114 (Pelew, Uap, Ualan); Griiffe, Journ. Mus. Godeffroy, 2, 1873, p. 123 (Yap); 
Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 6, 43 (Palau); idem, Journ. Mus. Godeffroy, 
12, 1876, pp. 18, 40 (Ponape); idem, Proc. Zool. Soc. London, 1877 (1878), p. 782 
(Ponape); idem., Journ. f. Ornith., 1880, pp. 295, 309 (Ponape, Kuschai); idem, Proc. 
Zool. Soc. London, 1880, p. 577 (Ruk); idem, Ibis, 1881, pp. 105, 106, 109, 115, 246, 
247 (Kushai, Ponape); Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, pp. 
299, 330, 353 (Mortlock, Nukuor, Ruk); Finsch, Mitth. Ornith. Ver. Wien, 1884, p. 52 
(Kuschai); Wiglesworth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890- 
1891 (1891), p. 78 (Pelew, Uap, Luganor, Nukuor, Ruk, Ponape, Ualan); Oustalet 
(part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 8, 1896, p. 58 (Palaos, Carolines; 
Baker (part), Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 57 (Peleliu, Ulithi, Truk, 
Kusaie). 

Gygis alba kittlitzi Hartert, Katalog Vogelsamml. Senckenb., 1891, p. 237 (Type 
; locality, Ulea = Wolea) ; idem, Novit. Zool., 7, 1900, p. 10 (Ruk); Dubois, Syn. 

Avium, 2, 1904, p. 1020 (Carolines); Mathews (part). Birds Australia, 2, 1912, p. 443 
\ (Carolines); Ridgway (part), Bull. U. S. Nat. Mus., 50, pt. 8, 1919, p. 559 (Caro- 

lines); Kuroda (part), Avifauna Riu Kiu, 1925, p. 193 (Carolines); Hand-list Japanese 
Birds (part), rev., 1932, p. 196 (Pelew, Yap, Wolea, Luganor, Ruk, Ponape, Kusaie); 
Yamashina (part). Tori, 10, 1940, p. 678 (Babelthuap). 



Baker: The Avifauna of Micronesia 181 

Gygii albus kittlitzi Kuroda (part), in Momiyama, Birds Micronesia, 1922, p. 50 
(Pelews, Yap, Wolea, Luganor, Nukuor. Ruk, Ponape, Kusaie). 

Leucanous albus kittlitzi (Mathews (part), Syst. Avium Australasianarum, 1, 1927, 
p. 143 (Carolines). 

Gygi3 alba Candida Peters, Check-list Birds World, 2, 1934, p. 349 (Carolines); 
Hand-list Japanese Birds (part), 3d ed., 1942, p. 219 (Babelthuap, Koror, Angaur, Yap, 
Wolea, Truk, Lukunor, Nukuoro, Ponape, Kusaie). 

Geographic range. — Central and southern Pacific from Carolines southeast 
through Melanesia and east through Samoa to Tuamotus, Easter to Cocos and 
Clipperton (see figure 12). In Micronesia: Palau Islands — Angaur, Peleliu, 
Garakayo, Koror, Babelthuap, Kayangel; Caroline Islands — Yap, Ulithi, Wolea, 
Truk, Lukunor, Ponape, Kusaie. 

Characters.— Resembles G. a. Candida, but size larger, wing length of adult 
males and females 236-253 (245) ; length of exposed culmen 38-44 (42). 

Measurements. — Measurements are listed in table 24. 

Specimens examined. — Total number, 36 (22 males, 12 females, 2 unsexed), as follows: 
Palau Islands, USNM — Peleliu, 1 (Sept. 1); AMNH — exact locality not given, 1 (Nov. 13); 
Caroline Islands, USNM— Ulithi, 12 (Aug. 14, 15, 16, 20, 21)— Truk, 1 (Dec. 13); AMNH 
—Truk, 7 (Mar. 8, May 7, June 8, Nov. 11, 26) — Ponape, 1 (undated) — Kusaie, 10 (Jan., 
Feb., March 20-30, April 1-10); MCZ — Yap, 3 (Jan. 13). 

Nesting. — The NAMRU2 party learned that in May and June, 1945, several 
young White Terns were seen at Asor, Ulithi Atoll, by service personnel. 
These young were observed in breadfruit trees within a recreational area; the 
presence of the service personnel seemingly had little disturbing effect on the 
terns. At Bulubul. another island of this atoll, a downy young was obtained 
on August 22. Hartert (1900:10) reports that eggs of the White Tern were 
found on the ground and in forks of branches of trees at Truk in June. 

Food Habits. — The author (1948:58) reports that stomachs of birds taken 
at Ulithi and Peleliu contained fish, insects and marine crustaceans. Probably 
the birds feed to a large extent along the edge of the tidal reef. They almost 
certainly obtain food also on the islands as indicated by the presence of insects 
in stomach contents; this is not surprising since the birds frequent woodland 
habitats. 

Rc?7iarks. — Gygis alba is one of the most characteristic birds in 
Micronesia. It is seemingly more numerous at the coral atolls than 
at the high, volcanic islands. At the latter islands the birds prefer 
the coastal coconut grove environment. At Pau and Bulubul, two 
small islands in the Ulithi Atoll, the WTiter counted approximately 
100 birds on August 21, 1945. Kittlitz was the first to publish an 
account of these birds in the Caroline Islands. Tetens, Peters, Sem- 
per and Kubary reported their presence in the Palaus. No doubt, 
these terns attract the attention of every traveler in the islands 
owing to their conspicuously white beauty and their seemingly 
friendly behavior toward man. Their habit of hovering in small 
flocks close over the head of the observer is indeed spectacular. 



182 University of Kansas Publs., Mus. Nat. Hist. 

Columba Hvia Gmelin 
Blue Rock Pis;eon 

Columba domestica P livia Gmelin, Syst. Nat., 1, pt. 2, 1789, p. 769. (No type 
locality = Europe.) 

Columba livia Bryan, Guam Rec, vol. 13, no. 2, 1936, p. 24 (Guam) ; Marshall, 
Condor, vol. 51, 1949, p. 221 (Tinian). 

Geographic range. — Europe and Asia Minor. Introduced to many parts of 
the world. In Micronesia: Mariana Islands — Guam, Tinian. 

Remarks. — In 1945, the NAMRU2 party observed pigeons about 
the towns on Guam, particularly at the town of Inarajan. Bryan 
(1936:24) writes that the birds were introduced by the United 
States Navy and Marine Corps at Guam ; the stock originating from 
escaped carrier pigeons. Marshall (1949:221) records this bird 
from Tinian. 

Ptilinopus porphyraceus ponapensis (Finsch) 
Crimson-crowned Fruit Dove 

Ptilonopus ponapensis Finsch, Proc. Zool. Soc. London, 1877 (1878), p. 779. (Type 
locality, Ponape.) 

Ptilinopus? fasciatus Finsch, Journ. Mus. Godeffroy, 12, 1876, pp. 18, 37 Ponape). 

Ptilopus fasciatus Elliot, Proc. Zool. Soc. London, 1878, p. 536 (Ponape) ; Tristram, 
Cat. Birds, 1889, p. 44 (Ponape). 

Ptilopus ponapensis Schmeltz, Verhandl. Ver. nat. Unterhatlung Hamburg, 1877 
(1879), pp. 178, 179 (Ponape); Finsch, Proc. Zool. Soc. London, 1880, p. 576 (Rule, 
Ponape); idem, Journ. f. Ornith., 1880, pp. 291, 303 (Ponape); idem, Proc. Zool. 
Soc. London, 1880, p. 578 (Ruk, Ponape): idtm, Ibis, 1881, pp. 113, 115 (Ponape); 
Wiglesworth, Ibis, 1891, p. 583 (Ponape, Rojk) ; idem, Abhandl. und Ber. Zool. Mus. 
Dresden, no. 6, 1890-1891 (1891), p. 50 (Ponape, Ruk); Salvador!, Cat. Birds 
British Mus., 21, 1893, p. 93 (Ponape, Ruk); Oustalet, Nouv. Arch. Mus. Hist. Nat., 
Paris, (3), 7, 1895, p. 222 (Ponape); Nehrkron, Kat. Eiers., 1899, p. 180 (Ruk); 
Dubois, Syn. Avium, 2, 1904, p. 736 (Ruck, Ponape); Reichenow, Die Vogel, 1, 
1913, p. 354 (Ruk, Ponape); Takatsukasa and Kuroda, Tori, 1, 1915, p. 52 (Ruk, 
Ponape); Wetmore, in Townsend and Wetniore, Bull. Mus. Comp. Zool., 63, 1919, 
p. 189 (Uala, Ponape). 

Ptilinopus ponapensis Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, 
p. 353 (Ruk); Hartert, Novit. Zool., 7, 1900, p. 7 (Ruk, Ponape); Seale, Occ. 
Papers Bernice P. Bishop Mus., 1, 1901, p. 42 (Ponape); Matschie, Journ. f. Ornith., 
1901, p. 113 (Ruck, Ponape) ; Mathews, Syst. Avium Australasianarum, 1, 1927, 
p. 32 (Ponape); Bequaert, Mushi, 12, 1939, pp. 81, 82 (Ponape); Mayr. Proc. 6th 
Pacific Sci. Congr., 4, 1939 (1941), p. 204 (Ponape); Bequaert, Occ. Papers Bernice 
P. Bishop Mus., 16, 1941, pp. 266, 290 (Ponape). 

Ptilnopus Ponapensis Christian, The Caroline Islands, 1899, p. 357 (Ponape). 

Ptilinopus ponepensis ponapensis Kuroda, in Moiniyama, Birds Micronesia, 1922, 
p. 57 (Ponape, Ruk); Yamashina, Tori, 7, 1932, p. 408 (Ponape); Hand-list Japanese 
Birds, rev., 1932, p. 190 (Ponape, Ruk); Peters, Check-list Birds World, 3, 1937, 
p. 31 (Ruk, Ponape); Hand-list Japanese Birds, 3d ed., 1942, p. 213 (Ponape, Truk). 

Ptilinopus porphyraceus ponapensis Ripley and Birckhead, Amer. Mus. Novit., no. 
1192, 1942, p. 7 (Ruk, Ponapi-) ; Mayr, Birds Southwest Pacific, 1945, p. 289 (Truk, 
Ponape); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 59 (Truk). 

Geographic range. — Micronesia: Caroline Islands — Truk, Ponape. 

Characters. — Adult male: A green fruit dove with forehead, anterior lores 
and crown near "pansy purple," faintly margined with j'ellow; occiput, sides 
of head, neck, upper breast grayish-green with bifid feathers of midbreast more 



Baker: The Avifauna of Micronesia 



183 



olivaceous; chin and midthroat light yellow; breast, sides and tibia green; 
midpart of lower breast dark bluish-green, tinged with dark purple; lower 
abdomen, vent, and undertail 3^ellow, under tail-coverts deeper yellow tinged 
with orange; upper parts dark green; wings metallic green on outer webs and 
tips, inner secondaries and some posterior scapulars with purple spots near 
tips; priraarieis and secondaries edged on outer webs with yellowish; under- 
wing gray with yellow edges on hind, under wing-coverts; upper side of tail 
metallic green with terminal, broad yellow band; under side of tail gray; bill 
lead-colored, feet wine-brown, iris whitish to pale brown. Adult female re- 
sembles adut male, but slightly smaller and duller. 

Immature : Resembles adult, but entirely green with yellow edgings on 
feathers and lacking crimson crowTi and colored breast patch. 

Measurements. — Measurements of subspecies of P. -porphyraceus in Micro- 
nesia are presented in table 25. 

Table 25. Me-asuremexts of Ptilinopiis porphyraceus in Micronesia 



Subspecies 


Number 


Wing 


Exposed 
cu men 


Tarsus 


P. p. ponapensis .... 

P. p. hernsheimi 

P. p. pelewensis 


12 males 
11 females 

6 males 
5 females 

10 males 
4 females 


137 (133-141) 

133 (126-137) 

134 (130-138) 
127 (125-130) 

133 (131-134) 
133 (130-138) 


14 (13-15) 

14 (13-15) 

13 (12-14) 
13 (12-13) 

15 (13-15) 
15 (14-15) 


25 (24-27) 
25 (24-26) 

25 (24-26) 
25 (24-25) 

25 (23-26) 
24 (23-24) 



Specimens examined. — Total number, 81 (52 males, 26 females, 3 unsexed), as follows: 
Caroline Islands, USNM— Truk, 4 (Feb. 16, Dec. 24); AMNH— Truk, 24 (Jan., June, Oct.)— 
Ponape, 53 (Nov., Dec). 

Nesting. — Yamashina ( 1932a :408) reports on eggs taken at Ponape on the 
following dates: July 10, 12, August 1, 12, 15, 21. Only one egg was found to a 
ne.st. Hartert (1900:8) records nests containing eggs in May and June at Truk. 
Coultas (field notes) describes the nest as a flimsy affair. At Ponape in No- 
vember and December he found nests on low branches (10 to 20 feet from the 
ground) each containing a single egg. Nests were found also in the tops of tree 
ferns. Females taken in these months had enlarged gonads. 

Par(m7es.— Bequaert (1939:81, 82, and 1941:266, 290) records the two flies 
(Hippoboscidae), Omithoctona plicata and 0. pusilla, from the fruit dove at 
Ponape. 

Remarks. — McElroy of the NAIMRU2 party found the birds in 
mountainous areas at Truk in December, 1945. At Ponape in No- 
vember and December, 1931, Coultas (field notes) comments that 
the bird is rapidly disappearing owing to persistent hunting by the 
natives and, at that time, by the Japanese. He found the birds to be 
strictly forest-living and to frequent the larger fruit-bearing trees of 



184 University of Kansas Publs., Mus. Nat. Hist, 

the lowlands and the mountain sides. Coultas writes that the Japa- 
nese hunters attracted the doves by the use of calls. The natives 
catch the birds with a gum mixture obtained from bread-fruit gum 
and coconut oil. 

Ptilinopus porphyraceus hernsheimi (Finsch) 
Crimson-crowned Fruit Dove 

Ptilopus Hernsheimi Finsch., Joum. f. Ornith., 1880, p. 303. (Type locality, 
Kuschai.) 

Ptilopus hernsheimi Finsch, Proc. Zool. Soc. London, 1880, p. 577 (Kuschai); 
Reichenow and Schalow, Journ. f. Ornith., 1881, p. 75 (Kuschai); Finsch, Ibis, 1881, 
pp. 106, 107, 108 (Kushai); Wiglesworth, Ibis, 1891, p. 583 (Ualan) ; idem, Abhandl. 
und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 51 (Ualan); Salvador!, Cat. 
Birds British Mus., 21, 1893, p. 94 (Ualan); Oustalet, Nouv. Arch. Mus. Hist. Nat., 
Paris, (3), 7, 1895, p. 222 (Oualan) ; Dubois, Syn. Avium, 2, 1904, p. 736 (Kuschai); 
Reichenow, Die Vogel, 1, 1913, p. 355 (Kuschai); Wetmore, in Townsend and Wetmore, 
Bull. Mus. Comp. Zool., 63, 1919, p. 189 (Kusaie). 

Ptilinopus hernsheimi Matschie, Journ. f. Ornith., 1901, p. 113 (Ualan); Mathews, 
Syst. Avium Australasianarum, 1, 1927, p. 33 (Kusaie). 

Ptilinopus ponapensis hernsheimi Kuroda, in Momoyama, Birds Micronesia, 1922, p. 
57 (Kusaie); Hand-list Japanese Birds, rev., 1932, p. 190 (Kusaie); Peters, Check- 
list Birds World, 3, 1937, p. 31 (Kusaie); Hand-list Japanese Birds, 3d ed., 1942, p. 
212 (Kusaie). 

Ptilinopus marshallianus Peters and Griscom, Proc. New England Zool. Club, 10, 
1928, p. 104 (Type locality, Ebon); Hand-list Japanese Birds, rev., 1932, p. 190 
(Ebon). 

Ptilinopus ponapensis marshallianus Peters, Check-list Birds World, 3, 1937, p. 31 
(Ebon); Hand-list Japanese Birds, 3d ed., 1942, p. 213 (Ebon). 

Ptilinopus porphyraceus hernsheimi Ripley and Birckhead, Amer. Mus. Novit., no. 
1192, 1942, p. 6 (Kusaie, Ebon); Mayr, Birds Southwest Pacific, 1945, p. 289 (Ku- 
saie). 

Geographic range. — Micronesia: Caroline Islands — Kusaie; Marshall Is- 
lands — Ebon (extinct?). 

Characters. — Adults: Resembles P. p. ponapensis, but occiput, nape, sides 
of head more gray and less greenish-yellow; chin and midthroat paler; crown 
coloring very faintly margined with yellow; tail band brighter yellow; under 
tail-coverts more orange; abdominal spot may be present as a brownish-red 
tinge; abdomen slightly more yellowish. 

Immature : Resembles immature of P. p. ponapensis. 

Measurements. — Measurements are listed in table 25. Ripley and Birck- 
head (1942:7) give the measurements of the only known specimen from Ebon 
(Marshall Islands) as: wing, 124; tail, 74; bill from base, 15. 

Specimens examined. — Total number, 11 (6 males, 5 females), as follows: Caroline Islands, 
USNM— Kusaie, 1 (Feb. 9); AMNH— Kusaie, 10 (Jan., Feb., March, April). 

Remarks. — I am following Ripley and Birckhead (1942:6) in 
identifying the dove from Ebon Island as of the subspecies P. p. 
hernsheimi. This specimen from Ebon may, however, represent the 
final vestige of a formerly well-distributed population in the Mar- 
shall Islands. This distribution is of particular interest because it 
may show the pathway by which these small fruit pigeons invaded 
eastern Micronesia from Polynesia. 



Baker: The Avifauna of Micronesia 185 

The small fruit dove at Kusaie has apparently the same habitat 
requirements as others of the species. Coultas (field notes) com- 
ments that in 1931 the birds were "quite common." He found them 
in the high trees on the mountain sides away from the native vil- 
lages and gardens. 

Ptilinopus porphyraceus pelewensis Hartlaub and Finsch 
Crimson-crowned Fruit Dove 

Ptilinopus ■pelewensis Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, p. 7. 
(TjTDe locality, Pelew Islands.) 

Ptilinopus pelewensis Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, p. 118 
(Pelew); Gray, Hand-list Birds, 2, 1870, p. 225 (Pelew); Hartlaub and Finsch, Proc. 
Zool. Soc. London, 1872, pp. 89, 101 (Pelew); Graffe, Journ. Mus. Godeffroy, 1, 1873, 
pi. 7, fig. 5 (Pelew); Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 5, 24 (Palau); 
Finsch, Journ. Mus. Godeffroy, 12, 1876, p. 37 (Palau); Schmeltz and Krause, Eth- 
nogr. Abth. Mus. Godeffroy, 1881, p. 407 (Palau); Matschie, Journ. f. Ornith., 1901, 
p. 113, (Palau); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 56 (Pelew); Mat- 
hews, Syst. Avium Australasianarum, 1, 1927, p. 32 (Pelew) ; Hand-list Japanese Birds, 
rev., 1932, p. 190 (Palau); Peters, Check-list Birds World, 3, 1937, p. 31 (Babeltop, 
Korror); Hand-list Japanese Birds, 3d ed., 1942, p. 213 (Babelthuap, Koror). 

Ptilonopus pelewensis Finsch, Proc. Zool. Soc. London, 1874, p. 94 (Pelew). 

Ptilopus pelewensis Giebel, Thes. Ornith., 3, 1877, p. 366 (Pelew); Elliot, Proc. 
Zool. Soc. London, 1878, p. 531 (Palau); Schmeltz, Verhandl. Ver. nat. Unterhatlung 
Hamburg, 1877 (1879), p. 178 (Pelew); Tristram, Cat. Birds, 1889, p. 44 (Pelew); 
Wiglesworth, Ibis, 1891, p. 584 (Pelew); idem, Abhandl. und Ber. Zool. Mus. Dresden, 
no. 6, 1890-1891 (1891), p. 48 (Pelew); Salvadori, Cat. Birds British Mus., 21, 1893, 
p. 86 (Pelew); Dubois, Syn. Avium, 2, 1904, p. 736 (Pelew); Reichenow, Die Vogel, 
1, 1913 p. 354 (Palau); Takatsukasa and Kuroda, Tori, 1, 1915, p. 52 (Pelew). 

Ptilinopus porphyraceus pelewensis Ripley and Birckhead, Amer. Mus. Novit., no. 
1192, 1942, p. 7 (Palau); Mayr. Birds Southwest Pacific, 1945, p. 289 (Palau); Baker, 
Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 60 (Peleliu, Ngabad, Garakayo). 

Geographic range. — Micronesia: Palau Islands — Babelthuap, Koror, Gara- 
kayo, Peleliu, Ngabad, Anguar. 

Characters. — Adult male : A green fruit pigeon with anterior lores and crown 
purple, margined with pale j'ellow; forehead paler than crown; chin and mid- 
throat pale yellow; neck, sides of head, and breast greenish-gray, darker on 
occiput; feathers of upper breast cross-banded with partly concealed violet 
bands; abdomen orange, its lower part and region of vent yellow; sides green- 
ish; tibia grayish; under tail-coverts near "Indian lake" with yellowish-orange 
edgings; upper parts green; wings metallic green, secondaries and primaries 
margined on outer webs with yellow; inner secondaries spotted with violet- 
blue near tips; under wing gray; upper side of tail green with pale yellow 
terminal band; under side of tail gray; bill lead-colored; feet dark blood-red. 

Adult female: Resembles adult male, but upper parts greener with upper 
side of wing and upper tail-coverts washed with olivaceou.s-brown ; breast 
duskier. Immature resembles adult, but lacks purple crown, violet breast spot, 
orange abdomen and maroon under tail-coverts; upper and lower parts mar- 
gined with yellow; forehead pale green; superciliary stripe pale yellow. 

P. p. peleivensis resembles P. p. ponapensis, but crown more purple; yellow 
tail-bar narrower; bifurcated, central breast feathers violet; abdomen orange; 
and under tail-coverts near "Indian lake". 

Measurements. — Measurements are presented in table 25. 



186 University of Kansas Publs., Mus. Nat. Hist. 

Specimens examined.- — Total number, 14 (10 males, 4 females), as follows: Palau Islands, 
USNM— Koror, 3 (Nov. 14, Dec. 3)— Garakayo, 1 (Sept. 19)— Peleliu, 3 (Aug. 27, Sept. 1, 
4)— Ngabad, 2 (Sept. 11)— Pelew, 2 (Mar. 1, 2); AMNH— Palau, 3 (Oct., Dec). 

Nesting.— At Ngabad Island on September 11, 1945, the NAMRU2 party 
found a nest in jungle in a low tree about six feet above the ground. It was 
loosely constructed and contained a single white egg, size 31 by 23 mm. 
Another nest was found at Ngabad the same day. It was on the branch of a 
tree approximately 20 feet from the ground. The nest was not examined 
other than to observe a parent bird on the nest. Three males obtained in 
August and in September had enlarged testes. Males taken in December by 
Coultas had enlarged testes. 

Food Habits. — Stomachs examined by the NAMRU2 party contained fruit 
parts and seeds. This species seemingly obtains its foods from the large fruit- 
producing trees and to a lesser extent from the smaller shrubs or from ground 
berries. 

Remarks. — P. 'p. pelewensis was found in small numbers at all 
islands visited in the southern Palaus by the NAMRU2 party in 
1945. At Peleliu, the bird was restricted to undisturbed woodlands 
and thickets, although some were seen in the thickly growing vege- 
tation covering over the battle areas. The bird evidently lives a soli- 
tary existence; it was only rarely observed in pairs. It was often 
located by its calls. Coultas (field notes) reports that in 1931 the 
species was becoming rare in the Palaus, owing to persistent hunting 
by the Japanese, who sold the bird for 25 sen each. 

Ptilinopus roseicapillus (Lesson) 
Marianas Fruit Dove 

Columba roseicapilla Lesson, Traite d'Ornith., 6, 1831, p. 472. (Type locality, 
Marianne Islands.) 

Columba roseicapilla Lesson, Compl. de Buffon, 2d ed., 2, Oiseaux, 1838, p. 278 
(Mariannes). 

Columba purpurata Kittlitz, Kupfertaf. Naturgesch. Viigel, 3, 1833, p. 25, pi. 23, 
fig. 2 (Guahan); idem, Obser. Zool., in Lutke, Voy. "Le Seniavine," 3, 1836, p. 30.5 
(Guahan). 

Ptilinopus purpuratus Hartlaub, Journ. f. Ornith., 18.54, p. 167 (Mariannen) ; 
Hartert, Katalog Vogelsamml. Senckenb., 1891, p. 190 (Guaham). 

Ptilopus roseicapillus Bonaparte, Comptes Rendus Acad. Sci. Paris, 39, 1854, p. 
877 (Mariannes); idem. Icon. Pigeons, 1857, pi. 23 and desc. letterpress (Mariannes); 
Schlegel, Mus. Pays-Bas, 6, no. 35, 1873, p. 8 (Guam); Giebel, Thes. Ornith., 3, 1877, 
p. 368 (Mariannae); Elliot, Proc. Zool. Soc. London, 1878, p. 537 (Marianne); Ousta- 
let, Le Nat., 1889, p. 261 (Mariannes); Wiglesworth, Ibis, 1891, p. 584 (Marianne); 
idem, Abhandl. und Ber Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 48 (Mari- 
anne); Salvadori, Cat. Birds British Mus., 21, 1893, p. 108 (Marianne Islands); Ousta- 
let, Nouv. Arch. Mus. Hist. Nat., Paris, (3), 7, 1895, p. 218 (Saypan, Guam, Rota); 
Safford, The Plant World, 7, 1904, p. 264 (Guam); Dubois, Syn. Avium, 2, 1904, p. 
736 (Mariannes); Safford, Contr. U. S. Nat. Herb., 9, 1905, p. 78 (Guam); Schnee, 
Zeitschr. f. Naturwisch., 82, 1912, p. 465 (Marianen) ; Prowazek, Die deutschen Mari- 
anen, 1913, p. 101 (Marianen); Reichenow, Die Vogel, 1, 1913, p. 354 (Marianen); 
Cox, Island of Guam, 1917, p. 20 (Guam); Bryan, Guam Rec. vol. 13, no. 2, 1936, 
p. 24 (Guam) ; Thompson, Guam and its people, 1942, p. 23 (Guam). 

Kurukurv roseicapillus Prevost and Dcs Murs, Voy. "V'enus," Oiseaux, 1855, pp. 
221, 231, 257, 259, 269 (Guam). 



Baker: The Avifauna of Micronesia 187 

Ptilopus roseicapilla Bonaparte, Consp. Avium, 2, 1855, p. 21 (Mariannis). 

PtUonopus roseicapilliis Gray, Cat. Birds Trop. Is. Pacific Ocean, 1859, p. 31 
(Guam); Reichenbach, Tauben, 1861, p. 96 (Mariannen); Finsch, Proc. Zool. Soc. 
London, 1874, p. 94 (Mariannes). 

Ptilinopus roseicapilliLS Finsch and Hartlaub, Fauna Centralpolynesiens, 1867, pp. 
122, 127 (Mariannen) ; Gray, Hand-list Birds, 2, 1870, p. 225 (Ladrones) ; Hartert, 
Novit. Zool., 5, 1898, p. 60 (Guam, Rota, Saipan) ; Wheeler, Report Island of Guam, 
1900, p. 13 (Guam); Seale, Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 39 (Guam, 
Rota, Saipan); Matschie, Journ. f. Ornith., 1901, p. 113 (Guam, Saipan); Safford, 
Osprey, 1902, p. 68 (Marianas); idem, Amer. Anthro., 4, 1902, p. 711 (Guam); 
Kuroda, in Momiyama, Birds Micronesia, 1922, p. 56 (Guam, Rota, Saipan); Mat- 
hews, Syst. Avium Australasianarum, 1, 1927, p. 33 (Marianne) ; Hand-list Japanese 
Birds, rev., 1932, p. 190 (Tinian, Saipan, Rota); Hand-list Japanese Birds, 3d ed., 
1942, p. 212 (Guam, Rota, Tinian, Saipan); Mayr, Birds Southwest Pacific, 1945, p. 
288 (Marianas); Downs, Trans. Kansas Acad. Sci., 49, 1946, p. 95 (Tinian); Watson, 
The Raven, 17, 1946, p. 42 (Guam)); Strophlet, Auk, 63, 1946, p. 538 (Guam); 
Baker, Condor, 49, 1947, p. 125 (Guam); Stott, .Auk, 64, 1947, p. 526 (Saipan); 
Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 59 (Guam, Rota). 

Ptilopus diadematus Giebel, Thes. Ornith., 3, 1877, p. 363 (Marianae). 

Ptilinopus roseicapilla Peters, Cheek-list Birds World, 3, 1937, p. 31 (Saipan, Tin- 
ian, Rota, Guam); Ripley and Birckhead, Amer. Mus. Novit., no. 1192, 1942, p. 3 
(Guam, Rota, Tinian, Saipan). 

Geographic range. — Micronesia: Mariana Islands — Guam, Rota, Tinian, 
Saipan. 

Characters. — Adult male : A green dove with crown, forehead, anterior lores, 
and spot at base of mandible near "aster purple," margined with pale yellow 
especially on top of head; chin and throat pale yellow to white; sides of head 
greenish-gray, darker on occiput ; breast green with pearly-gray tinge on feath- 
ers of middle part ; lower breast with dark purple patch ; abdomen orange with 
yellowish-green coloring at midline; anal region and lower tail-coverts yellow, 
tinged with orange on lower tail-covcrts; sides and tibia greenish with yellow 
tinges; upper parts green, more yellowish-green on rump; wings glossy, upper 
wing-coverts brighter in middle and margined with yellow ; under side of wing 
and under side of tail gray; upper side of tail green with broad grayish termi- 
nal band margined with yellow; iris pale yellow; bill grass-green; legs and 
feet reddish-black. 

Adult female : Resembles male, but shghtly smaller with neck greener. Im- 
mature resembles adult, but lacking colored crown; body feathers edged with 
yellow. 

Birds from Guam, Rota, and Tinian exhibit no conspicuous differences. P. 
roseicapillus is closest to P. regina of southern Papua, Lesser Sunda Islands, 
and Australia being, according to Ripley and Birckhead (1942:3), "Similar to 
regina, but crown and abdominal band darker; malar apex concolorous with 
crown ; hind neck more grayish ; tail-bar wider and paler." 

Measurements. — Measurements of P. roseicapillus are presented in table 26. 

Weights.— In 1948 (1948:59) I listed the weights of 14 adult males as 
81-103 (90), of 4 adult females as 85-99 (92), and of one nestling in post natal 
molt as 44 grams. These were taken at Guam. 

Speci7nens examined. — Total number, 43 (32 males, 10 females, 1 unsexed), as follows: 
Mariana Islands, USNM — Guam, 28 (March 8, May 25, 27, June 3, 12, 14, July 2, 6, 10, 
18, 19, 29, Aug. 21)— Rota, 3 (Oct. 28, 31, Nov. 2)— Tinian, 1 (Oct. 26); AMNH— Guam! 
8 (Aug.)— Tinian, 3 (Sept.). 



188 University of Kansas Publs., Mus. Nat. Hist. 

Table 26. Measurements of Ptilinopiis roseicapillus 



Number 


Wing 


Tail 


Exposed 
cu men 


Tarsus 


32 adult males 

10 adult females .... 


127 (122-133) 
124 (121-130) 


80 (75-84) 
76 (75-79) 


14 (13-15.3) 
13 (12-13.7) 


25 (24-27) 
24 (22-25.5) 



Nesting. — At Guam, I obtained records of nests of fruit doves on March 1, 
27, and May 7, 1945. David H. Johnson observed a pair of fruit doves in 
the act of copulation on May 26, 1945. Birds with enlarged gonads were 
taken by the NAMRU2 party in March and July. A nestling in postnatal 
molt, just beginning to fly, was taken on July 6. Seale (1901:39) reports two 
nests, each containing one white egg, taken in the period from May to July. 
These nests were found in trees eight to ten feet above the ground. 

Food habits. — The Marianas Fruit Dove feeds on fruits and seeds of trees 
and shrubs. The birds are apparently strictly tree dwellers; I saw no birds on 
the ground. A favorite fruit is that of a flowering shrub known as the "ink 
berry." Birds were collected which contained stomachs full of these small 
black berries. The fruit of the papaya is also a favorite food. 

Remarks. — The NAMRU2 party found the Marianas Fruit Dove 
at Guam to be fairly numerous in undisturbed jungle, and more 
abundant in the heavy, second-growth, scrub-forest as was found 
on Amantes Point in 1945. The birds were secretive but were easily 
located by their calls. They were usually found as singles sitting 
quietly concealed in thick vegetation. Birds were seen flying rather 
infrequently, and then only for short distances. The removal of 
large tracts of jungle to provide space for the construction of air 
strips and installations in the late war has disturbed some of the 
habitat of these birds. Although vast tracts of forest were undis- 
turbed, the birds probably have decreased at Guam. Coultas (field 
notes) found the birds common at the northern end of Guam in 1931. 
He commented that natives catch them with snares and bird lime for 
the local markets. At Tinian in 1931, Coultas found few birds. 
Downs (1946:95) and Stott (1947:526) record the birds at Tinian 
and Saipan, respectively, in 1945. At Rota, the NAMRU2 party 
found the dove to be numerous. 

Evolutionary history of PtiUnopus in Micronesia. — Oceania is es- 
pecially rich in species and subspecies of the genus PtiUnopus. Rip- 
ley and Birckhead (1942) have made the most recent and most 
thorough contribution concerning these birds. They state that the 
center of distribution for the genus lies in the Papuan rfegion. 
Within the Oceanic region there are several species of PtiUnopus 



Baker: The Avifauna of Micronesia 189 

which in one way or another are rather closely related; Rensch 
(1938:277) uses these as examples of species which have been 
formed by isolation. These include P. peroiisii from Samoa, Fiji, 
and Tonga; P. mercierii from the Marqueasas; P. dupetithouarsii 
from the Marquesas; P. huttoni from Rapa; P. purpuratus from 
Henderson, Tuamotus, Societies; P. porphyraceus from Samoa, Fiji, 
Tonga, Carolines, Solomons, New Hebrides, New Caledonia, and 
adjacent areas; and P. roseicapillus from Marianas. In all of these 
birds the crown is wine-red except in P. dupetithouarsii in which it 
is whitish. P. porphyraceus appears to be more closely related to 
P. purpuratus than to any other species and is characterized by an 
often brightly washed spot of color of some shade of red or orange 
on the breast. These birds may have invaded Micronesia from 
the region of the Solomon Islands, although it appears more likely 
that they arose in the Samoa-Fiji-Tonga region and moved north- 
ward, probably by way of the Marshall Islands. P. p. hernsheimi 
from Kusaie and P. p. ponapensis from Ponape and Truk resemble 
P. p. faciatus Peale from Samoa more closely than they do any 
other subspecies. P. p. pelewensis from Palau, on the other hand 
shows little relation to these other two Micronesian subspecies and 
appears to be closest to P. p. porphyraceus of Fiji and Tonga or 
possibly to P. grayi from Melanesia. Ripley and Birckhead 
(1942:7) suggest that the subspecies at Palau owes its marked 
divergence to its isolated position at the periphery of the range of 
the species. P. p. pelewensis probably represents an independent and 
an earlier colonization, possibly from a stock different from that 
from which the two subspecies in the Carolines arose. The presence 
in the Palaus of subspecies singularly different from subspecies in 
the Carolines can also be observed in other genera, as for example, 
Rhipidura, and Myiagra. Figure 13 shows the inferred routes of 
colonization of Ptilinopus to Micronesia. 

P. roseicapillus seemingly represents a remnant, or perhaps a 
successful straggler, of an early invasion to Micronesia. Ripley 
and Birckhead (1942:2) classify this species as "Old Stock," 
along with P. monachus, P. coronulatus and P. regina. Its pathway 
of invasion to the Marianas was probably directly northward from 
the Papuan area and not by way of the Polynesian islands. Its 
resemblance to the species P. regina of southern Papua, Lesser 
Sundas, and Australia is most unusual, especially since there is a 
separation between the two species of some 1,400 miles; this is 
pointed out by Ripley and Birckhead (1942:4). As I have said 



190 



University of Kansas Publs., Mus. Nat. Hist. 




160 



20 



20 



160, 



Fig. 13. Geographic distribution of Ptilinopiis porphyraceus and routes of 
its dispersal. (1) P. p. porphyraceus ; (2) P. p. fasciatus; (3) P. p. hernsheimi ; 
(4) P. p. ponapensis ; (5) P. p. pelewensis. 



(1948:59) elsewhere, "On the basis of its characters the Mariana 
birds would merit only subspecific separation, but owing to the 
great distance between the two doves and the possibility of inde- 
pendent origin and subsequent convergence, it may be more advisable 
to continue to regard the two as separate species." 

Ducula oceanica monacha (Momiyama) 
Micronesian Pigeon 

Globicera oceanica monacha Momiyama, Birds Micronesia, March, 1922, p. 4. (Type 
locality. Yap.) 

Columba oceanica Lesson and Garnet (part). Diet. Sci. Nat., ed. Levrault, 40, 1826, 
p. 317 (Pelew); Lesson (part), Man. d'Ornith., 2, 1828, p. 166 (Pelevv) ; ide?n. (part), 
Voy. "La Coquille," Zool., 2, 1828, pp. 432, 709 (Pelew); idem, Compl. de Buffon, 2d 
ed,. 2, Giseaux, 1838, p. 292 (Pelew); Prevost and Knip, Les Pigeons, 2, 1838-43, p. 49 
(Pelew). 

Carpophaga oceanica Hartlaub (part), Archiv. f. Naturgesch., 18, 1852, p. 11.5 
(Pelewinseln); idetn, Proc. Zool. Soc. London, 1867 (1868), p. 830 (Pelew); Gray 
(part), Hand-list Birds, 2, 1870, p. 229 (Pelew); Hartlaub and Finsch (part), Proc. 
Zool. Soc. London, 1872, pp. 89, 101 (Pelew); Finsch, Journ. Mus. Godeffroy, 8, 1875, 
pp. 5, 26 (Palau); idem (part), Proc. Zool. Soc. London, 1877 (1878), pp. 775, 780 
(Palau) ; Salvadori (part), Cronaca del R. Liceo-Ginnasio Cavour, 1878, pp. 3, 8 
(Pelew); idem. Ibis, 1879, p. 364 (Pelew); Tristram, Cat. Birds, 1889, p. 42 (Pelew); 
Wiglesworth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), 
p. 52 (Pelew); Matschie (part), Journ. f. Omith., 1901, p. 113 (Palau); Dubois (part), 
Syn. Avium, 2, 1904, p. 743 (Pelew); Reichenow (part). Die Vogel, 1, 1913, p. 351 
(Palau). 

Globicera oceanica Bonaparte (part), Consp. Avium, 2, 1855, p. 31 (Pelew); 
Reichenbaeh (part), Tauben, 1861, p. 120 (Pelew); Salvadori (part). Cat. Birds British 



Baker: The Avifauna of Micronesia 



191 



Mus., 21, 1893, p. 176 (Pelew) ; Takatsukasa and Kuroda (part). Tori, 1, 1915, p. 52 
(Pelew); Uchida, Annot. Zool. Japon., 9, 1918, pp. 486, 489 (Palau). 

Carpophaga {Globicera) oceanica Gray (part), Cat. Birds Trop. Is. Pacific Ocean, 
1859, p. 41 (Pelew). 

Carpophaga pacifica Finsch and Hartlaub (part), Faiina Centralpolynesiens, 1867, 
p. 145 (Pelew); Hartlaub and Finsch, Proc. Zool. See. London, 1868, pp. 7, 118 
(Pelew); Finsch and Hartlaub, Joum. f. Ornith., 1870, p. 134 (Pelew). 

Globicera oceanica monacha Kuroda, in Momiyama, Birds Micronesia, 1922, p. 55 
(Yap); Mathews, Syst. Avium Australasianarum, 1, 1927,, p. 46 (Yap); Yamashina, 
Tori, 7, 1932, p. 408 (Yap); Hand-list Japanese Birds, rev., 1932, p. 189 (Uap, Palau, 
Current = Palo Anna). 

Globicera oceanica momiyamai Kuroda, in Momiyama, Birds Micronesia, March, 
1922, pp. 25, 56 (Type locality, Angaur) ; Mathews, Syst. Avium Australasianarum, 1, 
1927, p. 46 (Pelew); Kuroda, Ibis, 1927, p. 719 (Pelew). 

Muscadivora oceanica winkleri Neumann, Verhandl. Ornith. Ges. Bayern, 25, Sept. 1, 
1922, p. 234 (Type locality, Palau). 

Ducula oceanica monacha Peters, Check-list Birds World, 3, 1937, p. 43 (Yap, 
Babelthuap, Koror, Angaur, Current); Hand-list Japanese Birds, 3d ed., 1942, p. 211 
(Yap, Babelthuap, Koror, Angaur, Current); Amadon, Amer. Mus. Novit., no. 1237, 
1943, p. 11 (Yap, Palau); Mayr, Birds Southwest Pacific, 1945, p. 289 (Palau, Yap); 
Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 66 (Peleliu, Garakayo, Babel- 
thuap). 

Geographic range. — Micronesia: Palau Islands — Babelthuap, Koror, Gara- 
kayo, Peleliu, Angaur, Pulo Anna; Caroline Islands — Yap. 

Characters. — Adult: Resembles D. o. oceanica from Kusaie but throat, 
breast, head, and neck light ashy-gray; feathers around bill grayish-white; 
abdomen and under tail-coverts tipped with light brown. 

Immature: Resembles adult, but underparts paler; back lacking dark 
bluish spots; back feathers and wing feathers edged with light brown. 

Measurements. — Measurements of D. oceanica are listed in table 27. 



Table 27. Me.asurements of Ducula oceanica 



Subspecies 


Number 


Wing 


Exposed culmen 


Tarsus 


D. 


0. monacha . . . 


8 males... . 
6 females . . 


228 (219-233) 
221 (214-228) 




36 (34-37) 




22.5 (22.0-23.0) 


31 (29-33) 


D. 


0. teraokai . . . 


5 males .... 
8 females . . 


230 (225-237) 

231 (221-238) 


23.5 (23.0-25.0) 
23.0 (21.5-24.5) 


34 (33-35) 
34 (33-35) 


D. 


0. townsendi. . 


8 males .... 
5 females . . 


226 (211-234) 
226 (215-233) 


24.0 (23.5-25.0) 
24.0 (23.0-24.5) 


34 (32-35) 
33 (32-34) 


D. 


0. oceanica . . . 


4 males .... 
13 females. . 


222 (217-228) 
219 (213-226) 


25.0 (24.5-26.0) 
24.0 (23.0-25.0) 


35 (34-36) 
32 (30-34) 


D. 


0. ratakensis * 


6 males .... 
3 females . . 


(211-217) 
(208-213) 


(25 -27 ) 
(25 -26 ) 













*From Takatsukasa and Yamashina (1932:221). 

Specimens examined. — Total number, 17 (9 males, 8 females), as follows: Palau Islands, 
USNM— Garakayo, 1 (Sept. 19)— Peleliu. 7 (Aug. 27, 28, 29, Sept. 4, 5); AMNH— Palau, 
9 (Oct., Nov. 13, 15, 21, Dec. 1). 

iVesim^.— Yamashina (1932a :408) records the finding of one egg at Yap on 



192 University of Kansas Publs., Mus. Nat. Hist. 

December 3, 1930. The N'AMRU2 party obtained no evidence of breeding 
activity of these pigeons at the Palaus in August and September, 1945. Coul- 
tas, in November and December of 1931, obtained birds with enlarged gonads 
at Palau. Probably the nesting season begins in November or December. 

Food habits. — The pigeons feed on both fruits and green stuffs. The 
NAMRU2 party found berries, fruit parts and green plant materials in stom- 
achs of birds taken in September, 1945. The birds were found to be exceed- 
ingly fat at this time. 

Parasites. — Uchida (1918:486, 489) records the bird lice (Mallophaga), 
Goniocotes carpohagae and C olopocephalum unicolor, from this pigeon at 
Palau. 

Remarks. — The Micronesian Pigeon at Palau was first observed 
in 1783, when Captain Henry Wilson of the packet "Antelope" was 
shipwrecked in these islands. In his account of the islands, as com- 
piled by George Keate (Wilson, 1788), Wilson described the large 
pigeons, which were kept as pets by the natives and were eaten by 
only certain classes of people. In 1826, Lesson and Garnot made 
first reference to the birds found by Wilson. It was almost 100 
years after Wilson's visit that the bird was again observed; this 
time it was obtained by the sea captains, Tetens and Heinsohn, and 
by Kubary, the collector for the Godeffroy Museum. 

It is surprising that a pigeon as large and conspicuous as this 
one has not already been exterminated by man on these small is- 
lands. Every traveller to the islands, who has made observations, 
writes that the pressure of hunting on these birds is severe. Coultas 
(field notes) reports that in 1931 the birds were "very scarce and 
wild." He comments that the Japanese hunters obtained the birds 
and received the market price of 35 sen for each one. He writes, 
"There is a group of Japanese hunters in the islands who vie with 
one another to see who can obtain the most birds. They are all 
atrocious shots but some employ natives and since so many of them 
are in the business they are inflicting considerable damage to the 
bird life. During my stay there one Japanese was sentenced to six 
weeks hard labor for hiring native hunters. The native hunter who 
preferred charges claimed that money was due him for having shot 
some 3,500 birds and the account had been standing over a year." 
Price (1936b: 491) shows a picture of a captive pigeon at Palau. 
The natives used this bird as a calling decoy to attract others within 
range of their blowguns. 

The NAMRU2 party observed pigeons at all islands visited in 
August and September, 1945. At Peleliu, the pigeons were found 
to be restricted to relatively undisturbed areas where tall trees re- 



Baker: The Avifauna of Micronesia 193 

mained or where shrubs were present on the faces of overhanging 
cliffs. The shrubs on cliffs were favorite roosting places. Although 
the pigeons remained in these relatively inaccessible areas, they 
were not especially difficult to obtain with shotguns. I can see that 
it might be difficult for unarmed hunters to obtain the birds. The 
present writer (1946b: 210) has recorded the extensive utilization of 
pigeons, rails and megapodes by Japanese troops and by their pris- 
oners of war at Babelthuap and Koror during the latter part of the 
war. 

During our stay at Peleliu we were unable to learn whether the 
pigeon was still present at Pulo Anna (Current Island), a coral is- 
land some 160 miles southeast of Peleliu. The U. S. Navy fre- 
quently dispatched a ship to the island, but we did not learn of it 
until our stay at Peleliu was nearly over. Dr. C. K. Dorsey, then 
of the U. S. Naval Epidemiology Unit at Peleliu, reported that vari- 
ous kinds of birds were numerous at Pulo Anna, but he did not 
recall seeing the pigeon. This pigeon may occur also at Pais, a 
raised coral island west of Yap and Ulithi in the Carolines. I know 
of no reports dealing with the avifauna of this phosphate island, 
but I examined several pictures, taken by Navy landing parties and 
the Military Government personnel, which show the island to be 
covered with extensive and luxuriant vegetation. I suspect that an 
intensive survey of the island will reveal several new records for 

birds. 

Ducula oceanica teraokai (Momiyama) 

Micronesian Pigeon 

Globicera oceanica teraokai Momiyama, Birds Micronesia, 1922, p. 2. (Type 
locality, Tol, Truk Islands.) 

Columba oceanica Kittlitz (part), Kupfertaf. Naturgesch. V'ogel, 3, 1833, p. 25, 
pi. 33, fig. 1 (Lugunor); idem (part), Obser. Zool., in Lutke, Voy. "Le Seniavine," 
3, 1836, p. 299 (Lougounor) ; Hartlaub (part), Archiv f. Naturgesch., 18, 1852, pp 
115, 185, (Mordlockinseln). 

Carpophaga (Globicera) pacifica Gray (part), Cat. Birds Trop. Is. Pacific Ocean, 
1859, p. 41 (Mortlock's Island). 

Carpophaga pacifica Finsch and Hartlaub, Fauna Central poljTiesiensi, 1867, p. 146 
(Lugunor). 

Carpophaga oceanica Finsch, Proc. Zool. Soc. London, 1880, p. 576 (Ruk) ; 
Schmeltz and Krause (part), Ethnogr. Abth. Mus. Godeffroy, 1881, pp. 330, 353 
(Nukuor, Ruk); Wiglesworth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 
1890-1891 (1891), p. 52 (Luganor, Ruk, Nukuor); Hartert, Novit. Zool., 7, 1900, 
p. 8 (Ruk). 

Globicera oceanica Salvador! (part), Cat. Birds British Mus., 21, 1893, p. 176 
(Ruk); Takatsukasa and Kuroda (part). Tori, 1, 1915, p. 52 (Ruk). 

Globicera oceanica teraokai Kuroda, in Momiyama, Birds Micronesia, 1922, p. 55 
(Ruk, ?Mortlock, ?Nukuor) ; Mathews, Syst. Avium Australasianarum, 1, 1927, p. 45 
(Ruk); Hand-list Japanese Birds, rev., 1932, p. 189 (Truk). 

M [uscadivora] o[ceanica] oceanica Neumann (part), Verhandl. Omith. Ges. Bayern, 
25, 1922, p. 234 Ualam = Truk). 

13—8131 



194 University of Kansas Publs., Mus. Nat. Hist. 

Ducula oceanica feraokai Peters, Check-list Birds World, 3, 1937, p. 43 (Truk) ; 
Hand-list Japanese Birds, 3d ed., 1942, p. 212 (Truk); Amadon, Amer. Mus. Novit., 
no. 1237, 1943, p. 11 (Truk); Mayr, Birds Southwest Pacific, 194,5, p. 289 (Truk). 

Geographic range. — ^Micronesia: Caroline Islands — Truk, ?Lukunor, ?Nuku- 
oro. 

Characters. — Adult: Resembles D. o. monacha, but slightly darker on 
crown, nape, and mantle ; back more bluish and less greenish, underparts 
slightly darker chestnut. Differs from D.o. townsendi by being paler and gray 
on crown, nape, shoulder, side of neck, and upper breast; abdomen and under 
tail-coverts slightly deeper che.stnut. Differs from D. o. oceanica by larger size; 
upper parts paler; abdomen and under side of tail deeper chestnut. I agree 
with Amadon (1943:11) that this subspecies is only doubtfully distinct from 
D. a. monacha and that it might be advisable to unite these under one sub- 
specific name. 

Measurements. — Measurements are listed in table 27. 

Specimens examined. — Total number, 14 (5 males, 9 females, 1 unsexed) from Caroline 
Islands, AMNH— Truk (Nov., Dec). 

Remarks. — The Micronesian Pigeon at Truk was observed by 
Kittlitz (1836:299) and later by Kubary at the islands of Lukunor 
and Nukuoro. Momiyama (1922:4) remarks that he did not see 
specimens from these two islands but concludes that they probably 
belong to the subspecies named from Truk. It is possible that birds 
at these two atolls have been exterminated, although adequate field 
investigations have not been made. 

There is little information published concerning the natural history 
of this subspecies. McElroy, who visited Truk in December, 1945, 
did not find the bird; however, he did not visit all of the islands in 
the group during his stay. 

Ducula oceanica townsendi (Wetmore) 
Micronesian Pigeon 

Globicera oceanica townsendi Wetmore, in Townsend and Wetmore, Bull. Mus. 
Comp. ZooL, 63, 1919, p. 191. (Type locality, Ponape). 

Carpophaga oceanica Finsch (part), Proc. Zool. Soc. London, 1877 (1878), p. 780 
(Ponape); Nehrkorn, Journ. f. Ornith., 1879, p. 407 (Ponape); Finsch (part), Journ. f. 
Ornith., 1880, p. 292 (Ponape); idem, 1881, pp. 113, 115 (Ponape); Schmeltz and 
Krause (part), Ethnogr. Abth. Mus. Godeffroy, 1881, p. 281 (Ponape); Wiglesworth 
(part), Abhandl. und Ber. Zool. Mus. Dresden, no. 0, 1890-1891 (1891), p. 52 (Ponape); 
Matschie (part), Journ. f. Ornith., 1901, p. 113 (Guam, error = Ponape). 

Globicera oceanica Salvadori (part). Cat. Birds British Mus., 21, 1893, p. 176 
(Ponape). 

Globicera oceanica townsendi Momiyama, Birds Micronesia, 1922, p. 6 (Ponape); 
Kuroda, in Momiyama, Birds Micronesia, 1922, p. 55 (Ponape); Mathews, Syst. Avium 
Australasianarum, 1, 1927, p. 45 (Ponape); Hand-list Japanese Birds, rev., 1932, p. 190 
(Ponape). 

Ducula oceanica townsendi Peters, Check-list Birds World, 3, 1937, p. 44 (Ponape); 
Bequaert, Mushi, vol. 12, no. 2, 1939, pp. 81, 82 (Ponape); idem, Geo. Papers Bernice 
P. Bishop Mus., 16, 1941, pp. 266, 290 (Ponape); Hand-list Japanese Birds, 3d ed., 
1942, p. 212 (Ponape); Amadon, Amer. Mus. Novit., no. 1237, 1943, p. 11 (Ponape); 
Mayr, Birds Southwest Pacific, 1945, p. 289 (Ponape). 



Baker: The Avifauna of Micronesia 195 

Geographic range. — Micronesia: Caroline Islands — Ponape. 

Characters. — Adult: Resembles D. o. tcraokai, but darker. Rsemblcs closely 
D. o. oceanica but larger and darker on crown and nape; lower parts slightly 
paler but chin more cream-buff in color. As Adamon (1943:11) states, there is 
little difference between D. o. toicnscndi and D. o. oceanica except in size. 

Measurements. — Measurements are listed in table 27. 

Specimens examined. — Total number 21 (11 males, 9 females, 1 imsexed), as follows: 
Caroline Islands, USNM— Ponape, 2 (Feb. 11, 12); AMNH— Ponape, 19 (Nov. 22, 29, Dec. 
1, 2, 3). 

Nesting. — Coultas (field notes) writes that the pigeon at Ponape nests the 
year around, probably two or three times a year. He describes the nest as being 
made of loose twigs and as placed on a fork of a limb in a tall tree. One egg is 
laid. Coultas saw "two or three" females nesting in December. 

Parasites.— Bequaevt (1939:81, 82 and 1941:266, 290) found the flies (Hippo- 
boscidae), Ornithoctona plicata and 0. pmilla, on pigeons from Ponape. 

Remarks. — Coultas (field notes) writes that in 1930 several Jap- 
anese made a livelihood as professional hunters of pigeons at Ponape. 
He notes, ''Two or three j^ears ago, 4 or 5 Japanese, each, averaged 
from 75 to 100 birds per day, which they sold to the inhabitants for 
35 sen (I7I/2 cents) per bird. . . . Now these same hunters are 
fortunate if they obtain 4 or 5 Diicula each per day and are able to 
do so only by starting before daylight and covering great distances. 
Other birds are now replacing Ducula on the market." Coultas fur- 
ther records in his notes that the hunters used calls to attract the 
pigeons. In 1930, Coultas regarded the pigeon at Ponape as a rap- 
idly disappearing species; he found it only in small areas in remote 
regions of the mountains. With the shipping of supplies cut off to 
the Japanese garrison forces at Ponape, as well as at Kusaie, Truk, 
and Yap by the effective American blockade during the latter part 
of the war, it is probable that the pigeons were hunted more inten- 
sively by the Japanese hunting parties than ever before. Richards 
obtained two specimens at Ponape in the period from August, 1947, 
to January, 1948. 

Ducula oceanica oceanica (Lesson and Garnot) 
Micronesian Pigeon 

Columha oceanica Lesson and Garnot, Diet. Sci. Nat., ed., Levrault, 40, 1826, p. 
316. (Type locality, Oualan — Kusaie.) 

Columha oceanica Lesson (part), Voy. "La Coquille," Zool. ; Atlas, 1826, pi. 41; 
vol. 2, 1828, pp. 432, V08 (Oualan or Strong); idem, (part), Man, d'Ornith., 11, 1828, 
p. 166 (Oualan); Kittlitz (part), Kupfertaf. Naturgesch. Vogel, 3, 1833, p. 25, pi. 
23, fig. 1 (Ualan); idem (part), Observ. Zool., in Lutke, Voy. "Le Seniavine," 3, 
1836, p. 284 (Ualan); Lesson, Compl. de Buffon, 2d ed., 2, Oiseaux, 1839, p. 292 
(Oualan); Prevost and Knip (part), Les Pigeons, 2, 1838-43, p. 47, pi. 24 (Oualan); 
Bonaparte, Comptes Rendus Acad. Sci. Paris, 39, 1854, p. 1072 (Oualan); Kittlitz, 
Denkw. Reise russ. Anier. Micron, und Kamchat., 1, 1858, pp. 39, 49, 62 (Ualan). 



196 University of Kansas Publs., Mus, Nat. Hist. 

Carpohaga oceanica Hartlaub (part), Archiv f. Naturgesch., 18, 1852, pp. 115, 185 
(Ualan); idem, Journ. f. Ornith., 1854, p. 168 (Carolinen=Kusaie) ; Hartlaub and 
Finsch (part), Proc. Zool. Soc. London, 1872, p. 101 (Ualan); Schlegel, Mus. Pays- 
Bas, 6, no. 35, 1873, p. 87 (Oualan) ; Salvadori (part). Cronaca del R. Liceo-Gin- 
nasio Cavour, 1878, pp. 3, 8 (Oualan); Finsch (part). Ibis, 1880, pp. 220, 331, 332 
(Taluit); idem (part), Journ. f. Ornith., 1880, pp. 292, 304 (Kuschai) ; idem. Ibis, 
1881, p. 108 (Kuschai); idem, Mitth. Ornith. Ver. Wien, 1884, p. 50 (Kuschai, Ja- 
luit) ; Hartert, Katalog Vogelsamml, Senckenb., 1891, p. 190 (Ualan) ; Wiglesworth 
(part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 52 (Ualan, 
Taluit); Matschie (part), Journ. f. Ornith., 1901, p. 113 (Ualan). 

Globicera oceanica Bonaparte (part), Consp. Avium, 2, 1855, p. 31 (Oualan); Idem, 
Comptes Rendus Acad. Sci. Paris, 43, 1856, p. 835 (Oualan); Reichenbach (part), 
Tauben, 1861, p. 120 (Oualan); Salvadori (part), Cat. Birds British Mus., 21, 1893, 
p. 176 (Kushai). 

Carpophaga pacifica Finsch and Hartlaub (part). Fauna Centralpolynesiens, 1867, p. 
145 (Ualan). 

Carpophaga (Globicera) oceanica Gray (part). Cat. Birds Trop. Is. Pacific Ocean, 
1859, p. 41 (Oualan). 

Globicera oceanica oceanica Wetmore, in Townsend and Wetmore, Bull. Mus. Comp. 
Zool., 63, 1919, p. 191 (Kusaie) ; Momiyama (part). Birds Micronesia, 1922, p. 6 
(Kusaie, Taluit); Kuroda (part), in Momiyama, Birds Micronesia, 1922, p. 55 (Kusaie, 
Taluit) ; Mathews, Syst. Avium Australasianarum, 1, 1927, p. 45 (Kusaie) ; Takatsu- 
kasa and Yamashina, Dobutsu. Zasshi, 44, 1932, p. 221 (Jaluit, Iringlob, Kusaie); 
Hand-list Japanese Birds, rev., 1932, p. 190 (Kusaie, Jaluit, Elmore). 

Muscadivora oceanica oceanica Neumann (part), Verhandl. Ornith. Ges. Bayern, 25, 
1922, p. 234 (Kushai). 

Ducula Oceanica oceanica Peters, Check-list Birds World, 3, 1937, p. 44 (Kusaie, 
Jaluit, Elmore) ; Bequaert, Mushi, 12, 1939, p. 81 (Kusaie) ; idem, Occ. Papers Ber- 
nice P. Bishop Mus., 16, 1941, p. 266 (Kusaie); Hand-list Japanese Birds, 3d ed., 
1942, p. 212 (Kusaie, Jaluit, Elmore) ; Amadon, Amer. Mus. Novit., no. 1237, 1943, 
p. 11 (Kusaie, Jaluit, Elmore) ; Mayr, Birds Southwest Pacific, 1945, p. 289 (Kusaie, 
Jaluit, Elmore). 

Geographic range. — Micronesia: Caroline Islands — Kusaie; Marshall Is- 
lands — Jaluit, Elmore. 

Characters. — Adult male: A large knob-billed pigeon with breast gray, 
washed with buff; head and neck dark gray; feathers at base of bill and on 
chin buff-white; abdomen and under tail-coverts near "burnt sienna," sides 
grayer; mantle, back, rump, upper tail-coverts, wings and tail bronze-green 
edged with a dark bluish sheen; under side of wing and under side of tail 
brown; bill and knob black; feet blackish-red; iris reddish-brown. Adult 
female resembles adult male but smaller and possibly a little darker bluish- 
green on back, wings, and tail. 

D. o. Oceania resembles D. o. toitmsendi, but is smaller with upper parts 
slightly darker and abdomen and under side of tail lighter. 

Measurements. — Measurements are presented in table 27. 

Specimens examined. — Total number, 47 (25 males, 22 females), as follows: Caroline 
Islands, USNM— Kusaie, 2 (Feb. 8, 9,); AMNH— Kusaie, 45 (Jan., Feb., March). 

Parasites.— Bequaert (1939:81 and 1941:266) obtained the fly (Hippo- 
boscidae) Ornithoctona plicata from the pigeon at Kusaie. 

Remarks. — The Micronesian Pigeon at Kusaie has been known 
since 1824, when from June 5 to June 15 of that year personnel from 
the corvette "La Coquille" visited the island and observed the bird. 
Kittlitz visited Kusaie and observed the pigeon in December, 1827, 
and January, 1828. Finsch (1880c and 1880d) found the bird in the 



Baker: The Avifauna of Micronesia 197 

Marshalls at Jaluit. Takatsiikasa and Yamashina (1932:221) 
record the bird from Elmore in the Marshalls. Coultas (field notes) 
writes that the pigeon was numerous at Kusaie in 1931. He remarks 
that they appear stupid and are easily killed by the natives, who use 
a call to attract them. With regard to their habits he writes, "About 
four o'clock in the afternoon these birds begin congregating in the 
high trees trees of the lowlands close to the salt water where they 
roost for the night. At daybreak they begin migrating to the high 
mountain sides and peaks where they spend the time feeding." 

Ducu]a oceanica ratakensis (Takatsukasa and Yamashina) 
Micronesian Pigeon 

Globecera oceanica ratakensis Takatsukasa and Yamashina, Dobutsu. Zasshi, 44 
1932, p. 221. (Type locality, Aruno.) 

Columba australis Chamisso, in Kotzebue's, Vcy. "Rurick," 3, 1821, p. 157 
(Radak). 

Carpophaga oceanica Finsch, Ibis, 1880, p. 331 (Arno) ; Wiglesworth (part), 
Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 52 (Arno); idem. 
Ibis, 1893, p. 211 (Marshalls). 

Globicera oceanica oceanica Momiyama (part). Birds Micronesia, 1922, p. 5 (Arno); 
Kuroda (part), in Momiyaina, Birds Micronesia, 1922, p. 55 (Arno). 

Globicera oceanica ratakensis Hand-list Japanese Birds, rev., 1932, p. 190 (Arlino, 
Wotze); Mathews, Ibis, 1933, p. 87 (Aruno, Wozzie). 

Ducala oceanica ratakensis Peters, Check-list Birds World, 3, 1937, p. 44 (Arno, 
Wotje); Hand-list Japanese Birds, 3d ed., 1942, p. 212 (Arhno, Wotze); Amadon, 
Amer. Mus. Novit., no. 1237, 1943, p. 12 (Arno, Wotje); Mayr. Birds Southwest 
Pacific, 1945, p. 289 (Arno, Wotje). 

Geographic range. — Micronesia: Marshall Islands (Radak Chain) — Wotje, 
Arhno. 

Characters. — Takatsuka-'a and Yamashina (1932:221) describe this sub- 
species as follows, "This form differs from all other forms of Globicera oceanica 
by its smaller size, more bronze-sheen on the back, more vinaceous grey on the 
breast and duller brown on the abdomen." On examining two specimens from 
Arno in the collection of the Museum of Comparative Zoology, Amadon 
(1943:12) writes that he finds no distinguishing color characters between D. o. 
oceanica and D. o. ratakensis. He also questions whether there is any difference 
in size between the two populations. 

Measurements. — Measurements are listed in table 27. 

Remarks. — Chamisso (1821), the naturalist on board the ship 
"Rurick," was the first person to write of the pigeon in the Radak 
Chain of the Marshall Islands. The ship visited this area in 1817, 
Finsch (1880b) published an account of the bird when he visited the 
area about 1880. Takatsukasa and Yamashina (1932:221) de- 
scribed this bird as new on the basis of an examination of nine skins 
taken at Arhno and Wotje. 

Evoluntionary history of Ducula oceanica in Micronesia. — The 
distribution and evolutionary history of Ducula oceanica have been 
treated by Mayr (1940) and Amadon (1943). These authors place 



198 University of Kansas Publs., Mus. Nat. Hist. 

D. oceanica within a superspecies containing D. pacifica (Melanesia 
to Samoa and Cook Islands), D. aurwea (Society Islands), D. 
galeata (Marquesas Islands), and possibly other species in Papua 
and Malaysia. According to Mayr (1942b:fig. 7), D. pacifica is the 
species which is ancestral to other species of pigeons in Oceania. 
Apparently D. oceanica was derived from this ancestral stock and 
reached Micronesia via the Ellice and Gilbert islands. Records of 
Ducula were obtained in the Gilbert Islands in the days of explora- 
tion; Amadon (1943:11) tentatively refers these to D. o. oceanica. 

The irregular distribution of D. oceanica in the islands of Micro- 
nesia and the fact that the bird exists on both "high" volcanic islands 
as well as on "low" coral atolls suggest that the present population 
may be a remnant of a once more widely distributed one. The fact 
that D. oceanica may be divided into several subspecies shows that a 
greater amount of geographic variation has occurred as compared 
with its probable ancestral stock, D. pacifica, which is virtually un- 
differentiated over most of its extensive range. The pigeon of Micro- 
nesia has a more rounded wing than that of D. pacifica, which might, 
as Amadon has suggested, cause the bird to be more sedentary and 
lend itself more readily to differentiation through geographic isola- 
tion. D. pacifica is known to fly from island to island. As shown by 
the measurements in table 27, the length of wing of D. oceanica dif- 
fers, in the various insular populations, being longer in the west and 
shorter in the east. This cline has been discussed by Amadon 
(1943:11). 

It is interesting that Ducula or some other large pigeon has not 
become established in the Mariana Islands. Ducula is present at 
Yap and Truk, which are not very distant from Guam. Another 
tropical pigeon, Columba vitiensis, has extended its range northward 
and reached the Bonin Islands; probably it arrived there via the 
Philippines or the Riu Kiu Islands. Thus, there are representatives 
of large pigeons on islands to the southeast, south, west and north- 
west of the Marianas, but none has become established in the Mari- 
anas themselves. 

Streptopelia bitorquata dusumieri (Temminck) 

Philippine Turtle Dove 

Columba dusumieri Temminck, PI. col., livr. 32, 1832, p. 188. (Type locality, 
Vicinity of Manila, Luzon, Philippine Islands.) 

Colombe Dussumier Quoy and Gaimard, Voy. "Uranie," Zool., 1824, pp. 35, 680 
(Mariannes); idem,, Ann. Sci. Nat. Paris, 6, 1825, p. 148 (Mariannes). 
■: Columba dusumieri Wagler, Syst. Avium Columba, 1827, p. 206, sp. 99 (Marianis). 

Columba Dussumieri Kittlitz, Obser. Zool., in Lutke, Voy. "Le Seniavine," 3, 1836, 
p. 305 (Guahan). 



Baker: The Avifauna of Micronesia 199 

Streptopelia gaimardi Bonaparte, Consp. Avium, 2, 1854, p. 66 (Type locality, 
Mariannes) ; idem, Comptes Rendus Acad. Sci. Paris, 40, 1855, p. 18 (Mariannes) ; 
Reichenbach, Tauben, 1862, p. 76 (Mariannen). 

TurtUT (Streptopelia) Giamardi Gray, Cat. Birds Trop. Is. Pacific Oceon, 1859, p. 
43 (Guam). 

Turtur gaimardi Gray, Hand-list Birds, 2, 1870, p. 239 (Marian). 

TuTtur dussumieri Schlegel, Mus. Pays-Bas, 6, no. 35, 1873, p. 120 (Mariannes); 
Wiglesworth, Abhandl. Und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 54 
(Marianne); Salvadori, Cat. Birds British Mus., 21, 1893, p. 423 (Mariannes); Ousta- 
let, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, p. 222 (Guam); Hartert, Novit. 
Zool., 5, 1898, p. 60 (Guam, Saipan) ; Wheeler, Report Island of Guam, 1900, p. 13 
(Guam); Seale, Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 43 (Marianas); Saf- 
ford, Osprey, 1902, p. 08 (Marianas); idem, Amer. Anthro., 4, 1902, p. 711 (Guam); 
idem. The Plant World, 7, 1904, p. 264 (Guam) ; Dubois, Syn. Avium, 2, 1904, p. 760 
(Marianne); Safford, Contr. U. S. Nat. Herb., 9, 1905, p. 78 (Guam); Schnee, Zei- 
tschr. f. Naturwisch., 82, 1912, p. 466 (Marianen) ; Prowazek, Die deutschen Marianen, 
1913, p. 101 (Marianen); Reichenow, Die Vogel, 1, 1913, p. 341 (Marianen); Cox, 
Island of Guam, 1917, p. 20 (Guam). 

Streptopelia dussumieri Kuroda, in Momiyama, Birds Micronesia, 1922, p. 54 (Guam, 
Saipan); Mathews, Syst. Avium Australasianarurn, 1, 1927, p. 62 (Marianas); Hand- 
list Japanese Birds, rev., 1932, p. 189 (Saipan, Tinian, Rota). 

Tuttur dessumieri Bryan, Guam Rec, vol. 13, no. 2, 1936, p. 24 (Guam). 

Streptopelia bitorquata dusumieri Peters, Check-list Birds World, 3, 1937, p. 96 
(Marianne); Hand-list Japanese Birds, 3d ed., 1942, p. 211 (Saipan, Tinian, Rota); 
Mayr, Birds Southwest Pacific, 1945, p. 289 (Marianas); Watson, The Raven, 17, 
1946, p. 41 (Guam); Downs, Trans. Kansas Acad. Sci., 49, 1946, p. 96 (Tinian); 
Strophlet, Auk, 1946, p. 538 (Guam); Stott, Auk, 1947, p. 526 (Saipan); Baker, 
Sniithson. Misc. Coll., vol. 107, no. 15, 1948, p. 00 (Guam, Rota). 

Streptopelia bitorquata Baker, Trans. 11th N. American Wildlife Conf., 1946, p. 
208 (Guam); idem. Condor. 49, 1947, p. 125 (Guam). 

Geographic range. — Philippine Islands, Sula Archipelago, northern Borneo. 
In Micronesia: Mariana Islands (introduced) — Guam, Rota, Tinian, Saipan. 

Characters. — Adult: A medium-sized dove with head and nape near ''light 
Quaker drab" with a vinous tinge; chin and upper throat whitish becoming 
near "vinaccous buff" on lower throat and to near "vinaceous-faun" on breast 
and upper abdomen; lower abdomen, vent, and under tail-coverts white; tibia 
grayish ; neck feathers dark with grayish centers and metallic greenish-slate 
edges; color near "Japan rose"; back, rump, upper tail-coverts, scapulars, 
upper wing-coverts, and inner secondaries dark "drab" ; sides, upper wing 
coverts, outer secondaries, and under wing-coverts lead colored; primaries 
blackish edged with light gray; central tail feathers like back but paler, outer 
feathers of tail darker with brownish tinge on edges; outermost tail feathers 
blackish tipped with gray and with outer webs whiti.sh; bill dark; feet reddish; 
iris orange. 

Measurements. — Measurements of 15 adult males from Guam, Rota and 
Tinian: wing, 154-162 (158); tail, 127-135 (130); culmen, 16.2-18.1 (17.2); tar- 
sus, 24-27 (25.5); of 10 adult females from Guam and Rota: wing, 150-162 
(156); tail, 120-130 (127); culmen, 16.2-19.1 (17.5); tarsus, 24-26 (25). No 
differences in measurements were found between populations from Guam, Rota 
and Tinian. 

Weights. — The author (1948:61) reports the weights of five adult males as 
130-167 (152) and of six adult females as 135-159 (146). These birds were 
taken at Guam. 



200 University of Kansas Publs., Mus. Nat. Hist. 

Specimens examined. — Total number, 27 (16 males, 11 females), as follows: Mariana Is- 
lands, USNM— Guam, 21 (Feb. 7, May 25, 2C, June 9, July 6, 7, 10, 18, 23, Aug. 2, 11, 
Sept. 8, Oct. 8)— Rota, 4 (Oct. 18, 22, 23, Nov. 2)— Tinian, 2 (Oct. 24, 25). 

Nesting. — The NAMRU2 party found evidences of nesting by this dove at 
Guam in February, March, April, and June. Nests were observed on May 29 
and June 28. On the latter date a nest containing one nestling and one un- 
hatched egg was found near Mount Santa Rosa. The nest was situated ap- 
proximately five feet from the ground in a low bush. Two eggs taken by 
Necker at Rota on October 31, 1945, are white and measure 29.6 by 23.0 and 
30.1 by 23.0. Strophlet (1946:538) observed a bird carrying nest materials at 
Guam on November 13. Hartert (1898:60) reports on nests found at Guam 
in April and May. Each nest contained one egg. It is probable that this 
bird nests at all times of the year. The nuptial flight of these birds reminds 
one very much of that of the mourning dove of North America. 

Remarks. — The Philippine Turtle Dove was introduced from the 
Philippines to Guam and other islands of the southern Marianas 
by the Spanish probably in the 18th Century; it was in 1771-1774 
that the Philippine deer (Riisa) was introduced to Guam. Perhaps 
these birds were initially introduced as caged birds or possibly were 
liberated to offer hunting for the colonial governors. They have 
been a very successful introduction and are well established. At 
Guam (see Baker 1947b: 124), this species comprised 15.5 percent 
of all birds seen along roadways. Although open areas appear to 
be preferred by this dove and although it may be on the increase 
owing to the clearing operations of the war effort, it appears to be 
equally adapted to forested areas and coconut groves. It feeds on 
the ground to a large extent, fitting into an ecologic niche which 
few other species of birds of the islands occupy. It was even ob- 
served feeding on sandy beaches and tidal fiats at low tide. 

In 1931, Coultas found the dove to be numerous at Guam, but 
thought that it was in danger of extinction at Tinian and Saipan 
owing to extensive hunting. Downs (1946:96) reported that in 1945 
the dove was abundant at Tinian. Gleise (1945:22) estimated the 
population of these doves at 300 on Tinian in 1945. From the re- 
marks of Stott (1947:526), we may assume that the population at 
Saipan is in no immediate danger of extinction. 

A comparison of specimens from the Marianas with those from 
the Philippines reveals no significant difference between the two. 
Bonaparte described the dove in the Marianas as new, naming it 
Sterptopelia gaimardi. The name Turtur prevostianus has been 
used by some authors to denote the dove in the Marianas, but this 
was through error as explained by Salvadori (1893:410). This name 
refers to a dove found on Marianne, an island of the Seychelles in 
the Indian Ocean. 



Baker: The Avifauna of Micronesia 201 

Gallicolumba canifrons (Hartlaub and Finsch) 
Palau Ground Dove 

Phlegoenas caiujrans Hartlaub and Finsch, Proc. Zool. Soc. London, 1872, p. 101. 
(Type locality, Pelew Islands.) 

Phlegoenas canifrons Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 5, 27, pi. 5, fig. 
1 (Palau); Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, p. 407 (Palau); 
Wliglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 57 
(Pelew); Hartert; Novit. Zool., 5, 1898, p. 61 (Pelew); Matschie, Journ. f. Ornith., 
1901, p. 113 (Palau); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 53 (Pelews). 

Phlegoenas canifrons .Sclater, Proc. Zool. Soc. London, 1877, p. 112 (Pelew); Sal- 
vadori, Ornith, Papuasia, 3, 1882, p. 169 (Pelew); idem. Cat. Birds British Mus., 21, 
1893, p. 592 (Pelew); Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, p. 
227 (Palaos); Bolau. Mitteil. Naturhist. Mus. Hamburg, 1898, p. 68 (Palau); Dubois, 
Syn. Avium, 2, 1904, p. 772 (Pelew). 

Phaps canifrons Giebel, Thes. Ornith., 3, 1877, p. 89 (Pelew). 

Gallicolumba canifrons canifrons Mathews, Syst. Avium Australasianarum, 1, 1927, 
p. 74 (Pelew). 

Gallicolumba canifrons Hand-list Japanese Birds, rev., 1932, p. 189 (Palau); Mayr, 
Amer. Mus. Novit., no. 828, 1936, p. 4 (Palau); Peters, Check-list Birds World, 3, 
1937, p. 136 (Palau); Hand-list Japanese Birds, 3d ed., 1942, p. 211 (Babelthuap) ; 
Mayr, Birds Southwest Pacific, 1945, p. 290 (Palau); Mayr, Audubon Mag., 47, 1945, 
p. 282 (Palau); Baker, Smithson. Misc. Oil., vol. 107, no. 15, 1948, p. 62 (Gara- 
kayo, Peleliu). 

Geographic range. — Micronesia: Palau Islands — Babelthuap, Koror, Gara- 
kayo, Peleliu, Ngabad, Angaur. 

Characters. — Adult male: A small, ground dove with forehead, crown, sides 
of head, chin, throat, and breast ashy gray, lighter on forehead, chin, and 
throat, and washed with "light vinaceous-faun" on breast; occiput, nape and 
mantle dark "ferruginous"; rest of upper parts glossed with bronze-olive; 
lesser and middle wing-coverts tipped with metallic purple; wings reddish- 
brown with dark brown tips; under side of wing reddish-brown to brown; 
abdomen, vent and under tail-coverts dark grayish-brown ; tail colored like 
back, outer feathers have a paler brown terminal band rather obscure; bill 
horn colored; feet red; iris brown. 

Female : A female molting into adult plumage is cinnamon colored, mottled 
with dark brown; on the back an olive-green sheen is beginning to appear; tail 
brown with some greenish sheen; tips of tail edged with light brown; bill and 
feet light brown. 

Measurements. — Measurements of six adult males are: wing, 112-119 (115); 

tail, 65-72 (70) ; exposed culmen, 15.3-16.1 (15.7) ; tarsus, 30.1-31.2 (30.8) ; of one 

female in postjuvenal molt: wing, 107; tail, 69; exposed culmen, 17.1; tarsus, 30. 

Specimens examined. — Total number, 8 (7 males, 1 female), as follows: Palau Islands, 
USNM— Koror, 1 (Nov. 18)— Garakayo, 2 (Sept. 17, 19)— Peleliu, 2 (Sept. 1, Dec. 5)— 
Ngabad, 1 (Sept. 11); AMNH — exact locality not given, 1 (Dec. 1). 

Food haibts. — Stomachs of specimens taken by the NAMRU2 party at 
Peleliu and Garakayo contained one and one-half to two cc. of hard seeds and 
seed parts. 

Remarks. — The Palau Ground Dove, according to Amadon (1943: 
19), is a member of a superspecies containing G. hoedti (Wetar), 
G. beccarii (New Guinea, Bismarcks, Solomons), G. sanctaecrucis 
(Santa Cruz, New Hebrides), and G. stairi (central Polynesia). 



202 University of Kansas Publs., Mus. Nat. Hist. 

G. canifrons apparently came to the Palaiis from either New 
Guinea or the region of the Bismarck Archipelago, evolving from 
G. beccarii or some related form. The Palau Ground Dove has a 
copper-colored occiput, nape, and shoulder patch, but otherwise it 
resembles this Melanesian species, G. becarii. Amadon (1943:20) 
discusses two types of plumage of females in G. stairi; one is a male 
type of plumage. The lack of female specimens prevents me from 
determining whether this characteristic is present in G. canifrons. 

Coultas (field notes) had difficulty in obtaining even one specimen 
of G. canifrons in the Palaus in 1931. He concluded that either the 
bird was practically extinct or that he just could not find it. From 
the experience of the NAMRU2 party in the southern Palaus in 1945, 
I would think that he merely did not find the bird. Although it is 
probably rare in comparison with some other members of the family 
Columbidae of these islands, we found this bird on most of the islands 
visited. 

The NAMRU2 party arrived at Palau expecting to find the ground 
dove a fairly conspicuous member of the avifauna and expecting to 
see it sitting in trees and flying across the roads much in the same 
manner as did the ground dove at Guam, G. x. xanthonura. At first, 
we did not find the bird, but in the dense jungles a low, penetrating, 
and intermittent, call was heard which may be described as a moan. 
This was the call of the ground dove. The bird was difficult to dis- 
cover because its color blended so well with the shadows and dark 
background of the coral rocks and forest litter. The bird was very 
active and moved along rapidly pecking at food particles. Also it 
was wary. Once the distinctive call note was recognized, it was not 
difficult to locate the area in which the bird was living; however, 
finding the bird was difficult. On one occasion I stalked a dove for 
at least a half an hour knowing that it was always within fifty yards 
of me. A bird that was flushed, flew about twenty-five feet and 
dropped down in open forest litter and disappeared. On the basis of 
specimens collected and call notes heard, I estimate that the popula- 
tion of the Palau Ground Dove on the islands visited in 1945 was as 
follows: Peleliu — a minimum of 15 (found in most forested areas 
which were not greatly damaged by the invasion operations) ; Gara- 
kayo — a minimum of 10 (the doves were found to live equally well 
on the steep hillsides or in flat jungle on this islet) ; Ngabad — 5 to 10 
(doves were heard in several areas on this small islet) ; Angaur — not 
estimated (one call was heard in brush near the edge of a fresh water 
lake). 



Baker: The Avifauna of Micronesia 203 

Gallicolumba xanthonura xanthonura (Temmiack) 
White-throated Ground Dove 

Columba xanthonura Temminck, PI. col., livr. 32, 1823, pi. 190. (Type locality, 
Mariannes.) 

Columba xanthonura Lesson, Compl. de Buffon, 2nd ed., 2, Oiseau.K, 1838, p. 281 
(Mariannes). 

Columba Pampusan Quoy and Gaimard, Voy. "Uranie," Zool., 182-1, pp. 121, 681, 
pi. 30 (Mariannes) ; Dumont, Diet. Sci. Nat., ed. Levrault, 40, 1826, p. 345 (Guam) ; 
Lesson, Traite d'Ornith., 1831, p. 471 (Mariannes); Hartlaub, Journ. f. Ornith., 1854, 
p. 167 (Mariannen). 

Columba erythroptera Lesson, Traite d'Ornith., 1831, p. 471 (Mariannes); Kittlitz, 
Obser. Zool., in Lutke, Voy. "Le Seniavine," 3, 1836, p. 305 (Guahan); Hartlaub, Journ. 
f. Ornith., 1854, p. 167 (Mariannen). 

Columba xanthura Prevost and Knip, Los Pigeons, 2, 1838-43, p. 45, pi. 23 (Guam). 

Pampusana xanthua Bonaparte, Consp. Avium, 2, 1854, p. 89 (Mariannis); idem, 
Comptes Rendus Acad. Sci. Paris, 40, 1855, p. 207 (Mariannes); Reichenbach, Tauben, 
1861, p. 39 (Guam). 

Caloenas (Pampusana) xanthura Gray, Cat. Birds Trop. Is. Pacific Ocean, 1859, p. 
45 (Guam). 

Phlegoenas erythroptera Reichenbach, Tauben, 1861, p. 41 (Mariannen). 

Caloenas xanthura Gray, Hand-list Birds, 2, 1870, p. 247 (Marian). 

Phlegoenas yapensis Hartlaub and Finsch, 1872, p. 102 (TjTJe locality, Uap); 
Graffe, Journ. Mus. Godeffroy, 2, 1873, pp. 122, 123 (Yap); Schmeltz and Krause, 
Ethnogr. Abth. Mus. Godeffroy, 1881, p. 391 (Yap); Wiglesworth, Abhandl. und Ber. 
Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 57 (Yap); Hartert, Novit. Zool., 5, 
1898, p. 61 (Yap); Matschie, Journ. f. Ornith., 1901, p. 113 (Yap). 

Pampusana rousseaui Hartlaub and Finsch, Proc. Zool. See. London, 1872, p. 103 
(Marianne). 

Phaps erythroptera Giebel (part), Thes. Ornith., 3, 1877, p. 89 (Marianne). 

Phaps xanthura Giebel, Thes. Ornith., 3, 1877, p. 91 (Marianne). 

Phaps yapensis Giebel, Thes. Ornith., 3, 1877, p. 91 (Uap). 

Phlegoenas virgo Reichenow. Journ. f. Ornith., 1885, p. 110 (Type locality, Palau- 
Inseln, error =: Guam). 

Phlogaenas erythroptera Oustalet, Le Nat., 1889, p. 261 (Mariannes). 

Phlegoenas pampusan Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 
1890-1891 (1891), p. 55 (Marianne); Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, 
(3), 7, 1895, p. 224 (Saypan, Guam, Rota). 

Phlogoenas yapensis Salvador!, Cat. Birds British Mus., 21, 1893, p. 593 (Uap); 
Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, p. 227 (Mackensie); Bolau, 
Mitteil. Naturhist. Mus. Hamburg, 1898, p. 68 (Yap); Dubois, Syn. Avium, 2. 1904, 
p. 772 (Uap). 

Phlogoenas pampusan Salvadori, Cat. Birds British Mus., 21, 1893, p. 602 (Mari- 
anne). 

Phlegoenas xanthonura Hartert, Novit. Zool., 5, 1898, p. 60 (Guam, Saipan) ; 
Wheeler, Report Island of Guam, 1900, p. 13 (Guam) ; Matschie, Journ. f. Ornith., 
1901, p. 113 (Guam, Saipan) ; Safford, Amer. Anthro., 4, 1902, p. 711 (Guam) ; idem, 
Osprey, 1902, p. 68 (Mariannas); idem. The Plant Worid, 7, 1904, p. 264 (Guam); 
idem, Contr. U. S. Nat. Herb., 9, 1905, p. 78 (Guam); Cox, Island of Guam, 1917, 
p. 20 (Guam). 

Phlogoenas xanthonura Seale, Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 42 
(Marianas) ; Reichenow, Die Vogel, 1, 1913, p. 331 (Mariannen) ; Brj'an, Guam Rec., 
vol. 13, no. 2, 1936, p. 24 (Guam). 

Phlegoenas xanthonura xanthonura Kuroda, in Momiyama, Birds Micronesia, 1922, 
p. 54 (Guam, Rota, Saipan). 

Phlegoenas xanthonura yapensis Kuroda, in Momoyama, Birds Micronesia, 1922, p. 
54 (Yap). 

Gallicolumba xanthonura Mathews, Syst. Avium Australasianarum, 1, 1927, p. 75 
(Marianas, Mackenzie); Hand-list Japanese Birds, rev., 1932, p. 189 (Pagan, Almagan, 
Saipan, Tinian, Rota, Mackenzie); Mayr, Amer. Mus. Novit., no. 828, 1936, p. 4 



204 



University of Kansas Publs., Mus. Nat. Hist. 



(Marianne); Peters, Check-list Birds World, 3, 1937, p. 136 (Marianne, Yap); Hand- 
list Japanese Birds, 3d ed., 1942, p. 211 (Yap, Assongsong, Pagan, Almagan, Saipan, 
Tinian, Rota); Strophlet, Auk, 1946, p. 538 (Guam); Wharton, Ecol. Monogr., 16, 
1946, p. 174 (Guam); Baker, Condor, 49, 1947, p. 125 (Guam). 

Gallicolumba canifrons yapensis Mathews, Syst. Avium Australasianarum, 1, 1927, 
p. 74 (Yap). 

Terricolumba xanthonura Yamashina, Tori, 10, 1940, p. 677 (Assongsong). 

Gallicolumba xanthonura xanthonura Mayr, Birds Southwest Pacific, 1945, p. 290 
(Marianas, Yap); Watson, The Raven, 17, 1946, p. 41 (Guam); Stott, Auk, 1947, p. 
526 (Saipan); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 61 (Guam, Rota 
Yap). 

Gallecolumba xanthonura xanthonura Downs, Trans. Kansas Acad. Sci., 49, 1946, 
p. 96 (Tinian). 

Geographic range. — Micronesia: Mariana Islands — Asuncion, Pagan, Alma- 
gan, Saipan, Tinian, Rota, Guam; Caroline Islands — Yap. 

Characters. — Adult male: Forehead, face, chin, throat, and upper breast 
white, lightly washed with pale buff; crown, occiput, sides of head, and nape 
rusty brown to dark brown; rest of upper parts dark bronze-olive; feathers of 
mantle and upper wing-coverts broadly edged with metallic purple-violet; pri- 
maries, under wing-coverts and axillaries brown; tail, lower breast and rest of 
underparts dark brown; bill and feet dark brown. 




Fig. 14. Geographic distribution of Gallicolumba of Micronesia and Eastern 
Polynesia and routes of its dispersal. (1) G. jobiensis; (2) G. x. kubaryi; 
(3) G. X. xanthonura; (4) G. erythroptera; (5) G. rubescens. 



Adult female: Resembles adult male, but smaller and with underparts 
colored between "ochraceous-tawny" and "cinnamon brown" instead of dark 
brown and white; head and neck darker and with more rufous than underparts; 
remainder of upper surface resembles underparts but with striking olive green 
sheen, especially on upper wing-coverts; primaries brown but outer webs 
lighter; tail rufous-brown, with a broad, black subterminal band. 

The male type of plumage in the adult female is: breast light drab tinged 
with light brown and darkening anteriorly; crown resembles that of normal 
female although darker and becoming lighter and grayer on neck and nape; 



Baker: The Avifauna of Micronesia 



205 



shoulder and wing-coverts compare favorably with that of adult male although 
lighter and with yellowish tinge ; back bronzed olive-green as in normal female 
but mantle with a few purplish feathers characteristic of male; abdomen near 
"olive brown" with buffy-brown edges to feathers. 

Immature male: Resembles adult male, but head and nape darker brown; 
throat and upper breast may be more brown and less white. 

Immature female: Resembles adult female, but with more rufous coloring; 
olive-green sheen on feathers reduced in amount or absent. 

Measurements. — Measurements are found in table 28. 



Table 28. Me-asurements of Gallicolumba xanthonura 



Subspecies 


Number 


Wing 


Tail 


Culmen 


Tarsus 


G. 


X. xanthonura . . 


43 males .... 


146 
(139-153) 


102 
(97-111) 


22.0 
(21.0-23.0) 


32 

(31-33) 






31 females. . 


136 
(131-141) 


94 
(90-98) 


20.5 

(20.0-21.5) 


30 

(28-32) 


a 


X. kubaryi 


7 males . . . 


157 

(152-160) 




23.0 

(20.5-23.5) 


35 






(33-35) 






7 females . . 


148 
(145-151) 




23.0 

(22.5-23.5) 


33 






(32-34) 









There is little difference in the measurements of specimens from Guam, 
Rota, Tinian, Saipan, and Asuncion. No specimens from Yap were available 
for examination. 

Weights. — The NAMRU2 party obtained weights of this ground dove from 
Guam as follows: seven adult males 119-154 (130); seven adult females 96- 
150 (118). 

Specimens examined. — Total number, 96 (50 males, 38 females, 8 unsexed) as follows: 
Mariana Islands, USNM — Guam, 29 (Mar. 18, April 4, 17, May 20, 28, June 2, 9, 13, 14, 
15, 20, 23, 24, 27, 28, July 2, 6, 10, 23, Aug. 11, 21)— Rota, 6 (Oct. 20, 22, 25, 26, Nov. 1. 
2)— Tinian, 4 (Oct. 24, 26); AMNH— Guam, 40 (Jan. 17, 30, Feb. 12, 20, March 3, 5, 7, 11, 
13, 23, April 13, 19, June 13, 15, July 10, 25, Aug. 4, 10, 11, 13, 15, 18, 20, 21, 22, Sept. 
4, Dec. 26, 30)— Tinian, 8 (Sept. 7, 10, 11, 12, 13) — Saipan, 6 (July 13, 15, Aug. 24, Sept. 
7, 8) — Asuncion, 3 (Jan. 18, Feb. 7, June). 

Nesting. — The NAMRU2 party found the ground dove nesting at Guam 
in the winter and spring months beginning in late Januar>\ Nests were ob- 
served in tall trees, many of which were well isolated from other trees and 
vegetation. On February 10 a nest was discovered in a breadfruit tree near 
one of the NAMRU2 barracks on Oca Point. It was approximately 50 feet 
above the ground. On February 26 I found pieces of egg shell beneath the 
tree. Occasionally during the day, the male, but never the female, was ob- 
served sitting on this nest. On February 10, a dove (the male) was observed 
building a nest in a large banyan tree at Oca Point. Another nest was being 
constructed by a female on March 7. On March 17 a young female dove, 
just beginning to fly, was taken; another was found on April 3. Adult birds 



206 Univeesity of Kansas Publs., Mus. Nat. Hist. 

with enlarged gonads were taken in April, May, June, and July. Marche, ac- 
cording to Oustalet (1895:224), obtained eggs in May, 1887. 

Food habits. — Stomachs of doves taken at Guam contained fruits and fruit 
parts. On March 9, a dove was observed feeding on the berries of the shrub 
known as ''inkbush." This appeared to be a favorite food. Scale (1901:42) 
also mentions that this berry is a preferred food. 

Parasites. — Wharton (1946:174) lists the chigger (Acarina), Trombicula sp., 
from the ground dove at Guam. 

Remarks. — At Guam, the NAMRU2 party observed the ground 
dove to be fairly common in 1945. Along roadways, the present 
author (1947b: 124) found that individuals of this species comprised 
2.5 percent of the total population of birds observed, and the ground 
dove was seen on 31.2 percent of 125 road counts made. The male 
was much more in evidence than the female and was frequently 
seen flying high over the roadways and jungle areas; eighty percent 
of the ground doves seen while road-counts were being made were 
males. The female was found less frequently; it was a less conspic- 
uous bird and was seen only occasionally in flight. Neither sex 
appeared to have the secretive, terrestrial habits of G. canijrons of 
the Palau Islands. On the basis of our observations at Guam, I 
would say that the name "ground dove" for the bird at Guam is 
not descriptive. The birds were found to spend considerable time 
in tall trees; the closest that I saw them to the ground was when 
they were feeding only three to four feet from the ground in the 
ink berry bushes. 

The call note of this dove is much like that of the Palau Ground 
Dove; Scale (1901:42) describes it as follows, "These pigeons seem 
to prefer the deep jungle, from whence their deep low moan, like 
the sound of a man dying in great distress, comes with a wierd un- 
canny effect, heightened by the gloom and darkness of the unknown 
forest. , . . This sound, which always seems to come from a long 
distance, is very misleading, and one is considerably surprised to 
find he is perhaps within a few feet of the bird." Scale writes that 
they were very common on Guam in 1900. In 1931, Coultas found 
the dove "quite common at the north end of the island." The bird 
apparently prefers the dense forest or second growth brushy areas, 
but was found also in the partly cleared areas surrounding the 
NAMRU2 headquarters at Oca Point in 1945. At Rota, the 
NAMRU2 party found the birds to be numerous in 1945. Coultas 
observed only a few birds on Tinian in 1931; Downs (1946:96) 
found only a small population at this island in 1945. The extensive 
cultivation and clearing activities at Tinian have removed much of 



Baker: The Avifauna of Micronesia 207 

the habitat suitable for these, as well as other birds. At Saipan, 
Stott (1947:526) writes that the bird is common on "brush-covered 
hillsides and semi-wooded country." There is little information pub- 
lished regarding the status of this dove in the northern Marianas. 

Gallicolumba xanthonura kubaryi (Finsch) 
White-throated Ground Dove 

Phlegoenas Kubaryi Finsch, Journ. f. Omith., 1880, p. 292. (Type locality. Ruck 
and Ponape.) 

Phlegoenas erythroptera Bonaparte, Consp. Avium, 2, 1854, p. 89 (Carolines) ; 
Reichenbach, Tauben, 1862, p. 41 (Carolines); Finsch, Proc. Zool. See. London, 1877 
(1878), p. 780 (Ponape); idem, Ibis, 1881, p. 115 (Ponape); Schmeltz and Krause, 
Ethnogr. Abth. Mus. Godeffroy, 1881, pp. 281, 353 (Ponape, Ruk) ; Tristram, Cat. 
Birds, 1889, p. 41 (Ruk). 

Phlegoenas kubaryi Reichenow and Schalow, Journ. f. Omith., 1881, p. 75 (Ruk, 
Ponape); Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), 
p. 55 (Ruk, Ponape); Hartert, Novit. Zool., 7, 1900, p. 8 (Ruk, Ponape); Matschie, 
Journ. f. Oniith., 1901, p. 113 (Ruck, Ponape); Kuroda. in Momiyama, Birds Micro- 
nesia, 1922, p. 53 (Ruk, Ponape). 

Phlogoenas erythroptera Finsch, Proc. Zool. Soc. London, 1880, p. 576 (Ponape, 
Ruk); Takatsukasa and Kuroda, Tori, 1, 1915, p. 52 (Ruk). 

Phlogoenas kubaryi Salvadori, Cat. Birds British Mus., 21, 1893, p. 599 (Ruk, 
Ponape); Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, p. 227 (Caroline 
= Truk) ; Bolau, Mitteil. Naturhist. Mus. Hamburg, 1S98, p. 68 (Ruck); Reichenow, 
Die Vogel, 1, 1913, p. 331 (Karolinen). 

Phlegoenas kubaryi Christian, The Caroline Islands, 1899, p. 357 (Ponape). 

Gallicolumba kubaryi Mathews, Syst. Avium Australasianarum, 1, 1927, p. 74 
(Caroline Is.); Hand-list Japanese Birds, rev., 1932, p. 189 (Truk, Ponape); Peters, 
Check-list Birds World, 3, 1947, p. 136 (Ruk, Ponape); Mayr, Proc. 6th Pacific Sci. 
Congr., 4, 1941, p. 204 (Ponape); Bequaert, Mushi, 12, 1939, p. 81 (Ponape); idem 
Occ. Papers Bernice P. Bishop Mus., 16, 1941, p. 266 (Ponape); Hand-list Japanese 
Birds, Sd ed., 1942, p. 211 (Truk, Ponape). 

Gallicolumba xanthonura kubaryi Mayr, Birds Southwest Pacific, 1945, p. 290 (Truk, 
Ponape); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 62 (Truk). 

Geographic range. — Micronesia: Caroline Islands— Truk, Ponape. 

Characters. — Adult male: Resembles adult male of G. x. xanthonura, but 
larger with crown, nape, and hind neck sooty-black; upper back and lesser 
upper wing-coverts purplish-violet, extending lower on back than in G. x. 
xanthonura. 

Adult female : Resembles adult male, but smaller and paler with upper back 
glossy, bronze-green margined with purplish-violet; lower back and rump glossy, 
olive-green; upper tail-coverts greenish-brown; central tail feathers blackish- 
brown; innermost secondaries bright, glossy green tinged with bluish. 

Measurements. — Measurements are presented in table 28. 

Specimens examined. — Total number, 21 (9 males, 11 females, 1 unsexed), as follows: 
Caroline Islands, USNM— Truk, 1 (July); AMNH— Ponape, 13 (Nov. Dec.)— Truk, 7 (Jan., 
Feb., May). 

Nesting. — At Ponape in November and December, Coultas obtained speci- 
mens which had enlarged gonads. He did not find the nest of this bird but 
writes (field notes) that the natives told him that the nest is placed in the 
top of the tree fern 10 or 15 feet above the ground. In contrast, the ground 
dove at Guam may select a nesting site considerably higher in the tree. Coul- 
tas reports that one egg is laid by C. x. kubaryi. 



208 University of Kansas Publs., Mus. Nat. Hist. 

Food habits. — Coultas (field notes) writes that the bird feeds and Uves on 
the ground at Ponape. He lists food as small snails, seeds, and worms. 

Parasites. — Bequaert (1939:81 and 1941:266) records the fly (Hippobosci- 
dae), Ornithoctona plicata, from the ground dove at Ponape. 

Remarks. — Coultas (field notes) writes that in 1930 the ground 
dove at Ponape was rare in the forested areas and generally found 
more along the sea coast and in the upland valleys. Coultas de- 
scribes its call as an infrequent shrill, whistle-like call. He writes 
that hunting by the Japanese and natives was reducing the popu- 
lation of G. X. kubaryi at Ponape in 1930. In 1945, McElroy of the 
NAMRU2 party found the dove at Truk on forested slopes in tall 
trees, and reported that its habits at Truk were similar to those of 
C. X. xanthonura at Guam. In 1947-1948, Richards noted {in litt.) 
that the dove at Ponape was rare (he saw only one specimen). At 
Truk, he found the bird to be "rather common" in thickets, dry 
gullies, and flying over grassy slopes. He found the bird near sea 
level, never in country above 300 feet in altitude and not in deep 
forest. I offer no explanation for the conflicting reports concerning 
the habits of this species, unless it be that the bird is capable of 
varying its habits to fit particular habitats; for example, in jungle 
areas it may be ground-living and in open woodlands it may be 
tree-living. 

Evolutionary history of Gallicolumba in Micronesia. — There have 
been two unrelated invasions of Micronesia by the genus Galli- 
colimiba. One invasion established G. canifrons at the Palau Is- 
lands. The other established the populations of G. xanthonura in 
the Caroline and Mariana islands, Mayr (1936:4) points out that 
G. xanthonura is related to G. jobiensis (New Guinea and Northern 
Melanesia), G. erythroptera (Society and Tuamotu islands), and 
G. rubescens (Marquesas Islands). This group may be regarded as 
a superspecies. The adults of G. jobiensis, the male and female, re- 
semble one another. In both, the head, neck, and auriculoloral 
stripes are sooty-black; the eye stripe, chin, throat, and breast are 
white; the abdomen is dark; and the upper parts are blackish with 
a coppery sheen. Immatures are rusty-brown. G. xanthonura is 
closely related to G. jobiensis, and they conceivably, along with G. 
erythroptera, might be considered conspecific. The close relation- 
ship between the G. xanthonura in Micronesia and G. erythroptera 
has been noted by Oustalet (1896:71). Among named kinds, G. x. 
kubaryi most closely resembles G. jobiensis with sooty-black color- 
ing present on the head. The male and female of G. x. kubaryi 



Bakeb: The Avifauna of Micronesia 209 

closely resemble each other, although immature type of plumage 
may occur in adult females as indicated by the immature plumage 
of a bird containing well-developed eggs taken at Ponape by Coul- 
tas. 

In G. X. xanthonura the male lacks the sooty-black head and has 
lost some of the coppery sheen from the middle of the back. The 
female has taken on the immature type of plumage, except for 
occasional near-male type plumage. In G. erythroptera the male 
has lost some of the sooty-black coloring on the forehead, anterior 
crown, and loral area and some of the coppery sheen in the middle 
of the back. The female of G. erythroptera resembles the female 
of G. X. xanthonura except that the throat and breast are faintly 
outlined by the brownish color. The head and malar stripe are also 
outlined in this manner. Some females have some coppery gloss on 
the shoulder and a few white feathers on the breast; these may be 
considered as in the near-male type of plumage. 

The tendencies in the evolution of these insular populations of 
Gallicolumba include a reduction of sooty-black on the head and 
a reduction of coppery gloss on the back of the male and the reduc- 
tion of malelike plumage in the female. G. rubescens of the Mar- 
quesas Islands is smaller and darker. It retains the coppery gloss 
on the back and has, in addition, a white bar on the tail and one 
on the wing. On the basis of color and structural characters, it is 
apparent that this superspecics of Gallicolumba has evolved from 
a center of evolution in the region of New Guinea (as shown in 
figure 14) with a colonization of Micronesia, from which (probably 
from G. x. kubaryi) an invasion of eastern Polynesia occurred es- 
tablishing G. erythroptera in the Society and Tuamotu islands, al- 
though it is also possible that G. erythroptera may have reached 
Polynesia by way of a more direct route from Melanesia. Such a 
pathway of colonization as that just described is not unusual since 
representatives of other general including Acrocephalus, Myzomela, 
and Zosterops may have followed similar paths of dispersal from 
Micronesia into Polynesia. Apparently a population isolated in the 
Marquesas has evolved the distinctive G. rubescens. 

Caloenas nicobarica pelewensis Finsch 
Nicobar Pigeon 

Caloenas nicobarica var. pelewensis Finsch, Journ. Mus. Godeffroy, 8, 1875, p. 159 
(in reprint p. 27). (Type locality, Palau.) 

Caloenas nicobarica pelewensis Mathews, Syst. Avium Australasianarum, 1, 1927, p. 
77 (Pelew); Hand-list Japanese Birds, rev., 1932, p. 188 (Palau); Peters, Check-list 
Birds World, 3, 1937, p. 139 (Palau); Hand-list Japanese Birds, 3d ed., 1942, p. 210 

14—8131 



210 University of Kansas Pitbls., Mus. Nat. Hist. 

(Babelthuap, Koror); Mayr, Birds Southwest Pacific, 1945, p. 291 (Palau) ; Baker, 
Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 62 (Garakayo). 

Caloenas nicobarica Salvadori, Ornith. Papuasia, 3, 1882, p. 211 (Pelew); Wigles- 
worth, Abhandl. und. Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 57 (Pelew). 

Caloenas pelewensis Salvadori, Cat. Birds British Mus., 21, 1893, p. 618 (Pelew); 
Bolau, Mitteil. Naturhist. Mus. Hamburg, 1898, p. 69 (Palau); Matschie, Journ. f. 
Ornith., 1901, p. 113 (Palau); Reichenow, Die Vcigel, 1, 1913, p. 328 (Palauinseln) ; 
Takatsukasa and Kuroda, Tori, 1, 1915, p. 52 (Pelew). 

Caloenas nicobaricus pelewensis Kuroda, in Momiyania, Birds Micronesia, 1922, p. 
53 (Pelew). 

Geographic range. — Micronesia: Palau Islands — Babelthuap, Koror, Grar- 
kayo. 

Characters. — Adult: A large heavy-bodied pigeon with head, neck, and upper 
breast blackish; rest of plumage metallic bluish-green with coppery sheen; 
wings glossy green ; tail and under tail-coverts white ; feathers of hind-neck long 
and lanceolate; bill heavy and slightly hooked with lump at base. 

Resembles C. n. nicobarica (Linnaeus), but slightly .smaller and with upper 
parts metallic bluish-green and underparts darker and less green. 

Measurements. — One adult female measures: wing, 232; tail. 82; culmen, 31; 
tarsus, 44; one immature female: wing, 236; tail. 89; culmen, 32; tarsus, 45. 

Specimens examined. — Total number, three females from Palau Islands, AMNH — exact 
locality not given (undated). 

Remarks. — C. nicobarica is (iistributed from the Nicobar Islands 
east through Malaysia to Melanesia as a single undifferentiated form. 
In the northeasternmost part of its range, in the Palau Islands, it 
exhibits geographic variation and is considered to be subspecifically 
distinct from the rest of the population. C. nicobarica appears to 
have no close relatives. It may represent the last remnant of some 
ancient group of pigeons. 

The Nicobar Pigeon is rare. Coultas, who visited the islands in 
1931, did not obtain the bird. The only specimens available for study 
are those in the collections of the American Museum of Natural His- 
tory taken by Kubary in the period between 1870 and 1880. The 
NAMRU2 party did not obtain specimens but saw the bird on five 
occasions at the island of Garakayo in the middle Palaus. The 
writer expected the bird to be ground-living in habit, but the individ- 
uals, which I saw at Garakayo, were either perched on scrubby vege- 
tation on high and inaccessible cliffs or were flying high overhead. 
In its flight overhead, the short, white tail was a particularly con- 
spicuous mark of identification. The flight reminded me very much 
of that of the Black Vulture (Corogyps atrdtus) of North America. 
No birds were found at Peleliu or Angaur, and the small population 
of this pigeon that remains is probably restricted to uninhabited 
coral islets, as Mayr (1945a: 291) has already noted. Marshall 
(1949: 207) saw one bird on Peleliu and one on Koror in November 
and December, 1945. This endemic subspecies is probably on the 



Baker: The Avifauna of Micronesia 211 

road to extinction unless governmental protection can be established 
and enforced. 

Trichoglossus rubiginosus (Bonaparte) 

Ponape Lory 

Chalcopsitta rubiginosus Bonaparte, Comptes Rendus Acad. Sci. Paris, 30, February, 
1850, p. 134; Consp. Avium, 1, after April 15, 1850, p. 3. (Type locality, "ex Insulis 
Barabay et Guebe," error = Ponape.) 

Chalcopsitta rubiginosus Bonaparte, Proc. Zool. Soc. London, 1850, p. 26, pi. 16 
("Ins. Barabay et Guebe," error = Ponape); Pelzeln, Reise "Novara," Vogel, 1865, pp. 
99, 162 (Puynipet); Reichenow, Journ. f. Omith., 1881, p. 162 ("Nordwestl. Poly- 
nessische subregion Carolinen" = Ponape); Tristram, Cat. Birds, 1889, p. 73 (Ponape); 
Finsch, Deut. Verein zum Schultze der Vogelwelt, 18, 1893, p. 458 (Carolinen = 
Ponape); Matschie, Journ. f. Omith., 1901, p. 112 (Ponape). 

Domicella rubiginosa Finsch, Die Papageien, 2, 1868, p. 781 (Puynipet); Hartlaub 
and Finsch, Proc. Zool. Soc. London, 1872, p. 88 (Puinipet). 

Lorius rubiginosus Gray, Hand-list Birds, 2, 1870, p. 153 (Puynipet); Schlegel, 
Mus. Pays-Bas, 3, no. 38, 1874, p. 58 (Puynipet). 

Loriuis rubiginosa Giebel, Thes. Ornith., 2, 1875, p. 502 (Senjawin = Ponape). 

Trichoglossus rubiginosus Finsch, Journ. Mus. Godeffroy, 12, 1876, pp. 17, 18 
(Ponape); idem, Proc. Zool. Soc. London, 1877 (1878), p. 778 (Ponape); idem, Journ. 
f. Ornith., 1880, p. 284 (Ponape); Schmeltz and Krause, Ethnogr. Abth. Mus. Godef- 
froy, 1881, p. 281 (Ponape); Finsch, Ibis, 1881, pp. 110, 111, 114 (Ponape); idem, 
Mitth. Ornith. Ver. Wien, 1884, p. 49 (Ponape); Hartert, Kat. Vogelsamml. Senckenb., 
1891, p. 161 (Puypinet); Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6 
1890-1891 (1891), p. 8 (Ponape); Peters, Check-list Birds World, 3, 1937, p. 151 
(Ponape); Mayr, Proc. Sixth Pac. Sci. Congr., 4, 1941, p. 204 (Ponape); Hand-list 
Japanese Birds, 3d ed., 1942, p. 201 (Ponape); Mayr, Birds Southwest Pacific, 1945, 
p. 291 (Ponape). 

Eos rubiginosa Salvadori, Ornith. Papua.sia, 1, 1880, p. 267 (Puynipet); idem. Cat. 
Birds British Mus., 20, 1891, p. 29 (Ponape); Christian, The Caroline Islands, 1899, 
p. 357 (Ponape); Finsch, Notes Lcyden Mus., 22, 1900, p. 142 (Ponape): Dubois, 
Syn. Avium, 1902, p. 29 (Puinipet); Uchida, Annot. Zool. Japon., 9, 1918, pp. 484, 
493 (Ponape) ; Wetmore, in Town^^end and Wetmore, Bull. Mus. Comp. Zool., 63, 
1919, p. 192 (Ponape). 

Chalcopsittacus rubiginosus Finsch, Sammlung wissensch. Vortrage, 14th Ser., 1900, 
p. 639 (Ponape). 

Oenopsittacus rubiginos^^s Reichenow, Die Vogel, 1, 1913, p. 443 (Karolinen =: 
Ponape); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 58 (Ponape); Mathews, 
Syst. Avium Australasianarum, 1, 1927, p. 295 (Ponape); Hand-list Japanese Birds, 
rev., 1932, p. 181 (Ponape). 

Eos rubiginosus Takastukasa and Kuroda, Tori, 1, 1915, p. 53 (Ponape). 

Geographic range. — Micronesia: Caroline Islands — Ponape. 

Characters. — Adult : A medium-sized, dark raspberry-red lory with head 
and nape deep purplish-red; upper back, scapulars, and upper wing-coverts 
raspberry-red, edged with blackish; lower back, rump, and upper tail-coverts 
more purplish; tail yellowish-green becoming more yellow and less green to- 
ward tip; wings black with outer webs olivaceous-green; outer edges of pri- 
maries more yellowish; lores, chin, auriculars, sides of head, and neck deep 
purplish-red, chin feathers faintly barred with raspberry and edged with 
blackish; throat, breast, abdomen, and flanks raspberry-red, feathers edged 
with blackish except on lower abdomen; under tail-coverts orange-red, under 
wing-coverts deep purple with black edges; bill of male orange, of female 
paler yellow; feet black; iris of male light yellowish-orange, of female gray- 
ish-white. 



212 



University of Kansas Publs., Mus. Nat. Hist, 



Immature: Resembles adult, but with narrow and more sharply pointed 
tail feathers. 

Measurements. — Measurements are presented in table 29. 

Table 29. Measurements of Trichoglossus ruhiginosus 



Sex 


No. 


Wing 


Tail 


Culmen 
from 
cere 


Tarsus 


Adult males 


18 
13 


147 
(143-153) 

142 
(141-146) 


105 
(100-110) 

101 
(98-104) 


20 

(19-20) 

19 
(18-19) 


16 


Adult females 


(15-17) 
16 




(15-17) 



Specimens examined. — Total number, 31 (18 males, 13 females), as follows: Caroline Is- 
lands, USNM— Ponape, 2 (Feb. 12); AMNH— Ponape, 29 (Nov.). 

Nesting. — According to Coultas (field notes) the nest is placed in the top 
of a coconut tree or in a hollow of a large forest tree. He says that one 
egg is laid, but does not record dates of nesting. Four of the birds taken 
by Coultas at Ponape in November had swollen gonads. 

Molt. — Specimens taken in November by Coultas were either in fresh 
plumage or were completing the molt when obtained. 

Parasites.— Vchida. (1918:484, 493) found the bird lice (Mallophaga), Pdtta- 
conirmus harrisoni and Eomenopon denticulatus, on the Ponape Lory. 

Remarks. — There is little written information concerning the 
habits of the Ponape lory. Mayr (1945a :291) describes the bird 
as being "very noisy" and with "habits apparently similar to T. 
haematodus." Coultas made a number of observations on this spe- 
cies; some of these unpublished notes are essentially as follows: 
Trichoglossus is common on Ponape. It is found everywhere on the 
island, preferring the coconut palms; it is noisy and quarrelsome. 
The parrot travels usually in small groups of two to six or eight birds, 
keeping up a continuous chatter all of the time. This chatter quiets 
down into a very pleasant-sounding crooning-tone after sunset. 
Trichoglossus is a continual nuisance to the hunter, inquisitive and 
easily attracted by the slightest noise, to which the bird responds with 
a frantic yapping that frightens everything within a radius of a mile. 
One sometimes finds a bird alone working quietly about among the 
low trees of the high mountain ridges. The natives' name for the 
bird, "se ridt," means "always hide out in rain." The bird stays 
under a big leaf and keeps dry during the rain. This lory is intelli- 
gent, easily tamed, and sometimes learns to repeat a few words. 



Baker: The Avifauna of Micronesia 213 

Evolutionary history of Trichoglossus rubiginosus. — The Ponape 
Lory is the only native parrot in Micronesia. It is an aberrant spe- 
cies and seemingly is of long residence on the island, as indicated by 
its differences from related forms to the southward and southwest- 
ward. The bird shows some relationships to T. ornatus (Linnaeus) 
of Celebes, but the plumage of T. rubiginosus lacks the brilliant red, 
green, and yellow of this bird. The plumage of the Ponape Lory is 
also softer in texture; this is a character exhibited also by other 
Micronesian birds, for example, Cleptornus and Colluricincla. T. 
rubiginosus and T. ornatus correspond, however, in having the feath- 
ers of the breast edged with blackish. T. rubiginosus resembles also 
T. flavovirides of Celebes and Sula in that the edges of the feathers 
of the breast are dark, no markings are present on the inner w'eb of 
the wing, and feathers of the ujiper back are edged with dark color- 
ing. T. rubiginosus may have been derived from either of these two 
species; however, it shows a close relationship also to the T. haemat- 
odus group from the Papuan region. In any case, the Ponape Lory, 
isolated in Micronesia, has not the multicolored plumage of its rela- 
tives and has, instead, a rather uniformly colored plumage. The 
presence of this parrot at only a single island in Micronesia is difficult 
to explain; perhaps at one time the bird was more widely distributed 
in Micronesia, or it may be that the population represents a single 
successful invasion to Ponape. Like Aplonis pelzelni, another en- 
demic species at Ponape, this lory may have reached the island as a 
straggler, perhaps being carried north by the prevailing winds in the 
post-nesting season. 

Cuculus canorus telephonus Heine 
Common Cuckoo 

Cuculus telephonus Heine, Joum. f. Ornith., 1863, p. 332. (Type locality, Japan.) 

Cuculus canorus Hartlaub and Finsch, Proc. Zool. Soc. London, 1872, pp. 89, 100 

(Pelew); Finsch, Joum. Mus. Godeffroy, 8, 1875, pp. 4, 12 (Palau) ; Wiglesworth, 

Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 10 (Pelew); 

Takatsukasa and Kuroda, Tori, 1, 1915, p. 63 (Pelew). 

Cuculus canorus telephonus Kuroda, in Momiyama, Birds Micronesia, 1922, p. 57 
(Pelew); Hand-list Japanese Birds, rev., 1932, p. 181 (Palau); Hand-list Japanese 
Birds, 3d ed., 1942, p. 201 (Palau); Mayr, Birds Southwest Pacific, 1945, p. 302 
(Palau). 

Geographic range. — Breeds in northeastern Asia and Japan. Winters south 
to India, Malaysia, and Melanesia. In Micronesia: Palau Islands — exact local- 
ity not given. 

Remarks. — The Common Cuckoo is a straggler on winter migra- 
tion to the Palau Islands. 



214 University of Kansas Publs., Mus. Nat. Hist. 

Cuculus saturatus horsfieldi Moore 
Oriental Cuckoo 

Cuculus horsfieldi Moore, in Moore and Horsfield, Cat. Birds Mus. Hon. East-India 
Co., 2, 1856-58 (1857), p. 703. (Type locality, Java.) 

Cuculus striatus Hartlaub and Finsch, Proc. Zool. See. London, 1872, pp. 89, 100 
(Pelew); Finsch. Joum. Mus. Godeffroy, 8, 1875, pp. 4, 12 (Palau) ; Takatsukasa and 
Kuroda, Tori, 1, 1915, p. 63 (Pelew). 

Cuculus intermedius Wiglesworth. Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 
1890-1891 (1891), p. 10 (Pelew). 

Cuculus optatus optatus Kuroda, in Momiyama, Birds Micronesia, 1922, p. 57 (Pelew); 
Hand-list Japanese Birds, rev., 1932, p. 181 (Palau). 

Cuculus saturatus horsfieldi Hand-list Japanese Birds, 3d ed., 1942, p. 201 (Babel- 
thuap, Koror) ; Mayr, Birds Southwest Pacific, 1945, p. 302 (Palau). 

Geographic range. — Breeds in eastern Asia and Japan. Winters south to 
India, Malaysia, and Melanesia. In Micronesia : Palau Islands — Babelthuap, 
Koror. 

Remarks. — The Oriental Cuckoo reaches the Palau Islands as a 
winter visitor. On November 11 an(i 25 of 1931, Coultas obtained 
four immature birds at Palau near taro swamps. The natives told 
him that the cuckoo visited the islands each year from December to 
June. On September 21 at Angaur the NAMRU2 party saw one bird 
which may have been this cuckoo. 

Eudynamis taitensis (Sparrman) 
Long-tailed New Zealand Cuckoo 

Cuculus taitevsis Sparrman, Mus. Carls., fasc, 2, 1787, pi. 32. (No type locality 
= Tahiti.) 

Eudynamis tahitiensis Griiffe, Journ. Mus. Godeffroy, 2, 1873, p. 123 (Yap). 

Eudynamis taitiensis Finsch, Journ. Mus. Godeffroy, 8, 1875, p. 49 (Palau); idem, 
Jouni. Mus. Godeffroy, 12, 1876, pp. 17, 20 (Ponape) ; idem, Proc. Zool. Soc. London, 
1877 (1878), p. 778 (Ponape); idem, Journ. f. Ornith., 1880, pp. 284, 298 (Ponape, 
Kuschai, Palaos, Marshalls) ; idem. Ibis, 1880, pp. 331, 332 (Taluit) ; idem. Ibis, 1881, 
pp. 104, 108, 113, 114 (Kushai, Uleai, Ponape); Schmeltz and Krause, Ethnogr. Abth. 
Mus. Godeffroy, 1881, pp. 281, 299, 353 (Ponape, Mortlock, Ruk) ; Christian, The 
Caroline Islands, 1899, p. 358 (Ponape). 

Urodynamis taitensis Finsch, Mitth. Omith. Ver. Wien, 1884, p. 53 (Jaluit, Ponape, 
Palau); Bogert, Amer. Mus. Novit., no. 933, 1937, p. 9 (Palau, Ruk, Kusaie, Ponape, 
Truk, Iringlove, Wozzie, Auru, Jaluit, Ratak) ; Peters, Check-list Birds World, 4, 1940, 
p. 40 (Palaus, Carolines, Marshall); Hand-list Japanese Birds, 3d ed., 1942, p. 201 
(Palau, Truk, Lukunor, Ponape, Kusaie, Jaluit, Elmore, Aurh, Wotze). 

Urodynam.is taitiensis Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 
1890-1891 (1891), p. 11 (Pelew, Ualan, Ponape, Luganor, Taluit); idem. Ibis, 1893, p. 
212 (Marshalls); Hartert, Novit. Zool., 7, 1900, p. 7 (Ruk); Finsch, Notes Leyden 
Mus., 22, 1900, p. 120 (Ponape, Palau, Kuschai, Ruk, Mortlock, Uleai, Jaluit); 
Takatsukasa and Kuroda, Tori, 1, 1915, p. 52 (Ruk); Kuroda, in Momiyama, Birds 
Micronesia, 1922, p. 58 (Pelew, Ualan, Ponape, Luganor, Ruk, Taluit); Hand-list 
Japanese Birds, rev., 1932, p. 180 (Palau, Kusaie, Ponape, Luganor, Truk, Jaluit, 
Elmore, Aurh, Wotze). 

Urdynamis taitiensis Finsch, Sammulung wissensch. Vortriige, 14th ser., 1900, p. 
659 (Palau). 

Eudynamis taitiensis Schnee, Zool. Jahrbiicher, 20, 1904, p. 389 (Marshalls) ; Mayr, 
Birds Southwest Pacific, 1945, p. 302 (Micronesia). 

Geographic range. — Breeds in New Zealand and adjacent islands. Winters 
chiefly in Polynesia, also Melanesia and Micronesia. In Micronesia: Palau 



Baker: The Avifauna of Micronesia 215 

Islands — exact locality unknown; Caroline Islands — Yap, Lukunor, Truk, Po- 
nape, Kusaie; Marshall Islands — Jaluit, Elmore, Auru, Wotze, Bikini. 

Characters. — Adult: A large, long-tailed cuckoo with upper parts dark brown; 

top of head spotted with white; wings, upper back and tail barred with rufous; 

underparts pale rufous or buffy-rufous with shafts of feathers streaked with 

brown. 

Specimens examined. — Total number, 4 (2 males, 2 females), as follows: Caroline Islands, 
AMNH— Truk, 1 (Jan. 7)— Kusaie, 2 (March); Marshall Islands, USNM— Bikini, 1 (May 1). 

Remarks. — Bogert (1937) has summarized the information known 
concerning the migration of the New Zealanci Long-tailed Cuckoo. 
Its principal winter range is in eastern and central Polynesia: Fiji, 
Samoa, Tonga, Union, Cook, Society, and Tuamotu islands. The 
bird reaches the northern extent of its range in the INIarshall and 
Caroline islands (see map in Bogert, 1937:3-4). There are no rec- 
ords for the Marianas and only one record from the Palaus (taken by 
Peters, as recorded by Finsch, 1875:49). The bird is seemingly 
much more numerous as a winter visitor in the Marshall Islands than 
in the Caroline Islands. Coultas (field notes) writes that the cuckoo 
appears at Kusaie about the first of February. Bogert (1937) re- 
marks that the cuckoo arrives at New Zealand for the breeding pe- 
riod in October or November and leaves for the northern wintering 
grounds in February or March. 

Bogert (1937:11) discusses briefly the history of migration of this 
bird. She presents as a possible reason for the migration the fact 
that the cuckoo feeds principally on caterpillars and that as a conse- 
quence it moves northward to the tropics during the winter months 
because this food is not available at the breeding grounds in the 
winter months. Perhaps this cuckoo in developing its ability to fly 
long distances over water on migration has expanded the breadth of 
its range eastward into the oceanic islands, rather than westward 
through Malaysia and Melanesia, because it has found less competi- 
tion from resident birds and from other migrants for feed and hab- 
itat. On many of the islands and atolls of the Pacific Basin, this 
species is the only land bird known. 

Otus podarginus (Hartlaub and Finsch) 
Palau Scops Owl 

Noctua podargina Hartlaub and Finsch, Proc. Zool. Soc. London, 1872, p. 90. 
(Type locality, Pelew Islands.) 

Noctua podargina Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 4, 8, pi. 1, fig. 1 and 
2 (Palau) ; Giebel, Thes. Ornith., 2, 1875, p. 720 (Pelew) ; Schmeltz and Krause, Eth- 
nogr. Abth. Mus. Godeffroy, 1881, p. 407 (Palau). 

Ninox podargina Sharpe, Cat. Birds British Mus., 2, 1875, p. 151 (Palau); Bolau, 
Mitteil. Naturhist. Mus. Hamburg, 1898, p. 51 (Palau); Kuroda, in Momiyama, Birds 
Micronesia, 1922, p. 61 (Pelew); Hand-list Japanese Birds, rev., 1932, p. 181 (Palau). 



216 University of Kansas Publs., Mus. Nat. Hist. 

Scops podargina Sharpe, Cat. Birds British Mus., 2, 1875, p. 313 (Palau) ; Nehr- 
korn. Journ. f. Ornith., 1879, p. 394 (Palau); Wiglesworth, Abhandl. und Ber. Zool. 
Dresden, no. 6, 1890-1891 (1891), p. 3 (Pelew); Matschie, Joum. f. Ornith., 1901, p. 
112 (Palau); Dubois, Syn. Avium, 2, 1904, p. 883 (Pelew). 

P[isorhina] podargina Reichenow, Die Vogel, 1913, p. 424 (Palau). 

Otus podarginus Mathews, Syst. Avium Australasianarum, 1, 1927, p. 2G8 (Palau); 
Mayr. Amer. Mus. Novit., no. 1269, 1944, p. 3 (Palau); idem. Birds Southwest Pa- 
cific, 1945, p. 291 (Palau). 

Pyrroglaux podargina Yamashina, Tori, 10, 1938, p. 1 (Pelew); Peters, Check-list 
Birds World, 4, 1940, p. 109 (Babelthuap, Koror) ; Hand-list Japanese Birds, 3d ed., 
1942, p. 202 (Palau). 

Geographic range. — Micronesia: Palau Islands — Koror, Babelthuap, Angaur. 

Characters. — Adult male: A small owl with forehead and superciHary area 
whitish tinged with buff and narrowly barred blackish-brown ; feathers at base 
of upper mandible with long, blackish shafts, crown and back rufous-brown; 
some feathers on neck narrowly barred ochraceous and black; some scapulars 
with outer webs barred dark brown and white ; rump and upper tail-coverts 
dark rufous, barred white and dark brown ; tail rufous, barred indistinctly dark 
brown, inner webs barred white and dark brown; wings sandy rufous, outer 
edges of all but first primary spotted bufTy-white ; lores rufous, shafts white ; 
indistinct eye ring rufous; ear-coverts whitish with rufous tips, chin white; 
throat white nan-owly barred with wavy dark lines and tipped with rufous; 
breast pale rufous, feathers barred with white and black; abdomen paler 
rufous; under tail-coverts often barred with black and white without rufous 
wash ; under wing-coverts white barred with dark brown ; bill and feet whitish ; 
iris brown. 

Adult female: Resembles adult male, but darker brown above with fine 
vermiculations of blackish color; underparts may be pale or dark rufous with 
slight or heavy white and brown barrings and spots. 

Immature : Resembles adult male, but upper parts darker brown ; forehead, 
crown, and back barred ochraceous and black; scapulars with white shaft 
streaks and spots of white; underparts more heavily barred. 

Measurements. — Eight males measure: wing, 155-163 (159) ; tail, 82-88 (84); 
culmen, 22.0-23.5 (23.0); tarsus, 32-35 (33); two females measure: wing, 158, 
165; tail, S3, 90; culmen, 23.5, 24.0; tarsus, 33, 35. 

Specimens examined. — Total number, 11 (9 males, 2 females), as follows: Palau Islands, 
USNM — Koror, 1 (Nov. 3); AMNH — exact locality not given, 10 (Oct., Nov., Dec). 

Remarks. — Coiiltas (field notes) found tlie Palau Scops Owl 
fairly common around villages on the island of Koror. He obtained 
specimens at night with the use of a flashlight. He writes that the 
bird moves about considerably remaining on one perch and calling 
for only approximately three minutes. The bird stays in the man- 
grove thickets in the daylight hours. Marshall (1949:207) also 
found the owl at Koror as well as at Peleliu in 1945. He observed 
33 pairs on Koror (approximately one-half of the total population) 
and four pairs on Peleliu. The NAMRU2 party did not find the 
owl in the southern Palaus in 1945. 

Yamashina (1938:1) gave the Palau Scops Owl the generic name, 



Baker: The Avifauna of Micronesia 217 

Pyrroglaux. Mayr (1944b: 3) has reviewed this treatment and pre- 
sents evidence to show that the name Pyrroglaux should not be 
recognized and that the bird correctly belongs in the genus Otus. 
He presents a detailed discussion to show its relationship to 0. spilo- 
cephalits, and that the characters possessed by 0. podargimis are no 
more different or imusual than those found in other members of this 
widespread genus. It is pointed out that the reduction of the 
feathering is probably caused by the change in habitat — from a 
colder one in Asia to a warmer, tropical one in the Palaus. The 
bird is probably derived from 0. spHocephahis of Asia and Malaysia. 

Asio flammeus flammeus (Pontoppidan) 
Short-eared Owl 

Strix Flammea Pontoppidan, Danske, Atlas, 1, 1763, p. 617, pi. 25. (T>'pe locality, 
Sweden.) 

Strix stridula Quoy and Gaimard, Voy. "Uranie," Zool., 1824, pp. 680, 696 
(Mariannes); idem, Ann. Sci. Nat. Paris, 6, 1825, p. 149 (Mariannes). 

Otxis brachyotus Hartlaub, Joum. f. Omith., 1854, p. 167 (Mariannen); Finsch 
(part), Journ. Mus. Godeffroy, 12, 1876, pp. 17, 18 (Mariannen?). 

Asio accipitrinus Wiglesworth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 
1890-1891 (1891), p. 3 (Marianne); Oustalet (part), Nouv. Arch. Mus. Hist. Nat. 
Paris, (3), 7, 1895, p. 168 (Mariannes); Hartert, Novit. Zool., 5, 1898, p. 51 (Mari- 
anne); Seale, Occ. Papers Bemice P. Bishop Mus., 1, 1901, p. 44 (Mariannes); Safford, 
Osprey, 1902, p. 68 (Marianas); idem, Ajner. Anthro., 4, 1902, p. 711 (Guam); idem. 
The Plant World, 7, 1904, p. 263 (Tinian); idem, Contr. U. S. Nat. Herb., 9, 1905, 
p. 79 (Tinian); Prowazek, Die deutschen Marianen, 1913, p. 88 (Marianen). 

Asi accipitrimus Wheeler, Report Island of Guam, 1900, p. 12 (Guam). 

Asio flammeus sandwichensis Kuroda (part), in Momiyama, Birds Micronesia, 1922, 
p. 61 (Marianne); Hand-list Japanese Birds (part), rev., 1932, p. 182 (Marianas). 

Asio flammeiLS ponapensis Hand-list Japanese Birds (part), 3d ed., 1942, p. 202 
(Pagan). 

Asio flammeus flammeus Mayr, Birds Southwest Pacific, 1945, p. 292 (Marianas). 

Geographic range. — Breeds in Europe, Asia, and North America. Winters to 
tropics. In Micronesia: Mariana Islands — Pagan, Tinian. 

Remarks. — The Short-eared Owl was taken at Tinian by Quoy 
and Gaimard (1824:680, 696) and in recent years has been recorded 
at Pagan. The committee which prepared the Hand-list of Japanese 
Birds (Hachisuka et al., 1942:202) writes that the bird taken at 
Pagan has a short wing (288) and indicates that it belongs to A. f. 
ponapensis. In the present work this bird is considered to be .4. /. 
jiammeus, a migrant from Asia; possibly, however, there is an un- 
recorded resident population of the Short-eared Owl in the northern 
Marianas, which may be closely related to A. f. ponapensis of Po- 
nape. Owls may have at one time been resident in the southern 
Marianas. At Guam, for instance, owls are well known to the native 
peoples, and there is suitable habitat for the owl in the extensive 
grassland areas of the island. Perhaps an owl was resident at Guam 
and at other islands but has been eliminated partly by the overgraz- 
ing and burning of the grassy habitats preferred by the owl. 



218 University of Kansas Publs., Mus. Nat. Hist. 

Asio flammeus ponapensis Mayr 

Short-eared Owl 

Asio flammeiis ponapensis Mayr, Amer. Mus. Novit., no. 609, 1933, p. 1. (Type 
locality, Ponape. ) 

Otus brachyotus Finsch (part), Journ. Mus. Godeffroy, 12, 1876, pp. 17, 18 (Po- 
nape); idem, Proc. Zool. Soc. London, 1877 (1878), p. 778 (Ponape); idem, Journ. f. 
Ornith., 1880, p. 283 (Ponape); idem, Mitth. Ornith. Ver. Wien, 1884, p. 47 (Ponape); 
idem., Sammlung wissensch. Vortrage, 14 ser., 1900, p. 659 (Ponape). 

Asio brachyotus Finsch, Ibis, 1881, pp. 113, 114 (Ponape). 

Asio accipitrinus Ridgway, Proc. U. S. Nat. Mus., 4, 1882, p. 367 (Strong'3 Island 
= Kusaie); Wiglesworth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890- 
1891 (1891), p. 3 (Ponape); Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 
1895, p. 169 (Ponapi). 

Asio flammeus sandwichensis Kuroda (part), in Momiyama, Birds Micronesia, 1922, 
p. 61 (Ponape); Hand-list Japanese Birds (part), rev., 1932, p. 182 (Ponape). 

Asio flmnmeus ponapensis Kelso, Oologist, 1938, p. 183 (Kusaie) ; Peters, Check-list 
Birds World, 4, 1940, p. 170 (Ponape); Hand-list Japanese Birds (part), 3d ed., 1942, 
p. 202 (Ponape); Mayr, Birds Southwest Pacific, 1945, p. 291 (Ponape). 

Geographic range. — Micronesia: Caroline Islands — Ponape, Kusaie? 

Characters. — Adult: a large, short-eared owl, dark brown above streaked with 
buff and lighter below streaked with dark brown. An adult female has upper 
parts dark brown, outer webs of feathers buffy to give a streaked appearance; 
rump pale buff, feathers edged subterminally with darker brown; scapulars like 
head and back; wing-coverts dark brown tipped and edged with splotches of 
buffy to buffy-rufous; primaries and secondaries brown with large spots of pale 
rufous; tail brown barred with whitish buff spots, webs with dark centers; fore- 
head whitish tinged with buff; region below and behind eye dark; chin pale 
with rufous tinged sides; throat and breast rufous-buff with heavy streaks of 
brown, becoming narrower on abdomen and under tail ; under wing-coverts 
buffy streaked with dark brown; axillaries buffy; feathering of tibia and tarsus 
pale buff; bill dark slate; feet grey-brown; iris yellow. 

Resembles A. j. jiammeus, but wing shorter and color darker. 

Measurements. — Mayr (1933:2) lists the following measurements for two 
adult females: wing. 295, 307; tail, 135, 139; culmen, 17, 17.5; and tarsus, 48. 51. 

Specimens examined. — Total number, 2 females, from Caroline Islands, AMNH-Ponape 
(Dec). 

Nesting. — Coultas (field notes) writes that the Short-eared Owl at Ponape 
builds its nest in the grass on the ground. He did not observe the nest but 
received reports of it from the natives. 

Remarks. — The owl at Ponape has been known since the time of 
Kubary. Coultas, visiting the island in 1930, was the first naturalist 
to record very much concerning the habits. According to him (field 
notes) the bird inhabits the open grasslands of Ponape and appar- 
ently has somewhat the same habits as other members of the species. 
He estimated tlie population in 1930 as two dozen or more. He 
found the birds extremely secretive during the daylight hours. They 
were observed flying over the patches of grassland at twilight and on 
moonlight nights. He comments that the catlike call of this owl is 
heard occasionally in the night. Richards writes {in litt.) that twice 



Baker: The Avifauna of Micronesia 219 

in late December, 1947, he saw this owl in a forested area near the 
summit of Jokaj Island (900 feet). 

Kelso (1938:138) records the Short-eared Owl from Kusaie on the 
basis of a specimen taken by Gulick, which Ridgway (1882:367) 
thought came from the West Indies. The specimen is labeled Strong's 
Island, which is an old name for Kusaie. Kelso gives the measure- 
ments of this bird as: wing, 275; tail, 141; culmen from cere, 19.5, 
and comments that the wings are shorter than those of specimens 
from Asia. The skin is in the U. S. National Museum. 

The Short-eared Owl at Ponape closely resembles A. f. fiammeus 
but is slightly smaller and darker. Apparently the owl came to 
Ponape as a straggler on migration from Asia, and becoming accli- 
mated and adapted to the grassy areas at Ponape remained as a 
resident. The occurrence of A. f. fiammeus in the Marianas on 
migration offers evidence as to how the bird originally reached 
Ponape. 

Caprimulgus indicus jotaka Temminck and Schlegel 

Jungle Nightjar 

Caprimulgus jotaka Temminck and Schlegel, in Siebold's Fauna Japonica, Aves, 1847, 
p. 37, pi. 12, 13. (Type locality, Japan.) 

Caprimvlgus indicus jotaka Hand-list Japanese Birds, rev., 1932, p. 179 (Palau); 
Hand-list Japanese Birds, 3d ed., 1942, p. 199 (Palau); Mayr, Birds Southwest Pacific, 
1945, p. 302 (Palau). 

Geographic range. — Breeds in eastern Asia and Japan. Winters south to 
tropics. In Micronesia: Palau Islands — exact locality unknown. 

Remarks. — According to the committee who prepared the Hand- 
list of Japanese Birds (Hachisuka et al, 1942:199), one female was 
obtained by Oba in the Palaus in November, 1930. The skin was 
placed in the Kuroda collection. Coultas obtained a male on De- 
cember 9, 1931, in the Palaus, which is in the American Museum 
of Natural History-. The bird is apparently an occasional migrant 
to western Micronesia. 

Caprimulgus indicus phalaena Hartlaub and Finsch 
Jungle Nightjar 

Caprimulgu^s phalaena Hartlaub and Finsch, Proc Zool. See. London, 1872, p. 91. 
(Type locality, Pelew.) 

Caprimulgus phalaena Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 4, 13, pi. 2, 
fig. 1, 2 (Palau); Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, p. 407 
(Palau); Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 
(1891), p. 17 (Pelew); Hartert, Cat. Birds British Mus., 16, 1892, p. 545 (Pelew): 
idem. Das Tierreich, no. 1, 1897, p. 51 (Palau); Bolau, Mitteil. Naturhist. Mus. 
Hamburg, 1898, p. 65 (Palau); Matschie, Journ. f. Ornith., 1901, p. 112 (Palau); 
Dubois, SjTi. A^^um 1, 1902, p. 124 (Pelew); Reichenow, Die Vogel, 2, 1914, p. 154 
(Palau); Mathews, Syst. Avium. Australasianarum, 1. 1927, p. 396 (Pelew); Hachi- 
suka, Birds Philippines, 2, 1934, p. 120 (Pelew). 



220 University of Kansas Publs., Mus. Nat. Hist. 

Caprimulgus indicus phalaena Kuroda, in Momiyama, Birds Micronesia, 1922, p. 61 
(Pelew); Hand-list Japanese Birds, rev., 1932, p. 179 (Palau) ; Peters, Check-list 
Birds World, 4, 1940, p. 204 (Palau); Hand-list Japanese Birds, 3d ed., 1942, p. 199 
(Babelthuap, Koror) ; Mayr, Birds Southwest Pacific, 1945, p. 292 (Palau). 

Geographic range. — Micronesia: Palau Islands — Babeltuap, Koror, Gara- 
kayo. 

Characters. — Adult male: "Above grayish-brown, very finely vermiculated, 
more rufous on the back, with large longitudinal streaks and a few cross mark- 
ings; scapulars partly with pale buff bands, mostly pale gray at the basal 
portion; primaries deep brown, with a white spot to the inner web of the 
first primary not extending to the shaft, second and third primary with fine 
spots to the inner web extending to the shaft and obsolete white spots to the 
outer web, fourth primary with a smaller and less pure white spot; chin and 
throat blackish brown, barred with rufous, with two white spots on the throat; 
breast brownish gray, vermiculated and spotted with brown and blackish; 
abdomen dirty ochraceous buflf barred with brown, the bars wider on the 
lower tail-coverts; retrices rufous-brown with blackish bars, outer ones with 
broad white terminal spots." (Hartert, 1892:545.) Bill basally whitish with 
black tip; feet blackish pink; iris dark brown. 

Adult female: According to Hartert (1892:545) similar to male, but with 
small, more or less obsolete, rufous-buff (not white) spots on the primaries; 
rectrices without white spots. 

Immature: Resembles adult but paler and less distinctly marked. 

C. i. -phalaena resembles C. i. jotaka, but is paler; the male is more broadly 
barred and more buffy on abdomen and under side of tail; the female has 
paler spots on wing. 

Meamrements. — Measurements of four males: wing, 161-168 (165); tail, 
118-129 (124); culmen, 22; tarsus, 14.0-15.1 (14.5); of four females: wing, 161- 
165 (163); tail, 118-127 (123); culmen, 22; tarsus, 14.5-15.6 (15.1). 

Specimens examined. — Total number, S (4 males, 4 females), as follows: Palau Islands, 
USNM— Koror, 3 (Nov. 3, 20, 29); AMNH— exact locality not given, 6 (Oct., Nov., Dec). 

Remarks. — This subspecies of the Jungle Nightjar is restricted to 
the Palau Islands and particularly to those islands possessing damp, 
shady forests and mangrove swamps. In September, 1945, two 
birds were observed at the edge of a mangrove swamp at Garakayo 
at twilight by the NAMRU2 party, but neither of them was taken. 
Coultas (jEield notes) found the nightjar in mangrove swamps. He 
writes that they remain quiet there during the daylight hours. He 
took specimens both in the evening and at dawn. He considers the 
bird as not very common. Marshall (1949:208) obtained specimens 
at Koror in 1945. 

Among the races of C. indicus, the coloration of C. i. -phalaena 
resembles most closely that of C. i. jotaka; probably C. i. phalaena 
was derived from C. i. jotaka of Asia. Apparently this bird arrived 
at the Palaus by way of the Philippines. It is found only in these 
islands of Micronesia and maybe another one of that group of spe- 



Baker: The Avifauna of Micronesia 



221 



cies which reached the Palaus without expanding their ranges far- 
ther into Micronesia. 

Collocalia inexpectata pelewensis Mayr 

Edible Nest Swiftlet 

Collocalia pelewensis Mayr, Amer. Mus. Novit., no. 820, 1935, p. 3. (Type locality, 
Palau Islands.) 

Collocalia vanicorensis Hartlaub, Proc. Zool. See. London, 1867 (1868), p. 829 
(Pelew); Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 4, 116, 118 (Pelew) ; 
idem, Proc. Zool. Soc. London, 1872, p. 89 (Pelew); Finsch, Journ. Mus. Godeffroy, 8, 
1875, pp. 4, 15 (Palau); idem (part), Journ. Mus. Godeffroy, 12, 1876, pp. 17, 24 
(Palau); idem (part), Proc. Zool. Soc. London, 1880, p. 575 (Palaos) ; idem (part). 
Ibis, 1881, p. 104 (Pelew); Tristram, Cat. Birds, 1889, p. Ill (Pelew); Wiglesworth 
(part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 18 (Pelew); 
Matschie (part), Journ. f. Omith., 1901, p. 112 (Palau). 

Collocalia vanikorensis Gray, Hand-list Birds, 1, 1869, p. 66 (Pelew); Giebel, Thes. 
Omith., 1, 1872, p. 737 (Pelew). 

Collocalia fuciphaga Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 
1895, p. 189 (Palaos); Reichenow, Die Vogel, 2, 1914, p. 161 (Palau). 

Collocalia francica Takatsukasa and Kuroda, Tori, 1915, p. 53 (Pelew). 

Collocalia fuciphaga inquieta Kuroda (part), in Momiyama, Birds Micronesia, 1922, 
p. 62 (Pelew). 

Collocalia unicolor amelis Kuroda (part), in Momiyama, Birds Micronesia, 1922, 
p. 63 (Pelew). 

Collocalia fuciphaga amelis Hand-list Japanese Birds, rev., 1932, p. 179 (Palau). 

Collocalia (,vanikorensis) pelewensis Mayr, Amer. Mus. Novit., no. 828, 1936, p. 11 
(Palau). 

Collocalia germani pelewensis Mayr, Amer. Mus. Novit., no. 915, 1937, p. 18 (Palau). 

Collocalia inexpectata pelewensis Peters, Check-list Birds World, 4, 1940, p. 224 
(Palau); Mayr, Birds Southwest Pacific, 1945, p. 292 (Palau); Baker, Smithson. Misc. 
Coll., vol. 107, no. 15, 1948, p. 63 (Garakayo, Peleliu). 

Collocalia vanikorensis pelewensis Hand-list Japanese Birds, 3d ed., 1942, p. 199 
(Babelthuap, Koror). 

Geographic range. — Micronesia: Palau Islands — Babelthuap, Koror, Gara- 
kayo, Peleliu, Angaur. 

Characters.— AduM, according to Mayr (1935:3): ''Small; tarsus naked; 
upper parts dark fuscous-green, with a brownish tone on back; crown not much 
darker than back; rump pale but no distinct light gray bar across rump as in 
C. spodiopygia ; color of the rump showing much individual variation, bases of 
feathers always being pale gray, but tips sometimes strongly glossy green ; inner 
margins of wing-feathers not particularly light; feathers of chin and throat soft, 
with fuscous bases and rather sharply defined silvery-gray edges, but no shaft- 
streaks; abdomen dull gray, slightly darker than throat, inconspicuous shaft- 
streaks on breast and abdomen, more pronounced shaft-streaks on under tail- 
coverts; longest under tail-coverts fairly glossj' green; white loral spot incon- 
spicuous." 

Measurements. — Measurements are listed in table 30. 

Table 30. Me^^surements of Collocalia inexpectata in Micronesia 



Subspecies 



C. i. pelewensis. 
C. i. bartschi. . . 



No. 



14 
13 



Wing 



111 (109-113) 
108 (105-108) 



Tail 



50 (47-51) 
54 (52-57) 



222 University of Kansas Publs., Mus. Nat. Hist. 

Specimens examined. — Total number, 20 (12 males, 8 females), as follows: Palau Islands, 
USNM— Peleliu, 1 (Sept. 13)— Garakayo, 2 (Sept. 18)— Koror, 3 (Nov. 5, 6, 7); AMNH— 
exact locality not given, 14 (Oct., Dec). 

Remarks. — The NAMRU2 party founci the swiftlet to be numer- 
ous on islands in the southern Palaus in 1945. The birds were ob- 
served flying in clearings and about the cliffs. Coultas writes (field 
notes) that they nest in caves on the smaller islands. 

Collocalia inexpectata bartschi Mearns 
Edible Nest Swiftlet 

Collocalia bartschi Mearns, Proc. U. S. Nat. Mus., 36, 1909, p. 476. (Type locality, 
Guam.) 

Cypselus inquietus Kittlitz (part), Obser. Zool., in Lutke., Voy. "Le Seniavine," 3, 
1836, p. 304 (Guahan) ; idem (part), Denkw. Reise russ. Amer. Micron, and Kamchat., 
2, 1858, p. 26 (Guahan). 

Collocalia nidifica Gray (part), Ann. Mag. Nat. Hist., (3), 17, 1866, p. 125 (Ma- 
rianne); idem (part). Hand-list Birds, 1, 1869, p. 65 (Marianne). 

Collocalia vanicorensis Finsch (part), Journ. Mus. Godeffroy, 12, 1876, p. 24 (Mari- 
anen); idem (part). Ibis, 1881, p. 105 (Guam); Oustalet, Le Nat., 1889, p. 260 
(Mariannes); Wiglesworth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890- 
1891 (1891), p. 18 (Marianne); Matschie (part), Journ. f. Ornith., 1901, p. 112 (Guam, 
Saipan). 

Collocalia fuciphaga Sclater, Proc. Zool. Soc. London, 1865, p. 616 (Marianne); 
Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, p. 187 (Mariannes); 
Hartert, Novit. Zool., 5, 1898, p. 53 (Rota, Guam, Saipan); Seale, Occ. Papers Bemice 
P. Bishop Mus., 1, 1901, p. 46 (Marianas); Safford, Osprey, 1902, p. 60 (Marianas); 
idem. The Plant World, 7, 1904, pp. 84, 263 (Guam); idem. Contr. U. S. Nat. Herb., 
9, 1905, p. 79 (Guam); Prowazek, Die deutschen Marianen, 1913, p. 102 (Marianen); 
Cox, Island of Guam, 1917, p. 21 (Guam); Bryan, Guam Rec, vol. 13, no. 2, 1936, p. 
25 (Guam). 

Collocalia fuchphaga Wheeler, Report Island of Guam, 1900, p. 13 (Guam). 

Collocalia fuciphaga fuciphaga Oberholser (part), Proc. Acad. Nat. Sci. Phila., 1906, 
p. 186 (Guam). 

Collocalia unicolor amelis Oberholser, Proc. Acad. Nat. Sci. Phila., 1906, p. 193 
(Guam); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 63 (Guam). 

Collocalia fuciphaga tachyptera Obersolser, Proc. U. S. Nat. Mus., 42, 1912, p. 20 
(Type locality, Guam); Stresemann, Verhandl. Ornith. Gesellsch. Bayern, 12, 1914, p. 
11 (Guam); Takatsukasa and Kuroda, Tori, 1, 1915, p. 63 (Marianas); Kuroda, in 
Momiyama, Birds Michnoseia, 1922, p. 62 (Guam, Saipan, Rota). 

Collocalia unicolor bartschi Kuroda, in Momiyama, Birds Micronesia, 1922, p. 63 
(Guam). 

Collocalia fuciphaga bartschi Mathews, Syst. Avium Australasianarum, 1, 1927, p. 
402 (Guam); Hand-list Japanese Birds, rev., 1932, p. 178 (Marianas). 

Collocalia vanikorensis bartschi Mayr, Amer. Mus. Novit., no. 828, 1936, p. 11 
(Marianne); Hand-list Japanese Birds, 3d ed., 1942, p. 198 (Saipan), Rota, Guam). 

Collocalia germani bartschi Mayr, Amer. Mus. Novit., no. 915, 1937, p. 18 (Mari- 
anne). 

Collocalia inexpectata bartschi Peters, Check-list Birds World, 4, 1940, p. 224 
(Marianne); Mayr, Birds Southwest Pacific, 1945, p. 292 (Marianas); Watson, The 
Raven, 17, 1946, p. 41 (Guam); Downs, Trans. Kansas Acad. Sci., 49, 1946, p. 105 
(Tinian); Stott, Auk, 64, 1947, p. 526 (Saipan); Baker, Smithson. Misc. Coll., vol. 
107, no. 15, 1948, p. 63 (Guam, Rota). 

Collocalia inexpectata Strophlet, Auk, 63, 1946, p. 538 (Guam); Baker, Condor, 
49, 1947, p. 125 (Guam). 

Geographic range. — Micronesia: Mariana Islands — Guam, Rota, Tinian, 
Saipan. 



Baker: The Avifauna of Micronesia 223 

Characters. — Resembles C. i. peleuxnsis, but with wing shorter; upper parts 
lighter; underparts more brownish and lacking dark shaft-streaks on breast 
and abdomen; feathers on lores whiter basally. 

Measurements. — Measurements are presented in table 30. 

Weights. — The present author (1948:63) lists the weights of seven adult 
males as 6.4-7.3 (6.8) ; of three adult females as 6.8-7.6 (7.1). These birds were 
taken at Guam. 

Specimens examined. — Total number, 48 (17 males, 19 females, 12 unsexed), as follows: 
Mariana Islands, USNM— Guam, 21 (Jan. 29, May 20, June 21, July 20, 29)— Rota, 1 (Oct. 
27); AMNH— Guam, 18 (Jan. 22, 29, Feb. 15, July 10, Aug. 11, 12)— Saipan, 8 (Sept. 17). 

Remarks. — The taxonomic relationships of the species and sub- 
species of the genus Collocalia are not fully known. The many 
different name combinations applied to the five kinds named from 
Micronesia are evidence of the lack of agreement among previous 
writers as to the correct systematic positions of the kinds. The 
genus is widely distributed in southeastern Asia and adjacent is- 
lands and is divisible into a number of species and subspecies. This 
diversity is apparently influenced by the restriction of the birds 
to local habitats caused, as Stresemann (1931b: 83) states, by the 
necessity of staying by their nesting areas which are in caves. 
Stresemann also points out that the birds are thus dependent on 
"narrowly limited ecological conditions." The birds are confined to 
certain areas and are, therefore, isolated from other populations. 
Most of the volcanic islands of Micronesia have numerous caves 
w^hich are suitable to the swiftlets for nesting. C. inexpectata 
evolved in the Malayan region and apparently spread to Micronesia 
via the Philippines to Palau and to the Marianas. The two sub- 
species of C. inexpectata in Micronesia resemble closely those to 
the westward but are smaller. I am following Peters (1940:224) in 
the treatment of these, and although some future reviser may re- 
arrange these species and subspecies, it appears to me that the 
Micronesian swiftlets fall into the two natural groups (C. inexpec- 
tata and C. inquieta) now recognized, even though their parent 
stocks in Malaysia, in my opinion, are inadequately known. 

At Guam and Rota, the NAMRU2 party found swiftlets concen- 
trated at cliff areas, flying about in large groups. Away from the 
cliffs fewer were seen and singles were observed in woodland open- 
ings, over fields, and in the coconut groves. On May 18, 1945, a 
colony of nesting birds was found approximately two miles east of 
Agaiia on Guam. This colony was in a coral sink-hole which was 
approximately 75 feet deep and 60 feet in diameter. The nests were 
grouped in clusters of 5 to 25 or more, on underhanging ledges, shel- 



224 University of Kansas Publs., Mus. Nat. Hist. 

tered from the light. The nests, which were fastened securely to the 
irregular ledges, were knocked down by shots from our collecting 
guns. Approximately 250 nests were found; no eggs were observed, 
the nests containing young birds. The young were in various stages 
of development; some were with little feather growth, others were 
completely feathered. Nests examined contained only one young 
each. The pile of guano below each cluster of nests was large; an 
estimate made at the time indicated that there were 10 or more tons 
in each pile. Guano deposits in large quantities were found also in 
caves at Amantes Point, Guam. 

CoIIocalia inquieta inquieta (Kittlitz) 
Carolines Swiftlet 

Cypselus inquietus Kittlitz, Obser. Zool., in Lutke, Voy. "Le Seniavine," 3, 1836, 
p. 285. (Type locality, Ualan.) 

Cypselus inquietus Kittliz (part), Denkw. Reise russ. Amer. Micron, und Kamchat., 
2, 1858, p. 26 (Ualan). 

CoIIocalia ualensis Streubel, Isis, 1848, p. 368 (no type locality = Kusaie?). 

CoIIocalia nidifica ualensis Gray, Ann. Nat. Hist., 17, 1866, p. 123 (Caroline Islands 
= Kusaie?). 

CoIIocalia vanicorensis Finsch (part), Joum. Mus. Godeffroy, 12, 1876, p. 24 
(Ualan); idem (part), Proc. Zool. Soc. London, 1880, p. 575 (Kuschai); idem (part), 
Journ. f. Omith., 1880, pp. 285, 298 (Kuschai); idem (part). Ibis, 1881, pp. 104, 108 
(Kushai); Wiglesworth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890- 
1891 (1891), p. 18 (Ualan); Matschie (part), Joum. f. Ornith., 1901, p. 112 (Ualan). 

CoIIocalia fuciphaga Hartert (part). Cat. Birds British Mus., 16, 1892, p. 498 
(Kuschai); Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, p. 190 
(Oualan). 

CoIIocalia fuciphaga fuciphaga Obersolser (part), Proc. Acad. Nat. Sci. Phila., 1906, 
p. 186 (Ualan). 

CoIIocalia fuciphaga vanikorensis Oberholser (part), Proc. U. S. Nat. Mus., 42, 1912, 
p. 20 (Kusaie). 

CoIIocalia fuciphaga inquieta Stresemann, Verhandl. Omith. Gesellsch. Bayern, 12, 
1914, pp. 9, 11 (Ualan); Kuroda (part), in Momiyama, Birds Micronesia, 1922, p. 62 
(Kusaie); Hand-list Japanese Birds (part), rev., 1932, p. 179 (Kusaie). 

CoIIocalia inquieta inquieta Mayr, Amer. Mus., Novit., no. 915, 1937, p. 11 (Kusaie); 
Peters, Check-list Birds World, 4, 1940, p. 225 (Kusaie); Mayr, Birds Southwest 
Pacific, 1945, p. 292 (Kusaie). 

CoIIocalia vanikorensis inquieta Hand-list Japanese Birds, 3d ed., 1942, p. 199 
(Kusaie). 

Geographic range. — Micronesia: Caroline Islands — Kusaie. 

Characters. — Adult: Upper parts dark (sooty-black) with a slight greenish 
gloss on head and back and a more conspicuous bluish-purple gloss on the wings 
and tail; feathers of lores white, tipped with black; underparts smoky-gray; 
feet brownish; bill black; iris dark brown. 

Measurements. — Measurements are presented in table 31. 

Specimens examined. — Total number, 42 (21 males, 20 females, 1 unsexed), as follows: 
Caroline Islands, USNM— Kusaie, 1 (Feb. 8); AMNH— Kusaie, 41 (Jan., Feb., March). 

Remarks. — Kittliz obtained this swiftlet when he visited Kusaie 
from December 8, 1827, to January 1, 1828. In 1931, Coultas found 



Baker: The Avifauna of Micronesia 225 

Table 31. Measurements of Collocalia inquieta 



Subspecies No. \ Win 




Collocalia i. inquieta 11 1 19 (116-125) 

Collocalia i. ponapensis 10 1 10 (107-114) 

Collocalia i. rukensis (1 12-1 19.5) * 

•(Mayr, 1935:3). 

the bird common at Kusaie. The name Collocalia ualensis, pub- 
lished by Streubel in Isis in 1848, p. 368, is without mention of a 
locality, but is later used by Gray to denote the swiftlet in the Caro- 
line Islands. 

Collocalia inquieta rukensis Kuroda 
Carolines Swiftlet 

Collocalia fuciphaga rukensis Kuroda, Tori, 1, 1915, pp. 58, 59, pi. 3, fig. 1. (Type 
locality, Ruk.) 

Collocalia vanicorensis Finsch (part), Proc. Zool. London, 1880, p. 575 (Ruk); 
Schtneltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, p. 353 (Ruk); Wigles- 
worth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 18 (Uap 
and Ruk); Hartert, Novit. Zool., 7, 1900, p. 11 (Ruk); Matschie, Journ. f. Ornith., 
1901, p. 112 (Yap, Ruk). 

Collocalia fuciphaga vanikorensis Oberholser (part), Proc. U. S. Nat. Mus., 42, 
1912, p. 20 (Uala = Truk). 

Collocalia fuciphaga rukensis Takatsukasa and Kuroda, Tori, 1, 1915, p. 53 (Ruk); 
Kuroda, in Momiyama, Birds Micronesia, 1922, p. 62 (Ruk, Yap); Kuroda, Ibis, 1927, 
p. 706 (Truk); Mathews, Syst. Avium Australasianaruni, 1, 1927, p. 402 (Ruk); 
Hand -list Japanese Birds, rev., 1932, p. 178 (Ruk). 

Collocalia fuciphaga inquieta Kuroda, in Momiyama, Birds Micronesia, 1922, p. 
62 (Ruk). 

Collocalia inquieta rukensis Mayr, Amer. Mus. Novit., no. 915, 1937, p. 11 (Ruk); 
Peters, Check-list Birds World, 4, 1940. p. 225 (Truk, Yap); Mayr, Birds Southwest 
Pacific, 1945, p. 292 (Yap, Truk). 

Collocalia vanikorensis rukensis Hand-list Japanese Birds, 3d ed., 1942, p. 198 
(Truk, Yap). 

Geographic range. — Micronesia: Caroline Islands — Truk, Yap. 
Characters. — Adult : Resembles C. i. inquieta but with wing shorter. 
Measurements. — Measurements are given in table 31. 
Specimen examined. — One unsexed bird from Caroline Islands, USNM — Truk (Feb. 16). 

Remarks. — Little is known concerning this swiftlet. The bird at 
Yap is referred to this race; I have not seen specimens from this 
island. McElroy reports seeing no swiftlets at Truk in December, 
1945. C. i. rukensis appears to be intermediate in size between C. i. 
inquieta and C. i. ponapensis. Richards writes {in litt.) that he 
found swiftlets common at Truk in 1948. He also noted a large 
swiftlike bird in "January or February," 1948, near the summit of 

15—8131 



226 University of Kansas Publs., Mus. Nat. Hist. 

Mount Tonachian on Moen Island. From his description, the bird 
may have been a large migratory swift, possibly Apus pacificus or 
Chaetura caudacuta, neither of which have been reported previously 
from Micronesia, 

Collocalia inquieta ponapensis Mayr 
Carolines Swiftlet 

Collocalia vanikorensis ponapensis Mayr, Amer. Mus. Novit., no. 820, 1935, p. 3. 
(Type locality, Ponape.) 

Collocalia vanicorensis Finsch (part), Journ. Mus. Godeffroy, 12, 1876, pp. 17, 23 
(Ponape); idem, Proc. Zool. Soc. London, 1877 (1878), p. 778 (Ponape); idern (part), 
Journ. f. Ornith., 1880, p. 285 (Ponape); idem. Ibis, 1881, p. 115 (Ponape); Wigles- 
worth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 18 
(Ponape); Matschie (part), Journ. f. Ornith., 1901, p. 112 (Ponape). 

Collocalia fuciphaga Hartert, Cat. Birds British Mus., 16, 1892, p. 498 (Ponape). 

Collocalia fuciphaga vanikorensis Takatsukasa and Kuroda, Tori, 1, 1915, p. 53 
(Ponape). 

Collocalia fuciphaga inquieta Kuroda (part), in Momiyama, Birds Micronesia, 1922, 
p. 62 (Ponape). 

Collocalia va7iikor£nsis ponapensis Mayr, Amer. Mus. Novit., no. 828, 1936, p. 12 
(Ponape) ; Hand-list Japanese Birds, 3d ed., 1942, p. 198 (Ponape). 

Collocalia inquieta ponapensis Mayr, Amer. Novit., no. 915, 1937, p. 11 (Ponape); 
Peters, Check-list Birds World, 4, 1940, p. 225 (Ponape); Mayr, Birds Southwest 
Pacific, 1945, p. 292 (Ponape). 

Collocalia inquieta Mayr, Proc. 6th Pac. Sci. Congr., 4, 1941, p. 204 (Ponape). 

Geographic range. — Micronesia: Caroline Islands — Ponape. 

Characters. — Adult: According to Mayr (1936:12), "Very similar to inquieta, 
but much smaller ; on the upper parts apparently somewhat less glossy, and not 
so dark, more brownish; under parts very variable, sometimes very dark (partly 
on account of greasing), sometimes quite silvery on the throat; very dark 
specimens show some greenish gloss not only on the longest under tail-coverts, 
but also on the entire under side, except on the throat ; rump of the same color 
as the back; tarsus unfeathered." 

Measurements. — Measurements are listed in table 31. 

Specimens examiited.— Total number, 37 (19 males, 18 females) from Caroline Islands, 
AMNH— Ponape (Nov., Dec). 

Nesting. — Coultas obtained young birds from nests in caves in November 
and December. 

Remarks. — I am following Mayr (1937:11) and Peters (1940: 
225) in this treatment of these Caroline swiftlets, even though the 
differences between C. inquieta and C. vanikorensis appear to be 
slight indeed. C. inquieta appears closest to the forms of C. vaniko- 
rensis in Northern Melanesia. The birds found in New Guinea and 
the Solomons are similar in size to the birds in the Carolienes, while 
those in the Moluccas, Admiralties and Lihir are larger. Color dif- 
ferences are slight with the pale color of the sides of the head and 
underparts being variable. All of these dark-rumped birds evidently 
evolved in the Melanesian area. 



Baker: The Avifauna of Micronesia 227 

Halc}'on cinnamomina cinnamomina Swainson 
Micronesian Kingfisher 

Halcyon cinnamomina Swainson, Zool. Illustr., 2, 1821, text to pi. 67. (No type 
locality = Guam.) 

Halcyon cinnamomina Hartlaub, Journ. f. Omith., 1854, p. 167 (Marianen = Guam); 
Gray, Cat. Birds Trop. Is. Pacific Ocean, 1859, p. 5 (Ladrone or Marian Islands = 
Guam); Sharpe (part), Monogr. Alced., 1868-71, pp. xxxii, 213, pi. 80 (Guam); Gray, 
Hand-list Birds, 1, 1869, p. 93 (Mariannes = Guam); Oustalet, Le Nat., 1889, p. 260 
(Mariannes =r Guam) ; Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 
1890-1891 (1891), p. 16 (Guam); Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 
1895, p. 175 (Guam); Hartert, Novit. Zool., 5, 1898, p. 53 (Guam); Matschie, Joum. 
f. Ornith., 1901, pp. 112, 113, 114 (Guam); Wharton, Ecol. Monogr., 16, 1946, p. 174 
(Guam); Strophlet, Auk, 63, 1946, p. 538 (Guam); Baker, Condor, 49, 1947, p. 125 
(Guam). 

Alcedo nificeps Dumont, Diet. Sci. Nat., 29, 1823, p. 273 (Mariannes = Guam) ; 
Pucheran, Rev. et Mag. de Zool., 1853, p. 387 (Mariannes = Guam); Hartlaub, Joum. 
f. Ornith., 1855, p. 423 (Mariannen = Guam). 

Dacela ruficeps Lesson, Traite d'Ornith., 1831, p. 247 (Mariannes = Guam). 

Halcyon cinnamomeus Kittlitz, Obser. Zool., in Lutke, Voy. "Le Seniavine," 3, 1836, 
p. 304 (Guahan). 

Dacelo cinnamomina Kittlitz, Denkw. Reise russ. Anier. Micron, und Kamchat., 2, 
1858, p. 131 (Guaham); Schlegel, Mus. Pays-Bas, 3, no. 17, 1863, p. 39; no. 39, 
1874, p. 29 (Mariannes = Guam); Giebel, Thes. Ornith., 2, 1875, p. 3 (Mariannae = 
Guam). 

Todiramphus cinnamominus Cassin, U. S. Expl. Exped. 1838-'42, 1858, pp. 220, 
225 (Ladrone or Marianna Islands = Guam). 

Sauropatis cinnamomina Cabanis, Mus. Hein., 2, 1859-'60, p. 159 (Marianen); 
Salvadori (part), Ornith. Papuasia, 1, 1880, p. 481 (Marianne =: Guam). 

Halcyon cinnam^ominus Finsch (part), Journ. Mus. Godeffroy, 12, 1876, pp. 17, 20 
(Marianen = Guam); Sharpe, Cat. Birds British Mus., 17, 1892, p. 259 (Marianne = 
Guam); Seale, Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 45 (Guam); Safford, 
Osprey, 1902, p. 69 (Guam); Dubois, Syn. Avium, 1, 1902, p. 108 (Guam); Safford, 
The Plant World, 7, 1904, p. 263 (Guam); idem, Contr. U. S. Nat. Herb., 9, 1905, 
p. 79 (Guam); Mearns, Proc. U. S. Nat. Mus., 36, 1909, p. 476 (Guam); Reichenow, 
Die Vogel, 2, 1914, p. 116 (Marianen = Guam) ; Takatsukasa and Kuroda, Tori, 1, 
1915, p. 63 (Mariannes = Guam); Cox, Islands of Guam, 1917, p. 21 (Guam); 
Thompson, Guam and its people, 1942, p. 23 (Guam). 

Halcyon rufigularis Sharpe, Cat. Birds British Mus., 17, 1892, p. 260 (No type 
locality := Guam). 

Halcyon cinnamanea Wheeler, Report Island of Guam. 1900, p. 12 (Guam). 

Halcyon cinnamonius Prowazek, Die deutschen Marianen, 1913, p. 102 (Marianen 
:= Guam). 

Souropatis cinnamominus Kuroda, in Momiyama, Birds Micronesia, 1922, p. 59 
(Guam). 

Hyposyma cinnamomina Mathews, Syst. Avium Australasianarum, 1, 1927, p. 384 
(Marianne — Guam). 

Halcyon cinnamomina cinnamomina Hand-list Japanese Birds, rev., 1932, p. 179 
(Guam); Hand-list Japanese Birds, 3d ed., 1942, p. 200 (Guam); Mayr, Birds South- 
west Pacific, 1945, p. 293 (Guam); Peters, Check-list Birds World, 5, 1945, p. 206 
(Guam) ; Watson, The Raven, 17, 1946, p. 41 (Guam) ; Baker, Smithson. Misc. Coll., 
vol. 107, no. 15, 1948, p. 63 (Guam). 

Halcyon cinnamomius Bryan, Guam, Rec, vol. 13, no. 2, 1936, p. 25 (Guam). 

Geographic range. — Micronesia: Mariana Islands — Guam. 

Characters. — Adult male: Head, neck, upper back, and entire under sur- 
face near "Sanford's brown" ; auriculars black with bluish wash ; narrow black 
line extending around nape ; orbital ring black ; lower back, lesser wing-coverts, 
and scapulars deep greenish-blue; outer webs of wing feathers and tail blue; 



228 



University of Kansas Publs., Mus. Nat. Hist. 



rump resembles tail but slightly lighter; under wing-coverts greenish-blue; feet 
dark brown; bill black, base of mandible paler; iris dark brown. 

Adult female: Resembles adult male, but chin, throat, and upper breast 
paler; rest of underparts and under wing-coverts white; a few cinnamon- 
tipped feathers on tibia and at bend of wing; back and scapulars darker olive- 
green and less blue. 

Immature: Resembles adult, but brown of crown mixed with greenish -blue; 
back and wing-coverts edged with pale cinnamon; chin and throat whitish; 
rest of underparts buffy-white in male and paler in female ; feathers on breast 
and nape with dark edgings. 

Measurements. — Measurements are listed in table 32. 

Table 32. Measurements of Halcyon cinnamomina 



Subspecies 


Number 


Wing 


Tail 


Exposed 
culraen 


Tarsus 


//. c. cinnamomina 


31 males 
25 females 


102 (96-105) 
102 (99-106) 


77 (73-83) 
79 (74-84) 


37 (35-39) 

38 (35-38) 


15 (14-17) 
15 (14-17) 


H. c. pelewensis . . . 


5 males 
4 females 


89 (88-89) 
88 (88-89) 


61 (58-64) 
64 (61-67) 


39 (38-40) 
39 (38-39) 


14 (13-14) 
14 (13-14) 


H. c. reichenbachii 


14 males 

15 females 


99 (96-101) 
100 (96-102) 


74 (72-77) 
74 (71-76) 


41 (39-43) 
41 (39-42) 


16 (16-17) 
16 (15-17) 



Weights. — The NAMRU2 party obtained the following weights: 11 adult 
males, 56-62 (59) ; 10 adult females, 58-76 (66). 

Specimens examined. — Total number, 72 (40 males, 32 females), as follows: Mariana Is- 
lands, USNM— Guam, 38 (Feb. 14, 24, March 8, May 25, 26, 30, June 2, 3, 4, 6, 13, 14, 
16, 18, 19, 28, 29, July 6, 7, 10, 18, 20, Aug. 24, 30, Nov. 19); AMNH— Guam, 34 (Jan., 
Feb., March, April, July, Aug., Sept., Nov., Dec.). 

Nesting. — In 1945, the NAMRU2 party found the kingfisher nesting in the 
months of March, April, May, and July. Nests were placed in hollows of 
trees, usually ten or more feet above the ground. On April 3, a nest was 
found in a banyan tree approximately 25 feet above the ground in a hollow 
limb. There were two entrances to the nest cavity and both the male and 
female were observed to feed the young. They did not enter the hollow 
but placed food in the protruding beaks of the young; the parents and nest- 
ling both were exceedingly noisy throughout most of the feeding period. On 
July 8, McElroy found a nest containing two white eggs, partly incubated, in 
a cavity of a felled coconut palm at Agfayan Bay. 

Molt. — Examination of specimens indicates that the time of molt is irregu- 
lar or that molting may occur at any time of the year. However, there may 
be a peak in molting in July, August and September; many of the adult birds 
taken then show evidence of molting of wing and tail. This is immediately 
following the period of greatest nesting activity. 

Food habits. — The Micronesian Kingfisher at Guam feeds on various kinds 
of animal life; lizards and insects are the principal items. Of three birds 
taken on February 14, the stomach of one contained a blue-tailed skink; one 



Baker: The Avifauna of Micronesia 229 

contained parts of insects and one contained parts of a gecko. I watched a 
kingfisher capture and swallow a skink on January 14. The bird remained 
motionless on its perch until the reptile approached within striking distance. 
Seale (1901:45) writes that the bird has a bad reputation as a chicken thief. 
He remarks, "I rather doubted his ability in this line until one day I actually 
saw him attack a brood of small chicks quite near me, and he would have 
undoubtedly secured one had not the mother hen rushed to the rescue." 

Parasites. — Wharton (1946:174) obtained the chigger (Acarina), Trombicula 
sp., from the Guam Kingfisher. 

Remarks. — In 1820, Quoy and Gaimard (1824:35) obtained five 
specimens of this kingfisher at Guam and called the bird "Martin- 
chasseur a teterouse." Kittlitz recorded the bird in ]\Iarch, 1828. 
Marche obtained a series of 57 skins at Guam in 1887 and 1888; 
these were sent to the Paris Museum. Sharpe described the female 
as a separate species in 1892. There is considerable variation in 
the coloration of adult birds, which is mostly due to fading, as 
suggested by Hartert (1898:52). Some individuals have the crown 
feathers much abraided as a result of rubbing the crown against 
the edge of the nest holes as the birds enter and leave them. 

The kingfisher is fairly common at Guam. It is primarily a bird 
of the forest, preferring particularly the marginal habitats between 
woodlands and openings. I saw only a few birds in open country; 
only rarely were birds seen sitting on the telephone lines along the 
roads. The writer (1947b: 124) found that of all the birds fre- 
quenting habitat along roadways on Guam, the kingfisher comprised 
only 1.2 percent. Thus, it can be said that it is not a bird of very 
conspicuous habits, although its noisy "rattle" may be heard in the 
day and at night. 

Halcyon cinnamomina pelewensis Wiglesworth 
Micronesian Kingfisher 

Halcyon pelewensis Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 
1890-1891 (1891), p. 15. (Type locality, Pelew Islands.) 

Halcyon reichenbachii Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 4, 
118 (Pelew); Sharpe, Cat. Birds British Mus., 17, 1892, p. 261 (Pelew). 

Halcyon cinnamomina Sharpe (part), Monogr. Alced., 1868-'71, pp. xxxii, 213, pi. 
SO (Pelew); Tristram (part). Cat. Birds, 1889, p. 92 (Pelew). 

Dacelo reichenbachii Schlegel, Mus. Pay-Bas, 3, no. 39, 1874, p. 29 (Pelew). 

Halcyon reichenbachi Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 4, 11 (Palau) ; 
Reichenow, Die Vogel, 2, 1914, p. 116 (Palau). 

Halcyon cinnamominus Finsch (part), Journ. Mus. Godeffroy, 12, 1876, pp. 17, 20 
(Palau). 

Sauropatis cinnamomina Salvador! (part), Ornith. Papuasia, 1, 1880, p. 481 (Pelew). 

Halcyon pelewensis Hartert, Novit. Zool., 5, 1898, p. 53 (Pelew); Matschie, Journ. 
f. Ornith., 1901, pp. 112, 113 (Palau); Takatsukasa and Kuroda, Tori, 1, 1915, p. 
53 (Pelew); Uchida, Annot. Zool. Japan., 9, 1918, p. 483 (Palau). 

Halcyon Reichenbachi var. pelewensis Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, 
(3), 7, 1895, p. 186 (Pelew). 



230 University of Kansas Publs., Mus. Nat. Hist. 

Halcyon cinnamominus var? pelewensis Dubois, Syn. Avium, 1, 1902, p. 108 (Pelew). 

Sauropatis reichenbachii pelewensis Kuroda, in Momiyama, Birds Micronesia, 1932, 
p. 60 (Angaur). 

Hyposyma cinnamomina pelewensis Mathews, Syst. Avium Australasianarum, 1, 
1927, p. 385 (Palau). 

Halcyon cinnamomina pelewensis Hand-list Japanese Birds, rev., 1932, p. 180 (Pa- 
lau); Hand-list Japanese Birds, 3d ed., 1942, p. 200 (Babelthuap, Koror) ; Mayr, 
Birds Southwest Pacific, 1945, p. 293 (Palau); Peters, Check-list Birds World, 5, 
1945, p. 206 (Babelthuap, Koror); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 
1948, pp. 63, 64 (Peleliu, Ngabad). 

Geographic range. — Micronesia: Palau Islands — Kayangel, Babelthuap, 
Koror, Garakayo, Ngabad, Angaur. 

Characters. — Adult: Resembles adult of H. c. cinnamomina, but smaller and 
with underparts white; auriculars with less bluish wash; outer webs of outer 
tail feathers edged with white. 

Immature: Resembles immature female of H. c. cinnamomina, but smaller 
with white underparts edged with black on throat, breast, and upper abdomen; 
outer webs of outer tail feathers edged with white. 

Measurements. — Measurements are presented in table 32. 

Specimens examined. — Total number, 17 (8 males, 8 females, 1 unsexed), as follows: Palau 
Islands, USNM— Babelthuap, 1 (Nov. 30)— Peleliu, 1 (Sept. 10)— Ngabad, 3 (Sept. 11); 
AMNH — exact locality not given, 12 (Oct., Nov., Dec). 

Food habits. — Stomachs of specimens obtained by the NAMRU2 party at 
Palau contained insects. One male had a large cicada in its stomach. Coultaa 
(field notes) writes that foods of this bird consist of grubs and ants. 

Parasites. — Uchida (1918:483) found the bird louse (Mallophaga), Docopho- 
rus alatoclypestus, on this bird at Palau. 

Remarks. — In 1945, the NAMRU2 party found this kingfisher in 
forested areas and at the edges of mangrove swamps on small islands 
near Peleliu. Only six birds were seen. The bird was located by 
listening for and determining the direction of its rasping call. After 
a search of the area of leafy foliage from where the call was coming, 
the bird would be seen sitting motionless on a near-by perch. Mc- 
Elroy of the NAMRU2 party saw a kingfisher with cinnamon under- 
parts at Bulubul Island at Ulithi Atoll on August 21, 1945. It was 
not taken. 

Halcyon cinnamomina reichenbachii (Hartlaub) 

Micronesian Kingfisher 

Todirhamphus Reichenbachii Hartlaub, Archiv f. Naturgesch., 18, 1852, p. 131. 
(Type locality, Ponape.) 

Halcyon cinnamominus Finsch (part), Journ. Mus. Godeffroy, 12, 1876, pp. 17, 19 
(Ponape); idem, Proc. Zool. Soc. London, 1877 (1878), p. 778 (Ponape); idem, Journ. 
f. Omith., 1880, p. 285 (Ponape); idem, Journ. f. Ornith., 1880, p. 285 (Ponape); 
idem, Mitth. Ornith. Ver. Wien, 1884, p. 47 (Ponape). 

Sauropatis cinnamomina Salvadori (part), Omith. Papuasia, 1, 1880, p. 481 (Ponape). 

Halcyon cinnamomina Finsch, Ibis, 1881, pp. 112, 114 (Ponape); Tristram (part), 
; Cat. Birds, 1889, p. 92 (Ponape). 

Halcyon mediocris Sharpe, Cat. Birds British Mus., 17, 1892, p. 260 (Type locality, 
Ponape); Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891). 



Baker: The Avifauna of Micronesia 231 

p. 16 (Ponape); Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, pp. 177, 
180, 181, 184, 185, 186 (Ponapi) ; Reichenow, Die Vogel, 2, 1914, p. 116 (Ponape). 

Halcyon reichenbachi Wiglesworth, Abhandl. und Bar. Zool. Mus. Dresden, no. 6, 
1890-1891 (1891), p. 15 (Ponape); Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris. (3), 
7, 1895, pp. 176, 180, 181, 182, 183, 184, 185, 186 (Ponapi); Hartert, Novit. Zool., 
5, 1898, p. 53 (Ponape); Matschie, Joum. f. Omith., 1901, pp. 112, 113 (Ponape); 
Mayr, Proc. 6th Pacific Sci. Congr., 4, 1941, p. 204 (Ponape). 

Halcyon cinnamominus var. reichenbachi Dubois, Syn. Avium, 1, 1902, p. 108 

(Ponape). 

Halcyon cinnamominus var. mediocris Dubois, Syn. Avium, 1, 1902, p. 108 (Po- 
nape). 

Halcyon reichenbachii Takatsukasa and Kuroda, Tori, 1, 1915, p. 53 (Ponape). 

Sauropatis mediocris Wetmore, in Townsend and Wetmore, Bull. Mus. Comp. Zool., 
63, 1919, p. 195 (Ponape). 

Sauropatis reichenbachii reichenbachii Kuroda, in Momiyama, Birds Micronesia, 
1922, p. 60 (Ponape). 

Hyposyma cinnamomina reichenbachii Mathews, Syst. Avium Australasianarum, 1, 
1927, p. 384 (Ponape). 

Halycyon cinnamomina reichenbachii Hand-list Japanese Birds, rev., 1932, p. 180 
(Ponape); Hand-list Japanese Birds, 3d ed., 1942, p. 200 (Ponape); Mayr, Birds 
Southwest Pacific, 1945, p. 293 (Ponape); Peters, Check-list Birds World, 5, 1945, 
p. 206 (Ponape). 

Halcyon cinnamomina reichenbachi Bequaert, Mushi, 12, 1939, p. 82 (Ponape); 
idem, Occ. Papers Bernice P. Bishop Mus., 16, 1941, p. 290 (Ponape). 

Geographic range. — Micronesia: Caroline Islands — Ponape. 

Characters. — Adult male: Resembles adult male of H. c. cinnamomina, but 
with slightly smaller wing and smaller tail; slightly longer bill; top of head 
paler cinnamon; feathers of back tipped with cinnamon and bordered by 
backish; underparts white. 

Adult female: Resembles adult male, but feathers forward of black nape 
band may be mixed white and cinnamon; back and scapulars duller and less 
olive. 

Immature: Resembles adult, but crown streaked with greenish-black; back 
and scapulars darker; wing-coverts edged with cinnamon, in male chin and 
throat creamj'', sides of throat, breast, and flanks cinnamon, and axillaries, 
under wing-coverts, abdomen, under tail-coverts paler cinnamon; in female 
chin and throat white and rest of underparts paler than in male. 

Measurements. — Measurements are presented in table 32. 

Specimens examined. — Total number, 49 (25 males, 24 females), as follows: Caroline Is- 
lands, USNM— Ponape, 1 (Feb. 12); AMNH— Ponape, 48 (Nov., Dec). 

Molt. — Most of the specimens taken by Coultas in November and Decem- 
ber are either worn or in molt. 

Parasites.— Bequaert (1939:82 and 1941:290) records a fly (Hippoboscidae), 
Ornithoica pusilla, from the Micronesian Kingfisher at Ponape. 

Remarks. — The difference in coloration between the adults and 
immatures has resulted in considerable confusion concerning the 
taxonomy of this subspecies. According to Wiglesworth (1891a:15), 
the name Halcyon reichenbachii was established by Gustav Hart- 
laub in 1852 for a kingfisher with a white abdomen in the Dresden 
Museum, which had been figured by Reichenbach (Synopsis Avium, 



232 University of Kansas Publs., Mus. Nat. Hist. 

Alcedineae, 1851) and called Todiramyhus cinnamomina. This 
specimen had been mislabeled and Hartlaub and Finsch (1868a :4), 
noting a resemblance between this bird and specimens from the 
Palau Islands, used the name H. reichenbachii for the birds from 
the Palaus. Later, when specimens from Ponape were taken, Hart- 
laub's bird was found to be identical with them; thus the name H. 
reichenbachii could be restricted to the bird at Ponape, and Wigles- 
worth supplied the new name H. pelewensis for the population at 
Palau. H. mediocris was used by Sharpe to designate the cinnamon- 
breasted birds at Ponape, because they were assumed to belong to 
a species different from the white-breasted ones. This confused 
situation was not clarified until additional collections were obtained 
by the Japanese. 

Coultas (field notes) comments on the conspicuously different 
field characters of the two color types in this bird. In 1930, he 
found the bird common and usually in marginal habitat in the 
lowlands and at the edges of mangrove swamps. 

Evolutionary history of Halcyon cinna7nomina. — The three races 
of kingfishers belonging to the species H. cinnamomina have been 
derived from H. chloris. The principal distinction between the two 
species is the presence of the cinnamon coloring in H. cinnamomina, 
although within H. chloris there are some subspecies possessing 
traces of this coloration. The link between these two species, as 
pointed out to me by Mayr, appears to be H. chloris matthias Hein- 
roth of the St. Matthias and Squally islands, which is colored like 
H. chloris except that on the head, especially on the occiput, there 
is a faint wash of color ranging from buff to ochre. This coloration 
of the head is a step toward the condition in the Micronesian popu- 
lations of H. cinnamomina. 

In H. c. pelewensis and H. c. reichenbachii, the adult birds resem- 
ble each other, although the former subspecies is slightly smaller. 
The immatures of H. c. reichenbachii, however, possess cinnamon 
coloring on the cheeks, sides of body, and breast in addition to that 
present on the crown and nape. The crown and nape are of this same 
color in the adults. In the subspecies at Guam, H. c. cinnamomina, 
the adult male has the immature type of plumage found in H. c. 
reichenbachii. The female of H. c. cinnamomina has this cinnamon 
coloring on the throat, but the breast, abdomen and under tail are 
white. The original stock from which the Micronesian birds came 
may have invaded the area via the Palau Islands, although Mayr 
(1940) is of the opinion that they reached Micronesia via Ponape 



Baker: The Avifauna of Micronesia 233 

(eastern Carolines) and spread to Guam and Palaii. He states fur- 
ther (1942b: 181, 182) that the presence of H. cinnamomina and H. 
chloris as reproductively isolated groups in the Palaus may not indi- 
cate that they are distinct species, but that they represent the over- 
lap of terminal links of the same species, which have diverged to such 
an extent as to leave these terminal links reproductively isolated. 

Halcyon chloris teraokai Kuroda 
White-collared Kingfisher 

Halcyon chloris teraokai Kuroda, Tori, 1, 1915, p. 56, pi. 3, fig. 3. (Type locality, 
Pelew.) 

Halcyon albicilla Hartlaub, Proc. Zool. See. London, 1867 (1868), p. 828 (Pelew); 
Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 4, 118 (Pelew); Gray (part). 
Hand-list Birds, 1, 1869, p. 93 (Pelew); Finsch, Joum. Mus. Godeffroy, 8, 1875, p. 49 
(Palau, Mackenzie, Matetotas) ; Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, 
(3), 7, 1895, p. 171 (Pelew). 

Halcyon chloris Hartlaub and Finsch, Proc. Zool. Soc. London, 1872, pp. 89, 93 
(Pelew); Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 4, 10 (Palau); Wiglesworth, 
Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 14 (Pelew); Mayr, 
Amer. Mus. Novit., no. 469, 1931, p. 3 (Pelew). 

Dacelo albicilla Giebel (part), Thes. Ornith., 2, 1875, p. 1 (Pelew). 

Halcyon sanctus Finsch, Journ. Mus. Godeffroy, 8, 1875, p. 50 (Palau); Sharpe, Cat. 
Birds British Mus., 17, 1892, p. 267 (Pelew). 

Dacelo albicilla Giebel (part). This. Ornith., 2, 1875, p. 1 (Pelew). 

Sauropatis chloris Salvadori, Ornith. Papuasia, 1, 1880, p. 470 (Pelew). 

Halcyon chloris teraokai Uchida, Annot. Zool. Japon., 9, 1918, p. 482 (Palau); 
Kuroda, Ibis, 1927, p. 707 (Pelew) ; Takatsukasa and Yamashina, Dobutsu. Zasshi, 43, 
1931, p. 484 (Pelew); Hand-list Japanese Birds, rev., 1932, p. 180 (Palau); Bequaert, 
Mushi, 2, 1939, p. 82 (Palau); idem, Occ. Papers Bernice P. Bishop Mus., 16, 1941, 
p. 290 (Palau); Hand-list Japanese Birds, 3d ed., 1942, p. 201 (Babelthuap, Koror, 
Angaur); Mayr, Birds Southwest Pacific, 1945, p. 293 (Palau); Peters, Check-list Birds 
World, 5, 1945, p. 209 (Babelthuap, Koror, Angaur); Baker, Smithson. Misc. Coll., 
vol. 107, no. 15, 1948, p. 64 (Peleliu, Garakayo). 

Sauropatis chloris teraokai Oberholser, Proc. U. S. Nat. Mus., 55, 1919, p. 357 
(Pelew); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 59 (Angaur); Mathews, 
Syst. Avium Australasianarum, 1, 1927, p. 381 (Palau). 

Geographic range. — Micronesia: Palau Islands — Kayangel, Babelthuap, 
Koror. Garakaj'o, Peleliu, Angaur. 

Characters. — Adult male: Dorsal surface bluish, head slightly darker, back 
and scapulars more greenish, rump lighter blue ; outer webs of feathers of wing 
and of tail dark blue, entire first primary blue, inner webs of other primaries 
black; collar and underparts white; ariculars black with bluish wash, the black 
extending around neck above white band; spot on upper lores and narrow line 
above eye white ; orbital ring and lower part of lores black ; under wing-coverts 
white; under tail black; feet black; bill black, mandible with whitish base; iris 
dark brown. 

Adult female: Resembles adult male, but crown and back more green and 
less blue; auriculars with greenish-blue wash. 

Immature: Resembles adult, but feathers of forehead edged with buff; spot 
on lores and underparts buffy margined with dusky. 

H. c. teraokai resembles closely H. c. chloris (Boddaert), but more greenish 
and less bluish, especially on tail. 



234 



University of Kansas Publs., Mus. Nat. Hist. 



Measurements. — Measurements are listed in table 33. Adult males and 
females have similar measurements and are treated together. 

Table 33. Measurements of Halcyon chloris in Micronesia 



Subspecies 


No. 


Wing 


Tail 


Exposed 
cu men 


Tarsus 


H. c. teraokai 

H. c. orii 


17 
9 

17 
3 


113 (110-116) 
111 (109-116) 
116 (109-119) 
115 (114-116) 


76 (72-81) 

80 (78-83) 

81 (78-84) 
81 (80-82) 


45 (41-52) 
44 (42-45) 

46 (42-49) 
44 (42-45) 


14 (13-16) 
16 (15-16) 


H. c. albicilla 

H. c. owstoni 


16 (14-17) 

17 (16-17) 



Specimens examined. — Total number, 53 (25 males, 28 females), as follows: Palau Islands, 
USNM— Garakayo, 3 (Sept. 20)— Peleliu, 14 (Aug. 27, 29, 30, 31, Sept. 1, 5, 6, Nov. 7); 
AMNH — exact locality not given, 36 (Oct., Nov., Dec). 

Food habits. — Unlike H. cinnamomina, H. chloris obtains much of its food 
by fishing in inland waters or in tidal flats and lagoons. It does, however, ob- 
tain terrestrial foods also. Stomachs of birds taken by the NAMRU2 party at 
Palau contained insects, fish, crab, and shrimp. One stomach contained 3 cc. 
of fragments of crab, another 2 cc. of shrimp and other Crustacea, and another 
2 cc. of grasshoppers. Marshall (1949:210) records the house mouse as a food 
of this bird. 

Parasites. — Uchida (1918:483) records the bird louse (Mallophaga), Docopho- 
rus alatoclypeatus, from this bird at at Palau. Bequaert (1939:82 and 1941: 
290) lists the fly (Hippoboscidae), Omithoica pusilla, from H. c. teraokai. 

Remarks. — The White-collared Kingfisher at Palau is a showy 
and conspicuous bird. It cannot be classed as a forest bird but seems 
to prefer openings and marginal woodlands. Its range does not over- 
lap that of the secretive and inconspicuous H. cinnamomina pelewen- 
sis, which prefers the denser forests. In 1945, the NAMRU2 party- 
found H. c. teraokai to be numerous in the cleared battle areas at 
Peleliu and Angaur. A favorite perch of this bird was the telephone 
lines, from which a number of our specimens were shot. Usually the 
bird was observed singly ; occasionally two birds were found together. 
A pair was seen in copulation on August 29. The call of this bird, a 
loud and harsh rattle, is noticeably different from the low rasping 
note of H. c. pelewensis. Coultas found H. c. teraokai to be numer- 
ous in 1931. He comments (field notes) that the bird frequents salt 
water areas, especially the mangrove swamps. He noted the bird 
fishing at the outer reef. 



Baker: The Avifauna of Micronesia 235 

Halcyon chloris orii Takatsukasa and Yamashina 
White-collared Kingfisher 

Halcyon chloris orii Takatsukasa and Yamashina, D'obutsu. Zasshi, 43, 1931, p. 484. 
(Type locality, Rota.) 

Halcyon albkillus Sharpe (part). Cat. Birds British Mus., 17, 1892, p. 249 (Ma- 
rianne =: Rota). 

Halcyon albicilla Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, 
p. 169 (Rota); Hartert (part), Novit. Zool., 5, 1898, p. 53 (Rota). 

Sauropatis albicillug Kuroda (part), in Momiyama, Birds Micronesia, 1922, p. 58 
(Rota). 

Halcyon chloris orii Hand-list Japanese Birds, rev., 1932, p. 180 (Rota); Hand-list 
Japanese Birds, 3d ed., 1942, p. 200 (Rota, Saipan as straggler); Mayr, Birds South- 
west Pacific, 1945, p. 293 (Rota); Peters, Check-list Birds World, 5, 1945, p. 210 
(Rota); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 64 (Rota). 

Geographic range. — Micronesia: Mariana Islands — Rota. 

Characters. — Adult: Resembles H. c. teraokai, but loral spot larger and more 
buffy; occiput lightly streaked with white and white line above eye; top of 
head and back more oily green and less blue, darker in female. 

Immature : Resembles adult, but underparts and loral spot buffy with dusky 
edges; feathers of forehead tipped with buff; remainder of upper parts slightly 
darker. 

Measurements. — Measurments are listed in table 33. 

Weights. — The author (1948:64) lists the weights of two adult females as 84 
and 85. 

Specimens examined. — Total number, 11 (4 males, 6 females, 1 unsexed), from Mariana 
Islands, USNM— Rota (Oct. 18, 19, 22, 26, Nov. 2). 

Molt. — The 11 specimens taken by the NAMRU2 party at Rota in October 
and November are in molt. 

Remarks. — The kingfisher at Rota was taken by Marche in June 
and July, 1888, and reported by Oustalet (1895:169). It was taken 
later by the Japanese and described by Takatsukasa and Yamashina 
as a new subspecies. Apparently, no other specimens were taken 
until the NAMRU party visited Rota in October and November, 
1945, and obtained 11 skins. The bird is conspicuous and common 
at Rota. 

The color characters of white feathers intermingled with the bluish 
coloring of the crown and the occiput and the large, whitish loral spot 
place this subspecies as intermediate between H. c. teraokai and the 
two subspecies known from the more northern Marianas. 

Halcyon chloris albicilla (Dumont) 

White-headed Kingfisher 

Alcedo albicilla Dumont, Diet. Sci. Nat., ed. Levrault, 29, 1823, p. 273. (Type 
locality, Marianne = Tinian.) 

Alcedo albicilla Pucheran, Rev. et Mag. Zool., 1853, p. 388 (Marianne = Tinian) ; 
Hartlaub, Journ. f. Ornith., 1855, p. 423 (Mariannen = Tinian) ; Cassin, U. S. Expl 
Exped. 1838-'42, 1858, p. 225 (Mariannes = Tinian). 

Todiramphus albicilla Reichenbach, Syn. Avium, Alcedineae, 1851, p. 30 (Mariannen 
= Tinian). 



236 University of Kansas Publs., Mus. Nat. Hist. 

Halcyon albicilla Hartlaub, Journ. f. Ornith., 1854, p. 167 (Mariannen = Tinian) ; 
Gray, Cat. Birds Trop. Is. Pacific Ocean, 1859, p. 5 (Ladrone or Marian Islanas = 
Tinian); Gray (part). Hand-list Birds, 1, 1869, p. 93 (Mariannes = Tinian) ; Ousta- 
let, Le Nat., 1889, p. 260, (Saypan) ; Wiglesworth (part), Abhandl. und Ber. Zool. 
Mus. Dresden, no. 6, 1890-1891 (1891), p. 14 (Marianne = Tinian) ; Oustalet (part), 
Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, p. 169 (Saypan); Hartert (part), 
Novit. Zool., 5, 1898, p. 52 (Saipan) ; Matschie, Journ. f. Ornitii., 1901, pp. 112, 113, 
114 (Saipan); Seale, Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 45 (Saipan). 

Dacelo albicilla Giebel (part), Thes. Omith., 2, 1875, p. 1 (Marianne = Tinian). 

Sauropatis albicilla Salvadori, Ornith. Papuasia, 1, 1880, p. 470 (Marianne = Tin- 
ian). 

Halcyon albicillus Sharpe (part), Cat. Birds British Mus., 17, 1892, p. 249 (Mari- 
anne = Saipan). 

Halcyon saurophagus Schnee, Zeitschr. f. Naturwisch., 82, 1912, p. 4G3 (Saipan). 

Sauropatis albicillus Kuroda (part), in Momiyama, Birds Micronesia, 1922, p. 58 
(Saipan). 

Leucalcyon albicilla albicilla Mathews (part), Syst. Avium Australasianarum, 1, 
1927, p. 376 (Saipan). 

Halcyon chloris albicilla Hand-list Japanese Birds, rev., 1932, p. 180 (Saipan, 
Tinian); Hand-list Japanese Birds, 3d ed., 1942, p. 200 (Saipan), Tinian, Yap?); 
Mayr, Birds Southwest Pacific, 1945, p. 293 (Saipan, Tinian); Peters, Cherk-hst 
Birds World, 5, 1945, p. 210 (Saipan, Tinian); Downs, Trans. Kansas Acad. Sci., 49, 
1946, p. 97 (Tinian); Stott, Auk, 64, 1947, p. 526 (Saipan). 

Geographic range. — Micronesia: Mariana Islands — Saipan, Tinian. 

Characters. — Adult: Resembles H. c. teraokai, but slightly larger; pileum 
white; white collar broad; black band on nape narrow and faint in some 
individuals; back and scapulars more oily green and less blue. 

Immature: Resembles adult, but pileum pale buff streaked with bluish- 
green; back and scapulars darker; upper wing-coverts edged with white; 
breast feathers edged with dusky black. 

Measurements. — Measurements are listed in table 33. 

Specimens examined. — Total number, 20 (12 males, 8 females, as follows: Mariana Is- 
lands, USNM— Saipan, 1 (Sept. 27)— Tinian, 4 (Oct. 18, 23, 26); AMNH— Saipan, 11 (July 
8, 9, 11, 13, 15, 17, August 5, 21, 26)— Tinian, 4 (Sept. 7, 8, 10). 

Nesting. — Hartert (1898:42) records an egg found in a hole of a tree at 
Saipan on July 31, 1895. He writes that the egg "is only slightly glossy, very 
thin, pure white, but soiled all over with deep brown spots, evidently from 
the decaying wood in the nest hole. It measures 33:25 mm." 

Molt. — Most of the birds taken in July, August, September, and October 
are in molt. 

Remarks. — Quoy and Gaimard, who visited the Marianas while 
on the expedition in the "Uranie," obtained this kingfisher at Tinian. 
Additional material was taken by Marche in 1887 at Saipan and by 
Owston's Japanese collectors in 1895. In 1932, Coultas (field notes) 
found the bird to be common on both Tinian and Saipan, especially 
in open country. At Saipan, Stott (1947:526) found the birds as 
singles or in pairs on wooded hillsides. At Tinian, Gleise (1945: 
220) estimated the population in 1945 as 150 . 

The completely white head in H. c. albicillu closely resembles that 
in H. s. saurophaga Gould of Melanesia. These two species resemble 



Baker: The Avifauna of Micronesia 237 

each other in several other respects. H. saurophaga is smaller than 
H. chloris with black or greenish blue on the anterior part of the 
ear-coverts and the color of the back, wings, and tail is more green- 
ish. The presence of both H. saurophaga and H. chloris on the 
same islands in Melanesia is an indication that the two groups are 
specifically distinct. 

Halcyon chloris owstoni Rothschild 
White-collared Kingfisher 

Halcyon owstoni Rothschild, Bull. British Ornith. Club, 15, 1904, p. 6. (Type lo- 
cality, Asuncion.) 

Halcyon albicillus Sharpe (part), Cat. Birds British Mus., 17, 1892, p. 249 (Mari- 
anne = Pagan, Agrigan). 

Halcyon albicilla Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, 
pp. 169, 170 (Pagan, Agrigan); Hartert, Novit. Zool., 5, 1898, p. 52 (Pagan, Agrigan). 

SauTopatis chloris owstoni Kuroda, in Momiyama, Birds Micronesia, 1922, p. 59 
(Asuncion). 

Leucalcyon albicilla owstoni Mathews, Syst. Avium Australasianarum, 1, 1927, p. 
376 (Asuncion). 

Halcyon chloris owstoni Takatsukasa and Yamashina, Dobutsu. Zasshi, 43, 1931, 
p. 484 (Asuncion); Hand-list Japanese Birds, rev., 1932, p. ISO (Asuncion); Hand-list 
Japanese Birds, 3d ed., 1942, p. 200 (Assongsong, Pagan, Almagan) ; Mayr, Birds 
Southwest Pacific, 1945, p. 293 (Almagan, Pagan, Agrigan, Asuncion); Peters, Check- 
list Birds World, 5, 1945, p. 209 (Asuncion, Pagan, Alamagan); Borror, Auk, 64, 1947, 
p. 417 (Agrighan). 

Geographic range. — Micronesia: Mariana Islands — Asuncion, Agrigan, 
Pagan, Almagan. 

Characters. — Adult: Resembles H. c. albicilla, but hind part of crown blue- 
green and black collar broader. 

Immature: Resembles adult, but forehead buflfy and edges of feathering on 
anterior crown, upper wing-coverts, and tips of secondaries brownish. 

Measurements. — Measurements are listed in table 33. 

Specimens examined. — Total number, 4 (2 males, 1 female, 1 unsexed), as follows: Mari- 
ana Islands, AMNH — Asuncion, 4 (Jan., July). 

Remarks. — Marche obtained specimens of this bird at Pagan in 
November, 1887, and at Agrigan in December, 1888, and in Febru- 
ary, 1889. Owston's Japanese collectors obtained birds at Asuncion 
in 1904, which were named as new by Rothschild. Apparently he 
used an immature specimen in preparing the diagnosis of his new 
subspecies. Borror (1947:417) visited Agrigan in 1945 and obtained 
specimens of this kingfisher. He reports that the bird is a ''common 
and abundant species and probably nests on the island." 

Evolutionary history of Halcyon chloris in Micronesia. — Halcyon 
chloris is distributed from eastern Africa at the Red Sea eastward 
through southern Asia to INIalaysia, Australia and the Pacific islands. 
Peters (1945:207-213) recognized 47 subspecies within this species. 

In its colonization of Micronesia, H. chloris apparently arrived 
first at the Palaus probably from the Philippines or the Moluccas. 



238 University of Kansas Publs., Mus. Nat. Hist. 

Whether H. cinnamomina was established at Palau prior to the 
arrival of H. chloris is unknown. H. chloris teraokai dominates most 
of the available habitats at Palau, although it has differentiated but 
little from subspecies to the west and southwest of Palau. Among 
named kinds it most closely resembles H. c. chloris (Boddaert) of 
the Moluccas, Lesser Sundas and adjacent areas in color and struc- 
ture. The species did not succeed in establishing itself in the Caro- 
lines or at Guam, but did so in the Marianas at Rota and northward. 
In comparison with other subspecies of H. chloris those in the Mari- 
anas are characterized by a slight increase in size and a replacement 
of the bluish-green coloring of the head either partly or wholly by 
white. It is noteworthy that on the islands of Tinian and Saipan, 
which occupy a geographically intermediate position in the Mariana 
chain, the bird has an almost completely white head, whereas the 
birds on islands to the north and south have only partly white heads. 
The geographic ranges of H. chloris and H. cinnamomina in Micro- 
nesia overlap only at Palau as shown by Mayr (1942b: 181). Even 
here each is restricted to a different habitat. Possibly the present 
ranges resulted from competition between each group, and both may 
have had more extensive ranges in Micronesia in the past. Another 
possibility is that the original stock of H. chloris arrived in Micro- 
nesia via the Palaus and that of H. cinnamomina via Ponape (east- 
ern Carolinas), and that the resulting successful colonizations were 
a matter of chance. If this were the case the present day ranges may 
represent the total amount of dispersal that has taken place. The 
absence of kingfishers from Kusaie, Yap, Truk and other apparently 
suitable islands favors this possibility. 

Eurystomus orientalis connectens Stresemann 
Dollar Bird 

Eurystomus orientalis connectens Stresemann, Novit. Zool., 20, 1913, p. 302. (Type 
locality, Moa.) 

Eurystomus orientalis connectens Yamashina, Tori, 10, 1940, p. 675 (Babelthuap) ; 
Hand-list Japanese Birds, 3d ed., 1942, p. 199 (Babelthuap). 

Eurystomus orientalis pacificus Mayr, Birds Southwest Pacific, 1945, p. 302 (Palau). 

Geographic range. — Celebes and adjacent islands, Lesser Sunda Islands from 
Lombock to Damar, Southeastern Islands. In Micronesia: Palau Islands — 
Babelthuap. 

Remarks. — Yamashina (1940:675) records an adult male taken 
at Babelthuap in 1938. He assigns it to E. o. connectens, comparing 
it with a series of 15 specimens of this race from Celebes, Halma- 
hera and Batchian. Mayr (1045a: 302) refers this visitor to Palau 
to E. 0. pacificus (Latham) ; this form is migratory and may fly 



Baker: The Avifauna of Micronesia 239 

north from Australia to the Melanesian area between breeding 
seasons. 

Hirundo rustica gutturalis Scopoli 

Eastern Barn Swallow 

Hirundo gutturalis Scopoli, Del. Flor. et Faune, Insubr., 2, 1786, p. 96. (Type 
locality, "in Nova Guinea," error = Panay, Philippine Islands.) 

Hirundo rmtica Graffe, Journ. Mus. Godeffroy, 2, 1873, p. 112 (Yap); Schmeltz 
and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, p. 391 (Yap). 

Hirundo rustica gutturalis Hand-list Japanese Birds, rev., 1932, p. 178 (Koror); 
Hand-list Japanese Birds, 3d ed., 1942, p. 198 (Koror); Mayr, Birds Southwest Pa- 
cific, 1945, p. 302 (Palau); Baker, Smithson. Mus. Coll., vol. 107, no. 15, 1948. p. 
65 (Guam, Angaur, Ngesebus). 

Geographic range. — Breeds in northeastern Asia, winters south to Australia 
and Pacific islands. In Micronesia: Mariana Islands — Guam, Tinian; Palau 
Islands — Babelthuap, Koror, Ngesebus, PeleHu, Angaur; Caroline Islands- 
Yap. 

Specimens examined. — Total number, 13 (9 males, 3 females, 1 unsexed), as follows: 
Mariana Islands, USNM — Tinian, 10 (Oct. 23, 25); Palau Islands, USNM— Babelthuap, 1 
(Nov. 27) — Angaur, 1 (Sept. 21); AMNH — exact locality not given, 1 (Oct. 26). 

Remarks. — This swallow is a winter migrant to western Micro- 
nesia from Asia. In the Palau Islands in September, 1945, the 
NAMRU2 party saw the swallow at Ngesebus and Angaur in small 
flocks. At Guam, the NAMRU2 party saw one bird on October 7 
and four birds flying near Agana River on October 11. Strophlet 
(1946:535) saw one bird on October 28, 1945, and six birds on 
November 16 at Guam. Marshall (1949:221) found swallows at 
Tinian, Saipan and Palau from October to February. He found 
only immature birds. 

Edolisoma tenuirostre monachum (Hartlaub and Finsch) 

Cicada Bird 

Campephaga monacha Hartlaub and Finsch, Proc. Zool. Soc. London, 1872, p. 99. 
(Type locality, Pelew Islands.) 

Volvocivora monacha Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 4, 19, pi. 3, fig. 
2-3 (Palau); idem, Journ. Mus. Godeffroy, 12, 1876, p. 28 (Palau); Schmeltz and 
Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, p. 407 (Palau). 

Lalage monacha Sharpe, Cat. Birds British Mus., 4, 1879, p. 105 (Pelew); Trist- 
ram, Cat. Birds, 1889, p. 186 (Pelew); Wiglesworth, Abhandl. und Ber. Zool. Mus. 
Dresden, no. 6, 1890-1891 (1891), p. 25 (Pelew); Bolau, Mitteil. Naturhist. Mus. 
Hamburg, 1898, p. 53 (Palau); Matschie, Journ. f. Ornith., 1901, pp. 112, 113 (Pa- 
lau); Dubois, Syn. Avium. 1, 1902, p. 303 (Pelew); Reichenow, Die Vogel, 2, 1914, 
p. 276 (Palau); Takatsukasa and Kuroda, Tori, 1, 1915, p. 54 (Pelew); Kuroda, in 
Momiyama, Birds Micronesia, 1922, p. 68 (Pelew); Hand-list Japanese Birds, rev., 
1932, p. 175 (Palau); Hand-list Japanese Birds, 3d ed., 1942, p. 194 (Babelthuap, 
Koror). 

Edolisoma monacha Mathews, Syst, Avium Australasianarum, 2, 1930, p. 541 (Pe- 
lew). 

Edolisoma tenuirostre monacha Stresemann, Omith. Monatsber., 47, 1939, p. 126 
(Palau); Mayr, Birds Southwest Pacific, 1945, p. 294 (Palau); Baker, Smithson. Misc. 
Coll., vol. 107, no. 15, 1948, p. 65 (Peleliu). 



240 



University of Kansas Publs., Mus. Nat. Hist. 



Geographic range. — Micronesia: Palau Islands — Babelthuap, Koror, Peleliu. 

Characters. — Adult male: Forehead, crown, nape, back, and underparts near 
"Tyrian blue"; auriculars darker than back; lores and chin black; throat black 
washed with blue gray; wing feathers black, margined with pale blue; black 
tail tipped with whitish, and basal part of middle two rectrices colored like 
back; under wing dark except for whitish inner margins of secondaries; bill and 
feet black; iris dark brown. 

Adult female: Resembles adult male, but forehead and under eye pale buff; 
superciliary stripe darker buff; crown, nape, and sides of neck dark slate-blue; 
mantle brown, feathers with buffy centers; back brown washed with burnt 
brown; feathers of rump and upper tail-coverts with terminal black bar edged 
with buff; wing and tail brownish-black, primaries margined with buff, inner- 
most three secondaries and upper wing-coverts broadly edged with lighter buff, 
tail tipped with buff, more broadly so on outermost tail feathers, two outermost 
tail feathers with outer edge buff ; two central tail feathers basally dark ochre ; 
ear-coverts buff, tinged with black; chin, throat, and under wing-coverts deep 
buff; breast, abdomen, and flanks buff, feathers with subterminal blackish bar; 
under tail buff. 

Immature : Resembles adult female, but crown, nape, and sides of neck 
brown; back faintly mottled with buff; tail feathers and primary wing-coverts 
tipped with white ; younger birds may have upper parts margined with pale buff. 

Measurements. — Measurements are listed in table 34. 

Specimens examined. — Total number, 23 (13 males, 10 females), as follows: Palau Islands, 
USNM— Koror, 4 (Nov. 6, 14, 26, Dec. 5)— Peleliu, 2 (Aug. 29, 30) ; AMNH— exact locality 
not given, 17 (Oct., Nov., Dec.). 

Table 34. Melasurements of Edolisoma tenuirostre in Micronesia 



W Subspecies 


No. 


Wing 


Tail 


Exposed 
culmen 


Tarsus 


E. t. monachum 


10 
35 


98 
96-103 

109 
107-112 


80 
76-83 

86 
82-91 


21.0 
20.0-22.5 

23.0 
22.0-24.0 


23.0 


E. I. insperatum 


22.5-24.0 

24.0 
23.0-25.0 



Molt. — Molt in this bird appears to take place in tlie period from August to 
December. Most of the specimens taken in August, October, November and 
December were in molt. None was taken in other months. 

Food habits. — This bird feeds principally on insects. A female taken on 
August 29 had in its stomach about one and a half cc. of parts of grasshopper. 
Marshall (1949:212) records both animal and vegetable matter in the stomach 
of this bird. 

Remarks. — The Cicada Bird at Palau inhabits the jungles, espe- 
cially the marginal areas between the thick jungle and the more open 
woodlands. In 1945, the NAMRU2 party observed only two birds, 
both of which were obtained. These were found at Peleliu in a small 
area of undisturbed woodland at the edge of a mangrove swamp. 



Baker: The Avifauna of Micronesia 241 

Each bird was perched approximately 25 feet above the ground on 
the outer branches of a densely foliated tree. The bird is thought not 
to be so rare as our records indicate; probably its secretive habits 
conceal it from man except as he makes special search for it. Coultas 
(field notes) describes the bird as one of the true forest. He found it 
shy and retiring and possessing a very weak voice. 

It may be noted that Delacour (1946:2) does not accept the genus 
Edolisoma but places birds which are currently assigned to it in the 
genus Coracina. 

Edolisoma tenuirostre nesiotis (Hartlaub and Finsch) 

Cicada Bird 

Campephaga nesiotis Hartlaub and Finsch, Proc. Zool. Soc. London, 1872, p. 98. 
(Type locality, Uap.) 

Camvehaga nesiotis Griiffe, Journ. Mus. Godeffroy, 2, 1873, p. 123 (Yap) ; Schmeltz 
and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, p. 391 (Yap). 

Volvocivora nesiotis Finsch, Journ. Mus. Godeffroy, 12, 1876, p. 28 (Yap). 

Edoliisoma nesiotis Sharpe, Cat. Birds British Mus., 4, 1879, p. 56 (Yap); Wigles- 
worth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 25 (Uap); 
Bolau, Mitteil. Naturhist. Mus. Hamburg, 1898, p. 53 (Yap); Matschie, Journ. f. 
Ornith., 1901, p. 112 (Yap); Dubois, Syn. Avium, 1, 1902, p. 299 (Uap); Reich- 
enow, Die Vogel, 2, 1914, p. 274 (Karolinen = Yap) ; Kuroda, in Momiyama, Birds 
Micronesia, 1922, p. 68 (Mackenzie, Yap). 

Edolisoma nesiotis Mathews, Syst. Avium Australasianarum, 2, 1930, p. 542 (Mac- 
kenzie group); Hand-list Japanese Birds, rev., 1932, p. 174 (Yap); Hand-list Japa- 
nese Birds, 3d ed., 1942, p. 194 (Yap). 

Edolisoma tenuirostre nesiotis Stresemann, Ornith. Monatsber., 49, 1939, p. 126 
(Yap); Mayr, Birds Southwest Pacific, 1945, p. 294 (Yap). 

Geog-raphic range. — Micronesia: Caroline Islands — Yap. 

Characters. — Adult male: Resembles adult male of E. t. monachum. Adult 
female: Resembles adult female of E. t. monachum, but wings and upper parts 
less buffy and more rufous; eye-stripe rufous; breast barred on sides only. 

Remarks. — No specimen of the Cicada Bird from Yap has been 
examined by me. For a long time this bird was thought to be a 
species distinct from any other member of this genus, but Strese- 
mann (1939:126) arranged it as a subspecies of Edolisoma tenu- 
rostre. The type specimen is an immature, and the adult is un- 
known. The presence of rufous coloring shows a relationship with 
E. t. insperatum of Ponape, but Mayr, who has examined the type 
of E. t. nesiotis in the Hamburg Museum, and has obligingly showed 
me his notes on the bird, says that it has a greater resemblance to 
the Cicada Bird at Palau especially because of the amount of bar- 
ring on the underparts. The true status of this bird, as well as 
that of other members of the avifauna of Yap, will be incompletely 
known until such time as good collections are available from this 
island group. 

16—8131 



242 University of Kansas Publs., Mus. Nat. Hist. 

Edolisoma tenuirostre insperatum (Finsch) 

Cicada Bird 

Volvocivora inseperata Finsch, Proc. Zool. Soc. London, 1875, (1876), p. 644. 
(Type locality, Ponape.) 

Volvocivora insperata Finsch, Joum. Mus. Godeffroy, 12, 1876, pp. 17, 27 (Po- 
nape); idem, Proc. Zool. Soc. London, 1877 (1878), p. 779 (Ponape); idem, Ibis, 
1881, pp. 110, 112, 115 (Ponape); Schmeltz and Krause, Ethnogr. Abth. Mus. Godef- 
froy, 1881, p. 281 (Ponape). 

Volvozivora insperata Finsch, Journ. f. Ornith., 1880, p. 289 (Ponape). 

Lalage insperata Sharpe, Cat. Birds British Mus., 4, 1879, p. 108 (Ponape); 
Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 25 
(Ponape); Bolau, Mitteil. Naturhist. Mus. Hamburg, 1898, p. 53 (Ponape); Mat- 
schie, Journ. f. Ornith., 1901, pp. 112, 113 (Ponape); Reichenow, Die Vogel, 2, 1914, 
p. 276 (Karolinen = Ponape) ; Kuroda, in Momiyama, Birds Micronesia, 1922, p. 68 
(Ponape) ; Hand-list Japanese Birds, rev., 1932, p. 174 (Ponape) ; Hand-list Japanese 
Birds, 3d ed., 1942, p. 194 (Ponape). 

Lisomada insperata Mathews, Novit. Zool., 24, 1928, p. 372 (new generic name) ; 
idem, Syst. Avium Australasianarum, 2, 1930, p. 545 (Ponape). 

Edolisoma tenuirostre insperata Stresemann, Ornith. Monatsber., 47, 1939, p. 126 
(Ponape); Mayr, Birds Southwest Pacific, 1945, p. 294 (Ponape). 

Edolisoma tenuirostre Mayr, Proc. 6th Pacific Sci. Congr., 4, 1941, p. 204 (Ponape). 

Geographic range. — Micronesia: Caroline Islands — Ponape. 

Characters. — Adult male: Resembles adult male of E. t. monachum, but 
larger; upper parts more grayish-blue; wings with outer edges bluish-gray and 
inner webbings grayish-white; central tail feathers with subterminal, roundish, 
black spots; two outermost tail feathers black tipped with broad, pale bluish- 
gray coloring; lores more bluish-gray and less black; ear-coverts pale bluish- 
gray; chin, throat, breast, abdomen, flanks, under wing, and under tail-coverts 
grayish-blue; bill and feet black; iris dark brown. 

Adult female: Resembles adult female of E. t. monachum, but larger; fore- 
head slate-gray; crown brownish-gray, browner on nape; back chocolate-brown; 
rump rufous; upper tail-coverts more cinnamon; wing and tail brownish-black, 
outer margins or primaries edged with buff; outer margins of secondaries and 
upper wing-coverts except primary wing-coverts edged with rufous ; central tail 
feathers like back but tipped with buff, other tail feathers more broadly tipped 
with buff; lores grayish-black; malar stripe to auriculars darker and more 
brownish-black with lighter shafts; underparts rufous, under wing paler and 
more buffy. 

Immature: Resembles adult female, but forehead grayish tinged with ochre; 
crown and neck brown becoming slightly more reddish on back and more burnt 
reddish-brown on rump; tail edged and tipped with buff; primaries tipped with 
whitish, secondaries broadly edged with buff, primary wing-coverts tipped with 
buffy-white; lores blackish; ear-coverts rufous with lighter shafts; tail feathers 
pointed while in adult more rounded. Younger birds resemble older ones, but 
plumage except wings and tail may be spotted or barred with buff and black 
with whitish margins. 

Measurements. — Measurements are listed in table 34. 

Specimens examined. — Total number, 46 (23 males, 23 females), from Caroline Islands, 
AMNH— Ponape (Nov., Dec). 

Nesting. — Coultas (field notes) writes that the nest is cup-shaped, made of 
grasses and strands of hair fern, and placed at low elevations in small trees and 
bushes. He was told that two eggs are laid. He comments that the nesting 



Baker: The Avifauna of Micronesia 243 

season had just been completed in November and December (the time of his 
visit to Ponape), because he noted juveniles being attended and fed by the 
adults. 

Molt. — Most of the specimens taken by Coultas in November and December 
are in fresh plumage or in the final stages of molt, indicating that the molt was 
initiated possibly in September and would be completed possibly in January. 
This time of molt appears to be approximately one month later than the time 
of molt of E. t. monachum of Palau. Probably the bird at the Palau Islands 
breeds slightly earlier in the year than the subspecies on Ponape. 

Examination of the large series of birds taken by Coultas at Ponape shows 
the presence of three types of plumages. The writer has not made a thorough 
diagnosis of these plumages, but suspects that the phenomenon obtained here 
is the same as was found by Mayr (1933e) in his study of Neolalage banksiana 
(Gray), which is a related bird. Immatures of E. t. insperatum seemingly pre- 
sent two plumages, which, if Mayr's arrangement is followed, may be inter- 
preted as a more primitive or "retarded" t3'pe in one case, with less striking 
plumage, barred with black and buff, and a more advanced or "progressive" 
type in the other case, with plumage of the latter resembling more the adult 
type, especially the adult female. It was not ascertained whether any of these 
specimens represented adult birds in "retarded" plumage. 

Remarks. — The Cicada Bird at Ponape resembles in habits its 
related subspecies at Palau. Coultas (field notes) writes that it is 
a forest bird, with retiring habits. He observed the birds in small 
groups, and describes their musical call notes as "to-to-wee, to-to- 
wee" repeated several times. 

Evolutionary history of Edolisoma tenuirostre in Micronesia. — 
Mayr (in Stresemann, 1939:126) first pointed out the close relation- 
ship between the cicada birds of Micronesia and Edolisoma tenui- 
rostre of the Solomon Islands. Up to that time the Micronesian 
birds were considered to belong to the genus Lalage. The cicada 
birds probably invaded Micronesia along two independent routes 
from a dispersal center in the Papuan area. The form at Palau, 
E. t. monachum, resembles closely several of the subspecies to the 
south and southwest, particularly those in the New Guinea area. 
Aside from the smaller size of the Palau form there are differences 
in coloration between this bird and those of Melanesia. In the 
adult female and the juvenile there are differencs in the amount 
of barring on the underparts and in the shade of color on the upper 
parts. In the adult male there are differences in the marginal color- 
ing of the primaries and secondaries. E. t. nesiotis may have ar- 
rived at Yap from Palau. Little is known concerning the taxonomic 
position of this bird. On the basis of the information available, it 
appears closer to the Palau bird than the Ponape bird in color; 
however, in size it probably more closely approaches the latter sub- 
species. 



244 University of Kansas Publs., Mus. Nat. Hist. 

The Ponape Cicada Bird, E. t. insperatum, appears to represent 
a colonization distinct from that which established the populations 
at Yap and Palau. This conclusion is based on the fact that the 
adult female of E. t. insperatum has distinctive reddish coloring 
and lacks the barring on the underparts, and that it may have been 
derived from an ancestral stock, which was reddish and not barred, 
such as E. t. remotum of the New Ireland area. The three sub- 
species in Micronesia may represent remnants of a single coloniza- 
tion, since additional material from Yap may prove that this island 
population has characters intermediate between those of the other 
subspecies of Micronesia. 

Dicrurus macrocercus harterti S. Baker 
Black Drongo 

Dicrurus ater harterti S. Baker, Novit. Zool., 26, 1918, p. 299. (Type locality, 
Formosa.) 

Dicrurus macrocercus Baker, Trans. 11th N. Amer. Wildlife Conf., 1946, p. 211 
(Rota). 

Dicrurus macrocercus harterti Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, 
p. 65 (Rota). 

Geographic range. — Formosa. In Micronesia: Mariana Islands — Rota 
(introduced). 

Specim.ens examined. — Total number, 7 (4 males, 3 females), from Mariana Islands, USNM 
—Rota (Oct. 18, 19, Nov. 2). 

Remarks. — This drongo was introduced from Formosa to Rota 
by the Japanese South Seas Development Company (Nanyo Ko- 
hatsu Kabushiki Kaisha) apparently in 1935. An illustrated book- 
let, printed by this organization and seen by members of the 
NAMRU2 party at the Rota Civil Government headquarters, 
showed pictures of the captive birds before release and indicated 
that they had been brought to Rota for the purpose of controlling 
destructive insects. Dr. Charles Vaurie has examined these birds 
and compared them with a series of drongos from Formosa in the 
collection of the American Museum of Natural History. 

The drongo appears well adapted at Rota, where it prefers culti- 
vated areas and the bombed village sites to thick woodlands. Birds 
were found in small flocks often perched in large shade trees in 
village areas. Weights of two immature males are 53 and 61 grams. 
One adult male measures: wing, 144, tail, 153, culmen, 26, tarsus, 

22. 

Corvus kubaryi Reichenow 
Marianas Crow 

Corvua Kubaryi Reichenow, Joum. f. Ornith., 1885, p. 110. (Type locality, Palau. 
error = Guam.) 

Corvus solitarius Kittlitz, Obser. Zool., in Lutke, Voy. "Le Seniavine," 3, 1836, p. 305 
(Guahan) ; Bonaparte, Comptes Rendus Acad. Sci. Paris, 37, 1853, p. 830 (Mariannes) ; 



Baker: The Avifauna of Micronesia 



245 



Kittlitz, Denkw. Reise russ. Amer. Micron, und Kamchat., 2, 1858, p. 143 (Guahan) ; 
Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, p. 216 (Guam and Rota). 

Corvus spec. Hartlaub, Journ. f. Ornith., 1854, p. 167 (Mariannen) ; Gray, Hand- 
list Birds, 2, 1870, p. 12 (Marianne). 

Corvus kubarpi Hartert, Novit. Zool., 5, 1898, p. 59 (Guam, Rota) ; Wheeler, Report 
Island of Guam, 1900, p. 13 (Guam); Matschie, Journ. f. Ornith., 1901, p. 112 (Guam); 
Seale, Occ. Papers Bemice P. Bishop Mu?., 1901, p. 55 (Guam); Safford, Osprey, 1902, 
p. 69 (Guam) ; idem. The Plant World, 7, 1904, pp. 3, 264 (Guam) ; idem, Contr. U. S. 
Nat. Herb., 9, 1905, p. 79 (Guam); Prowazek, Die deutsehen Marianen, 1913, pp. 87, 
102 (Marianen); Reichenow, Die Vogel, 2, 1914, p. 306 (Palau); Takatsukasa and 
Kuroda, Tori, 1, 1915, p. 64 (Marianne); Cox, Island of Guam, 1917, p. 21 (Guam); 
Kuroda, in Momiyama, Birds Micronesia, 1922, p. 69 (Guam, Rota); Meinertzhagen, 
Novit. Zool., 33, 1926, p. 73 (Guam); Hand-list Japanese Birds, rev., 1932, p. 169 
(Guam, Rota); Br>'an, Guam Rec, vol. 13, no. 2, 1936, p. 25 (Guam); Hand-list 
Japanese Birds, 3d ed., 1942, p. 187 (Guam, Rota) ; Mayr, Birds Southwest Pacific, 
1945, p. 298 (Guam, Rota); Watson, The Raven, 17, 1946, p. 41 (Guam); Wharton, 
Ecol. Monogr., 16, 1946, p. 174 (Guam); Strophlet, Auk, 1946, p. 540 (Guam); Baker, 
Ecol. Monogr., 16, 194G, p. 408 (Guam); idem, Condor, 49, 1947. p. 125 (Guam); 
idem, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 66 (Guam, Rota). 

Corone phillipina Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890- 
1891 (1891), p. 46 (Marianne). 

Corone kubaryi Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890- 
1891 (1891), p. 46 (Pelew, error = Guam). 

Geographic range. — Micronesia: Mariana Islands — Guam, Rota. 

Characters. — Adult: A small, black crow with a slight groenish-black gloss on 
head; back, wings, and tail with bluish-black gloss; underparts with dull, 
greenish-black gloss; bases of feathers light grayish, more nearly white on 
neck, producing a somewhat ragged appearance; nasal bristles short but ex- 
tending over nostrils and base of cidmen; bill and feet black; iris dark brown. 
Female smaller. 

Immature: Resembles adult, but feathers with less gloss; wings and tail 
browner. 

Measurements. — Measurements of Corvus kubaryi are listed in table 35. 

Weights. — The NAMRU2 party obtained weights of the Marianas Crow as 
follows: from Guam, 5 males. 231-270 (256), 11 females 205-260 (242); from 
Rota, 1 male, 256; 1 female, 260 grams. 

Table 35. Me,^surements of Corvus kubaryi 



Location 


Number 
and 
sex 


Wing 


Tail 


Full 
culmen 


Tarsus 


Guam . . . 
Rota .... 


9 males 
19 females 

3 males 


236 (229-244) 
227 (222-241) 

235 (233-236) 


165 (158-170) 
151 (143-166) 

167 (166-169) 


55 (51-57) 
50 (47-54) 

54 (53-56) 


51 (49-52) 
50 (46-54) 

50 (49-51) 



Specimens examined. — Total number, 49 (20 males, 27 females, 2 unsexed), as follows: 
Mariana Islands, USNM — Guam, 26 (May 25, 29, June 4, 7, 8, 9, 18, 28, 29, July 10, 12, 18, 
Sept. 5, 11)— Rota, 4 (Oct. 22, 25, 29); AMNH— Guam, 19 (Jan., Feb., March, Aug., Sept., 
Dec.). 

Nesting. — In the spring of 1945, the NAMRU2 party obtained records of nest- 
ing activities by crows. One nest was observed on March 8 in a banyan tree. 
Specimens collected from May to September were not in breeding condition, and 



246 University of Kansas Publs., Mus. Nat. Hist. 

it is thought that the nesting period is concentrated in the winter and spring 
months. Watson (1946:41) reports finding a young crow being fed on May 8 by 
an adult. 

Molt. — The Marianas Crow molts in the period from May to August or Sep- 
tember. Most of the birds taken by the NAMRU2 party in this period were in 
the process of molt. Skins obtained at Rota in late October also exhibit signs of 
molt. Specimens taken in December, January and February are in fresh or 
slightly worn plumage. The crow presents an exceedingly shabby appearance in 
molt, because the grayish and whitish basal parts of the feathers are exposed. 

Food habits. — The crow is an omnivorous feeder. Stomachs examined con- 
tained both plant and animal food. Both Seale (1901 :55) and Safford (1905:79) 
comment on the damage which the crow does to the com crop at Guam. Seale 
remarks that the crow has a reputation for plundering nests of other birds. The 
NAMRU2 party saw crows being chased by starlings on several occasions. 

Parasites. — Wharton (1946:174) obtained the chigger (Acarina), Trombicula 
sp., from the crow at Guam. 

Remarks. — The Marianas Crow is confined to the forested areas 
and to the coconut plantations at Guam. The birds were seen as sin- 
gles or in small flocks, often along the roadways. In a count of the 
number of birds seen along the roadways of Guam, the author (1947: 
124) found crows to constitute 2.4 per cent of the total population of 
birds counted and observed the crow on 21.6 per cent of the 125 road- 
way counts made. Coultas (field notes) noted the birds at the north- 
ern part of Guam. The NAMRU2 party found the birds distributed 
in most parts of the island but usually they were infrequent near 
areas where large numbers of service personnel were stationed. The 
birds were often noisy when flying in small flocks or in pairs ; Seale 
(1901:55) also notes this. When observed in jungle areas, the birds 
were generally quiet, feeding and perching in dense foliage. At Rota, 
the NAMRU2 party found the bird to be fairly numerous and with 
habits resembling those of the crow at Guam. No differences in 
color or structure could be found between the specimens of crows 
obtained at the two islands. 

Kittlitz (1836:305) was the first person to write an account of 
the crow at Guam. He called it Corvus solitarius and remarked 
that he later found the same species in the Philippines. Wiglesworth 
(1891a :46) also considered the crow at Guam to resemble one 
found in the Philippines and called it Corone phillipina. Later 
Reichenow named the bird Corvus kubaryi with the type locality 
as the Palau Islands. This locality proved to be erroneous and the 
bird was judged to be from Guam by Hartert (1898:59), who did 
not use the name C. solitarius because it was a nomen nudum, and 
recognized C. kubaryi as the correct name. 



Baker: The Avifauna of Micronesia 247 

Evolutionary history of Corvus kubaryi. — Meinertzhagen (1926: 
59) writes that "Environmental influences seem to be mainly, if 
not entirely, responsible for geographic differences in the genus 
Corvus." Such may be the case in C. kubaryi, which is a small, 
dull-colored crow with a relatively unmodified bill. In structure, 
it has little resemblance to other crows found in the Pacific area. 
Kittlitz was the first to note a resemblance between the bird at 
Guam and one in the Philippines. Oustalet (1896:70) wrote that 
the bird at Guam is related to crows of the Moluccas and New 
Guinea. Although not closely related to the Hawaiian Crow, C. 
tropicus, both have little gloss on their feathers, a character which 
is common to many of the insular populations of crows. Mayr 
(1943:46) is of the opinion that the Hawaiian bird was derived from 
a North American ancestor, although Brj^an (1941:187) suggests 
that it is related to C. macrorhynchus of southeastern Asia and 
remarks that the Hawaiian Crow, "has some relation to the Guam 
Crow." In looking for the ancestral stock of C. kubaryi, the several 
species of crows which occur to the north, west and south of the 
Marianas have been examined. In size and general structure, C. 
kubaryi appears to be closest to the C. enca group, and not as 
closely related to the C. macrorhynchus group. The small size, the 
shape of the culmen, the lack of pointed feathers on the breast, and 
the presence of white on the basal parts of the feathers of the nape 
are characters which C. kubaryi has in common with C. enca. Nasal 
bristles cover the frontal base of the culmen in C. kubaryi; this 
character is found also in C. enca florensis. C. kubaryi differs from 
the C. enca group by lacking the purple sheen on the upper parts; 
this sheen is conspicuous in the latter species. C. kubaryi appears 
to have little in common with C. meeki of the Solomons and C. orru 
of the Moluccas and New Guinea area. There is apparently no 
close relation between the Marianas Crow and the crow which 
reaches the Bonins. The latter crow, according to the Hand-list 
of Japanese Birds (Hachisuka et al., 1932:1), is called C. coronoides 
hondoensis Momiyama and is apparently now extinct in the Bonins. 
In summary, it may be said that C. kubaryi is an isolated and 
modified species of crow, which probably has been living at Guam 
and Rota for a considerable length of time. Whether it once lived 
on other islands in Micronesia is unknown, but it is entirely possible 
that the present population may represent a remnant of one which 
formerly had a more extensive distribution. The characters which 
show its distinctness from possible ancestral species include its 



248 University of Kansas Publs., Mus. Nat. Hist. 

small size, its slender bill, and its dull coloration. It is thought to 
have been derived from the C. enca group, C. e. pusillus of the 
Philippines or C. e. celebensis of the Celebean area. 

Luscinia calliope calliope (Pallas) 
Siberian Rubythroat 

Motacilla Calliope Pallas, Reise durch versch. Prov. russ. Reichs, 3, 1776, pp. 261, 
325, 697. (Type locality, Yenesei.) 

Luscinia calliope calliope Hand-list Japanese Birds, rev., 1932, p. 178 (Koror) ; 
Hand-list Japanese Birds, 3d ed., 1942, p. 197 (Koror); Mayr, Birds Southwest Pacific, 
1945, p. 302 (Palau). 

Geographic range. — Breeds in northeastern Asia. AVinters south to Malaysia. 
In Micronesia: Palau Islands — Koror. 

Remarks. — The Siberian Rubythroat is considered to be a casual 
winter visitor to the Palau Islands. 

Monticola solitaria philippensis (Muller) 
Chinese Blue Rock Thrush 

Turdus philippensis Miiller, Natursystem Supplements-und Register-Band, 1776, p. 
145. (Type locality, Philippine Islands, ex Buffon.) 

Monticola philippensis philippensis Hand-list Japanese Birds, rev., 1932, p. 177 
(Koror); Mayr, Birds Southwest Pacific, 1945, p. 302 (Palau). 

Monticola solitarius philippensis Hand-list Japanese Birds, 3d ed.. 1942, p. 197 
(Koror). 

Geographic range. — Breeds in northeastern Asia and Japan. Winters south 
to Malaysia. In Micronesia : Palau Islands — Koror. 

Remarks. — The Chinese Blue Rock Thrush is apparently an infre- 
quent winter visitor to the Palau Islands. 

Turdus obscurus obscurus Gmelin 
Dusky Thrush 

Turdus obscuras Gmelin, Syst. Nat., 1, 1789, p. 816. (Type locality. Lake Baikal.) 

Turdus obscuras Hartlaub and Finsch, Proc. Zool. Soc. London, 1872, pp. 89, 96 
(Pelew); Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 5, 22 (Palau); Kuroda, in Momi- 
yama. Birds Micronesia, 1922, p. 66 (Pelew). 

Merula obscura Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890- 
1891 (1891), p. 39 (Pelew). 

Turdus obscuras ooscuras Hand-list Japanese Birds, rev., 1932, p. 177 (Koror); 
Hand-list Japanese Birds, 3d ed., 1942, p. 197 (Koror); Mayr, Birds Southwest Pacific, 
1945, p. 302 (Palau). 

Geographic range. — Bi'eeds in northeastern Asia. Winters south to Malaj'-- 
sia. In Micronesia: Palau Islands — Koror. 

Remarks. — The Dusky Thrush is considered to be a casual winter 
visitor to the Palau Islands. It was first taken there by Captain 
Heinsohn, according to Hartlaub and Finsch (1872:96). 



Baker: The Avifauna of Micronesia 



249 



Psamathia annae Hartlaub and Finsch 
Palau Bush-warbler 

Psamathia annae Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, p. 5, pi. 2. 
(Type locality, Pelew Islands.) 

Psamathia annae Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 116, 
118 (Pelew); idem, Proc. Zool. Soc. London, 1872, pp. 89, 94 (Pelew); Finsch, Joum. 
Mus. Godeffroy, 8, 1875, pp. 5, 22 (Palau); Nehrkorn, Journ. f. Omith., 1879, pp. 399, 
404 (Palau); Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, p. 407 
(Palau); Sharpe, Cat. Birds British Mus., 7, 1883, p. 101 (Pelew); Tristram, Cat. 
Birds, 1889, p. 155 (Pelew) ; Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 
6, 1890-1891 (1891), p. 40 (Pelew); Bolau, Mitteil. Naturhist. Mus. Hamburg, 1898. 
p. 57 (Palau); Matschie, Journ. f. Ornith., 1901, p. 112 (Palau); Reichenow, Die 
Vogel, 2, 1914, p. 536 (Palau); Takatsukasa and Kuroda, Tori, 1, 1915, p. 54 (Pelew); 
Kuroda, in Momiyama, Birds Micronesia, 1922, p. 67 (Pelew); Mathews, Syst. Avium 
Australasianarum, 2, 1930, p. 629 (Pelew); Hand-list Japanese Birds, rev., 1932, p. 177 
(Palau) ; Hand-list Japanese Birds, 3d ed., 1942, p. 196 (Babelthuap, Koror, Peleliu) ; 
Delacour, Ibis, 84, 1942, p. 514 (Palau); Mayr, Birds Southwest Pacific, 1945, p. 294 
(Palau); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 197 (Peleliu, Ngabad). 

Calamodyta annae Gray, Hand-list Birds, 1, 1869, p. 208 (Pelew). 

Geographic range. — Micronesia: Palau Islands — Babelthuap, Koror, Gara- 
kayo, Peleliu, Ngabad. 

Characters. — Adult: A mcdium-sizod warbler with a rather long bill and 
tail ; upper parts near ''buff olive," slightly lighter on head ; lores olive-gray 
to olive-green; supraloral stripe and orbital ring pale yellow-buf?; auriculars 
yellow-brown; underparts lighter and more olive-yellow than back, especially 
in midsection ; chin paler ; sides, tibia and under tail-coverts darker and more 
olivaceus; wings and tail dark brown with outer edges olive; under wing- 
coverts light yellow; axillaries more whitish; upper mandible horn-colored,, 
darker at base; lower mandible yellowish, darker at base; legs and feet light 
yellowish-brown; iris grayish -brown. Adult female resembles adult male but 
is slightly smaller. Immature: Re.sembles adult but forehead and crown 
slightly lighter and more yellowish ; back and rump more brownish. 

Measurements. — Measurements are listed in table 36. 

Table 36. Measurements of Psamathia annae 



Sex 


No. 


Wing 


Tail 


Exposed 
culmen 


Tarsus 


Adult males 


7 
11 


74 

(72-77) 

69 

(65-74) 


64 

(62-68) 

58 
(55-61) 


21.0 
(19.5-22.5) 

21.0 

(19.5-22.0) 


28.5 


Adult females 


(27.0-30.0) 
26.5 




(25.0-29.0) 



SpeciTnens examined. — Total number, 23 (9 males, 14 females), as follows: Palau Islands, 
USNM— Koror, 5 (Nov. 7, 9, 11, 18, 19)— Peleliu, 4 (Aug. 29, 30, Sept. 4, Dec. 5)— Ngabad, 
1 (Sept. 11); AMNH — exact locality not given, 13 (Nov., Dec). 

Nesting. — Nehrkorn (1879:399, 404) records the egg of Psamathia from 
Palau. The NAMRU2 party obtained no evidence of nesting of this bird in 



250 University of Kansas Publs., Mus. Nat. Hist. 

August and September, 1945. In 1931, Coultas secured birds in November 
and December, which had enlarged gonads. Marshall (1949:219) records 
breeding in November and December. 

Molt. — Most of the skins taken from August to December have worn or 
molting feathers. Apparently there is a high point in the molting process 
in autumn and early winter. 

Food habits. — Stomachs obtained from birds taken by the NAMRU2 party 
in August and September contained parts of insects and small seeds. One 
stomach contained about one-half cc- of parts of insects. Coultas (field notes) 
found the bird scratching "on the ground for seeds as well as working in the 
low trees and bushes." Marshall (1949:212) records insects and snails as 
food items. 

Remarks — Psamathia has the habit of a typical bushwarbler, 
occurring in jungle undergrowth and along woodland margins. In 
1945, specimens were obtained by the NAMRU2 party in the scrub 
vegetation which was growing over the devastated battle areas of 
Peleliu. The bird was not common in this habitat, nor was it very 
numerous on the smaller offshore islands. Coultas (field notes) 
found the bird to be rather tame and frequently to live close to 
human habitation. Its call, as noted by Coultas, is a loud whistle 
that breaks off into a beautiful song. The bird is quick in its move- 
ments; one seen by the writer at Ngabad was constantly moving 
about in low, second-growth vegetation and was making a low, 
whistling call. The resemblance of Psamathia to Rukia palauensis 
is noteworthy. These two unrelated birds live together in jungle 
areas, although Psamathia is perhaps confined more to the forested 
undergrowth and is more solitary in its habits. Aside from its 
longer legs and bill, Psamathia closely resembles Rukia in shape 
and coloration. They appear to have developed along somewhat 
similar evolutionary lines with regard to structure, color and eco- 
logic requirements. 

The Palau Warbler was first discovered by Captain Tetens and 
described as belonging to a new genus by Hartlaub and Finsch 
(1868a: 5). In the original description the authors remark that, 
"The generic position of this new form is in the Calamoherpe group; 
the feet are the same as in Calamoherpe; but the beak is weaker and 
slenderer, and the wings are very different. Calamoherpe has the 
first quill quite spurious, the third is the longest, and the second and 
sixth are subequal. In Calamoherpe there are twelve tail-feathers; 
in Psamathia I can find only ten. Tatare is a very different form, 
with a scutellated tarsi, a very different structure of the plumage, a 
much more elongated beak, and a twelve-feathered tail. Tatare 
syrinx is a typical Calamoherpe. In the structure of the wing of 



Baker: The Avifauna of Micronesia 251 

Psamathia, there seems to be a great resemblance to the genus Arun- 
dinax of Blyth, a form with which it is not in my power to compare." 
The genera Calmyioherpe and Tatare are now included in Acroceph- 
alus; the describers were comparing the Palau Warbler with the 
reed-warblers of Micronesia and Polynesia. 

Sharpe (1883:93) writes that the Palau Bush-warblers are "Aber- 
rant reed warblers, and should, in my opinion, be placed in future 
classifications of the Cichlomorphae near the genera Cettia and 
Acrocephalus, from which they are separated by their larger first 
primary only. Through Megalurus and Sphenoeacus they approach 
the grass-warblers and Cisticolae especially." 

Mayr (1941b :203) cites Psamathia as an example of "restricted 
endemism" and points out that the nearest relative occurs in the 
Philippines. Delacour (1942:514), in a discussion of the bush- 
warblers of the genera Cettia, Bradypterus and related forms, says, 
"Psamathia annae, from Palau Islands, is related to Cettia, differing 
mainly in its much longer bill and legs." 

Psamathia is a specialized bush- warbler and has followed a pat- 
tern of evolution which characterizes some of the other island birds 
in that the bill and legs arc long and the wing is rather short and 
rounded. Psamathia resembles many of the bush-warblers, as well 
as the reed-warblers [Acrocephalus) ; in general, body coloring being 
paler below and darker above. It differs from Acrocephalus by hav- 
ing a longer tenth primary, smaller second and third primaries, only 
ten tail feathers, a more rounded wing, differently shaped nostrils, 
and by much softer plumage (the latter character is found also in 
Collurcincla tenebrosus and Cleptornis viarchei of Micronesia). 
Rather than being related to the reed-warblers, as was supposed by 
Hartlaub and Finsch, Psamathia seems closest to Cettia, especially 
to Cettia (Horeites) diphone seebohmi of the Philippine Islands. 
Psamathia has a longer bill than this bird, but the general appear- 
ance and structure of the feet, tail, wing, body and bill are the same. 

Acrocephalus luscinia luscinia (Quoy and Gaimard) 
Nightingale Reed-warbler 

Thryothorus luscinivs Quoy and Gaimard, Voy. "I'Astrolabe," Zool., 1, 1830, p. 202, 
pi. 5, fig. 2. (Type locality, Marian Is. = Guam.) 

Sylvia syrinx Kittlitz (part), Obser. Zool., in Lutk^, Voy. "Le Seniavine," 3, 1836, 
p. 306 (Guahan) ; idem (part), Denkw. Reise russ. Amer. Micron, und Kamchat, 2, 
1858, p. 141 (Guaham). 

Tatare luscinia Gray, Genera Birds, 3, 1849, App. 8 (Marian Is.= Guam); Hart- 
laub, Journ. f. Ornith., 1854, p. 167 (Mariannen = Guam); Gray, Cat. Birds Trop. Is. 
Pacific Ocean, 1859, p. 14 (Ladrone or Marian Is. = Guam) ; Finsch, Journ. Mus. 
Godeffroy, 12, 1876, p. 31 (Guaham); Wiglesworth, Abhandl. und Ber. Zool. Mus. 
Dresden, no. 6, 1890-1891 (1891), p. 41 (Guam); Biittikofer, Notes Leyden Mus., 14, 



252 



University of Kansas Publs., Mus. Nat. Hist. 



1892, p. 16 (Guam); Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, p. 209 
(Guam). 

Tatare luscinius Bonaparte, Consp. Avium, 1, 1850, p. 224 (Guam); idem, Comptes 
Rendus Acad. Sci. Paris, 41, 1855, p. 1111 (Mariannes = Guam) ; Gray, Hand-list 
Birds, 1, 1869, p. 194 (Ladrone = Guam). 

Hybristes [luscinia] Reichenbach, Syst. Avium, 1850, pi. 57, fig. 7 (no locality ^ 
Guam). 

Acrocephalus orientalis Pelzeln, Reise, "Novara," Vogel, 1865, p. 64 (Guaham). 

Tatares luscinitis Giebel, Thes. Ornith., 3, 1877, p. 599 (Marianae). 

Acrocephaltis mariannae Tristram, Ibis, 1883, p. 45 (Type locality, Guam). 

Tatare mariannae Sharpe, Cat. Birds British Mus., 7, 1883, p. 528 (Marianne = 
Guam); Oustalet, Le Nat., 1889, p. 260 (Mariannes = Guam). 

Acrocephalus luscinia Hartert, Novit. Zool., 5, 1898, p. 57 (Guam, Saipan) ; Seale, 
Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 53 (Guam, Saipan); Matschie, Joum. 
f. Ornith., 1901, pp. 112, 113 (Guam, Saipan); Safford, Osprey, 1902, p. 69 (Guam); 
Dubois, Syn. Avium, 1, 1902, p. 369 (Marianne); Safford, Amer. Anthro., 4, 1902, p. 
711 (Guam); idem. The Plant Worid, 7, 1904, p. 264 (Guam); idem, Contr. U. S. 
Nat. Herb., 9, 1905, pp. 30, 79 (Guam); Reichenow, Die Vogel, 2, 1914, p. 545 
(Marianen); Cox, Island of Guam, 1917, p. 21 (Guam); Kuroda, in Momiyama, Birds 
Micronesia, 1922, p. 67 (Guam, Saipan); Bryan, Guam Rec, vol. 13, no. 2, 1936, p. 
25 (Guam); Thompson, Guam and its people, 1942, p. 23 (Guam); Strophlet, Auk, 
1946, p. 539 (Guam). 

Conopoderas luscinia Mathews, Syst. Avium Australasianarum, 2, 1930, p. 594 
(Marianas); Hand-list Japanese Birds, rev., 1932, p. 177 (Marianas). 

Conopoderas luscinia hivae Yamashina, Bull. Biogeogr. Soc. Japan, 12, 1942, p. 81 
(Type locality, Saipan); Hand-list Japanese Birds, 3d ed., 1942, p. 196 (Almagan, 
Saipan). 

Conopoderas luscinia luscinia Hand-list Japanese Birds, 3d ed., 1942, p. 197 
(Guam). 

Acrocephalus luscinia luscinia Mayr (part). Birds Southwest Pacific, 1945, p. 294 
(Guam, Saipan, Almagan) ; Stott, Auk, 1947, p. 526 (Saipan) ; Baker, Sniithson. Misc. 
Coll., vol. 107, no. 15, 1948, p. 67 (Guam). 

Geographic range. — Micronesia: Mariana Islands — Guam, Saipan, Almagan. 

Character. — Adult: A rather large warbler with long, curved bill; upper 
parts near "Saccardo olive"; feathers of head grayer because of darker shafts; 
rump paler and browner; lores dark; supraloral stripe light buffy-yellow ; au- 
riculars, cheeks, and sides of neck slightly darker; chin, throat, breast, and 
abdomen pale buffy-yellow; tibia darker and more olivaceous-brown; under 
tail-coverts pale yellow-buff; wing and tail feathers brown, edged with ochra- 
ceous; under wing grayish, inner edges lighter; axillaries pinkish-white; upper 
mandible dark horn colored; lower mandible lighter yellow; feet light gray; 
iris brown. Female resembles male but is slightly smaller. 

Table 37. Measurements of Acrocephalus luscinia 



Subspecies 


No. 


Sex 


Wing 


Tail 


Exposed 
culmon 


Tarsus 


A. I. luscinia 

A . l. syrinx 


11 

1 

31 

12 


males 

female 

males 

females 


84 

(81-86) 

78 

78 
(76-80) 

75 
(74-78) 


83 

(80-86) 

73 

71 

(68-75) 

68 
(65-70) 


36.0 

(35.5-39.0) 

37.0 

26.5 
(25.0-27.0) 

25.5 
(24.0-27.0) 


30.5 
(30.0-31.0) 

28.5 

26.5 
(25.0-29.0) 

26.0 
(24.0-26.0) 



Baker: The Avifauna of Micronesia 253 

Immature: Resembles adult, but upper parts duller and more brown and 
Jess olive; underparts less yellow; wing and tail feathers lighter brown- 
Measurements. — Measurements are listed in table 37. 

Weights. — The weights of three adult males obtained at Guam by the 
NAMRU2 party are 30, 30, and 31 grams. An adult female from Guam 
weighed 27 grams. 

Specimens examined. — Total number, 12 (11 males, 1 female), as follows: Mariana Islands, 
USNM— Guam, 6 (June 2, 13, July 2, 18) — Saipan, 6 (Sept. 27, 30). 

Nesting. — Oustalet (1895:209) writes that Marche found nests at Guam in 
June, 1887. The NAMRU2 party obtained two males with enlarged gonads in 
June, 1945. 

Molt. — Specimens taken in June, July, and September are either in worn 
plumage or in molt. Birds in worn plumage become a faded straw-brown above. 
Oustalet apparently interpreted this coloring of the worn plumage as a seasonal 
coloration. 

Food habits. — Seale (1901:53) reports that four stomachs which he exam- 
ined contained insects and larvae. Marshall (1949:21) lists as food items: 
lizards, snails, spiders, and insects. 

Remarks. — The Nightingale Reed-warbler at Guam is restricted 
to cane thickets and adjacent areas in and near fresh and brackish 
water marshes. In 1945, the NAMRU2 party found the bird fairly 
numerous in some of these habitats. Seale (1901:53) writes, "This 
bird is now quite scarce on the island of Guam. It lives exclusively 
among the reedy swamps, and those swamps are now being drained 
to make room for the Chinaman's rice paddies." Mayr (1945a :295) 
also notes the rarity of the species. As a result of the late war, the 
cultivation of rice was reduced and the reed-warbler probably has 
been able to increase in some of the now fallow areas. The most 
extensive range of this bird at Guam is found in the Agaiia Swamp, 
where there is a large area consisting of thick cane. Here, and in the 
other large cane patches, the chief hazard to the bird population ap- 
pears to be fire. In dry periods, the entire habitat might be easily 
destroyed by fire. The birds are extremely shy; their melodious 
songs may be heard in the reeds, but their active movements in the 
thick cane are difficult to observe. While hunting for these birds 
along the edges of Agafia Swamp on June 2, the writer observed, or 
located the calls of, at least six or seven individuals but could only 
get within shooting range of three birds. Within the cane thickets, 
these birds feed and move about near the ground or the surface of 
the water. Rarely do they perch in a conspicuous manner in the 
upper parts of the cover. Their color patterns blend perfectly with 
the coloration of the dry cane stalks. Perhaps failure to find many 
of the birds because of their secretive habits has caused many ob- 



254 University of Kansas Publs., Mus. Nat. Hist. 

servers to assume that the bird is near extinction. Nevertheless, it is 
my contention that the bird, being restricted to these limited areas, 
has never been very abundant at Guam. The absence of natural 
enemies, especially snakes, may be one of the principal reasons why 
they have been able to survive. 

Reed-warblers were not found by the NAMRU2 party at Rota in 
1945, nor have they been reported from Tinian. Yamashina in 1942 
described the populations at Saipan and Almagan as distinct. I have 
not seen this description, but on the basis of examinations of speci- 
mens from Saipan, I can see no recognizable differences between 
these and birds from Guam. 

Acrocophalus luscinia syrinx (Kittlitz) 
Nightingale Reed-warbler 

Sylvia syrinx Kittlitz, Mem. Acad. Imp. Sci. St. Petersbourg, 2, 1835, p. 6, pi. 8. 
(Type locality, Lugunor and Ulcei = Woleai.) 

Sylvia syrinx Kittlitz (part), Obser. Zool., in Lutke, Voy. "Le Seniavine," 3, 
1836, p. 297 (Lougounor) ; idem. Denkw. Reise russ. Amer. Micron, und Kamchat., 
2, 1858, p. 92 (Ualan, Lugunor, Ulea). 

Eparnetes [syrinx] Reiclienbach, Syst. Avium, 1850, pi. 57 (no locality := Caro- 
lines) ; Bonaparte, Comptes Rendus Acad. Sci. Paris, 41, 1855, p. 1111 (Carolines). 

Tatare syrinx Hartlaub, Archiv f. Naturgesch., 18, 1852, p. 131 (Ualan, Lugunor); 
Pucheran, Voy. Pole Sud, 3, 1853, p. 92 (Hogoleu = Truk) ; Hartlaub, Journ. f. 
Ornith., 1854, pp. 164, 168 (Hogoleu); Gray, Cat. Birds Trop. Is. Pacific Ocean, 1859, 
p. 14 (Ualan); Sharpe, Cat. Birds British Mus., 7, 1883, p. 527 (Carolines); Wigles- 
■worth, Abhandl. und Bar. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 41 (Ruk, 
Ualan, Luganor. Uleei, Nukuor, Ponape) ; Oustalet (part), Nouv. Arch. Mus. Hist. 
Nat. Paris, (3), 7, 1895, p. 210 (Ruk, Ponapi, Mortlock, Kusaie, Uleei, Nukuor). 

Acrocephalus orientalis Pelzeln, Reise "Novara," Vogel, 1865, pp. 63, 162 (Puynipet, 
Lugunor, Ulcei). 

Calamodyta syrinx Gray, Hand-list Birds, 1, 1869, p. 208 (Ualan); Giebel, Thes. 
Ornith., 1, 1872, p. 529 (Carolin.). 

Calamoherpe syrinx Finsch, Journ. Mus. Godeffroy, 12, 1876, p. 17 (Ponape, 
Lugunor, Ruck, Ualan, Uleei); idem, Proc. Zool. Soc. London, 1877 (1878), p. 778 
(Ponape); idem, Journ. f. Ornith, 1880, pp. 287, 297 (Ponape, Ruck, Mortlocks, 
Kuschai); idem, Proc. Zool. Soc. London, 1880, p. 575 (Ruk, Ponape); idem, Ibis, 
1881, pp. 108, 112, 115, 247 (Kuschai, Ruck, Ponape, Mortlocks); Schmeltz and 
Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, pp. 298, 330, 353 (Ponape, Mortlocks, 
Nukor, Ruk); Finsch, Ibis, 1883, p. 143 (Ruck); idem, Mitth. Ornith. Ver. Wien, 
1884, p. 49 (Ponape); idem, Sammlung wissensch. Vortrage, 14 ser., 1900, p. 659 
(Carolinen). 

Acrocephalus syrinx Seebohm, Cat. Birds British Mus., 5, 1881, p. 100 (Ponape); 
Tristram, Ibis, 1883, p. 44 (Ponape, Ruk, Mortlock, Lugunor, Uleei); idem. Cat. Birds, 
1889, p. 152 (Ponape, Ruk); Nehrkorn, Kat. Eiers., 1899, p. 33 (Ponape, Ruk); 
Hartert (part), Novit. Zool., 5, 1898, p. 58 (Carolines); idem, Novit. Zool., 7, 1900, 
p. 3 (Ruk); Scale (part), Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 53 (Po- 
nape); Matschie, Journ. f. Ornith., 1900, pp. 112, 113 (Ruk, Ponape, Ualan); Dubois, 
Syn. Avium, 1, 1902, p. 369 (Ponape); Reichenow, Die Vogel, 2, 1914, p. 545 (Po- 
nape); Takatsukasa and Kuroda, Tori, 1, 1915, p. 54 (Ponape, Ruk); Mayr, Proc. 6th 
Pacific Sci. Congr., 4, 1941, p. 204 (Ponapi). 

Conopoderas syrinx Wetmore, in Townsend and Wetmore, Bull. Mus. Comp. Zool., 
63, 1919, p. 214 (Ponape, Truk); Takatsukasa and Yamashina, Dobotsu. Zasshi, 43, 
1931, p. 485 (Caroline Is.); Yamashina, Tori, 7, 1932, p. 405 (Ponape); Hand-list 
Japanese Birds, rev., 1932, p. 177 (Carolines). 



Baker: The Avifauna of Micronesia 255 

Acrocephalus stentoreus syrinx Kuroda (part), in Momiyama, Birds Micronesia, 
1922, p. 67 (Ruk, Ualan, Lugunor, Wolea, Nukuoro, Ponape). 

Conopoderas luscinia syrinx Hand-list Japanese Birds, 3d ed., 1942, p. 197 (Wolea, 
Lamotrek, Truk, Lukunor, Nukuoro, Ponape, Kusaie). 

Acrocephalus luscinia syrinx Mayr, Birds Southwest Pacific, 1945, p. 294 (Caro- 
lines); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 67 (Truk). 

Geographic range. — Micronesia: Caroline Islands — Woleai, Lamotrek, Truk, 
Lukunor, Nukuoro, Ponape, Kusaie. 

Characters. — Adult: Resembles A. I. luscinia, but smaller; with shorter, 
straighter bill; head and neck more reddish-brown; back, rump, wing, and tail 
edged with cinnamon; flight feathers faintly tipped with white. 

Immature: Resembles adult, but lighter and more rufous in color; wings 
and rump paler, wings edged with rufous buff. 

Measurements. — Measurements are listed in table 37. 

Specimens examined. — Total number, 62 (35 males, 20 females, 7 unsexed), as follows: 
Caroline Islands, USNM— Ponape, 1 (Feb. 12)— Truk, 4 (Feb. 16, Mar. 15); AMNH— 
Ponape, 35 (Nov., Dec.) — Truk, 22 (Feb., March, May, June, Nov.). 

Nesting. — Birds nest in reedy swamps and scrub vegetation in the Caroline 
Islands, although Finsch (1881b: 115), recording a field note by Kubary, states 
that nests were found in trees at Mortlock Atoll (= Lukunor). Yamashina 
( 1932a :405) reports the collecting of seven sets of eggs at Ponape in July and 
August, 1931. The sets consisted of one or two eggs each. McElroy of the 
NAMRU2 party obtained specimens with enlarged gonads at Truk in Decem- 
ber and noted that birds were carrying nest materials to cane swamps. Of the 
birds secured by Coultas in November and December at Ponape, only a small 
number had enlarged gonads. He also found nests containing no eggs in low 
bushes at Ponape. Hartert (1900:3) reports that at Truk Owston's Japanese 
collectors obtained "many nests" from the end of May to the beginning of July. 
These nests contained one or two eggs and were found 7 to 20 feet above the 
ground in breadfruit, coconut and ivory-nut palm trees. Hartert writes, "The 
eggs are white, covered with darker and lighter brown patches, and underlying 
ashy grey or lavender-grey spots. These spots are generally thicker near the 
broad end, sometimes forming a loose ring, and they are sometimes equally 
spread over the whole surface." He lists measurements of 48 eggs. 

Molt. — Of the specimens examined by me, those taken in the spring and 
summer are in fresh or worn plumage; those taken in fall and winter are in 
molt, with a few skins exhibiting worn or fresh plumage in the latter period. 
Apparently the peak in the molting process occurs from September to De- 
cember. 

Food habits. — The reed-warbler is an insect feeder. Coultas, in his observa- 
tions of the bird at Ponape, relates that he was able to locate the warbler by 
listening for the "snapping of the mandibles as the bird is catching food." 

Remarks. — From the observations of Kittlitz, Kubary, Coultas, 
McElroy, and others, it is apparent that the Nightingale Reed- 
warbler in the Caroline Islands is restricted to the lower elevations 
of the islands. Whereas the reed-warbler at Guam seems closely 
associated with cane swamps and adjacent vegetation, the bird in the 
Carolines may range more extensively into brush lands, forest mar- 



256 University of Kansas Publs., Mus. Nat. Hist. 

gins and grass lands. Coultas (field notes) notes that the reed- 
warbler at Ponape is a "common bird of the small bush and grass- 
lands. One is attracted by its warbler-like song. The bird spends 
hours perched on a stem of a bush caroling the time of day. When 
feeding, one finds it on the ground or working away quietly among 
the bushes. Acrocephalus is a friendly bird who does not become 
frightened easily. He responds to man-made calls." 

The Nightingale Reed-warbler is found on many of the islands in 
the Caroline Chain, including both the "high" volcanic islands (Po- 
nape and Truk) and the "low" coral islands (Lukunor and Nuku- 
nor) . Although the bird has been recorded at Kusaie by Kittlitz and 
Finsch, it was not taken there by Coultas in 1931. Reed-warblers 
are unknown at Yap, Ulithi, Fais or at other islands of the extreme 
western Carolines, or in the Palau Archipelago. 

They are unrecorded also in the Marshall Islands, but at Nauru 
in the Gilbert Islands, to the southeast, an isolated population of this 
bird occurs and has been named A. I. rehsei (Finsch). 

Acrocephalus luscinia yamashinae (Takatsukasa) 
Nightingale Reed-warbler 

Conopoderas yamashinae Takatsukasa, Dobutsu. Zasshi, 43, 1931, p. 485. (Type 
locality, Pagan.) 

Tatare syrinx Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, p. 
210 (Pagan). 

Acrocephalus syrinx Hartert (part), Novit. Zool., 5, 1898, p. 58 (Pagan); Seale 
(part), Occ. Papers Bemice P. Bishop Mus., 1, 1901, p. 53 (Pagan). 

Acrocephalus stentoreus syrinx Kuroda (part), in Momiyama, Birda Micronesia, 1922, 
p. 67 (Pagan). 

Conopoderas yamashinae Hand-list Japanese Birds, rev., 1932, p. 177 (Pagan). 

Conopoderas luscinia yamashinae Hand-list Japanese Birds, 3d ed., 1942, p. 196 
(Pagan). 

Acrocephalus luscinia yamashinae Mayr, Birds Southwest Pacific, 1945, p. 294 
(Pagan). 

Geographic range. — Micronesia: Mariana Islands — Pagan. 

Characters. — Resembles A. I. syrinx, but duller and more brownish and less 
olive-rufous on back, rump and tail; bill shorter and more curved. 

Takatsukasa (1931:485) gives the following description: "Upperparts dark 
olive brown, paler on the lower rump; remiges and rectrices dark olive-brown, 
margined with brown. Superciliary stripe distinct and buff; chin, throat, breast 
and abdomen pale brown; ear-coverts, sides of neck, sides of breast and flanks 
dusty greyish brown, belly and under tail-coverts pale buff. Bill clove brown, 
legs grey, and iris Van Dyke brown." He continues, "It differs from Conopo- 
deras syrinx of Caroline Islands by its colouration and the shape of the bill, 
namely in the new form the culmen is more curved and more stout, and the tail 
is less roundish and nearly square." 

Measurements. — Takatsukasa and Yamashina (1931b :485) lists the follow- 
ing measurements: 13 adult males — wing, 75-80; tail, 65-70; culmen, 20-22; 6 
adult females— wing, 73-77; tail, 60-65; culmen, 20-22. 



Baker: The Avifauna of Micronesia 257 

Mayr examined seven specimens from Pagan in the Paris Museum. His 
measurements are: five males — wing. 76-79; tail, 66-69; bill from nostril, 14-14.5; 
two females — wing, 75, 77; tail, 66, 67; bill from nostril, 14.5, 15. 

Remarks. — No specimens have been examined by me. Oustalet 
(1895:210) was the first to note the difference between the reed- 
warblers from Pagan and those from Guam and Saipan (A. I. lus- 
cinia). He regarded those from Pagan as similar to the population 
in the Carolines, calling them Tatare syrinx. Hartert, Seale, and 
Momiyama followed Oustalet in this regard, and it was not until 
1931 that the population at Pagan was recognized as distinct, when 
further collections were made by the Japanese. 

Acrocephalus luscinia nijoi (Yamashina) 
Nightingale Reed-warbler 

Conopoderas luscinia nijoi Yamashina, Tori, 10, 1940, p. 674. (Type locality, 
Agiguan.) 

Conopoderas luscinia nijoi Hand-list Japanese Birds, 3d ed., 1942, p. 196 (Agiguan). 

Acrocephalus luscinia luscinia Mayr (part). Birds Southwest Pacific, 1945, p. 294 
(Agiguan). 

Geographic range. — Micronesia: Mariana Islands-Agiguan. 

Characters. — Adult: Resembles A. I. lusciana, but with shorter bill. Yama- 
shina (1940:674) describes the birds as, "upper parts much less rusty in 
colour and the flanks and bellies are darker and more brownish than those 
of the specimens from Almagan and Saipan." 

Measurements. — Yamashina (1940:674) gives the measurements of five 
adult birds from Agiguan as: exposed culmen 27-29, bill from nostril 17.0- 
20.0; as compared with 27 adult birds from Almagan and Saipan as: exposed 
culmen 30-34, bill from nostril 21.2-24.5. 

Remarks. — No specimens have been examined by me. The island 
of Agiguan is a very small one lying offshore from Tinian and not 
far from Saipan, where A. I. luscinia occurs. A. I. nijoi is given 
tentative recognition, on the basis of the measurements of the five 
adult specimens given by Yamashina. These indicate that the 
population has a distinctly shorter bill. 

Evolutionary history of Acrocephalus luscinia. — The species of 
Acrocephalus in Micronesia and Polynesia have received several 
taxonomic treatments. In regard to the Micronesian forms, Quoy 
and Gaimard called the population at Guam Thryothorus while 
Kittlitz called the population in the Carolines, Sylvia. Evidently 
to emphasize the distinctness of these two birds, Reichenbach 
in 1850 renamed the bird in the Marianas as Hybristes and the 
bird in the Carolines as Eparnetes. The birds were later placed 
in the genus, Tatare, by Hartlaub, Gray, Sharpe and other workers. 
Gray also used the name, Calamodyta, for the bird in the Carolines. 
17—8131 



258 University of Kansas Publs., Mrs. Nat. Hist. 

The generic term, Calamoherpe, was employed also by a number of 
workers for the Caroline population. Sharpe (1883:525) placed 
the reed-warblers in the family Timelidae and retained the name, 
Tatare, for the Micronesian and Polynesian forms. In distinguish- 
ing Acrocephalus from Tatare he has the following to say of Acro- 
cephalus: "besides having a much shorter bill, possesses a very 
much more pointed wing, the distance between the primaries and the 
secondaries being much more than the length of the hind toe and 
claw; whereas in Tatare the wing is much more obtuse, and the 
distance between the primaries and the secondaries is less than the 
length of the hind toe and claw." More recent authors have fol- 
lowed Shrape using the generic name, Conopoderas {=: Tatare, old 
name preoccupied). However, Tristram (1883:38-46) regarded the 
separation of these oceanic forms from Acrocephalus as a taxonomic 
error. He said that this is "one of the very few links (the others 
being the solitary Hirundo tahitica and the Merulae) between the 
avifauna of Oceania and our own; and it has a much wider range 
east and west than either of the other links, extending from the 
Carolines in the east to the Marquesas in the west." Mayr has 
pointed out (orally to the writer) that the separation of the Oceanic 
reed-warblers from Acrocephalus is an unnatural one, although it 
is perfectly true that the extreme members {A. caffra and A. I. lus- 
cinia) have a very long bill, but forms with shorter bills like A. I. 
syrinx point to the close affinity between the continental species 
and these insular birds. This has also been noted by Hartert (1898: 
58). Mayr {in litt.) comments that "There is no difference between 
Acrocephalus and Conopoderas in regard to the wing formula, pro- 
vided that we compare the Polynesian species with the tropical 
forms of Acrocephalus (such as toxopei and cervinus). The char- 
acter mentioned by Sharpe is very artificial and merely indicates 
the difference in the wing between a migrant of the temperate zone 
and a resident of the tropics. There is no denying that some of the 
warblers of eastern Polynesia are no longer reed-warblers but have 
become dwellers of trees and bushes. However, this same tendency 
prevails among some of the unquestionable species of Acrocephalus 
(scirpaceus and palustris) and at any rate a slight change in habits 
is not sufficient for generic separation." Earlier, Mayr (1942b: 169) 
used Conopoderas as one of the several genera that is based on 
"morphologically distinct geographic forms." The degree of modi- 
fication that has occurred in these oceanic reed-warblers, would, if 
the birds were in a continental area, undoubtedly be considered 



Baker: The Avifauna of Micronesia 259 

worthy of specific or even generic rank by some authors; however, 
as Mayr (1942b: 162) points out, "the majority of well-isolated 
subspecies have all the characters of good species and are indeed 
considered to be such by the more conservative systematists." Ow- 
ing to their differentiation, the Micronesian and Polynesian reed- 
warblers might not be considered by some ornithologists as belong- 
ing to a single superspecies; however, all evidence seems to point 
to the origin of this group by a single invasion from Asia." 

Tristram (1883:41) was the first worker to recognize the relation- 
ship of the Micronesian and Polynesian reed-warblers to the conti- 
nental forms, when he placed them within the genus Acrocephalus. 
Rothschild (1893:2) further stated, "Tatare cannot be separated 
generically from Acrocephalus." In discussing the status of the Ha- 
waiian species, A. jamiliaris, Hartert (1898:58) also follows this 
treatment. Bryan (1941:187) also comments on the relationship of 
the "miller" birds of Laysan and Nihoa to species at Guam, Christ- 
mas and other islands of the Pacific. 

The reed-warblers of Polynesia and Micronesia represent an an- 
cient invasion from Asia. The continental form, Acrocephalus arun- 
dinaceous, is apparently closest to the ancestral stock of these oce- 
anic birds. This species resembles the oceanic populations in size, 
general coloring, shape of bill, and wing and tail structure. Some of 
the continental races of this species have a shorter first primary 
which is similar to that in the oceanic forms. How rapid the spread 
was of the reed warbler through the large insular area that it now 
occupies is unknown. A. syrinx of Micronesia has a shortened wing 
and some populations have a long bill. Species in Polynesia have 
stronger wings than the one in Micronesia, but have become differen- 
tiated in other ways, as, for example, by the long bill of A. caffra and 
the small size of A. aequinoctialis. In addition, call notes have 
become varied, as noted by Chapin (in Mayr, 1942b:54). Also cer- 
tain of the reed-warblers have become bush and tree-living birds. 
The Hawaiian birds are reduced in size and have become tree-living 
in a manner similar to that of other Polynesian species. These modi- 
fications of the reed-warblers of the Oceanic area appear, according 
to Murphy and Mathews (1929), to indicate their long-time resi- 
dence in the islands, as compared with subspecies of A. arundina- 
ceous that are found in Melanesia. The latter birds, which are not 
ancestral to the Polynesian birds, resemble closely their Asiatic an- 
cestors and have also retained their swamp-living habits. This 
would seem to indicate that the birds in Melanesia may be of more 



260 



University of Kansas Publs., Mus. Nat. Hist. 



recent occurrence. Stresemann (1939b: 324) presents a map of the 
distribution of A. arundinaceous in southeastern Asia and adjacent 
islands. The original stock came from a point in China, north of 
Indochina, spreading to the Philippines and to Celebes, from where 
it reached the Solomons and New Guinea via the Lesser Sundas and 
Australia. 




Fig. 15. Geographic distribution of Acrocephalus in the Pacific area and 
routes of its dispersal. (1) A. arundinaceus; (2) A. luscinia; (3) ranges of A. 
atypha, A. caffra, and A. vaughani; (4) A. aequinoctialis; (5) A. familiaris. 



The path of invasion of Oceania by the reed-warbler is pictured in 
figure 15. Probably the birds became established in Micronesia by 
an invasion from the Bonins, where A. arundinaceus orientalis is 
known to occur today. From the Marianas and Carolines, the birds 
spread to Polynesia; A. I. rehsei of the Gilbert Islands (Nauru) 
might well be a connecting link. Possibly, the Hawaiian birds came 
as a separate invasion via the Volcano and Bonin islands or through 
the Micronesia Chain, or through the Line and Christmas islands 
from the south. It seems evident, however, that owing to their geo- 
graphic proximity and comparative structural similarity, the species 
in Hawaii is closest to A. luscinia of Micronesia. The absence of 
reed-warblers from the western Carolines and Palaus seems to reduce 
the possibility of an invasion from the Philippine region. However, 
reed-warblers are absent from the Marshall and the northern Gilbert 
islands, where there is undoubtedly suitable habitat for their occur- 
rence. Possibly these islands were once occupied by the birds but 
they were eliminated by natural causes or by man and his land uses. 



Baker: The Avifauna of Micronesia 261 

Rhipidura rufifrons uraniae Oustalet 
Rufous-fronted Fantail 

Rhipidura Uraniae Oustalet, Bull. Soc. Philom. Paris, (7), 5, 1881, p. 76. (Type 
locality, Mariannes = Guam.) 

Rhipidura pectoralis Gray, Cat. Birds Trop. Is. Pacific Ocean, 1859, p. 17 (Ladrone 
or Marian Is. = Guam). 

Rhipidura uraniae Reichenow and Schlow, Journ. f. Ornith., 1884, p. 398 (Mariannes 
=:Guam); Hartert, Novit. Zool., 5, 1898, p. 53 (Guam); AViglesworth, Abhandl. und 
Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 20 (Marianne = Guam) ; But- 
tikofer. Notes Leyden Mus., 15, 1893, p. 76 (Guam); Wheeler, Report Island of Guam, 
1900, p. 13 (Guam); Matschie, Journ. f. Ornith., 1901, pp. 112, 113 (Guam); Seale, 
Occ. Papers Bemice P. Bishop Mus., 1, 1901, p. 48 (Guam); Safford, Osprey, 1902, p. 
69 (Guam); Dubois, Syn. Avium, 1, 1902, p. 277 (Guam); Safford, The Plant World, 
7, 1904, p. 263 (Guam); idem, Contr. U. S. Nat. Herb., 9, 1905, p. 79 (Guam); 
Mearns, Proc. U. S. Nat. Mus., 36, 1909, p. 477 (Guam) ; Schnee, Zeitschr. f. Natur- 
wisch., 82, 1910, p. 464 (Marianen =: Guam) ; Reichenow, Die Vogel, 2, 1914, p. 267 
(Marianen = Guam); Cox, Island of Guam, 1917, p. 21 (Guam); Kuroda, in Momi- 
yama. Birds Micronesia, 1922, p. 65 (Guam); Bryan, Guam Rec, vol. 13, no. 2, 1936, 
p. 25 (Guam). 

Rhipidura atrigularis Reichenow, Journ. f. Ornith., 1885, p. 110 (Type locality, 
Palau, error = Guam); Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 
1890-1891 (1891), p. 21 (Pelew, error = Guam); Takatsukasa and Kuroda, Tori, 1, 
1915, p. 63 (Marianne = Guam). 

Rhipidura versicolor Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 
1895, p. 190 (Guam). 

Rhipidura rufifrons uraniae Mathews, Syst. Avium Australasianarum, 2, 1930, p. 490 
(Marianne r= Guam); Hand-list .Japanese Birds, rev., 1932, p. 176 (Guam); Hand-list 
Japanese Birds, 3d ed., 1942, p. 195 (Guam); Mayr, Birds Southwest Pacific, 1945, p. 
295 (Guam); Watson, The Raven, 17, 1946, p. 42 (Guam); Mayr and Moynihan, Amer. 
Mus. Novit., no. 1321, 1946, pp. 3, 9 (Guam); Baker, Proc. Biol. Soc. Washington, 59, 
1946, p. 77 (Guam); idem, Smithson, Misc. Coll., vol. 107, no. 15, 1948,, p. 67 (Guam). 

Rhipidura rufifrons Wharton, Ecol. Monogr., 16, 1948, p. 174 (Guam); Strophlet, 
Auk, 1946, p. 339 (Guam). 

Geographic range. — Micronesia: Mariana Islands-Guam. 

Characters. — Adult: Forehead and anterior crown near "cinnamon-buff"; 
lores and orbital ring black, auriculars more brownish than lores; malar stripe 
white; a few feathers in posterior malar region tipped with "citrine drab"; 
anterior part of chin white; posterior part of chin, throat, and upper breast 
black; feathers on breast edged with white; lower breast, abdomen, sides, 
flanks, tibia, vent, and under tail-coverts near "roA'al brown," becoming lighter 
on breast and more rufous on under tail-coverts; sides of neck and back near 
"Dresden brown," becoming grayer on neck and crown where feathers have 
darker shafts; rump and upper tail-coverts near "orange rufous"; basal half 
of tail slightly lighter than rump ; terminal part of tail black, tipped with 
white; wings dark edged with coloring like back; under wing grayish with 
axillaries tipped with buffy-white ; bill black with base of upper mandible 
lighter; feet dark brown; iris dark brown. 

Immature: Resembles adult, but head, neck, scapulars, and secondaries 
edged with nifous; feathers of chin and throat edged with whitish. Younger 
birds may have less rufous on head but feathers of body more rufous with 
creamy edges. 

Measurements. — Measurements are listed in table 38. 



262 University of Kansas Publs., Mus. Nat. Hist. 

Table 38. Me.asurements of Rhipidura rufifrons in Micronesia 



Subspecies 


Number 
and 
sex 


Wing 


Tail 


Exposed 
culmen 


Tarsus 


R. 


T. uraniae 


11 males 
6 females 


66 

(64-69) 

65 
(61-68) 


78 
(75-82) 

76 
(73-81) 


13.6 
(13.1-14.5) 

12.3 
(11.6-12.5) 


16.6 

(15.6-17.2) 

16.8 
(16.1-17.6) 


R. 


r. saipanensis . . 


7 males 
6 females 


68 
(68-69) 

64 
(62-66) 


81 

(80-83) 

76 
(72-81) 


13 3 

(13.0-13.5) 

12.7 
(12.4-13.4) 


17.3 

(16.2-18.4) 

17.9 
(17.2-18.1) 


R. 


r. mariae 


2 males 


65,67 


82,82 


12.1, 12.4 


17.1, 17.2 


R. 


r. kubaryi 


14 males 
10 females 


77 
(75-79) 

72 
(69-75) 


88 
(82-95) 

87 
(83-90) 


14.4 
(13.6-15.0) 

14.5 
(14.0-15.0) 


20.0 
(19.0-21.0) 

20.0 
(20.0-20.5) 



Weights. — The NAMRU2 party recorded the weights of nine males as 
9.0-10.0 (9.0) ; of three females as 7.2-9.6 (8.S) grams- 

Specimens examined. — Total number, 41 (19 males, 14 females, 8 unsexed), as follows: 
Mariana Islands, USNM— Guam, 17 (May 29, 30, June 6, 14, 18, July 12, 20); AMNH— 
Guam, 24 (Jan., Feb., March, Aug., Sept., Dec.). 

Nesting. — Hartert (1898:54) recorded ne.sts taken at Guam in February 
and March. 

Molt. — On the basis of specimens examined, it is apparent that molt begins 
in August or September and continues through the months of the fall. 

Parasites — Wharton (1946:174) obtained the chigger (Acarina), Tromhicula 
sp., from this bird at Guam. 

Remarks. — The Rufous-fronted Fantail at Guam is a birci of the 
forest and forest scrub. It prefers the areas where leafy under- 
growth is present. It moves rapidly about continually fluttering 
its wings and spreading its long fanlike tail. The birds are usually 
observed in pairs. On January 21, 1945, E. W. Coleman of the 
NAMRU2 party killed a fantail but before he could retrieve it, a 
large toad [Bujo marinus) seized the fallen bird and carried it into 
a hole in the ground. 

Rhipidura rufifrons saipanensis Hartert 
Rufous-fronted Fantail 

Rhipidura saipanensis Hartert, Novit. Zool., 5, 1898, p. 54. (Type locality, Sai- 
pan). 

Rhipidura versicolor Oustalet, Le Nat., 1889, p. 260 (Mariannes = Saipan) ; 
Wiglesworth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), 
p. 21 (Marianne = Saipan) ; Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 
7, 1895, p. 190 (Saipan). 

Rhipidura saipanensis Matschie, Joum. f. Ornith., 1901, pp. 112, 113 (Saipan); 
Scale, Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 48 (Saipan) ; Dubois, Syn. 



Baker: The Avifauna of Micronesia 263 

Avium, 1, 1902, p. 277 (Saipan) ; Takatsukasa and Kuroda, Tori, 1, 1915, p. 63 
(Marianne = Saipan). 

Rhipidura Tufifrons saipanensis Kuroda, in Momiyama, Birds Micronesia, 1922, p. 
65 (Saipan); Mathews, Syst, Avium Australasianarum, 2, 1930, p. 490 (Saipan); 
Hand-list Japanese Birds (part), rev., 1932, p. 176 (Saipan, Tinian) ; Hand-list Japa- 
nese Birds (part), 3d ed., 1942, p. 195 (Saipan, Tinian); Mayr (part), Birds South- 
west Pacific, 1945, p. 295 (Saipan, Tinian); Mayr and Moynihan (part), Amer. Mus. 
Novit., no. 1321, 1946, p. 3 (Saipan, Tinian); Baker, Proc. Biol. Soc. Washington, 
59, 1946, p. 77 (Saipan, Tinian); Downs, Trans. Kansas Acad. Sci., 49, 1946, p. 98 
(Tinian). 

Rhipidura lepida saipanensis Stott, Auk, 64, 1946, p. 527 (Saipan). 

Geographic range. — Micronesia: Mariana Islands — Saipan, Tinian. 

Characters. — Adult: Resembles adult of R. r. uraniae, but forehead and 
anterior crown more rufous; posterior crown and nape lighter; rump and 
upper tail-coverts lighter and richer in color; white malar stripe broader; 
chin with white feathering more extensive, covering edge of upper throat. 

Measurements — Measurements are listed in table 38. 

Specimens examined. — Total number, 16 (9 males, 6 females, 1 unsexed), as follows: 
Mariana Islands, USNM — Saipan, 1 (Dec. 15) — Tinian, 3 (Oct. 16, 23); AMNH — Saipan, 
6 (July, Aug.) — Tinian, 6 (Sept.). 

Molt. — Molt begins in July and extends through the autumn. Most of the 
specimens examined, that were taken in this period, are in molt. 

Food habits. — Stott (1947:527) writes that the fantail forages for insects 
in the undergrowth and also while on the wing captures flying insects. Downs 
(1946:99) made similar observations concerning this bird at Tinian. 

Remarks. — In studying the collection of faintails obtained by 
Marche at Guam and Saipan, Oustalet (1895:191) reached the con- 
clusion that the birds from these two islands were the same as the 
bird from Yap, which he called R. versicolor. He thought that the 
white-throated birds were in breeding plumage, and that the black- 
throated birds (from Guam) were in autumn and winter dress. This 
error was corrected by Hartcrt (1898:53). 

Downs (1946:98-100) has published some interesting observations 
concerning the fantail at Tinian. He describes feeding behavior and 
the song which he says is "a beautiful rolling whistle, starting rather 
shrilly, then rolling on. Something like a meadow-lark and song 
sparrow combined." Gleise (1945:220) estimated the population of 
fantails at Tinian to be "40-50" in 1945. In 1931, Coultas found the 
bird at Tinian but not at Saipan. Stott (1947:527) observed the 
bird at Saipan 'in forested areas and vine-draped crevices in the lava 
above Magicienne Bay." 

Rhipidura rufifrons mariae R. H. Baker 
Rufous-fronted Fantail 

Rhipidura rufifrons ?nariae R. H. Baker, Proc. Biol. Soc. Washington, 59, 1946, p. 
7. (Type locality. Rota.) 

Rhipidura rufifrons saipanensis Takatsukasa and Yamashina, Dobutsu. Zasshi, 44, 
1932, p. 222 (Rota); Hand-list Japanese Birds (part), rev., 1932, p. 176 (Rota); 
Hand-list Japanese Birds (part), 3d ed., 1942, p. 195 (Rota); Mayr (part). Birds 



264 University of Kansas Publs., Mus. Nat. Hist. 

Southwest Pacific, 1945, p. 295 (Rota); Mayr and Moynihan (part), Amer. Mus. 
Novit., no. 1321, 1946, p. 3 (Rota). 

Rhipidura rufifrons mariae Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, 
p. 68 (Rota). 

Geographic range. — Micronesia: Mariana Islands — Rota. 

Characters. — Adult: Resembles adult of R. r. saipanensis, but with richer 
brown coloring on the breast and abdomen; darker above, especially the fore- 
head, rump, and basal part of tail; chin with .small mount of white; malar line 
of white thinner. 

Measurements. — Measurements are listed in table 38. 

Weights. — Baker (1946:78) records the weights of two adult males from 
Rota as 8.3 and 9.0 grams. 

Specimens examined. — Total number, 2 males, from Mariana Islands, USNM — Rota 
(Oct. 22). 

Remarks. — Takatsiikasa and Yamashina (1932:222) published 
the first account of the fantail from Rota although Coultas (field 
notes) obtained a report of its presence at Rota in 1931. The 
NAMRU2 party obtained the two specimens studied, and reported 
that the birds were numerous in the forested areas of Rota in 1945. 

Rhipidura rufifrons versicolor Hartlaub and Finsch 
Rufous-fronted Fantail 

Rhipidura versicolor Hartlaub and Finsch, Proc. Zool. Soc. London, 1872, p. 96. 
(Type locality, Uap.) 

Rhipidura versicolor Graffe, Journ. Mus. Godeffroy, 2, 1873, p. 123 (Yap); Sharpe, 
Cat. Birds British Mus., 4, 1879, p. 320 (Yap); Nehrkorn, Journ. f. Ornith., 1879, 
p. 402 (Yap); Oustalet, Bull. Soc. Philom. Paris, (7), 5, 1881, p. 76 (Uap); Schmaltz 
and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, p. 391 (Yap); Wiglesworth (part), 
Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 21 (Uap); 
Buttikofer, Notes Leyden Mus., 15, 1893, p. 78 (Uap); Oustalet (part), Nouv. Arch. 
Mus. Hist. Nat. Paris, (3), 7, 1895, p. 191 (Yap); Hartert, Novit. Zool., 5, 1898, p. 
54 (Yap) ; Bolau, Mitteil. Naturhist. Mus. Hamburg, 1898, p. 54 (Yap) ; Matschie, 
Journ. f. Ornith.. 1901, pp. 112, 113 (Yap); Dubois, Syn. Avium, 1, 1902, p. 277 
(Yap); Reichenow, Die Vogel, 2, 1914, p. 267 (Jap); Takatsukasa and Kuroda, Tori, 
1, 1915, p. 64 (Mackenzie = Yap) ; Kuroda, in Momiyama, Birds Micronesia, 1922, 
p. 66 (Yap); Hand-list Japanese Birds, rev., 1932, p. 176 (Yap). 

Rhipidura rufifrons versicolor Mathews, Syst. Avium Australasianarum, 2, 1930, p. 
489 (Uap); Hand-list Japanese Birds, 3d ed., 1942, p. 196 (Yap); Mayr, Birds 
Southwest Pacific, 1945, p. 295 (Yap) ; Mayr and Moynihan. Amer. Mus. Novit., 
no. 1321, 1946, p. 3 (Yap). 

Geographic range. — Mici'onesia: Caroline Islands — Yap. 

Characters. — Adult: Resembles R. r- uraniae, but chin and upper throat 
white; upper parts darker; abdomen whitish. 

The description of the adult given by Hartlaub and Finsch (1872:96) is 
"Upper parts a rich brown with a slight reddish tinge; forehead bright rufous; 
upper and under tail-coverts rufous; throat white, margined underneath by 
an irregular jugular band of pure black; pectoral plumes black, broadly mar- 
gined with yellowish white; middle of abdomen whitish, sides of a paler 
olive-brown under wing-coverts whitish; wing-feathers blackish brown; tail 
feathers brownish black, all largely tipped with white, the four middle ones 



Baker: The Avifauna of Micronesia 265 

rufous at the base, the white terminal spots becoming smaller towards the 
middle; beak fuscous, the under mandible paler except at tip; feet fuscous." 
Hartert (1898:54) writes that R. r. saipanensis differs from the bird at Yap 
"in having the bases of all rectrices rufous, the rump and upper tail-coverts 
rufous. The sides of the abdomen are not olive-brown, but rufous." 

Remarks. — No specimens of the Rufous-fronted Fantail of Yap 
have been seen by me. On the basis of published descriptions and 
comments, it appears that the bird is subspecifically distinct from 
the forms in the Marianas but shows close relationships to them. 
R. r. versicolor has the chin and throat white; R. r. saipanensis has 
the chin and part of the throat white and a heavy, white line in the 
malar region; R. r. mariae has the chin and only a small amount of 
the throat white and a thinner, white malar stripe; R. r. uraniae 
has only a small amount of white present on the chin and a very 
thin, white line in the malar region. 

Rhipidura rufifrons kubaryi Finsch 
Rufous-fronted FaJitail 

Rhipidura kubaryi Finsch, Proc. Zool. Soc. London, 187,') (1876), p. 044. (Type 
locality, Ponape.) 

Rhipidura kubaryi Finsch, Journ. Mus. Godeffroy, 12, 1870, pp. 17, 29, pi. 2, 
fig. 2 (Ponape); idem, Proc. Zool. Soc. London, 1877 (1878), p. 779 (Ponape); Nehr- 
korn, Journ. f. Ornith., 1879, p. 403 (Ponape); Finsch, Journ. f. Ornith., 1880, p. 
289 (Ponape); idem, Ihis, 1881, pp. 110, 112, 115 (Ponape); Schiiieltz and Krause, 
Ethnogr. Abth. Mus. Godeffroy, 1881, p. 281 (Ponape); Tristram, Cat. Birds, 1889, 
p. 198 (Ponai>e) ; Wiglosworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890- 
1891 (1891), p. 20 (Ponape); IMittikofer, Notes Leyden Mus., 15, 1893, p. 70 (Po- 
nape); Matschie, Journ. f. Ornith., 1901, pp. 112, 113 (Ponape); Duliois, Syn. Avium, 
1, 1902, p. 277 (Ponape); Takatsukasa and Kuroda, Tori, 1, 1915, pp. 54, 64 (Po- 
nape); Wetmore, in Townsend and Wetmore, Bull. Mus. Comp. Zool., 63, 1919, p. 
204 (Ponape); Kuroda, in Momiyania, Birds Micronesia, 1922, p. 05 (Ponape); 
Yamashina, Tori, 7, 1932, p. 403 (Ponaf)e) ; Hand-list Japanese Birds, rev., 1932. p. 
176 (Ponape); Mayr, Proc. 6th Pacific Sci. Congr., i. 1941, p. 204 (Ponape); Hand- 
list Japane.'^e Birds, 3d ed., 1942, p. 196 (Ponape). 

Rhipidura kubarii Sharpe, Cat. Birds British Mus., 4, 1879, p. 314 (Ponape); 
Bolau, Mitteil. Naturhist. Mus. Hamburg, 1898, p. 55 (Ponape). 

Rhipdura rufifrons kubaryi Mayr, Birds Southwest Pacific, 1945, \i. 295 (Ponape); 
Mayr and Moynihan, .A.mer. Mus. Novit., no. 1321, 1940, pp. 3, 0. 9, 11, 12, 15, 
10 (Ponape). 

Geographic range. — Micronesia: Caroline Islands — Ponape. 

Characters. — Adult: Upper parts smoky olivaceous-brown, less smoky on 
rump and upper tail-coverts; anterior forehead and supraloral region nar- 
rowly edged with white; lores and orbital ring black; auriculars brown; 
feathers of chin and malar region tipped with white; rest of chin and throat 
black, lower feathers of throat edged with white ; abdomen dark olivaceous- 
brown with whitish mid-portion anteriorily; sides and under tail-coverts ashy, 
the latter broadly tipped with white ; wings and tail dark, tail tipped with 
white and outer rectrices more broadly so; axillaries and under wing-coverts 
gray, broadly tipped with white; bill and feet black, mandible basally whitish; 
iris dark brown. 



266 University of Kansas Publs., Mus. Nat. Hist. 

R. r. kubaryi resembles R. r. uraniae, but larger; lacking rufous coloring; 
smaller and shorter, white malar stripe ; white on chin reduced. 
Measurements. — Measurements are listed in table 38. 

Specimens examined. — Total number, 40 (24 males, 15 females, 1 unsexed), as follows: 
Caroline Islands, USNM — Ponape, 1 (Feb. 12); AMNH — Ponape, 39 (Nov., Dec). 

Nesting. — Yamashina (1932a :403) records nests containing one or two eggs 
taken at Ponape in 1931 on the following dates: July 11, August 2, 14, 19, 
22, 30. Coultas (field notes) obtained reports that the eggs are two in number 
and laid in a cup-shaped nest of grass and fern, which is placed near the 
ground. 

Molt. — Many of the specimens examined that were taken in November and 
December are in fresh or slightly worn plumage. Only a few are molting. 
Apparently molt occurs earlier, perhaps beginning in August and continuing 
until October or November. 

Remarks. — Coultas obtained a large series of these bircis at Po- 
nape in 1931. He writes (field notes) that the fantail is a common 
bird and is found in forest and brush lands. This bird has a nervous 
behavior similar to that of other fantails and is constantly "wagging 
its long tail." Coultas describes it as an aggressive bird, chasing 
honey-eaters and white-eyes. 

Rhipidura lepida Hartlaub and Finsch 
Palau Fantail 

Rhipidura lepida Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, p. 6. (Type 
locality, Pelew Islands.) 

Rhipidura lepida Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 117, 118 
(Pelew Islands) ; Gray, Hand-list Birds, 1, 1869, p. 331 (Pelew) ; Hartlaub and Finsch, 
Proc. Zool. Soc. London, 1872, pp. 89, 97 (Pelew); Finsch, Journ. Mus. Godeffroy, 8, 
1875, pp. 4, 21, pi. 4, fig. 2-3 (Palau); Sharpe, Cat. Birds Britich Mus., 4, 1879, 
p. 322 (Pelew); Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, p. 407 
(Palau); Tristram, Cat. Birds, 1889, p. 198 (Pelew); Wiglesworth, Abhandl. und 
Bar. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 21 (Pelew); Buttikofer, Notes 
Leyden Mus., 15, 1893, p. 81 (Pelew); Bolau, Mitteil, Naturhist. Mus. Hamburg, 
1898, p. 55 (Palau); Matschie, Journ. f. Ornith., 1901, pp. 112, 113 (Palau); Dubois, 
Syn. Avium, 1, 1902, p. 278 (Pelew); Reichenow, Die Vogel, 2, 1914, p. 267 (Palau); 
Takatsukasa and Kuroda, Tori, 1, 1915, p. 54 (Pelew); Kuroda, in Momiyama, Birds 
Micronesia, 1922, p. 66 (Pelew) ; Mathews, Syst. Avium A9ustralasianarum, 2, 1930, 
p. 484 (Pelew); Hand-list Japanese Birds, rev., 1932, p. 176 (Palau); Hand-lipt 
Japanese Birds, 3d ed., 1942, p. 196 (Babelthuap, Koror, Peliliu); Mayr and Moyni- 
han, Amer. Mus. Novit., no. 1321, 1946, pp. 3, 5, 8, 10, 12, 19 (Palau); Baker, 
Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 68 (Peleliu). 

Geographic range. — Micronesia: Palau Islands — Babelthuap, Koror, Gara- 
kayo Peleliu, Ngabad. 

Characters. — Adult: Upper parts near "cinnamon-rufous," slightly lighter on 
the upper wing-coverts, scapulars, edges of inner secondaries, and rump; lores 
blackish ; orbital ring and auriculars dark brown ; chin, upper throat, and malar 
region white; lower throat and upper breast black with ashy-gray sides; lower 
breast whitish; rest of underparts like back but slightly paler; wings dark; 
tail black with tips rufous, inner rectrices with rufous tips narrower than on 
tail; under wing-coverts and axillaries broadly edged with rufous; bill black- 



Baker: The Avifauna of Micronesia 



267 



ish, lower mandible whitish at base; feet brownish; iris dark brown. Female 
slightly smaller. 

Immature: Resembles adult, but head and neck brown; throat coloring 
dingy; patch on breast blackish cinnamon. Younger individuals may be more 
tinged with rufous above and below. 

Measurements. — Measurements are listed in table 39. 

T.ABLE 39. Me.\surements of Rhipidura lepida 



Number and Sex 


Wing 


Tail 


Culmen 


Tarsus 


7 adult males 


80 

(77-83) 

77 
( 76-79) 


89 
(85-94) 

86 

(83-88) 


15.5 
(14.5-16.0) 

15.5 
(14.5-15.5) 


23.3 


/ adult females 


(23.0-24.0) 
22.5 




(21.7-23.0) 



Speciviens examined. — Total number, 18 (9 males, 9 females), as follows: Palau Lslands, 
USNM— Koror, 2 (Nov. 6, 18)— Babelthuap, 1 (Nov. 27)— Peleliu, i (Aug. 29, 30, 31); 
AMNH — exact locality not given, 11 (Nov., Dec). 

Molt. — Some of the birds taken in August are in molt. Specimens taken 
in November and December are mostly in fresh plumage. Apparently this 
bird molts in late summer and early fall. 

Remarks.— In 1945 the NAMRU2 party found the Palau Fantail 
in small numbers at Peleliu, Garakayo and Ngabad. At Peleliu the 
birds were noted as singles or in pairs in brushy undergrowth in 
forested areas. The birds were observed also in the second growth 
vegetation in the battle areas. Coultas (field notes) found the bird 
to be rare and restricted to the true forest, when he visited the 
Palau Islands in 1931. The fantail is one of the most attractive 
birds found in the jungles of the Palau Islands. Its bright rufous 
coloring is conspicuously displayed by the rapid movements of the 
wings and tail as the bird moves and feeds in the undergrowth. The 
population is apparently not large, and the individual or pair of 
birds probably ranges in a relatively large home territory. 

Evolutionary History of Rhipidura in Micronesia. — The evolu- 
tionary history of Rhipidura in Micronesia has been studied con- 
siderably more than that of some of the other genera in the area. 
Oustalet (1896:70) notes a close relation between Rhipidura of the 
Marianas and R. rufi,frons of Australia. Mayr (1941b :202, 203) 
regards the genus Rhipidura as typical of the Polynesian area and 
remarks that speciation within this genus has proceeded at a rela- 
tively rapid rate. Mayr and Moynihan (1946) have devoted a 
21-page paper to a thorough discussion of the R. rufifrons group, 



268 University of Kansas Publs., Mus. Nat. Hist. 

based on the extensive collections at the American Museum of 
Natural History. They remark that no other genera are closely 
related to Rhipdura and that evolution has proceeded further in R. 
rufifrons than in any other species of the genus. These authors re- 
gard the Papuan area, probably New Guinea, as the original home 
of this group. From their study they point out that many of the 
subspecies of R. rufifrons of the Papuan area, especially those of 
the Louisiades and the Solomons, appear to be the least specialized 
of the species, and that this lack of specialization in these subspecies 
indicates that the ancestral stock of the species R. rufifrons acquired 
its specificity somewhere in that area. With regard to the kinds of 
Rhipdura in Micronesia, Mayr and Moynihan (1946: fig. 2) have 
logically found three separate colonizations within the area: one 
represented today by R. lepida at Palau; one of subspecies of R. 
rufifrons at Yap and in the Marianas; and one by R. r. kubaryi at 
Ponape. 

R. lepida, according to Mayr and Moynihan (1946), is a result 
of an early colonization by Rhipidura. It is related to R. dedemi, 
R. superfiua, and R. teijsinanni, which are mostly monotypic or have 
only two or three subspecies within the species. These three species 
are found in the region including Celebes and the Moluccas. R. 
lepida apparently invaded the Palau Islands from Celebes or an 
adjacent area and, among named species, most closely resembles 
R. teijs77ianm. Both of these species have a white chin and throat, 
black breast patch, and rufous abdomen. R. lepida has become 
differentiated chiefly by the presence of a rufous head and back, a 
more distinct breast band, and proportionately different amounts of 
rufous and black coloration of the tail feathers. 

Mayr and Moynihan (1946:6) give as the chief characters of 
R. rufifrons the following: "a rufous forehead, a grayish brown 
head and upper back, a well-defined rufous rump, a white chin and 
throat, a black breast band with scaling at its lower edge, and a 
dark brown tail with a distinct rufous base and a white tip." The 
Micronesian subspecies of R. rufifrons at Yap and in the Marianas 
display these characters. Of the four subspecies found in the area 
including Yap and the Marianas, R. r. versicolor, R. r. saipanensis, 
R. r. mariae and R. r. uraniae, the two first named most closely 
approach the ancestral stock, which may have been R. r. comnioda 
Hartert of the northern Solomons or some near relative in Mela- 
nesia. The amount of white on the chin and throat and on the 
malar stripe, in R. r. versicolor and R. r. saipanensis is probably 



Baker: The Avifauna of Micronesia 269 

nearer that which obtained in the ancestor. At Rota, R. r. mariae, 
exhibits less white on the throat and a thinner, white malar stripe, 
while at Guam, R. r. uraniae possesses only a small amount of 
white on the chin and only a very thin line of white in the malar 
region. This variation in coloration suggests that the birds may 
have originally become established at Yap, Saipan and Tinian and 
later, birds from Saipan and Tinian spread to Rota and lastly to 
Guam. 

R. r. kubaryi of Ponape, although considered as a subspecies of 
R. rufifrons by most workers, has lost the rufous coloring found in 
most members of the species. Mayr and Moynihan (1946:6) point 
to its evolution through subspecies in the Santa Cruz Islands, where 
in R. r. agilis Mayr the rufous of the lower back is restricted to the 
upper tail-coverts, and in R. r. melanolaema Sharpe and R. r. 
utupuae Mayr the rufous is absent. In the latter two subspecies, as 
well as in R. r. kubaryi, the forehead is white instead of rufous. 

The invasion of Micronesia by Rhipidura has undoubtedly been 
the result of abnormally long flights by a relatively weak flyer. The 
fact that Rhipidura has succeeded in establishing itself at only a few 
of the seemingly suitable islands in Micronesia may indicate that the 
possibilities for chance migration and resulting colonization are 
small, but that new colonization may be expected in the future. 

It is my opinion that the populations of Rhipidura, as I have 
observed them in Micronesia, are small because each individual or 
pair of birds is dependent on a relatively large area of woodland to 
satisfy its habitat requirements, especially for food. This suggestion 
needs to be tested by observation made in the field. In comparison 
with the insect fauna of New Guinea or some other large island, that 
of Micronesia is indeed small in number of kinds. Hesse, Allee and 
Schmidt (1937:524) explain the absence of insectivorous animals 
such as "swallows, swifts, flycatchers, and insectivorous bats" in 
island communities on the basis of the small number of flying insects 
in these communities. Probably Rhipidura is able to forage for 
sedentary insect life as well as for the flying forms. 

Metabolus rugensis (Hombron and Jacquinot) 
Truk Monarch 

Muscicapa Rugeiisis Hombron and Jacquinot, Ann. Sci. Nat. Paris, (2), 16, 1841, 
p. 312. (Type locality, Roug = Truk.) 

Monarcha rugensis Hartlaub, Archiv. f. Naturgesch., 18, 18.52, p. 133 (Gruppe 
Roug. = Truk); idem, Journ. f. Ornith., 1854, p. 168 (Carolinen = Truk); idem, Proc. 
Zool. Soc. London, 1867 (1868), p. 829 (Hogoleu = Truk) ; Gray, Hand-list Birds, 1, 
1869, p. 321 (Caroline = Truk) ; Giebel, Thes. Ornith., 2, 1875, p. 614 (Carolinae == 
Truk); Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, p. 353 (Ruk). 



270 



University of Kansas Publs., Mus. Nat. Hist. 



Colluricincla rugensis Pucheran, Voy. Pole Sud, Zool., 3, 1853, p. 62 (Ruk); Hart- 
laub, Journ. f. Ornith., 1854, p. 162 (Roug = Truk). 

Metabolus rugensis Bonaparte, Comptes Rendus Acad. Sci. Paris, 38, 1854, p. 650 
(no locality = Truk); Sharpe, Cat. Birds British Mus., 4, 1879, p. 238 (Ruk); Finsch, 
Proc. Zool. See. London, 1880, p. 575 (Ruk); Tristram, Cat. Birds, 1889, p. 197 (Ruk); 
Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 19 
(Ruk); Nehrkorn, Kat. Eiers., 1899, p. 26 (Ruk); Hartert, Novit. Zool., 7, 1900, p. 
4 (Ruk); Matschie, Journ. f. Ornith., 1901, p. 112 (Ruk); Reichenow, Die Vogel, 2, 
1914, p. 262 (Karolinen = Truk) ; Takatsukasa and Kuroda, Tori, 1, 1915, p. 54 
(Ruk); Wetmore, in Townsend and Wetniore, Bull. Mus. Comp. Zool., 63, 1919, p. 
203 (Truk); Kuroda, in Moniiyama, Birds Micronesia, 1922, p. 63 (Ruk); Mathews, 
Syst. Avium Australasianarum, 2, 1930, p. 470 (Ruk) ; Yamashina, Tori, 7, 1932, p. 
404 (Truk); Hand-list Japanese Birds, rev., 1932, p. 178 (Truk); Hand-list Japanese 
Birds, 3d ed., 1942, p. 197 (Truk); Mayr, Birds Southwest Pacific, 1945, p. 295 
(Truk). 

Monarcha (Metabolus) rugensis Gray, Cat. Birds Trop. Is. Pacific Ocean, 1859, p. 
19 (Caroline Islands). 

Geographic range — Micronesia: Caroline Islands — Truk. 

Characters. — Adult male: White feathers (with dark bases) throughout 
except for black ones on forehead, lores, chin, and throat; chin and throat 
with steel-blue gloss; tips of second to fifth or seventh primaries black, black 
coloring extending along inner webs; shafts of primaries and basal half of 
tail feathers black; bill and feet black. 

Adult female: Resembles adult male, but generally sooty-black, darker 
above ; under tail-coverts and sometimes rump splotched with white ; white 
coloring may also be present on tips of secondaries, on chin, and on tail. 

Immature: Resembles adult, but bright cinnamon on upper parts and on 
wings and tail; inner webs of primaries grayish or dark brown, shafts of pri- 
maries lighter on basal half; lores, chin and throat white or washed with 
creamy-buff; breast and abdomen whitish, washed with cinnamon, sides darker 
rufous, under wing-coverts, axillaries, and under tail-coverts usually light 
rufous although somewhat variable in color; basal part of bill yellow, tip of 
bill horn colored. Immatures may be observed in all stages of color change 
toward the adult condition. 

Measurements. — Measurements are listed in table 40. 



Table 40. Measurements of Metabolus rugensis 



Number and Sex 


Wing 


Tail 


Culmen 


Tarsus 


8 males 


103 

(98-105) 

100 

(97-101) 


91 

(88-93) 

87 
(86-89) 


27 
(26-28) 

27 
(26-28) 


26 


6 females 


(25-27) 
26 




(25-27) 



Specimens examined. — Total number, 27 (14 males, 13 females), as follows: Caroline 
Islands, USNM— Truk, 2 (Feb. 16, not dated); AMNH— Truk, 25 (Jan. 29, Feb. 1, 8, 
10, 11, May 6, 9, June 11, 13, 14, 15, Oct. 11, 31, Nov. 2, 11, Dec. 3, 12, 17, 20). 

Nesting. — Yamashina (1932a :404) reports on the taking of a nest contain- 
ing one egg at Natsushima, Truk Atoll, in May, 1931. According to Hartert 



Baker: The Avifauna of Micronesia 271 

(1900:5) Owston's collectors obtained nests on June 1, 4, and 12. Two were 
in breadfruit trees about twenty feet above the ground. Each nest contained 
one egg. Hartert writes, "The eggs are cream-coloured, speckled with brownish 
red, more frequently and often very thickly on the large end, and with some 
deeper lying pale purplish grey patches, and one has some very fine black 
lines on the large end." 

Molt. — A study of adult specimens obtained at various times of the year 
indicates that Metabolus normally molts in the period from about October 
through January. 

Mayr (1933e:l-10) has studied the variation of immature and adult plum- 
ages in Neolalage banksiana (Gray) and other birds pointing out the occur- 
rence of "retarded" and "progressive" plumages. Bogert has followed this work 
in interpreting the condition of the plumages in Metabolus, and through the 
kindness of Ernst Mayr I have examined Bogert's unpublished manuscript on 
the series of Metabolus at the American Museum of Natural History, from 
which the following account of the plumage is taken. 

In the series of skins, there are specimens of non-molting, immature males 
with "normal" plumage (that is to say, plumage with upper parts cinnamon- 
colored and lower parts whitish and darker buff) taken in October and in 
February. There are also specimens of non-molting, immature females with 
"normal" plumage taken in November and in May. These immatures are in 
fresh or slightly worn plumages. In addition, there is one non-molting, male 
specimen (November) which has some white on the crown and throat, some 
black on the lores and chin, but because the black feathers are fresh, the 
specimen is considered to be a "transition" bird and may be either a "retarded" 
adult or a "progressive" immature male. One non-molting female (October) 
shows some sooty-black mottling on the chin and throat and a few black 
feathers on the crown ; this is apparently a "progressive" immature because the 
lower mandible has a yellow basal part, characteristic of the immature. An- 
other female (June) shows black feathers on the crown, nape, chin, throat, and 
breast; this bird is in the process of molting with the black feathers representing 
new growth and is an immature assuming the adult condition — in "progressive" 
plumage. One non-molting male (January) has an intermingling of white 
feathers in the cinnamon coloring of the head and body, black on the forehead, 
chin and throat, primaries black with cinnamon edges, and bill similar in color 
to that of the adult; it is considered to be an adult with "retarded" plumage. 
Two molting males (December) resemble adults except for cinnamon coloring 
on shoulders, back, primaries, retrices and a slight cinnamon wash on breast 
feathers; these may be "retarded" adults. One molting female (June) has 
mixed cinnamon and sooty-black feathering; this may also be a "retarded" 
adult. Another molting female (December) with more sooty-black feathering 
and less cinnamon feathering is also con.sidered to be a "retarded" adult. In 
fully adult birds there is considerable individual variation, especially in the 
males where the amount of black on the throat, the extent of the black on the 
terminal part of the primaries, and the extent of the black on the basal part of 
the tail feathers is variable. Scattered white feathers may be present on adult 
females. 

Remarks. — Hombron and Jacquinot first obtained the Truk Mon- 
arch, but it was not until the time of Kubary and of the Japanese 



272 University of Kansas Publs., Mus. Nat. Hist. 

collectors of Owston that very much was learned concerning the 
bird. In 1945, McElroy of the NAMRU2 party reported that he 
found no birds at the several islands of Truk that he visited in 
December. Some of the Japanese residents of the islands told 
McElroy that they did not know the bird. Evidently, its numbers 
are low or it has been eliminated, at least on the islands then popu- 
lated by the Japanese. 

Metabolus belongs to a group of flycatchers including the genera 
Pomarea, Mayrornis, Neolalage, Monarcha, and Clytorhynchus. 
The different plumages of the adults and the immatures are not 
unusual in this group of genera, this feature being observed in 
many of the flycatchers of Oceania. Mayr (1933c :1) points out 
some of the relationships between Metabolus and some of these 
other genera; he comments that all of them have rather thin bills, 
in contrast to those of other flycatchers. 

Metabolus became established at Truk probably as the result of 
an independent colonization. It is a well differentiated genus show- 
ing little resemblance to Monarcha godeffroyi of Yap. In looking 
over the genera found in the Pacific area, it appears that Metabolus 
is closest to Clytorhynchus of the Melanesian region, especially to 
Clytorhynchus hamlini Mayr, which is resident at Rennell in the 
Solomon Islands. The bills of these two birds are similar, both 
being long and thin, with a pronounced hook. In coloration there 
is some resemblance; C. hamlini has the blackish forehead and chin 
like the male Metabolus and also the burnt-orange underparts. In 
C. hamlini, however, the sexes are similar, Metabolus also resembles 
C. nigrogularis. Like Metabolus, the immatures of this latter spe- 
cies are different in color from the adults. 

Monarcha godefifroyi Hartlaub 
Yap Monarch 

Monarcha godeffroyi Hartlaub, Proc. Zool. Soc. London, 1867 (1868), p. 829, pi. 
38. (Type locality. Yap.) 

Monarcha godeffroyi Finsch, Joiirn. Mus. Godeffroy, 8, 1875, p. 50 (Yap) ; Sharpe, 
Cat. Birds British Mus., 4, 1879, p. 432 (Yap); Nehrkorn, Journ. f. Ornith., 1879, 
p. 403 (Yap) ; Bolau, Mitteil. Naturhist. Mus. Hamburg, 1898, p. 56 (Yap) ; Dubois, 
Syn. Avium, 1, 1902, p. 289 (Yap); Reichenow, Die Vogel, 2, 1914, p. 261 (Yap); 
Mayr, Birds Southwest Pacific, 1945, p. 295 (Yap). 

Monarcha godeffroyi Gray, Hand-list Birds, 1, 1869, p. 321 (Yap); Hand-list 
Japanese Birds, rev., 1932, p. 175 (Yap) ; Hand-li.st Jajianese Birds, 3d ed., 1942, p. 
194 (Yap). 

Monarches godeffroyi Hartlaub and Finsch, Proc. Zool. Soc. London. 1872, pp. 89, 
97 (Yap); Graffe, Journ. Mus. Godeffroy, 2, 1873, p. 123 (Yap); Schmeltz and Krause, 
Ethnogr. Abth. Mus. Godeffroy, 1881, p. 391 (Yap); Kuroda, in Momiyama, Birds 
Micronesia, 1922, p. 63 (Yap). 



Baker: The Avifauna of Micronesia 273 

Pomarea godeffrot/i Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 
1890-1891 (1891), p. 19 (Yap); Matschie, Journ. f. Ornith., 1901, p. 112 (Yap). 

Manarcharses geojjroyii Mathews, Bull. British Ornith. Club, 45, 1925, p. 94 (new 
generic name) ; idem, Syst. Avium Australasianarum, 2, 19.30, p. 514 (Yap). 

Monarcharses godeffroyi Takatsukasa and Yamashina, Dobutsu. Zasshi, 43, 1931, 
p. 486 (Yap?). 

Geographic ran^e.— Micronesia: Caroline Islands — Yap. 

Characters. — Adult male: according to Sharpe (1879:432) "General colour 
above white, from the hind neck to the rump and including scapulars; wings 
black, the quills browner; upper tail-coverts and tail black; head all around 
black, including the lower throat; sides of neck and rest of under surface, 
from the foreneck downwards, pure white; thighs and under tail-coverts black; 
under wing-coverts black, quills ashy blackish below; white along the inner 
edge of the primaries; 'bill entirely blue; feet whitish blue; iris black' (Kubary 
M. S.)." 

Adult female: ''Entirely black, excepting the hind neck and upper mantle, 
sides of neck, lower throat, and fore neck, which are pure white" (Sharpe, 
1879:432). 

Immature : "Above brown, the head and hind neck ashy grey, the scapulars 
rufescent at the tips, the rump rufous, becoming paler and more fulvous on 
the upper tail-coverts; wing-coverts dusky brown, broadly edged externally 
with rufous-buff, becoming fulvous on the median and greater coverts; quills 
dark brown, externally edged with rufous, the primaries narrowly, the sec- 
ondaries more broadly, the innermost of the latter edged and tipped with buff; 
tail-feathers ashy brown, narrowly edged with ochraceous brown and tipped 
with white, more broadly on the outer feathers; lores and a broad eyebrow 
rufous-buff; ear-coverts rather deeper rufous, shading on to the sides of the 
throat; under surface of body light cinnamon-rufous inclining to rufous on 
the throat and under tail-coverts; under wing-coverts light cinnamon, like 
the breast; quills light brown below, whitish along the inner web; 'bill horn- 
colour, the point brown, under mandible paler, feet dirty white, iris black' 
(Kubary M. S.)-" (Sharpe, 1879:433). 

Remarks. — No specimens of this species have been seen by me. 
Most taxonomists have regardeci this bird as a member of the genus 
Monarcha, although Mathews did propose the name Monarcharses 
for this bird. On the basis of descriptions and pictures (especially 
plate 38 in Hartlaub, 1868:828) the bird appears to be related to the 
monarch flycatchers of the Melanesian area. It may be closest to 
Monarcha menckei from the Bismarcks, M. manadensis of the New 
Guinea region, M. barbatus from the Solomons or to M. leucurus 
from Buru. The drab color of the immatures and the black and 
white color of the adults are characteristics of the Yap Monarch 
which are shared with some of the other species of Monarcha. The 
connection between M. godeffroyi and Metabolus rugensis of Truk is 
not known, but they evidently represent separate colonizations. M. 
takatsukasae of Tinian appears to be an offshoot of M. godeffroyi of 
18—8131 



274 University of Kansas Publs., Mrs. Nat. Hist. 

Yap, in which the black and white plumage has been suppressed (or 
never developed). As indicated by the published descriptions, the 
immature of M. godeffroyi shows a close resemblance to the adult of 
M. takatsukasae. The latter also shows relationships to immature 
specimens of M. leucotis and to M. giittula of Melanesia. 

The relationship of the two species of Monarcha in Micronesia to 
the Hawaiian Flycatcher, Chasiempsis sanchvichensis is not known. 
It is apparent that this Hawaiian form was derived from some 
ancestor from Melanesia, which arrived in the Hawaiian Islands by 
way of either Polynesia or Micronesia. Mayr (1943:45) has already 
pointed out that Chasiempsis is '"related to the Monarcha group 
{Pomarea, Mayrornis, etc.)." 

Monarcha takatsukasae (Yamashina) 
Tinian Monarch 

Monarcharses takatsukasae Yamashina, in Takatsukasa and Yamashina, Dobutsu. 
Zasshi, 43, 1931, p. 485. (Type locality, Tinian.) 

Monarcha takatsukasae Yamashina, Tori, 7, 1932, p. 400 (Tinian); Hand-list 
Japanese Birds, rev., 1932, p. 175 (Tinian); Hand-list Japanese Birds, 3d ed., 1942, 
p. 195 (Tinian); Mayr, Birds Southwest Pacific, 1945, p. 296 (Tinian); Downs, Trans. 
Kansas Acad. Sci., 49, 1946, p. 100 (Tinian). 

Geographic range. — Micronesia: Mariana Islands — Tinian. 

Characters. — Adult male: Forehead, lores, eyering, auriculars, and under- 
parts rufous, chin paler; under tail-coverts white; crown and nape dark slate- 
gray ; back reddish-brown ; rump white ; wing and tail dark brown, outer edges 
of first three primaries white, tail with white tips, more broadly tipped on outer 
tail feathers; outer edges of scapulars and secondaries buffy but tips more whit- 
ish, forming two wing bars; under wing-coverts whitish; bill slate-blue, tip 
pearl; feet dark slate; iris dark brown. 

Adult female: Resembles adult male, but slightly smaller and crown more 
brownish. 

Immature: Resembles adult, but base of bill lighter and underparts paler. 

According to the original description by Yamashina, M. takatsukasae resem- 
bles closely the immature M. godefjroyi of Yap in coloration; however, the 
Tinian Monarch has a shorter wing. 

Measurements. — Measurements are listed in table 41. 

Table 41. Measurements of Monarcha takatsukasae 



Number and Sex 


Wing 


Tail 


Full culmen 


Tarsus 


6 males 


70 

(67-72) 

67 
(65-68) 


68 
(65-70) 

67 
(64-69) 


18.0 
(17.5-19.0) 

17.2 
(17.0-19.0) 


22.0 


10 females 


(21.0-23.0) 
22.5 




(21.7-23.0) 



Baker: The Avifauna of Micronesia 275 

Specimens examined. — Total number, 20 (10 males, 10 females), as follows: Mariana 
Islands, USNM— Tinian, 10 (Oct.); AMNH— Tinian, 9 (Sept.); KMNH— Tinian, 1 (Sept.), 

Nesting. — Yamashina (1932a :400, 401) records two nests of the Tinian Mon- 
arch. He writes of one nest containing two eggs taken at Churo, Tinian, on 
January 29, 1932, that was "hung on a fork of an upward pointing branch of a 
road side tree commonly called 'Oba' 1.5 m. high from the ground in a forest. 
. . . The ground color of the egg shells is white. The spots are pale reddish- 
brown and distributed all round the surface like small dots, being concentrated 
especially round the larger end." Another nest containing three eggs was found 
on January 29, 1932. Yamashina writes that the eggs measure 20.5 x 15, 21 x 
15, and 18 x 15 mm. In describing these nests Yamashina notes, "The shape of 
the two nests mentioned above is like a deep cup. The outer layer of them is 
made chiefly of dead leaves, fibers, cotton, wools and moss, and the inner layer 
of fine stems and fibers only." 

Downs (1946:101) writes that a nest found near Lake Hagoi at Tinian on 
August 31, 1945, "was about three feet from the ground carefully woven into the 
framework of a triangular crotch. ... It was composed exteriorly of small 
leaves, scattered white feathers, and heavy grass; interiorly of grasses only." 
In the nest he found a young bird which "was black-skinned, with ugly white 
quills and a few short dark feathers on its tail and wings. The back feathers 
were rusty brown as were the tufted head feathers." Marshall (1949:219) 
assumes that this bird breeds all year. 

Molt. — Birds taken by Coultas in September are in fresh plumage. 

Remarks. — The Tinian Monarch is known only from Tinian, 
where it was (iescribcd in 1931 by Yamashina. Downs (1946:100- 
103) presents a detailed account of this bird as he saw it in 1945. He 
found it living in brushy woodlands where other birds, including 
Rhipidura rufifrons, were observed. From his description, the ac- 
tions and food-catching behaviors of this bird must be much like 
those of Rhipidtira. Gleisc (1945:220) estimated the population of 
these birds to be 40 to 50 in 1945. 

Myiagra oceanica erythrops Hartlaub and Finsch 

Micronesian Broadbill 

Myiagra erythrops Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, p. 6. (Type 
locality, Pelew Islands.) 

Myiagra erythrops Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 117, 
118 (Pelew Islands); idem, Proc. Zool. Soc. London, 1872, pp. 89, 97 (Pelew); Finsch, 
Journ. Mus. Godeffroy, 8, 1875, pp. 4, 20 (Palau) ; Giebel, Thes. Ornith., 2, 1875, p. 658 
(Pelew); Nehrkom, Journ. f. Omith., 1879, pp. 399, 403 (Palau); Sharpe, Cat. Birds 
British Mus., 4, 1879, p. 383 (Pelew); Schmeltz and Krause, Ethnogr. Abth. Mus. 
Godeffroy, 1881, p. 407 (Palau); Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, 
no. 6, 1890-1891 (1891), p. 23 (Pelew); Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, 
(3), 7, 1895, p. 195 (Palaos) ; Bolau, Mitteil. Naturhist. Mus. Hamburg, 1898, p. 55 
(Palau); Matschie, Journ. f. Omith., 1901, pp. 112, 113 (Palau); Dubois, Syn. 
Avium, 1, 1902, p. 283 (Pelew); Reichenow, Die Vogel, 2, 1914, p. 260 (Palau); 
Takatsukasa and Kuroda, Tori, 1, 1915, p. 54 (Pelew); Kuroda, in Momiyama, Birds 
Micronesia, 1922, p. 64 (Pelew); Yamashina, Tori, 10, 1940, p. 674 (Palau); Hand- 
list Japanese Birds, 3d ed., 1942, p. 195 (Babelthuap, Koror) ; Baker, Smithson. Misc. 
Coll., vol. 107, no. 15, 1948, p. 69 (Peleliu, Ngabad, Garakayo). 



276 



University of Kansas Publs., Mus. Nat. Hist. 



Suhmyiagra erythrops Mathews, Syst. Avium Australasianarum, 2, 1930, p. 504 
(Palau); Hand-list Japanese Birds, rev., 1932, p. 176 (Palau). 
Myiagra oceanica erythrops Mayr, Birds Southwest Pacific, 1945, p. 296 (Palau). 

Geographic range. — Micronesia: Palau Islands — Babelthuap, Koror, Gara- 
kayo, Peleliu, Ngabad. 

Characters. — Adult male: Crown, occiput, nape, and auriculars dark "slate- 
blue"; forehead, lores and orbital ring dark ''cinnamon-rufous"; black and 
upper wing-coverts olive-brown ; rump more like crown ; underparts near "cin- 
namon," paler on middle of abdomen, sides, and under tail-coverts; wings and 
tail dark brown, edged with white; secondaries edged with brownish; under 
wing-coverts whitish with dusky bases; bill and feet black. 

Adult female : Resembles adult male, but shghtly smaller and paler in color. 

Immature: Resembles adult, but head and rump browner; forehead, lores, 
and orbital ring sandy in some individuals, more rufous in others; underparts 
usually paler than in adult; bill basally lighter. 

Measurements. — Measurements are listed in table 42. 

Table 42. Measurements of Adult Specimens of Myiagra oceanica 



Subspecies 


Number 
and 

sex 


Wing 


Tail 


Full culmen 


Tarsus 


M. 


0. erythrops 


14 males 


69 

(68-71) 


53 

(51-56) 


16.4 
(16.0-17.3) 


19.5 

(18.5-20) 






11 females 


66 

(64-68) 


51 

(48-53) 


16.1 

(15.5-17.0) 


19.5 

(18.5-20) 


M. 


0. freycineti 


25 males 


70 

(67-73) 


60 

(57-64) 


16.3 

(15.8-17.0) 


19.5 

(18.5-20) 






16 females 


67 

(65-70) 


57 

(55-62) 


16.0 

(15.5-17.0) 


19.0 
(18.0-19) 


M. 


0. oceanica 


11 males 


81 

(78-83) 


68 

(65-71) 


20.1 
(19.5-20.5) 


20.0 
(19.5-21) 






10 females 


79 

(77-81) 


66 

(65-68) 


20.0 
(20.0-20.5) 


20.0 
(19-20.5) 


M 


0. pluto 


14 males 


82 
(79-83) 


74 

(71-77) 


17.5 

(17.5-18.0) 


19.0 






(18.5-20) 






14 females 


80 

(78-84) 


73 

(69-77) 


17.5 

(17.0-18.0) 


19.0 

(18.5-20) 



Specimens examined. — Total number, 33 (17 males, 15 females, 1 unse.xed), as follows r 
Palau Islands, USNM— Babelthuap, 1 (Nov. 27)— Koror, 4 (Nov. 6, 19, 26)— Garakayo, 1 
(Sept. 18) — Peleliu, 2 (Aug. 30) — Ngabad, 2 (Sept. 11); AMNH — exact locality not given, 
23 (Oct., Nov., Dec). 

Molt. — Molt apparently takes place in fall and early winter. Of the speci- 
mens examined, there is little evidence of molt in those obtained in August 



Baker: The Avifauna of Micronesia 277 

and September while there is considerably more evidence of molt in those 
taken in November and December. 

Food habits. — A bird taken by the writer on September 17. 1945, at Gara- 
kayo had approximately one-half cc. of insect parts in its stomach. 

Remarks. — The Micronesian Broadbill at Palau is a friendly 
little bird and easily called-up to within a few yards of a person 
by imitating its note. It was seen by the NAMRU2 party in 1945 
as singles and in pairs in the dense underbrush of the undisturbed 
forested areas. The bird was seen at only one woodland area at 
Peleliu (Southeastern Peninsula), but it was observed more fre- 
quently on the smaller islands of Ngabad and Garakayo. Coultas 
(field notes) also notes that in 1931 this bird was found more fre- 
quently on the smaller islands. Myiagra was found to be much less 
conspicuous at Palau than Rhipidura lepida. Myiagra appears to 
be less active, more solitary in its habits, and possibly more re- 
stricted in the territory that it covers in feeding than Rhipidura. 

Myiagra oceanica freycineti Oustalet 
Micronesian Broadbill 

Myiagra freycineti Oustalet, Bull. Soc. Philom. Paris, (T), 5, 1881, p. 73. (Type 
locality, Mariannes = Guam.) 

Myiagra freycineti Reichenow and Sohalnw, Journ. f. Ornith., 1884, p. 395 (Mari- 
annes = Guam) ; Oustalet, Le Nat., 1889, p. 260 (Mariannes r= Guam) ; Wiglesworth, 
Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 24 (Marianne — 
Guam); Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, p. 194 (Guam); 
Hartert, Novit. Zool., 5, 1898, p. 54 (Guam); Wheeler, Report Island of Guam, 1900, 
p. 13 (Guam); Matschie, Journ. f. Ornith., 1901, pp. 112, 113 (Guam); Seale, Occ. 
Papers Bernice P. Bishop Mus., 1, 1901, p. 50 (Guam); Safford, Osprey, 1902, p. 69 
(Guam); idem, Amer. Anthro., 4, 1902, p. 711 (Guam); idem. The Plant World, 7, 
1904, p. 263 (Guam); idem, Contr. U. S. Nat. Herb., 9, 1905, p. 79 (Guam); Reich- 
enow, Die Vogel, 2, 1914, p. 260 (Marianen); Cox, Island of Guam, 1917, p. 21 
(Guam); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 65 (Guam); Bryan, 
Guam Rec, vol. 13, no. 2, 1936, p. 25 (Guam) ; Hand-list Japanese Birds, 3d ed., 
1942, p. 195 (Guam); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 68 
(Guam). 

Submyiagra freycineti Mathews, Syst. Avium Australasianarum, 2. 1930, p. 504 
(Guam); Hand-list Japanese Birds, rev., 1932, p. 176 (Guam). 

Myiagra oceanica freycineti Mayr, Birds Southwest Pacific, 1945, p. 296 (Guam). 

Myiagra oceanica Strophlet, Auk. 1946, p. 539. (Guam). 

Geographic range. — Micronesia: Mariana Islands — Guam. 

Characters. — Adult male: A small flycatcher with head and neck near 
"dark delft blue" with a metallic luster; lores and anterior forehead ashy-gray, 
more bluish and darker on auriculars and sides of neck than on lores; back 
and upper wing-coverts near "green-blue .slate" but darker and with metallic 
luster less apparent than on head; rump grayer than back; chin and throat 
white; breast light "cinnamon." fading to pale buff and white on abdomen, 
sides, and under tail-coverts; tibia smoky-gray, tips of feathers paler; wings 
dark brown edged with light bluish-gray; tail bluish-slate, especially middle 



278 University of Kansas Publs., Mus. Nat. Hist. 

rectrices, tips of tail feathers edged with white; bill and feet black; iris dark 
brown. 

Adult female: Resembles adult female of M. o. erythrops, but crown and 
neck near "deep Payne's gray," auriculars grayer than neck; anterior forehead 
and lores buffy and tinged with cinnamon; back browner than lores with upper 
wing-coverts and scapulars edged with slightly lighter brown; rump resembles 
crown but grayer; underparts paler than those of M. o. erythrops, especially 
chin and throat; tibia more brownish. 

Immature male : Resembles adult male, but back more brown and less blue- 
green, lacking luster; anterior forehead more rufous; scapulars, upper wing- 
coverts, and wings edged with light brown; underparts variable but generally 
more buffy than those of adult. 

Immature female: Resembles adult female, but more brown and less blue 
on head and back; underparts more buffy; base of bill paler. 

Measurements. — Measurements are listed in table 42. 

Weights. — The author (1948:68) records the weights of five adult males as 
10.5-12.5 (11.9), and those of two adult females as 11.4 and 12.0 grams. 

Specimens examined. — Total number, 64 (33 males, 22 females, 9 unsexed), as follows: 
Mariana Islands, USNM— Guam, 26 (Jan. 21, March 16, May 21, 29, 30, June 1, 3, 14, 24, 26, 
July 10, 12, 13, 20, 23, Aug. 30); AMNH— Guam, 38 (Jan., Feb., March, July, Aug.). 

Nesting. — The writer (1948:68) records a nest containing one egg found by 
Muennink at Guam near Mt. Santa Rosa en May 7, 1945. The nest was in a 
bamboo stump approximately six feet from the ground. The egg hatched on 
about May 21. Seale (1901:50) reports on a nest and egg taken in the period 
from May to July. The NAMRU2 party obtained a female on March 15 with 
an enlarged gonad. Strophlet (1946:539) observed a pair of broadbills building 
a nest on September 20, 1945; it was completed on October 4 and was approxi- 
mately seven feet above the ground. Hartert (1898:33) reports on a nest taken 
at Guam on February 14, 1895. 

Molt. — As shown by the specimens examined, molt begins in June or July. 
Food habits. — The stomach of a bird obtained on January 21, 1945, contained 
one unidentified bug (Hemiptera) and several parts of other insects. 

Remarks. — The Micronesia Broacibill at Guam is not a common 
bird, and like its relative Rhipidura rufifrons is an inhabitant of for- 
ested areas, especially those containing brushy undercover. It is an 
active bird, although less conspicuous than Rhipidura. The birds 
were found as singles or in pairs. The pair of birds which had a nest 
at the west base of Mount Santa Rosa in May, 1945, allowed the ob- 
servers to approach closely to them. The birds are easily attracted 
by squeaking sounds. There is considerbale variation in the amount 
of cinnamon coloring on the breasts of adult birds. 

The Micronesian Broadbill at Guam was first discovered by Quoy 
and Gaimard, who called it "Moucherolle a gorge rouge." Kittlitz 
(1836:304) evidently records two species of flycatchers from Guam, 
which he calls Muscicapa. I judge these birds to be Myiagra and 
Rhipidura. It was not until 1881 that Oustalet recognized this bird 



Baker: The Avifauna of Micronesia 279 

to be new. The first large series of specimens was obtained by 
Marche for the Paris Museum and was reported on by Oustalet 
(1895:194). Marche collected 12 skins in August and September, 
1887, and 4 additional skins in February, 1889. 

Myiagra oceanica oceanica Pucheraa 
IVIicronesian Broadbill 

Myiagra oceanica Pucheran, Voy. Pole Sud, Zool., 3, 1853, p. 77. (Type locality, 
Hogoleu =: Truk.) 

Myiagra oceanica Hartlaub, Journ. f. Ornith., 1854, p. 168 (Carolinen =: Tnik) ; 
Gray, Cat. Birds Trop. Is. Pacific Ocean, 1859, p. IS (Hogoleu = Truk) ; Finsch and 
Hartlaub, Fauna Centralpolynesiens, 1867, p. 94 (Hogoleu = Truk) ; Gray, Hand-list 
Birds, 1, 1869, p. 328 (Caroline Is. = Truk); Pelzeln, Journ. f. Omith., 1875, p. 51 
(Hogoleu = Truk) ; Sharpe, Cat. Birds British Mus., 4, 1879, p. 383 (Hogoleu == 
Truk); Nehrkorn, Journ. f. Ornith., 1879, p. 403 (Ruk) ; Finsch, Proc. Zool. Soc. 
London, 1880, p. 575 (Ruk); Oustalet, Bull. Soc. Philom. Paris, (7), 5, 1881, p. 73 
(Carolines = Truk); Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, p. 
353 (Ruk); Reichenow and Schalow, Journ. f. Ornith., 1884, p. 395 (Carolines = 
Truk); Tristram, Cat. Birds, 1889, p. 200 (Ruk); Wiglesworth, Abhandl. und Ber. 
Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 23 (Ruk); Oustalet, Nouv. Arch. Mus. 
Hist. Nat. Paris, (3), 7, 1895, p. 196 (Hogoleu = Truk) ; Nehrkorn, Kat. Eiers., 1899, 
p. 30 (Ruk); Hartert, Novit. Zool., 7, 1900, p. 5 (Ruk); Matschie, Journ. f. Ornith., 
1901, pp. Ill, 112, 113 (Ruck); Dubois, Syn. .\vium, 1, 1902, p. 283 (Hogoleu = 
Truk); Reichenow, Die Vogel, 2, 1914, p. 260 (Karolinen = Truk) ; Takatsukasa and 
Kuroda, Tori, 1, 1915, p. 54 (Ruk); Wetmore, in Townsend and Wetmore, Bull. Mus. 
Comp. Zool., 63, 1919, p. 204 (Truk); Kuroda, in Momiyama, Birds Micronesia, 1922, 
p. 64 (Ruk); Hand-list Japanese Birds, 3d ed., 1942, p. 195 (Truk); Baker, Smithson. 
Misc. Coll., vol. 107, no. 15, 1948, p. 68 (Truk). 

Myiagra albiventris Finsch and Hartlaub, Fauna Centralpolynesiens, 1867, p. 93 
(Hoguleu = Truk); Giebel, Thes. Ornith., 2, 1875, p. 658 (Carolinae = Truk). 

Submyiagra oceanica Mathews, Syst. Avium .^ustralasianarum, 2, 1930, p. 505 
(Ruk); Hand-list Japanese Birds, rev., 1932, p. 175 (Truk). 

Myiagra oceanica oceanica Mayr, Birds Southwest Pacific, 1945, p. 296 (Truk) 

Geographic range. — Micronesia: Caroline Islands — Truk. 
Characters. — Adult male : Resembles M. o. jreydneti, but larger with crown 
and nape less green and with less metallic luster; lores and anterior forehead 
darker gray; chin, throat, and .sides of neck more buffy-cinnamon; back, rump, 
upper wing-coverts, and scapulars less blue and more ashy gray; tibia, wings, 
and tail more brownish. 

Adult female: Resembles adult male, but smaller with less blue and more 
gray on crown; lores and anterior forehead lighter- 
Immature: Resembles adult, but crown and nape grayish, slate-blue; under- 
parts paler. 

Measurements. — Measurements are listed in table 42. 

Specimens examined. — TotaJ number, 23 (12 males, 10 females, 1 unsexed), as follows: 
Caroline Islands, USNM— Truk, 2 (Feb. 16); AMNH— Truk, 21 (Feb., June, Nov., Dec). 

Nesting. — Hartert (1900:5) reports the taking of several nests in the period 
from March to July by Owston's Japanese collectors. One nest contained two 
eggs, the other nests contained one. 

Remarks. — The broadbill at Truk was first taken by Hombron 
and Jacquinot, who called it 'Tlatyrhynque oceanien." Later, Ku- 
bary obtained material which was studied by Finsch (1880e:575). 



280 University of Kansas Publs., Mus. Nat. Hist. 

In December, 1945, McElroy of the NAMRU2 party examined two 
adults with enlarged gonads. Specimens obtained by him at Truk 
were lost in shipment to the United States. In coloration this sub- 
species is closest to M. o. freycineti; in size it is closest to M. o. 
pluto. 

Myiagra oceanica pluto Finsch 

Micronesian Broadbill 

Myiagra pluto Finsch, Proc. Zool. Soc. London, 1875 (1876), p. 644. (Type ]<i- 
cality, Ponape.) 

Myiagra pluto Finsch, Journ. Mus. Godeffroy, 12, 1876, pp. 17, 19 (Ponape); 
idem, Proc. Zool. Soc. London, 1877 (1878), p. 779 (Ponape); Sharpe, Cat. Bird.s 
British Mus., 4, 1879, p. 380 (Ponape); Nehrkorn, Journ. f. Ornith. 1879, p. 4f.4 
(Ponape); Finsch, Journ. f. Ornith., 1880, p. 288 (Ponape); idem, Proc. Zool. Soc. 
London, 1880, p. 576 (Ponape); ide7n. Ibis, 1881, pp. 110, 112, 115 (Ponape); 
Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, p. 280 (Ponape); Wigles- 
worth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 2.3 (Po- 
nape); Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, p. 195 (Ponapi); 
Bolau, Mitteil. Naturhist. Mus. Hamburg, 1898, p. 55 (Ponape); Nehrkorn, Kat. 
Eiers., 1899, p. 26 (Ponape); Christian, The Caroline Islands. 1899, p. 358 (Ponape); 
Mat.schie, Journ. f. Ornith., 1901, pp. Ill, 112, 113 (Ponape); Dubois, Syn. Avium, 
1, 1902, p. 283 (Ponapi); Reichenow, Die Viigel, 2, 1914, p. 260 (Ponape); Taka- 
tsukasa and Kuroda, Tori, 1, 1915, p. 54 (Ponape) ; Wetmore, in Tovvnsend and 
Wetmore, Bull. Mus. Comp. Zool., 63, 1919, p. 204 (Ponape); Kuroda, in Momi- 
yama. Birds Micronesia, 1922, p. 64 (Ponape) ; Mayr, Proc. 6th Pacific Sci. Congr., 
4, 1941, p. 204 (Ponape); Hand-list Japanese Birds, 3d ed., 1942, p. 195 (Ponape). 

Submyiagra pluto Mathews, Syst. Avium Australasianaruin, 2, 1930, p. 505 (Po- 
nape); Yamashina, Tori, 1, 1932, p. 401 (Ponape); Hand-list Japanese Birds, rev., 
1932, p. 176 (Ponape). 

Myiagra oceanica pluto Mayr, Birds Southwest Pacific, 1945, p. 296 (Ponape). 

Geographic range. — Micronesia: Caroline Islands — Ponape. 

Characters. — Adult male: A dark, bluish-gray broadbill with head, ear- 
coverts, and nape dark, metallic, steel-blue; back and rump darker and more 
slate-blue than head ; upper tail-coverts blackish ; tail black edged with greenish 
gloss ; wings dark brown, scapulars and secondaries with outer edges tinged with 
metallic bluish-gray; lores black; chin, throat, and upper breast dark with 
light metallic-blue wash; lower breast and abdomen slate-gray; under wing- 
coverts brownish-black; bill black; feet bluish-black; iris dark brown. Female 
resembles male, but slightly smaller and somewhat duller. Immature duller. 

Measurements. — Measurements are listed in table 42. 

Specimens examined. — Total number, 42 (23 males, 19 females), as follows: Caroline 
Islands, USNM— Ponape, 3 (Feb. 11); AMNH— Ponape, 39 (Nov., Dec). 

Nesting. — Yamashina (1932a :401) records nests and eggs of the Ponape 
broadbill. The nests were at heights of between .9 and 2.2 meters above the 
ground. Nests, each containing a single egg, were obtained on July 21, 25, and 
August 6. The eggs measure 19.5 by 16, 20.5 by 15.7, 20.5 by 16. and 20.2 by 16. 
Coultas (field notes) describes the nest as a cup-shaped structure, made of fine 
gras.ses and ferns, and placed in small trees and bushes at low elevations. Of 
specimens taken by Coultas in November and December, 1931, approximately 
fifty percent of the males had enlarged gonads. According to his specimen 
labels, none of the females was in breeding condition. 

Molt. — Of the large series of broadbills taken by Coultas, approximately 



Baker: The Avifauna of Micronesia 281 

twenty percent of tliose taken in November were in molt whereas only approx- 
imately ten percent of those taken in December were in molt. Specimens taken 
in February were not in molt. It is evident that molting takes place in the fall, 
possibly from August to December. 

Remarks. — The culoration of the IMicronesian Broadbill at Ponape 
is in marked contrast to that of other representatives of Myiagra in 
Micronesia, being dark, bluish-gray in color. Probably the bird has 
taken on melanistic characters, which is not unusual in birds which 
have become isolated; examples of this condition may be observed 
in Rhipidura, Terpsiphone and other genera. 

Coultas (field notes) writes that the bird is "Common everywhere 
on the island except in the grasslands. Two birds are working to- 
gether usually, darting around in the low trees, among the branches 
or on the ground. The birds are playful, friendly and inquisitive. 
I should not call them noisy as one or more will sit for many minutes 
watching the intruder without making a peep. Their call, "Que 
Que," is a spasmodic outburst that might be repeated many times 
or just once. The male, presumably, erects the long crown feathers 
when calling. Perhaps both male and female do this, I can't say. 
The bird flutters on the wing and displays the feathers as does Rhip- 
idura. When sitting, the bird often erects the crest and fluffs the 
tail and feathers." 

Evolutionary History of Myiagra oceanica. — According to Mayr 
(1933d :1) Myiagra "is easily recognizable by its broad bill and the 
color pattern which is similar in all species." The range of the 
genus Myiagra extends from Australia and Tasmania westward to 
Timor, northward to the Moluccas, and Micronesia, and eastward 
to Polynesia. Myiagra oceanica is restricted to Micronesia and 
consists of four subspecies, which until recently have been consid- 
ered as four separate species. Unlike many of the species of this 
genus, M. oceanica shows comparatively little sexual dimorphism. 
The male of M. oceanica has metallic coloring on the head and the 
upper back and often has rich, rufous coloring on the breast. The 
female is less brilliant in coloring, lacking the sheen. The four sub- 
species varj' from each other in size, color and even, to some extent, 
in basal breadth of the bill. M. oceanica resembles several broad- 
bills, including M. galeata of the Moluccas, M. rubecula of Aus- 
tralia, M. vanikorensis of Fiji, and M. rufi.collis of Australia and the 
Lesser Sundas; however, in my opinion, it has probably been de- 
rived from M. galesta of the Moluccan area or from a closely re- 
lated species. In Micronesia, M. o. oceanica and M. o. freycineti 
appear to resemble closely this parent stock, whereas M. o. ery- 



282 University of Kansas Publs., Mus. Nat. Hist. 

throps and M. o. pluto are more differentiated but are considered 
to have been derived from this same colonization. M. o. pluto bears 
some resemblance to M. atra of the Papuan area, particularly in 
the dark coloring; this is probably only a parallel evolution, since 
they have little else in common. M. vanikorensis of the Fiji area 
is close to M. oceanica in color and structure; the two species, I 
suspect, have been derived from a common source rather than from 
each other. Study of the evolutionary history of the entire genus 
is necessary before we can understand fully the derivation of the 
Micronesian and Polynesian species. It seems safe to say that the 
center of dispersal has been in the Australian region; the lack of 
diversity of this genus within the Papuan area is at present unex- 
plained. 

Muscicapa narcissina narcissina Temminck 
Narcissus Flycatcher 

Muscicapa narcissina Temminck, PI. Col., 3, 1835, pi. 577, fig. 1. (Type locality, 
Japan.) 

Musicapa narcissina narcissina Mayr, Birds Southwest Pacific, 1945, p. 302 (Palau). 

Geographic range. — Breeds in eastern Asia and Japan. Winters south to 
Malaysia. In Micronesia: Palau Islands — exact locality unknown. 

Reinarks. — Mayr (1945a: 302) records the Narcissus Flycatcher 
as a migrant visitor to the Palau Islands on the basis of two speci- 
mens in the Turloff collection, formerly in the Zoological Museum 
in Hamburg. 

Musicapa griseisticta (Swinhoe) 

Chinese Gray-spotted Flycatcher 

Hemichelidon griseisticta Swinhole, Ibis, 1861, p. 330. (Type locality, Amoy.) 
Hemichelidon griseisticta Hand-list Japanese Birds, rev., 1932, p. 175 (Koror) ; 
Hand -list Japanese Birds, 3d ed., 1942, p. 194 (Koror). 

Muscicapa griseisticta Mayr, Birds Southwest Pacific, 1945, p. 302 (Palau) ; Mar- 
shall, Condor, vol. 51, 1949, p. 221 (Palau). 

Geographic range. — Breeds in northwestern Asia and Japan. Winters south 
to Malaysia. In Micronesia: Palau Islands — Koror. 

Remarks. — The Chinese Gray-spotted Flycatcher is a casual 
winter visitor to the Palaus. Marshall (1949:221) took two speci- 
mens at Palau on November, 1945. 

CoIIuricincIa tenebrosa (Hartlaub and Finsch) 
Palau Morning Bird 

Rectes tenebrosus Hartlaub and Finsch, Proc. Zool. Soc. London, 18fi8, p. 6. (Type 
locality, Pelew Islands.) 

Rectes tenebrosus Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, p. 113 
(Pelew Islands); idem, Proc. Zool. Soc. London, 1872, pp. 89, 99 (Pelew); Finsch, 
Journ Mus Godeffroy, 8, 1875, pp. 4, 18, pi. 3, fig. 1 (Palau); Schmeltz and Krause, 
Ethnogr. Abth. Mus. Godeffroy, 1881, p. 407 (Palau) ; Wiglesworth, Abhandl. und 
Bar. Zool. Mus. Dresden, no 6, 1890-1891 (1891), p. 27 (Pelew). 



Baker: The Avifauna of Micronesia 



283 



Colluricinda tenebrosa Gray, Hand-list Birds, 1, 1869, p. 386 (Pelew) ; Dubois, 
Syn. Avium, 1, 1902, p. 496 (Pelew); Mayr, Amer. Mas. Novit., no. 1269, 1944, p. 
5 (Palau) ; idem. Birds Southwest Pacific, 1945, p. 297 (Palau) ; Baker, Smithson. 
Misc. Coll., vol. 107, no. 15, 1948, p. 69 (Pelelilu, Ngabad, Garakayo). 

Pinarolestes tenebrosus Sharpe, Cat. Birds British Mus., 3, 1877, p. 298 (Pelew); 
Matschie, Joum. f. Ornith., 1901, p. 112 (Palau); Reichenow, Die Vogel, 2, 1914, p. 
296 (Palau); Takasukasa and Kuroda, Tori, 1, 1915, p. 54 (Pelew); Kuroda, in 
Momiyama, Birds Micronesia, 1922, p. 69 (Pelew) ; Hand-list Japanese Birds, rev., 
1932, p. 174 (Palau). 

Myiolestes tenebrosus Tristram, Cat. Birds, 1899, p. 188 (Pelew). 

Caleya tenebrosus Mathews, Syst. Avium Australasianarum, 2, 1930, p. 649 (Pelew). 

Malacolestes tenebrosus Mayr, Amer. Mus. Novit., no. 590, 1933, p. 5 (Palau); 
Hand-list Japanese Birds, 3d ed., 1942, p. 193 (Babelthuap, Koror, Peliliu). 

Geographic range. — Micronesia: Palau Islands — Babelthuap, Koror, Gara- 
kayo, Peleliu, Ngabad. 

Characters. — Adult : Upper parts between "snuff brown" and "bister," head 
blacker; chin, throat, and upper breast like upper parts but darker; lower 
breast and abdomen lighter and more buffy, sides darker; feathers of under- 
parts with darker shafts producing a streaked appearance; underside of wing 
and under tail-coverts light-colored; bill dark brown; feet lighter brown; 
iris yellowish. Female smaller. 

Immature: Resembles adult, but head and neck lighter; ear-coverts, sides 
of neck, throat, upper breast darker; lower breast and abdomen paler. 

Measurements. — Measurements are listed in table 43. 

Table 43. Me.asurements of Colluricinda tenebrosus 



Number and Sex 


Wing 


Tail 


Full culmen 


Tarsus 


20 males 


104 
(100-107) 

97 
(94-101) 


76 

(73-79) 

73 

(71-76) 


23.5 

(22.5-24.5) 

23.0 
(22.0-24.0) 


31 


9 females 


(29-31) 
30 




(30-31) 



Specimens examined. — ^Total number, 32 (21 males, 11 females), as follows: Palau Islands, 
USNM— Koror, 6 (Nov. 5, 18)— Garakayo, 3 (Sept. 18)— Peleliu, 5 (Aug. 29, 30, Sept. 1. 6) 
—Ngabad, 2 (Sept. 11); AMNH — exact locality not given, 16 (Oct. 8, 13, 26, Nov. 11, 13, 
17, 19, 21, 23, Dec. 9). 

Molt — The molting process in this species seemingly takes place from 
August until December. Most of the birds taken by the NAMRU2 party in 
August and September were in molt. Molting specimens were obtained by 
Coultas in October, November and December. 

Food habits. — The Palau Morning Bird feeds on plant and animal mate- 
rials. Stomachs obtained by the NAMRU2 party contained green plant ma- 
terial, seeds, insect parts, and grit. The bird feeds principally on the ground 
or in low bushes. 

Remarks. — The Palau Morning Bird is a thrushlike bird which 
spends its time on or near the ground in areas where ground cover 
is thick. In 1945, the NAMRU2 party found the bird in the thick 



284 University of Kansas Publs., Mus. Nat. Hist. 

matting of vines which had covered over the battle-cleared areas. 
I did not find the bird at elevations of more than three to four feet 
above the ground. When flushed, it would flutter a short distance 
and disappear into the brush. It has a sweet song and may be 
considered as one of the finest singers in Micronesia. It heralds 
the break of day with its melodious carol, and its name is derived 
from its calling early in the morning. I heard the bird only infre- 
quently in the hot part of the day, although it would sing when the 
skies were overcast. Its song could be heard also as evening ap- 
proached. The bird is moderately common, and evidently is more 
abundant on the smaller islands than on Peleliu. Its occurrence on 
the smaller islands was noted also by Coultas. 

The taxonomic status of the Palau Morning Bird has been one 
of uncertainty as shown by the fact that the bird has been treated 
under six generic names since its discovery by Captain Tetens. 
Mayr (1933a: 5) erected a new genus, Malacolestes, for the morn- 
ing bird pointing to its differences from "Rhectes (^ Pitohui) and 
Pinarolestes (=z Myiolestes)." Later, he (1944b:5) disregards this 
name and places the bird in the genus Collurcincla stating that its 
special characters "are due to isolation." This treatment is followed 
here. The Palau Morning Bird is the most northern representative 
of a group of birds which have their center of dispersal in the New 
Guinea and Australian area. As Mayr has pointed out, C. tene- 
brosus appears closest to the C. m^garhynchus group of New Guinea. 
These species have bills of similar shape, coloration which is darker 
above and lighter below, soft feathers on underparts, and streaked 
appearance of throat and breast. The resemblances between C. 
tenebrosus and C. megarhynchus might be such as to indicate that 
these are merely subspecifically distinct from each other. 

Artamus leucorhynchus pelewensis Finsch 
White-breasted Wood-swallow 

Artamus pelewensis Finsch, Journ. Mus. Gocleffroy, 12, 1876, p. 41. (Type locality, 

Palau.) 

Artamus leucorhynchus Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 
116, 118 (Pelew); idein, Proc. Zool. Soc. London, 1872, pp. 89, 99 (Pelew) ; Finsch, 
Journ. Mus. Godeffroy, 8, 1875, pp. 4, 18 (Palau); Walden, Ibis, 1876, p. 188 (Pelew;. 

Artamus pelewensis Finsch, Proc. Zool. Soc. London, 1877 (1878), p. 739 (Pelew); 
Tweeddale, Ibis, 1878, p. 385 (Pelew); Sharpe, Cat. Birds British Mus., 13, 1890, p. 
9 (Pelew); Wiglesworth, AVihandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 
(1891), p. 26 (Pelew); Bolau, Mitteil. Natuihist. Mus. Hamburg, 1898, p. 02 (Palau); 
Matschie. Journ. f. Ornith., 1901, p. 112 (Palau); Dubois, Syn. Avium, 1, 1902, p. 
533 (Pelew); Reichenow, Die Vogel, 2, 1914, p. 346 (Pelew). 

Artamus leucorhynchus pelewensis Stresemann, Novit. Zool., 20, 1913, p. 293 (Pa- 
lau); Hand-list Japanese Birds, 3d ed., 1942, p. 193 (Babelthuap, Koror) ; Mayr, 
Birds Southwest Pacific, 1945, p. 297 (Palau). 



Baker: The Avifauna of Micronesia 



285 



Artamus melanoleucus pelewensis Kuroda, in Momiyama, Birds Micronesia, 1922, 
p. 69 (Pelew); Mathews, Syst. Avium Australasianarum, 2, 1930, p. 635 (Pelew) ; 
Hand-list Japanese Birds, rev., 1932, p. 174 (Palau). 

Geoffraphic range. — Micronesia: Palau Islands — Babelthuap, Angaur. 
Characters. — ^Adult: Upper surface black, except for back which is slightly 
brownish and for rump which is white; underparts white, except for chin, 
throat and upper breast which are black; wings with grayish tips; bend of 
wing black; bill milky blue, nostril and tip black; feet black; iris dark brown- 
Immature: Resembles adult, but black feathers with brownish tinges; pri- 
maries tipped with white. 

Measurements. — Measurements are listed in table 44. 

Table 44. Me.asurements of Artarmis leucorhynchiLS -pelewensis Finsch 



Number and Sex 


Wing 


Tail 


Culmen 


Tarsus 


5 males 


134 
(132-136) 

134 
(132-136) 


68 
(66-69) 

68 

(67-69) 


25 

(24-26) 

24 


16.5 






4 females 


17.0 




(16.5-17.0) 









Specimens examined. — Total number, 12 (7 males, 5 females), from Palau Island.*, AMNH 
— exact locality not given (March, Nov., Dec). 

Remarks. — Little is known concerning the habits and distribution 
of the white-breasted Wood-Swallow at Palau. Coultas obtained 
a series of eight birds in 1931; he writes (field notes) that his native 
hunter took every bird that he saw. The natives told him that they 
did not know the nest of the bird. Coultas concluded that the bird 
was not common. He commented that it may be found perched 
in the top of a tree on a dead branch or "even displaying in the 
air." The NAMRU2 party found no evidence of this bird in the 
southern Palaus in 1945. The specimens obtained by Coultas in 
November and December, 1931, were in molt and had small gonads. 

This wood-swallow is the only Micronesian representative of 
Artamus. le.ucorhynchus. Like several other species of birds it has 
become established only at the Palau Islands, and has either been 
unsuccessful in colonizing other parts of Micronesia or has not had 
the opportunity to do so. This bird had been compared with speci- 
mens representing ten subspecies of A. leucorhynchus in Melanesia 
and Malaysia. A. I. pelewensis differs from these subspecies ex- 
amined by its blacker appearance, with only a faint brownish wash 
on the back, and by its shorter, first primary. The curvature of the 
upper mandible of the bird in the Palaus is similar to that of P. I. 
leucorhynchus of the Philippines; the mandible is less curved than 



286 University of Kansas Publs., Mus, Nat. Hist. 

that of P. I. celebensis of Celebes; the mandible is slightly thicker 
than that of P. I. leucopygialis of the New Guinea and Australian 
region. In length of wing P. I. pelewensis resembles closely P. I. 
leucorhynchus ; P. I. celebensis has a longer wing and P. I. leuco- 
pygialis has a shorter one. Stresemann (1913:293) points to a close 
relationship between P. I. pelewensis and P. I. musschenbreeki of 
Tenimber and Babber islands and P. I. melaleucus of New Cale- 
donia; Mayr (1945a: 284) says the bird in the Palaus came from 
the Papuan area. Probably A. I. pelewensis has reached the Palau 
Islands from the New Guinea area by way of the Philippines. 

Aplonis opacus opacus (Kittlitz) 

Micronesian Starling 

Lamprothlornis] opaca Kittlitz. Kupfertaf. Naturgesch. Vogel, 2, 1833, p. 11, 
pi. 15, fig. 2. (Type locality, Ualan = Kusaie.) 

Turdus colombinus Lesson (part), Traite d'Ornith., 1832, p. 406 (Carolines- = 
Kusaie?). 

Lamprothlornis] opaca Kittlitz, Mem. Acad. Imp. Sci. St. Petersbourg, 2, 1935, 
p. 7 (Ualan); idem (part), Obser. Zool., in Lutke, Voy. "Le Seniavine," 3, 1836, pp. 
285, 297 (Ualan); Pelzeln, Reise "Novara," Vogel, 1865, p. 68 (Ualan). 

Lamprotomis columbinus Bonaparte (part), Consp. Avium, 1, 1850, p. 417 (Caro- 
linen = Kusaie?). 

Lamprotomis columbina Hartlaub, Archiv f. Naturgesch., 18, 1852, p. 133 (Ualan); 
idem (part), Joum. f. Ornith., 1854, p. 168 (Carolinen = Kusaie?) ; Kittlitz, Denkw. 
Reise russ. Amer. Micron, und Kamchat., 1, 1858, p. 376 (Ualan). 

Calornis opaca Gray (part), Cat. Birds Trop. Is. Pacific Ocean, 1859, p. 26 (Oua- 
lau=Kusaie); Tristram, Cat. Birds, 1889, p. 255 (Kusai) ; Hartert, Kat, Vogelsamml. 
Senckenb., 1891, p. 75 (Ualan). 

Calornis kittlitzi Finsch and Hartlaub (part). Fauna Central polynesiens, 1867, p. 
109 (Ualan, Puynipet, Marianen; type locality, by subsequent restriction, Ualan = 
Kusaie) ; Finsch (part), Journ. Mus. Godeffroy, 8, 1875, p. 23 (Ualan). 

Calornis kittlitzii Hartlaub, Proc. Zool. Soc. London, 1807 (1868), p. 830 (Ualan). 

Amadina Kittlitzi Gray, Hand-list Birds, 2, 1870, p. 58 (Ualan). 

Calornis pacifica Sharpe, Ibis, 1876, p. 47 (Caroline Is.=Kusaie?) ; Finsch (part), 
Mitth. Ornith. Ver. Wien, 1884, p. 49 (Kuschai). 

Calornis pacificus Finsch (part), Journ. Mus. Godeffroy, 12, 1876, p. 32 (Ualan) ; 
irfem (part), Journ. f. Ornith., 1880, pp. 289, 301 (Kuschai); idem, (part), Proc. 
Zool. Soc. London, 1880, p. 576 (Kuschai) ; idem, (part), Ibis, 1881, pp. 103, 104, 
108, 111 (Kuschai). 

Aplonis kittlitzi Sharpe (part). Cat. Birds British Mus., 13, 1890, p. 136 (Kuschai); 
Wiglesworth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), 
p. 44 (Ualan); Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, p. 
216 (Oualan); Hartert (part), Novit. Zool., 5, 1898, p. 59 (Ualan); Matschie (part), 
Journ. f. Ornith., 1901, p. 112 (Ualan); Takatsukasa and Kuroda, Tori, 1, 1915, p. 
64 (Kusaie). 

Lamprocorax kittlitzi Dubois (part), Syn. Avium, 1, 1902, p. 542 (Kuschai). 

Aplonis opaca Oberholser, Bull. U. S. Nat. Mus., 98, 1917, p. 59 (Ualan); Wetmore 
(part), in Townsend and Wetmore, Bull. Mus. Comp. Zool., 63, 1919, p. 219 (Kusaie). 

Aplonis kittlitzi kittlitzi Momiyama (part). Tori, 2, 1920, p. 1 (Kusaie). 

Aplonis opaca opaca Momiyama (part). Birds Micronesia, 1922, pp. 6, 12 (Kusaie); 
Kuroda (part), in Momiyama, Birds Micronesia, 1922, p. 70 (Kusaie); Mathews, Syst. 
Avium Australasianarum, 2, 1930, p. 847 (Kusaie); Takatsukasa and Yamashina, Tori, 
7, 1931, p. 109 (Kusaie); Hand-list Japanese Birds, rev., 1932, p. 170 (Kusaie). 

Aplomis opaca opaca Hand-list Japanese Birds, 3d ed., 1942, p. 188 (Kusaie). 

Aplonis opacus cfpacus Mayr. Birds Southwest Pacific, 1945, p. 298 (Kusaie). 



Baker: The Avifauna of Micronesia 



287 



Geographic range. — Micronesia: Caroline Islands — Kusaie. 

Characters. — Adult: Feathers black with dusky appearance caused by 
lighter bases; edges of feathers with slight amount of steel-green gloss; under- 
parts slightly duller than upper parts; bill black, with maxilla rather strongly 
curved; feet black, iris yellow. Females slightly smaller. 

Immature: Resembles adult, but upper parts more brown and less black; 
underparts dusky with edges of feathers tinged with smoky yellow producing 
a streaked appearance; base of bill horn-colored. 

Measurements. — Measurements are listed in table 45. 

Specimens examined. — Total number, 30 (18 males, 12 females), as follows: Caroline Is- 
lands, USNM— Kusaie, 5 (Feb. 8); AMNH— Kusaie, 25 (Jan., Feb., March). 

Remarks. — The Micronesian Starling at Kusaie was first taken 
by Kittlitz (1833:11) , who named it in the following manner: "Tur- 
dus columbinus Gm. L. 0(der Lamproth. opaca Lichenstein." The 
bird was later given the name of Calornis kittlitzi by Finsch and 
Hartlaub (1867:109). Oberholser (1917:59) has shown that the 
specific name opaca is applicable, since the manuscript name Lam- 





Table 45. Measurements of j 


\dtjlt Specimens of 


Aplonis opacus 


Subspecies 


Number 
and 
sex 


Wing 


Tail 


Full 
culmen 


Depth of 

culmen at 

nostril 


A 


0. opacus 


15 males 


124 
121-125 


80 

76-85 


24 
24-26 


9 5 






9.0-10.0 






12 females 


119 
115-125 


77 
72-82 


24 
23-26 


9.0 
8.5-9.0 


A. 


0. ponapensis 


17 males 


133 

130-138 


87 
85-91 


27 
26-29 


9.5 
9.0-10.0 






11 females 


126 
122-127 


83 
81-85 


27 
26-28 


9.0 
8.5-9.0 


A. 


0. anaus 


16 males 


129 
125-131 


88 
84-92 


28 
27-29 


95 






8.0-9.0 






7 females 


124 
121-129 


85 
83-88 


27 
25-28 


8.5 
8.0-9.0 


A. 


0. orii 


11 males 


128 
124-131 


86 
83-90 


27 
25-28 


75 






7.5-8.5 






7 females 


124 
121-126 


79 

77-82 


26 
25-27 


7.5 
7.5-8.0 


A. 


0. guami 


41 males 


128 
120-136 


86 
81-92 


27 
24-29 


9.5 






8.5-10.5 






32 females 


121 
117-126 


84 
78-89 


26 
24-30 


9.5 

8.5-10.5 



288 University of Kansas Publs., Mus. Nat. Hist. 

prothornis opaca of Lichtenstein is made available by Kittlitz's pub- 
lished description and figure, and since it is the earliest name used. 
Mathews (1938:342) reports that the name Aplomis appeared a 
few days before the name Aplonis. I have been unable to check his 
source of information. 

The Micronesia Starling is one of the most abundant birds at 
Kusaie. Coultas (field notes) observed the bird in all parts of the 
island, when he visited there in 1931. He found the bird in flocks 
of two to six or more and noted that birds in immature plumage 
seemed to outnumber the birds in adult plumage approximately five 
to one. This subspecies is characterized by the presence of only 
a slight amount of gloss on the black feathers of the adult. 

Aplonis opacus ponapensis Takatsukasa and Yamashina 
Micronesian Starling 

Aplonis opaca ponapensis Takatsukasa and Yamashina, Tori, 7, 1931, p. 109. (Type 
locality, Ponape.) 

Calonm columbina Pelzeln, Reise "Novara," Vogel, 1865, pp. 88, 162 (Puynipet;. 

Calomis kittlitzi Finsch and Hartlaub (part). Fauna Centralpolynesiens, 1867, p. 
109 (Puynipet); Schmeltz and Krause (part). Ethnogr. Abth. Mus. G'odeffroy, 1881, 
p. 298 (Ponape). 

Calomis opaca Gray (part). Hand-list Birds, 2, 1870, p. 27 (Seniavin = Ponape). 

Calomis pacificvs Finsch (part), .Journ. Mus. Godeffroy, 12, 1876, pp. 17, 32 (Po- 
nape); idem, Proc. Zool. Soc. London, 1877 (1878), p. 779 (Ponape); idem (part), 
Journ. f. Ornith., 1880, p. 289 (Ponape); idem, (part), Proc. Zool. Soc. London, 1880, 
p. 576 (Ponape). 

Calornis pacifica Finsch, Ibis, 1881, p. 115 (Ponape); idem, (part), Mitth. Ornith. 
Ver. Wien. 1884, p. 49 (Ponape). 

Aplonis kittlitzi Sharpe (part), Cat. Birds British Mus., 13, 1890, p. 136 (Ponape); 
Wiglesworth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891). 
p. 44 (Ponape); Bolau (part), Mitteil. Naturhist. Mus. Hamburg, 1898, p. 62 (Po- 
ape) ; Nehrkorn, Kat. Eiers., 1899, p. 122 (Ponape); Matschie (part), Journ. f. 
Ornith., 1901, p. 112 (Ponape); Takatsukasa and Kuroda (part). Tori, 1, 1915, p. 
55 (Ponape). 

Lamprocorax kittlitzi Dubois (part), Syn. Avium, 1, 1902, p. 542 (Ponape). 

Aplonis opaca Wetmore (part), in Townsend and Wetmore, Bull. Mus. Comp. 
Zoiil., 63, 1919, p. 219 (Ponape); Mayr. Proc. 6th Pacific Sci. Congr., 4, 1941, p. 
204 (Ponape). 

Aplonis kittlitzi kittlitzi Momiyama (part). Tori, 2, 1920, p. 1 (Ponape). 

Aplonis opaca opaca Momiyama (part). Birds Micronesia, 1922, p. 12 (Ponape); 
Kuroda (part), in Momiyama, Birds Micronesia, 1922, p. 70 (Ponape). 

Aplonis opaca ponapensis Yamashina, Tori, 7, 1932, p. 394 (Ponape); Hand-list 
Japanese Birds, rev., 1932, p. 170 (Ponape). 

Aplonis opaca ponapensis Hand-list Japanese Birds, 3d ed., 1942, p. 188 (Ponape). 

Aplonis opacus ponapensis Mayr. Birds Southwest Pacific, 1945. p. 297 (Ponape). 

Geographic range. — Micronesia: Caroline Islands— Ponape. 

Characters. — Adult: Resembles A. o. opacus. but larger with a longer bill 
and richer gieen luster on the back and breast. 

Immature: Resembles immature of A. o. opacus, but underparts more 
brightly streaked but still dingy in appearance. 



Baker: The Avifauna of Micronesia 289 

Measurements. — Measurements are listed in table 45. 

Specimens examined. — Total number, 47 (31 males, 16 females), as follows: Caroline Is- 
lands, USNM— Ponape, 1 (Feb. 11); AMNH— Ponape, 46 (Nov., Dec). 

Nesting. — Yamashina (1932a :394) reports the taking of an egg on August 
2, 1931, and two eggs on August 30, 1931, at Ponape. Coultas (field notes) 
writes that the nests of these birds are hidden in the tops of the tree-ferns 
and in holes in the trees. The natives told him that the starling lays two eggs. 

Molt. — Most of the adult specimens taken by Coultas in November and 
December, 1931, are in molting plumage. 

Remarks. — Coultas (field notes) writes that the starling is a com- 
mon bird at Ponape. He found it in flocks of from two to 12 or more 
birds. As at Kusaie he noted more birds in the immature plumage 
than in the adult plumage at Ponape. The starling occurs in large 
numbers even though the people of the island hunt this bird persist- 
ently for part of their food supply. 

The Micronesian Starling at Palau has the longest wing of any of 
the subspecies of Aplonis opacus. It most closely resembles A. o. 
opacus; both of these subspecies have only a faint amount of bronzy- 
green luster of the feathers, and the immatures have dingy yellow 
streaks on the abdomen. 

Aplonis opacus angus Momiyama 
Micronesian Starling 

Aplonis opaca anga Momiyama, Birds Micronesia, 1922, p. 6. (Type locality, 
Toroas, Ruk Island.) 

Lamprothlornis] opaca Kittlitz (part), Observ. Zool., in Lutke, Voy. "Le Seniavine," 
3, 1836, p. 297 (Lougounor = Lukunor). 

Lamprotornis columbinus Bonaparte (part), Consp. Avium, 1, 1850, p. 417 (Caro- 
linen =: Lukunor?). 

Lamprotornis columbina Hartlaub (part), Journ. f. Ornith., 1854, p. 168 (Carolinen 
= Lukunor?). 

Calomis kittlitzi Hartlaub and Finsch (part), Proc. Zool. Soc. London, 1872, pp. 89, 
100 (Mackenzie = Ulithi?); Finsch (part), Journ. Mus. Godeffroy, 8, 1875, p. 23 
(Mackenzie = Ulithi?); Schmeltz and Krause (part), Ethnogr. Abth. Mus. Godeffroy, 
1881, pp. 298, 330, 353 (Mortlock, Nukuor, Ruk). 

Calomis pacificus Finsch (part), Journ. Mus. Godeffroy, 8, 1875, p. 23 (Mackenzie 
= Ulithi?); idem (part), Journ. f. Ornith., 1880, p. 290 (Ruck, Mortlocks) ; idem 
(part), Proc. Zool. Soc. London, 1880, p. 576 (Ruk); idem (part). Ibis, 1881, p. Ill 
(Ruk). 

Calomis pacifica Finsch (part), Mitth. Ornith. Ver. Wien, 1884, p. 49 (Rukgruppe). 

Aplonis kittlitzi Sharpe (part). Cat. Birds British Mus., 13, 1890, p. 136 (Ruk, 
Lugunor); Wiglesworth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890- 
1891 (1891), p. 44 (Ruk or Luganor, Nukuor); Oustalet (part), Nouv. Arch. Mus. Hist. 
Nat. Paris, (3), 7, 1895, p. 216 (Ruk, Nukuor, Luganor); Hartert (part), Novit. Zool., 
5, 1898, p. 59 (Ruk, Luganor); idem, Novit. Zool., 7, 1900, p. 6 (Ruk); Matschie 
(part), Journ. f. Ornith., 1901, p. 112 (Ruck); Takatsukasa and Kuroda (part), Tori, 1, 
1915, p. 55 (Ruk). 

Lamprocorax kittlitzi Dubois (part), Syn. Avium, 1, 1902, p. 542 (Ruk, Luganor). 

Aplonis opaca Wetmore (part), in Townsend and Wetmore, Bull. Mus. Comp. Zool., 
63, 1919, p. 219 (Truk). 

Aplonis kittlitzi kittlitzi Momiyama (part). Tori, 2, 1920, p. 1 (Truk, Wolea). 

19—8131 



290 University of Kansas Publs., Mus. Nat. Hist. 

Aplonis opaca anga Kuroda, in Moniiyama, Birds Micronesia, 1922, p. 71 (?Luganor 
or Ruk, ?Nukuor, Wolea or Oleai) ; Takatsukasa and Yamashina, Tori, 32, 1930, p. 109 
(Ruk) ; Mathews, Syst. Avium Australasianarum, 2, 1930, p. 847 (Ruk) ; Hand-lisi 
Japanese Birds, rev., 1932, p. 170 (Uluthi, Feys, Wolea, Ifalik, Faraulep, Lamotrek, 
Truk, Nukuoro). 

Aplornis apaca anga Takatsukasa and Yamashina, Dobutsu. Zasshi, 43, 1931, p. 458 
(Truk?); Hand-list Japanese Birds, 3d ed., 1942, p. 188 (Uluthi, Feys, Wolea, Ifalik, 
Faraulep, Lamotrek, Truk, Nukuoro). 

Aplonis opacus angus Mayr, Birds Southwest Pacific, 1945, p. 297 (Truk and west- 
ern Carolines) ; Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, pp. 70, 71 (Ulithi 
Truk). 

Geographic range. — Micronesia: Caroline Islands — Ulithi, Fais, Wolea, Ifalik, 
Faraulep, Lamotrek, Truk, Nukuoro, Lukunor. 

Characters. — Adult: Resembles A. o. opacus, but larger and with bill less 
deep and feathers with distinct greenish luster both on the upper parts and the 
lower parts. Female smaller. 

Immature: Resembles immature of A. o. opacus, but underparts streaked 
with brighter, buffy-yellow coloring. 

Measurements. — Measurements are listed in table 45. 

Specimens examined. — Total number, 38 (24 males, 14 females), as follows: Caroline 
Islands, USNM— Ulithi, 27 (Aug. 15, 16, 19, 20, 21, 22)— Truk, 2 (Feb. 16, Dec. 13); AMNH 
—Truk, 9 (Jan. 29, Feb. 1, 28, June 14, Oct. 9, 14). 

Nesting. — Hartert (1900:6) reports that at Truk nests of the starling were 
obtained by Owston's Japanese collectors from May to July and one in March. 
Nests contained from one to three eggs each. 

Molt. — Adult birds taken by the NAMRU2 party at Ulithi in August are in 
molting plumage. 

Food habits. — The stomachs of starlings obtained in August at Ulithi con- 
tained pieces of fruit and seeds. Twelve stomachs contained between one and 
three cc. of these foods. Papaya and small berries were the foods most fre- 
quently observed in the stomachs. 

Remarks. — The Micronesian Starling of the central and western 
Carolines is one of the few land birds which lives on both the "high" 
islands and the "low" coral islands in Micronesia. It is found on 
several of the coral atolls in the Carolines. In the Hand-list of Japa- 
nese Birds (Hachisuka et al, 1932:170), the birds at Ulithi and Fais 
are placed in the subspecies A. o. angus, although these islands are 
only a short distance from Yap, at which place another subspecies, 
A. o. kurodai, occurs. Specimens from Yap are not available for 
comparison. Specimens from Ulithi and from Truk closely resemble 
one another. 

The NAMRU2 party found the starling to be numerous at Truk 
and at Ulithi in 1945. At both places the natives make use of the 
birds as food. At Truk, McElroy found a larger number of birds in 
immature plumage than that of birds in adult plumage. Similar 
observations have been made at several other islands in Micronesia. 

At Ulithi, the NAMRU2 party found the starling at all islands in 



Baker: The Avifauna of Micronesia 291 

the atoll visited in 1945. The bird was more numerous at the islands 
of Potangeras and Mangejang, and less numerous at the island of 
Losiep; the former two islands were occupied — at the time of the 
visit in 1945 — by service personnel and the vegetation was disturbed, 
whereas Losiep was uninhabited and rarely visited by people. I 
attribute the smaller population of starlings at Losiep to the fact that 
on this island the large monitor lizard, Varanus indicus, was numer- 
ous while at Potangeras and Mangejang it was apparently entirely 
absent. These large lizards depend principally on the birds, rodents, 
and insects for their food supply. At Potangeras the rat Rattus 
exulans was exceedingly numerous, while at Losiep no sign of rodents 
was found nor were any taken in traps set during the daytime. 

Aplonis opacus kurodai Momiyama 
Micronesian Starling 

Aplonis kittlitzi kurodai Momiyama, Tori, 2, 1920, p. 1. (Type locality, Yap.) 

Calomis kittlitzi Hartlaub and Finsch (part), Proc. Zool. Sec. London, 1872, p. 
100 (Uap); Griiffe, Journ. Mus. Godeffroy, 2, 1873, p. 123 (Yap); Finsch (part), 
Journ. Mus. Godeffroy, 8, 1875, pp. 5, 24 (Yap); Schnieltz and Krause (part), Eth- 
nogr. Abth. Mus. Godeffroy, 1881, p. 298 (Yap). 

Calomis pacificus Finsch (part), Journ. Mus. Godeffroy, 12, 1876, p. 32 (Yap). 

Aplonis kittlitzi Wiglesworth, Ahhandl. und Ber. Zool. Mus. Dresden, no. 6, 1800- 
1891 (1891), p. 44 (Yap); Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 
7, 1895, p. 216 (Yap); Hartert (part), Novit. Zool., 5, 1898, p. 58 (Yap); Bolau 
(part), Mitteil. Naturhist. Mus. Hamburg, 1898, p. 62 (Yap); Matsrliie (part), Journ. 
f. Ornith., 49, 1901, p. 112 (Yap); Takatsukasa and Kuroda (part). Tori, 1, 1915, 
p. 64 (Yap). 

Aplonis opaca kurodai Momiyama, Birds Micronesia, 1922, p. 11 (Yap); Kuroda, 
in Momiyama, Birds Micronesia, 1922, p. 71 (Yap); Mathews, Syst. Avium Austral- 
asianarum, 2, 1930, p. 848 (Yap); Hand-list Japanese Birds, rev., 1932, p. 170 (Yap). 

Aplo7iis opaca kurodai Takatsukasa and Yamashina, Dobutsu, Zasshi, 43, 1931, p. 
458 (Yap?); Hand-list Japanese Birds, 3d ed., 1942, p. 188 (Yap). 

Aplonis opacus kurodai Mayr, Birds Southwest Pacific, 1945, p. 297 (Yap); Baker, 
Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 71 (Yap). 

Geographic range. — Micronesia: Caroline Islands — Yap. 

Characters. — Adult: According to Momiyama (1922:11), "Similar to A. o. 
anga from Ruk group, but the bill thicker (9-10.5 mm.; that of the latter 8.5- 
9.5 mm.) and much longer (24-27.5 mm.; that of the latter 21.5-25 mm.) and 
the wing also longer in average (119-5-130 mm. instead of 116.5-129.5 mm.). 
It differs from typical opaca by the edge of feathers of both body .sides very 
distinctly tinged with a bronzy-green lustre, by the bill being longer and 
thicker (in typical opaca exposed culmen 21.5-24.5 mm., depth of bill 9-9.5 
mm.)." 

Immature: "Similar to the immature of the typical form, but both sides 
of body somewhat deeper in colour and the edge of feathers distinctly tinged 
with lustrous bronzy-green. It differs from the same stage of A. o. anga by 
the under-parts being without pale-yellowish area." Momiyama (1922:11). 

Young: "Similar to the young of typical bird, but differs from it by the 
mantle being very faintly tinged with bronzy-green and by the under-parts 



292 University of Kansas Publs., Mus. Nat. Hist. 

being somewhat tinged with brown. In the same stage of the typical form, 
the under-parts are much more greyish-ashy in colour." Momiyama (1922:11). 

Remarks. — No specimens have been examined. Momiyama 
(1920:1) regarded the birds at Yap and at Saipan as A. o. kurodai. 
Later (1922:10) he separated the birds at Saipan as A. o. harterti, 
remarking that the birds from Saipan differ ''from A. o. kurodai 
Momiyama from Yap islands, by the green lustre on both sides of 
body being less distinct and showing tendency to a purplish lustre, 
by the bill being decidedly shorter, and by the same thickness." 

Price (1936a :19) describes a method by which starlings and other 
birds are captured by the natives of Yap. The natives make slashes 
in the trunk of a breadfruit tree and allow the exuding juice to 
harden. This material is then chewed until soft and adhesive. It 
is then placed on a stick which has been secured directly under a 
papaya fruit. When the birds alight on this perch, they become 
stuck and are captured. 

Aplonis opacus orii (Takatsukasa and Yamashina) 
Micronesian Starling 

Aplornis opaca orii Takatsukasa and Yamashina, Dobutsu. Zasshi, 43, 1931, p. 4.58. 
(Type locality, Coror, Pelew Islands.) 

Calornis kittlitzii Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 7, 117, 
118 (Pelew). 

Calornis opaca Gray (part). Hand-list Birds, 2, 1870, p. 27 (Pelew). 

Calornis kittlitzi Hartlaub and Finsch (part), Proc. Zool. Soc. London, 1872, p. 89 
(Pelew); Finsch (part), Journ. Mus. G'odeffroy, 8, 1875, pp. 5, 23 (Palau) ; Schnieltz 
and Krause (part), Ethnogr. Abth. Mus. Godeffroy, 1881, p. 298 (Palau). 

Calornis kittlitzi Kubary, Journ. Mus. Godeffroy, 4, 1873, p. 225 (Palau-Inseln). 

Calornis pacificus Finsch (part), Journ. Mus. Godeffroy, 12, 1876, pp. 17, 32 (Pa- 
lau); idem (part), Journ. f. Ornith., 1880, p. 289 (Palau); idem (part), Proc. Zool. 
Soc. London, 1880, p. 576 (Palau); idem (part). Ibis. 1881, p. Ill (Pelew). 

Aplonis kittlitzi Wiglesworth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 
1890-1891 (1891), p. 44 (Pelew); Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, 
(3), 7, 1895, p. 212 (Palaos) ; Hartert (part), Novit. Zool., 5, 1898, p. 58 (Pelew); 
Matschie (part), Journ. f. Ornith., 1901, p. 112 (Palau); Takatsukasa and Kuroda 
(part). Tori, 1, 1915, p. 55 (Pelew). 

Aplonis opaca subsp nov.? Momiyama, Birds Micronesia, 1922, p. 13 (Pelew); Ku- 
roda, in Momiyama, Birds Micronesia, 1922, p. 72 (Pelew). 

Aplornis opaca orii Hand-list Japanese Birds, 3d ed., 1942, p. 188 (Babelthuap, 
Koror, Peliliu, Anguar). 

Aplonis opaca orii Hand-list Japanese Birds, rev., 1932, p. 169 (Palau); Yamashina, 
Tori, 10, 1940, p. 673 (Palau). 

Aplonis opacus orii Mayr, Birds Southwest Pacific, 1945, p. 297 (Palau); Baker, 
Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 71 (Peleliu, Ngesebus, Garakayo). 

Geographic range. — Micronesia: Palau Islands — Kayangel, Babelthuap, 
Koror, Garakayo, Ngesebus, Peleliu, Ngabad, Angaur. 

Characters. — Adult: Resembles adult of A. o. opacus, but slightly larger with 
bill longer and shallower, and feathers with distinct greenish gloss both on the 
upper parts and the lowerparts. Resembles A. o. angus in the amount of green- 
ish gloss on feathers, but bill shallower. Depth of bill of A. o. opacus measures, 



Baker: The Avifauna of Micronesia 293 

on the average, 9.5 for males and 9.0 for females; of ^. o. angus 8.5 for both 
males and females; of A. o. orii 7.5 for both males and females. 

Immature: Resembles immature of A. o. angus, but streaking on underparts 
duller. 

Measurements. — Measurements are listed in table 45. 

Specimens examned. — Total number, 40 (21 males, 19 females), as follows: Palau Islands, 
USNM— Koror, 3 (Nov. 6)— Garakayo, 2 (Sept. 19)— Ngesebus, 1 (Sept. 20)— Peleliu, 7 
(Aug. 28, 29, 30, 31, Sept. 5); AMNH — exact locality not given, 27 (Oct., Nov., Dec). 

Molt. — Many of the specimens taken in August and September show evi- 
dence of molt; most of the specimens taken in October, November and 
December are not in molt. 

Remarks. — The amount of greenish gloss on the feathers of ^. o. 
orii and A. o. angus appears to be the same, but the streaked under- 
parts of the immature of ^. o. orii are duller than those of the im- 
mature of A. 0. angus. The shallower bill in the Palau starling is 
caused by the lower edge of the mandible being generally straighter 
than that in A. o. angus and A. o. opacus. In comparing A. o. orii 
with A. o. kurodai, Takatsukasa and Yamashina (1931a :458) state 
that "the greenish gloss is less pronounced and of a duller shade than 
that of A. 0. kurodai Momiyama." 

The starling is probably the most abundant land bird in the 
Palaus. It was found as singles or in small flocks at all islands vis- 
ited by the NAMRU2 party in 1945. As at the other islands of 
Micronesia, the starling at Palau is noisy and conspicuous. It is a 
most inquisitive bird, often following the collector through the wood- 
lands. Apparently the starling prefers the open woodlands and mar- 
ginal areas to the thicker jungles; as a result of clearing operations 
during the war, the bird probably has increased. The starling is 
primarily a vegetarian; I found no animal matter in stomachs exam- 
ined at Palau or at Ulithi or Guam. At Palau, as at other islands, 
more of the starlings seen were in immature plumage than in adult 
plumage. Coultas (field notes) found the birds to be abundant at 
Koror and highly prized as food by the natives and Japanese. He 
writes, "It is surprising what a fine wholesome meal certain people 
can get out of handful of rice and a starling's breast." 

Aplonis opacus guami Momiyama 

Micronesia Starling 

Aplonis opaca guami Momiyama, Birds Micronesia, 1922, p. 9. (Type locality, 
Guam). 

Turdm columbinus Lesson (part), Traite d'Ornith., 1831, p. 406 (Mariannes = 
Guam). 

Lamprothlomis] opaca Kittlitz (part), Kupfertaf. Naturgesch. Vogel, 2, 1833, p. 11, 
pi. 15, fig. 2 (Marianen = Guam) ; idem (part), Obser. Zool., in Lutke, Voy. "Le 
Seniavine," 3, 1836, pp. 298, 304 (Guahan). 



294 University of Kansas Publs., Mus. Nat. Hist. 

Lamprotomis columbinus Bonaparte (part), Consp. Avium, 1, 1850, p. 417 (Mariann. 
= Guam). 

Lamprotomis columbina Hartlaub (part), Journ. f. Omith., 1854, p. 167 (Mariannen 
= Guam); Kittlitz, Denkw. Reise russ. Amer. Micron, und Kamchat., 1, 1858, pp. 367, 
376 (Guaham). 

Calornis opaca Gray (part), Cat. Birds Trop. Is. Pacific Ocean, 1859, p. 26 (La- 
drone or Marian Is.); idem, (part), Hand-list Birds, 2, 1870, p. 27 (Ladrone = 
Guam?). 

Calornis kittlitzi Finsch and Hartlaub (part). Fauna Centralpolynesiens, 1867, p. 
109 (Marianen = Guam?); Oustalet, Le. Nat., 1889, p. 261 (Mariannes). 

Calornis columbina Giebel (part), Thes. Ornith., 2, 1875, p. 427 (Marianae = 
Guam?). 

Calornis pacificus Finsch (part), Journ. Mus. Godeffroy, 12, 1876, pp. 17, 32 
Marianne). 

Aplonis kittlitzi Wiglesworth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 
1890-1891 (1891), p. 44 (Marianne; Oustalet (part), Nouv. Arch. Mus. Hist. Nat. 
Paris, (3), 7, 1895. p. 212 (Guam, Saypan); Hartert (part), Novit. Zool., 5, 1898, 
p. 58 (Guam, Saipan); Wheeler, Report Island of Guam, 1900, p. 13 (Guam); Seale, 
Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 54 (Marianas); Matschie, Journ. f. 
Ornith., 1901, p. 112 (Guam); Safford, Osprey, 1902, p. 69 (Guam); idem. The 
Plant World, 7, 1904, p. 264 (Guam); idem, Contr. U. S. Nat. Herb., 9, 1905, p. 79 
(Guam); Mearns, Proc. U. S. Nat. Mus., 36, 1909, p. 477 (Guam); Takatsukasa and 
Kuroda (part). Tori. 1, 1915, p. 64 (Marianas); Cox, Island of Guam, 1917, p. 21 
(Guam); Bryan, Guam Rec, vol. 13, no. 2, 1936, p. 25 (Guam). 

Aplonis opaca Wetmore (part), in Townsend and Wetmore, Bull. Mus. Comp. Zool., 
63, 1919, p. 219 (Guam). 

Aplonis kittlitzi kurodai Momiyama, Tori, 2, 1920, p. (Saipan). 

Aplonis opaca giiami Kuroda, in Momiyama, Birds Micronesia, 1922, p. 71 (Guam); 
Mathews, Syst. Avium Australasianarum, 2, 1930, p. 847 (Guam); Yamashina, Tori, 
7, 1932, p. 394 (Saipan, Rota); Hand-li.st Japanese Birds, rev., 1932, p. 169 (Guam, 
Rota, Tinian, Saipan). 

Aplonis opaca harterti Momiyama (part). Birds Micronesia, 1922, p. 10 (Type lo- 
cality, Saipan); Kuroda (part), in Momiyama, Birds Micronesia, 1922, p. 71 (Saipan); 
Mathews, Syst. Avium Australasianarum, 2, 1930, p. 847 (Saipan). 

Aplomis opaca harterti Takatsukasa and Yamashina, Dobutsu. Zasshi, 43, 1931, p. 
487 (Saipan). 

Aplomis opaca guami Takatsukasa and Yamashina, Dobutsu. Zasshi, 44, 1932, p. 221 
(Tinian, Rota); Hand-list Japanese Birds, 3d ed., 1942, p. 188 (Saipan, Tinian, Rota, 
Guam). 

Aplonis opacus guami Mayr, Birds Southwest Pacific, 1945, p. 297 (Guam, Rota, 
Tinian, Saipan); Watson, The Raven, 17, 1946, p. 41 (Guam); Downs, Trans. Kansas 
Acad. Sci., 49, 1946, p. 103 (Tinian); Stott, Auk, 1947, p. 527 (Saipan, Guam); 
Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 69 (Guam, Rota, Tinian, 
Saipan). 

Aplonis opacus Wharton, Ecol. Monogr., 16, 1946, p. 174 (Guam); Strophlet, Auk, 
1946, p. 540 (Guam); Baker, Condor, 49, 1947, p. 125 (Guam). 

Geographic range. — Micronesia: Mariana Islands — Guam, Rota, Tinian, 
Saipan. 

Characters. — Adult: Resembles closely A. o. angus in the amount of green- 
ish gloss present on the body feathers, but with slightly shorter and deeper 
bill. 

Immature: Resembles the immature of A. o. angxis but streaks on under- 
parts brighter and less-dingy yellow. 

Measurements. — Measurements are listed in table 45. The writer (1948:69) 
has given average measurements for the length of wing of adult males from 
Guam as 127, from Rota as 122, from Tinian as 131, and from Saipan as 131 ; 



Baker: The Avifauna of Micronesia 295 

for depth of bill of adult males from Guam as 9.0, from Rota as 9.0, from 
Tinian as 9.5, and from Saipan as 10.0. 

Weights. — The NAMRU2 party obtained weights of six adult males from 
Guam as 84-96 (87) ; of eight adult females from Guam as 78-108 (86) ; of two 
Juvenal males from Guam as 88 and 90; of five juvenal females from Guam 
as 77-87 (80) ; of two adult males from Rota as 70 and 83; and of five juvenal 
males from Rota as 64-80 (76). 

Specimens examined. — Total number, 95 (55 males, 37 females, 3 unsexed), as follows: 
Mariana Islands, USNM— Guam, 44 (Jan. 21, 22, Feb. 5, March 8, 13, April 12, May 18, 
22, 24, 27, 29, 30, June 3, 4, 6, 14, 16, 18, July 6, 7, 14, 20, Aug. 24, Oct. 8, Nov. 19, 23) 
—Rota, 12 (Oct. 18, 19, 26, 27, Nov. 2)— Tinian, 4 (Oct. 12, 18); AMNH— Guam, 16 
(Jan. 23, 24, 29, March 3, 12, 13, 24, May, Aug. 12, Nov. 23, 28, Dec. 26)— Tinian, 15 
(Sept. 7, 8, 10, 11, 12)— Saipan, 4 (July 9, 17, Aug. 26, Sept. 2). 

Nesting. — The NAMRU2 party found evidence of nesting by starlings at 
Guam as early as January 28, in 1945. On this date a bird was seen to carry 
food into a hollow tree at Oca Point. Signs of nesting activities were observed 
in the months that followed, the last record being obtained on June 11. Star- 
lings nest in cavities in trees, in holes in rocky cliffs, and probably in 
the tops of coconut palms. On June 2 a nest was found by Muennink in a 
cavity of a banyan tree at Oca Point, Guam. The nest was appro.ximately 12 
feet from the ground and consisted of a flattened mass of green foliage at the 
bottom of the cavity. Two eggs found in the nest have been described by the 
author (1948:69) as "Niagara green" with scattered, irregular spots of color, 
near "russet," "Mars brown" and "pallid purple-drab," most abundant near the 
large ends. Measurements are 32.1 by 22.1 and 32.0 by 22.4. 

Yamashina ( 1932a :394) records two eggs taken at Saipan on April 14, 1931; 
two eggs taken at Rota on March 10, 1931 ; and one egg taken at Rota on 
March 11, 1931. Seale (1901:54) writes that the starling nests in a hole in the 
dead trunk of the coconut palm and may lay three or four egg.« Hartert 
(1898:59) reports that two eggs were taken at Guam on March 11 

Food habits. — Probably the chief food of the starling at Guam is the fruit 
and seeds of the papaya. This plant grows in most parts of the island, espe- 
cially in the lowlands where land uses have disturbed the climax vegetation. 
Many of the garden plots lay fallow during the war and were allowed to grow 
up in thick stands of papaya. As a fruit began to ripen, the starlings would 
peck out one side of a ripe fruit, feeding on the tissues and the seeds. It was 
seldom that a fully ripe papaya fruit was found that had not been at least partly 
eaten by the starlings. Apparently the birds do not feed on the fruit before it 
is fully ripened. Seeds of other types of vegetation were also eaten by the birds. 

Parasites. — Wharton (1946:174) records the chigger (Acarina), Tromhicvla 
sp., from the starling at Guam. 

Remarks. — According to Oustalet (1895:212), the starling was 
taken in the Marianas by the expedition in the "Uranie" in 1820 and 
by the expedition in the "Astrolabe" in 1829. Kittlitz, who visited 
Guam from March 1-20, 1828, also recorded the starling. It was not 
until 1922, however, that the starling in the Marianas was recognized 
as subspecifically distinct from the birds in the Carolines and Palaus. 



296 University of Kansas Publs., Mus. Nat. Hist. 

The Japanese ornithologists named the bird at Guam as A. o. guami 
and the bird at Saipan as ^. o. harterti, but later regarded these as 
a single subspecies A. o. guami. Momiyama (1920:2) had, pre- 
viously to the naming of the new forms in the Marianas, considered 
the bird at Saipan as belonging to the same subspecies as that found 
at Yap. Among named kinds, A. o. guami found at Guam, Rota, 
Tinian, and Saipan appears to be most closely related to A. o. angus. 
These two subspecies differ in that the streaking of the underparts in 
the immatures is brighter in A. o. guami and duller in ^. o. angus. 
The bird at Saipan has a longer wing and a deeper bill than the bird 
at Guam ; however, birds at Tinian show intermediate measurements. 

At Guam, the starling is the most numerous land bird. The writer 
(1947b: 124), in counting birds along the roadways of Guam, re- 
corded the starling on all of the 125 counts and found the birds to 
include more than one-half (57.3 percent) of all the birds seen. 
Starlings may have increased during the years of the war, with the 
disruption of normal agricultural activities allowing the growth of 
papaya and other food plants in fallow areas; however, the use of 
the birds as food by the islanders probably increased during the war. 

As at other islands in Micronesia, the numbers of birds in imma- 
ture plumage at Guam seemingly exceeds the number of birds in 
adult plumage. Animals which may prey on the starling at Guam 
include the feral house cat, Rattus mindanensis, Corvus kubaryi, and 
the large lizard Varanus indicus. The starling spends little time on 
the ground; it feeds principally in the trees, which might limit the 
amount of damage done to it by the feral house cats which are nu- 
merous on the island. The rat, R. mindanensis, is a semi-arboreal 
animal and may feed on eggs and young birds in nest cavities of trees 
or on cliffs. The crow, C. kubaryi, has a reputation for stealing 
chicken eggs from poultiy yards and may prey on the eggs and 
young of the starling. The monitor lizard, V. indicus, is known to 
prey on the starling, as well as on the domestic chickens at farm 
houses. On January 31, 1945, one of these large lizards was seen 
descending a tree after robbing a nest of a starling; one of the star- 
ling's eggs was seen in the mouth of the lizard. The noise and com- 
motion set up by the parent birds and by other starlings, which had 
been attracted to the area, did not appear to perturb the uninvited 
guest. 

Downs (1946:103) writes that the starling at Tinian is less com- 
mon than the white-eye, Zosterops conspicillata saypani. Gleise 
(1945:220) estimated the population of starlings on Tinian at 200. 



Baker: The Avifauna of Micronesia 297 

Coultas (field notes) found the starling abundant at Tinian in 1931, 
but he did not find the bird at Saipan. According to Stott (1947: 
527), the starling was abundant at Guam but "appeared to be com- 
mon only locally on Saipan." He saw large flocks at the Marpi 
Point and Kingman Point areas on Saipan but found the bird less 
numerous elsewhere on the island. At Rota, the NAMRU2 party 
found the birds to be numerous and widely distributed over the island 
in 1945. 

At Guam, the present writer observed behavior of the starling on 
January 31, 1945, which may have been a courtship ceremony. Two 
adults were perched on a palm frond approximately 20 feet above the 
ground. The bird which was perched more distally on the frond 
opened its tail fan-fashion, spread its wings and at irregular intervals 
picked up in its beak a part of the frond and then released it. As 
this behavior was taking place, the birds would call in a sweet ascend- 
ing song, which reminded me very much of the song of the red- 
wing blackbird of North America. This was indeed a contrast to 
the usual squawking notes of this subspecies. 

Aplonis opacus aeneus (Takatsukasa and Yamashina) 
Micronesian Starling 

Aplornis opaca aenea Takatsukasa and Yamashina, Dobutsu. Zasshi, 43, 1931, p. 
487. (Type locality, Pagan.) 

Aplonis kittlitzi Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, 
p. 212 (Pagan, Agrigan). 

Aplonis opaca harterti Momiyama (part). Birds Micronesia, 1922, p. 11 (Pagan, 
Agiigan); Kuroda (part), in Momiyama, Birds Micronesia, 1922, p. 71 (Pagan, Agri- 
gan). 

Aplornis opaca aenea Hand-list Japanese Birds, 3d ed., 1942, p. 187 (Asongsong 
= Asuncion, Agrigan, Pagan, Almagan). 

Aplornis opaca aenea Takatsukasa and Yamashina, Dobutsu. Zasshi, 44, 1932, p. 
221 (Pagan, Almagan). 

Aplonis opaca aenea Hand-list Japanese Birds, lev., 1932, p. 169 (Agrigan, Pagan, 
Almagan); Yamashina, Tori. 10, 1940, p. 673 (Asongsong). 

Aplonis opacus aeneus Mayr, Birds Southewest Pacific, 1945, p. 297 (Agrigan, 
Pagan, Almagan); Borror, Auk, 64, 1947, p. 417 (Agrihan). 

Geographic range. — Micronesia: Mariana Islands — Alamagan, Pagan, Agri- 
han, Asuncion. 

Characters.— kdnW.: According to Takatsukasa and Yamashina (1931:487), 
A. o. aeneus resembles A. o. orii of Palau, but has a bronze rather than green 
luster. A. o. aeneus resembles A. o. opacus, but has a smaller bill. 

Remarks. — No specimens of this subspecies have been examined 
by me. Little information is available regarding the occurrence of 
this subspecies in the northern Marianas. Oustalet (1895:212) writes 
that Marche collected four specimens at Pagan and three at Agri- 
han. Borror (1947:417) writes that in 1945, it was a "common and 



298 University of Kansas Publs., Mus. Nat. Hist. 

abundant species" at Agrihan. He obtained one specimen between 
July 27 and August 14 and comments that it had a grasshopper in 
its stomach. 

Evolutionary history of Aplonis opacus. — Aplonis opacus is known 
from the Mariana, Palau, and Caroline islands in Micronesia. It 
consists of several subspecies, which have relatively few distinguish- 
ing characteristics. No starlings are known in the Marshall and 
Gilbert islands, although atolls occur in these island-chains that 
offer a habitat approximately the same as those in the western 
Carolines now occupied by A. o. angus. 

In regard to parental stock, Sharpe (1876:47) considered A. 
opacus as "nothing but a slightly more metallic race of C. mysolen- 
sis, with a still stouter bill." The species with which Sharpe com- 
pared A. opacus is known from My sol, Buru, and Ceram. Oustalet 
(1896:70) thought that the Aplonis in Micronesia belonged to a 
group of starlings whose members are scattered through the Pacific 
islands including Cook, Samoa, Tonga, Fiji, New Britain, New 
Guinea, Banta, Mysol, Salwatti, and Timor. Mayr (1941b: 204) is 
of the opinion that Aplonis in Micronesia was derived from central 
Polynesia. Amadon (1943:8), in his study of the genera of star- 
lings, places A. opacus within a superspecies containing A. cineras- 
cens, A. tabuensis, A. juscus, and possibly A. feadensis and A. can- 
toroides. All of these are blackish birds with greenish gloss with 
immatures having the underparts streaked. In comparing A. opacus 
with these mentioned species and with other species of Aplonis, I 
find that A. opacus more closely resembles A. feadensis and A. 
cantoroides than any others. Although there are differences in size 
of the bill, wing, and tail, these structures are proportionally the 
same. The streaked underparts of the immatures of A. cantoroides 
are much like that of the immatures of A. opacus, whereas the im- 
matures of A. feadensis are only faintly streaked with whitish below. 
The eye of A. cantroides is red, and that of A. opacus is more nearly 
yellow. The ancestral stock from which A. opacus developed in 
Micronesia seemingly reached the area from Melanesia. In Micro- 
nesia the birds dispersed to various groups of islands from some 
point in the Caroline Islands. The birds are absent from the Mar- 
shall Islands. Perhaps the birds never reached the Marshall Islands 
or they may have been present in former times and disappeared 
since then. 



Baker: The Avifauna of Micronesia 



299 



Aplonis pelzelni Finsch 
Ponape Mountain Starling 

Aplonis pelzelni Finsch, Proc. Zool. Soc. London, 1875 (1876), p. 644. (Type lo- 
cality, Ponape.) 

Aplonis pelzelni Finsch, Journ. Mus. Godeffroy, 12, 1876, pp. 17, 32, pi. 2, fig. 3 
(Ponape); idem, Proc. Zool. Soc. London, 1877 (1878), p. 779 (Ponape); idem, 
Journ, f. Ornith., 1880, p. 290 (Ponape); idem. Ibis, 1881, pp. 110, 112, 115 (Po- 
nape); Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, p. 281 (Ponape); 
Sharpe, Cat. Birds British Mus., 13, 1890, p. 136 (Ponape); Wiglesworth, Abhandl. 
und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 43 (Ponape); Oustalet, 
Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, p. 215 (Ponapi) ; Bolau, Mitteil. 
Naturhist. Mus. Hamburg, 1898, p. 62 (Ponape'); Matscliie, Journ. f. Ornith., 1901, 
pp. Ill, 112 (Ponape); Dubois, Syn. Avium, 1, 1902, p. 542 (Ponape); Reichenow, 
Die Vogel, 2, 1914, p. 355 (Ponape); Takatsukasa and Kuroda, Tori, 1, 1915, p. 64 
(Ponape); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 70 (Ponape); Mathews, 
Syst. Avium Australasianarum, 2, 1930, p. 849 (Ponape); Hand-list Japanese Birds, 
rev., 1932, p. 170 (Ponape); Bequaert, Mushi, 12, 1939, p. 82 (Ponape); Mayr, 
Proc. 6th Pacific Sci. Congr., 4, 1941, pp. 204, 213 (Ponape); Bequaert, Occ. Papers 
Bemice P. Bishop Mus., 16, 1941, p. 290 (Ponape); Mayr. Birds Southwest Pacific, 
1945, p. 298 (Ponape). 

Aplornis pelzelni Hand-List Japanese Birds, 3d ed., 1942, p. 189 (Ponape). 

Geographic range. — Micronesia: Caroline Islands — Ponape. 

Characters. — Adult: A small, dark starling with upper parts sooty-brown, 
darker on head with forehead and lores blackish ; wings, rump, upper tail- 
coverts, and tail lighter and more brownish than head; underparts paler and 
washed with olive-brown; bill and feet black; iris brown. 

Immature : Resembles adult, but lighter brown, especially the underparts. 

A. pelzelni diflfers from A. ojxicus by having no gloss on the feathers, smaller 
size, more slender bill, and a brown iris. 

Measurements. — Measurements are listed in table 46. 

Specimens examined. — Total number, 59 (32 males, 24 females, 3 unsexed), from Caroline 
Islands, AMNH — Ponape (Dec). 

Nesting. — Coultas (field notes) obtained reports that the Ponape Mountain 
Starling nests in cavities in trees and lays two eggs. 



Table 46 


. Me.'^surements of 


Aplonis pelzelni 




Number and Sex 


Wing 


Tail 


Exposed 
culmen 


Depth of 
bill a; 
nostril 


Tarsus 


10 adult males 


103 
101-105 

99 
97-102 


65 
63-67 

61 
57-64 


20.0 
19.0-21.0 

19.5 
19.5-20.5 


6.5 
6.0-7.0 

6.0 
6.0-6.5 


27 


10 adult females 


26-28 

27 
26-27 



Parasz7es.— Bequaert (1939:82 and 1941:290) records the fly (Hippoboscidae), 
Ornithoica ptisilla, from A. pelzelni. 



300 University of Kansas Publs., Mus. Nat. Hist. 

Remarks. — Coultas (field notes) writes that "the Mountain Star- 
ling is a bird of the true mountain forest. ... I did not record 
it below 1,400 feet. Natives tell me that the Mountain Starling for- 
merly covered the whole of the island and that now some individuals 
can be found on the low atoll of Ant, to the westward of Ponape. 
Unfortunately, I was not permitted to visit either Ant or Pakin." 
Coultas notes also that the birds are quiet and usually travel in 
pairs. They are easily attracted by squeaking the lips against the 
hand or by the cries of a wounded bird. Many of these starlings 
were taken in fruit trees. Coultas describes the call of A. pelzelni as 
"weaker and finer" than that of A. opacus. These two species may 
be found together, according to Coultas, but A. opacus is apparently 
the more aggressive and often drives A. pelzelni away. Richards 
{in litt.) found this bird to be "very rare" while on his visit to Po- 
nape in 1947-1948. He observed two individuals on January 15, 
1948, at an elevation of approximately 600 or 700 feet. A male was 
taken. 

Evolutionary history of Aplonis pelzelni. — The Ponape Mountain 
Starling is a distinctive bird which evidently represents an ancient 
and single colonization of Micronesia. It lacks the green gloss which 
is found on many of the other starlings of the Pacific region. It has 
a brown iris, and the immatures lack the streaked underparts which 
are characteristic of A. opacus and other species. The structure of 
its wing resembles that of A. opacus, but the primaries are more 
rounded. It is apparently better adapted to forested uplands, 
whereas A. opacus and its relatives, A. cantoroides and A. feadensis, 
appear to prefer lowland forests and coconut plantations. In habits 
and habitat preference, A. pelzelni seems to resemble A. santo- 
vestris, which is restricted to mountain environment on Espiritu 
Santo in the New Hebrides. The describers of this starling, Harris- 
son and Marshall (1937:149), write that ''Aplonis santovestris ap- 
parently most closely resembles A. pelzelni from Ponape, especially 
in bill and tarsus." According to the description, A. santovestris is 
approximately the size of A. pelzelni with brownish coloring, crown 
dark brown, lower back and rump dark rufous, wing and tail black- 
ish-brown, underparts rufous-brown, and iris grayish-green. These 
two birds are separated geographically and apparently exhibit evi- 
dences of parallel development. Possibly they came from a common 
ancestral stock. Mayr (1941b:204) writes that A. pelzelni belongs 
with the starlings of the Polynesian area. I have compared A. pel- 
zelni with other starlings of the Southwest Pacific, including A. 



Baker: The Avifauna of Micronesia 301 

feadensis, A. cantoroides, and A. zealandicus, but see no close re- 
semblances. 

Aplonis corvinus (Kittlitz) 

Kusaie Mountain Starling 

Lamvrothomis corvina Kittlitz, Kupfertaf. Naturgesch. Vogel, 2, 1833, p. 12, pi. 15, 
fig. 3. (Type locality, Ualan = Kusaie.) 

Lamprothomis corvina Kittlitz, Mem. Acad. Imp. Sci. St. Peterbourg, 2, 1835, p. 7, 
pi. 9 (Ualan); idem, Obser. Zool., in Lutke, Voy. "Le Seniavine," 3, 1836, p. 285 
(Ualan). 

Lamprotomis corvina Bonaparte, Consp. Avium, 1, 1850, p. 417 (Ualan); Hartlaub, 
Archiv. f. Naturgesch., 18, 1852, p. 133 (Ualan); Kittlitz, Denkw. Reise russ. Amer. 
Micron, und Kamchat., 2, 1858, pp. 25, 43, 59, 103 (Ualan); Finsch, Ibis, 1881, p. 104 
(Kuschai). 

Lamprocorax corvinus Hartlaub, Journ. f. Omith., 1854, p. 168 (Carolinen = 
Kusaie); Sclater, Ibis, 1859, p. 327 (Caroline = Kusaie) ; Dubois, Syn. Avium, 1, 1902, 
p. 543 (Kuschai). 

Calornis (Lamprocoraxf) corvina Gray, Cat. Birds Trop. Is. Pacific Ocean, 1859, p. 
25 (Oualan). 

Stumoides corvina Finsch and Hartlaub, Fauna Centralpolynesiens, 1867, p. 108 
(Ualan); Finsch, Journ. f. Omith., 1880, pp. 297, 302 (Kuschai). 

Calornis corvina Gray, Hand-list Birds, 2, 1870, p. 27 Caroline = Kusaie) ; Hart- 
laub and Finsch, Proc. Zool. Soc. London, 1872, p. 100 (Ualan); Giebel, Thes. Omith., 
2, 1875, p. 427 (Caroline = Kusaie) ; Sharpe, Cat. Birds British Mus., 13, 1890, p. 
137 (Kuschai); Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 
(1891), p. 46 (Ualan or Kushai); Matschie, Joum. f. Ornith., 1901, p. 112 (Ualan); 
Takatsukasa and Kuroda, Tori, 1, 1915, p. 64 (Kusaie). 

Stumoides corvinus Finsch, Ibis, 1881, pp. 107, 108 (Kushai). 

Kittlitzia corvina Hartert, Kat. Vogelsamml. Senckenb., 1891, p. 75 (Ualan) ; Ku- 
roda, in Momiyania, Birds Micronesia, 1922, p. 72 (Kusaie); Mathews, Syst. .\vium 
Australasianarum, 2, 1930, p. 853 (Kusaie); Hand-list Japanese Birds, rev., 1932, p. 
169 (Kusaie); Hand-list Japanese Birds, 3d ed., 1942, p. 187 (Kusaie). 

Aplonis corvina Reichenow, Die Vogel, 2, 1914, p. 356 (Ualan) ; Mayr, Proc. 6th 
Pacific Sci. Congr., 4, 1941, p. 213 (Kusaie). 

Aplonis corvinus Mayr, Birds Southwest Pacific, 1945, p. 298 (Kusaie). 

Geographic range. — Micronesia: Caroline Islands — Kusaie, probably ex- 
tinct for many years. 

Characters. — According to Sharpe (1890:137), "Shining black; each feather 
with a glossy margin, varying from steel-green to purplish red; bill and feet 
black (Kittlitz)." 

Remarks. — Kittlitz obtained two specimens of a unique starling 
at Kusaie when he visited the island in December and January, 
1827-'28. He named the birds as new and deposited the specimens 
in the museum in St. Petersburg. The bird has not been found at 
Kusaie since that time. Sharpe (1890:137-138, footnote) writes 
"This species I have never seen, and Dr. Finsch did not meet with 
it during his visit to Kuschai. He writes to me: — 'It no doubt exists 
on Kuschai, just as it did when Kittlitz visited the island. Nobody 
has reached the mountains in the interior since Kittlitz's time; and 
it is strictly a mountain bird.' " Coultas spent considerable time 
searching the higher areas of Kusaie for the bird in 1931. 

The Kusaie Mountain Starling apparently represents an early 



302 University of Kansas Publs., Mus, Nat. Hist. 

invasion of Micronesia, independent of that of any other starling 
in the area and perhaps the earliest of the three colonizations by 
starlings in Micronesia. The drawing of the bird as pictured by 
Kittlitz (1833: pi. 14, fig. 3) shows the long bill to be one of its 
distinctive characters. This suggests relationship to A. atrifuscus of 
Samoa, as noted by Mayr (1942a: 6). A. atrijuscus is larger than 
A. opacus with a longer bill and gloss on some of the feathering of 
the body; it looks a good deal like the drawing of A. corvinus by 
Kittlitz. A. corvinus may also have some relation to A. magnus of 
Biak, although this species has a longer tail and a shorter bill. 
A. corvinus probably has undergone an evolutionary development 
which parallels that of A. atrijuscus and possibly other species in 
the Polynesian and Melanesian areas. The ancestral stock from 
which A. corvinus was derived may have been close to A. grandis, 
which is found in the Solomon area. A. grandis is a forest bird, 
somewhat solitary in habits. 

Sturnus philippensis (Forster) 
Violet-backed Starling 

[Motacilla] philivperisis Forster, Ind. Zool., 1781, p. 41. (Type locality, Philip- 
pines.) 

Sttirnus philippensis Mayr, Birds Southwest Pacific, 1945, p. 302 (Palau). 

Geographic range. — Breeds in Japan. Winters to the Philippine Islands. In 
Micronesia: Palau Islands — exact locality unknown. 

Remarks. — Mayr (1945a: 302) records this starling as a migrant 
visitor to the Palau Islands. Coultas obtained an immature female 
of this species at Palau on October 13, 1931. 

Sturnus cineraceus Temminck 
Ashy Starling 

Sturnus cineraceus Temminck, PI. Col. 2, 1832, pi. 556. (Type locality, Japan.) 
Spodiopsar cineracca Kishida, Lansania, 1, 1929, p. 17 (Saipan) ; Hand-list Japa- 
nese Birds, 3d ed., 1942, p. 187 (Saipan). 

Geographic range. — Breeds in eastern Asia and Japan. Winters in southern 
China and Philippines. In Micronesia: Mariana Islands — Saipan. 

Remarks. — The Ashy Starling has been reported from Saipan by 
Kishida. It probably is a casual winter migrant. 

Cleptornis marchei (Oustalet) 
. , Golden Honey-eater 

Ptilotis Marchei Oustalet, Le Nat., 1889, p. 200. (Type locality, Saypan.) 
Cleptornis marchei Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 

1890-1891 (1891), p. 35 (Saypan); Hartert, Novit. Zool., 5, 1898, p. 56 (Saipan); 
■ . Matschie, Journ. f. Ornith., 1901, p. 112 (Saipan); Seale, Occ. Papers Bernice P. Bishop 

Mus., 1, 1901, p. 60 (Saipan); Dubois, Syn. Avium, 1, 1902, p. 722 (Marianne = 



Baker: The Avifauna of Micronesia 



303 



Saipan); Takatsukasa and Kuroda, Tori, 1, 1915, p. 64 (Marianne = Saipan) ; Kuroda, 
in Momiyama, Birds Micronesia, 1922, p. 75 (Saipan); Mathews, Syst. Avium Australa- 
sianarum, 2, 1930, p. 788 (Saipan); Hand-list Japanese Birds, rev., 1932, p. 171 
(Saipan); Hand-list Japanese Birds, 3d ed., 1942, p. 190 (Saipan); Mayr, Birds South- 
west Pacific, 1945, p. 298 (Saipan); Stott, Auk, 64, 1947, p. 527 (Saipan). 

Ptilotis (Cleptomis) marchei Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 
1895, p. 202 (Saypan). 

Geographic range. — Micronesia: Mariana Islands — Saipan. 

Characters. — Adult: A small honey-eater with head, rump, and underparts 
near "light cadmium" becoming lighter on the chin and darker on the nape; 
back near "orange-citrine"; wings and tail feathers brown with outer edges 
colored like back and inner edges whitish; orbital ring pale yellow; breast, 
belly, sides, and under tail- and upper tail-coverts near "raw sienna"; under 
wing-coverts pale yellow; axillaries yellow; bill and feet light yellow-brown, 
maxilla darker; iris chestnut-brown. Immature has lighter bill. 

Measurements. — Measurements are listed in table 47. 



Table 47. Measurements of Cleptomis marchei 



Number and Sex 


Wing 


Tail 


Full culmen 


Tarsus 


7 adult males 


79 

(77-80) 

73 

(72-75) 


64 

(61-66) 

58 
(56-59) 


19.5 
(19.0-20.0) 

18.0 
(17.5-18.5) 


26 


5 adult females 


(25-27) 
24 




(23-25) 



Specimens examined. — Total number, 17 (9 males, 8 females), as follows: Mariana Islands, 
USNM— Saipan, 4 (July 11, Dec. 15); AMNH— Saipan, 13 (July 8, Aug. 1, 10, 13, 14, 21, 
30, Sept. 3, 7, 9, 15). 

Nesting. — Hartert (1898:56) reports that one nest of the Golden Honey- 
eater was found on July 7. It was hung from a fork of a branch, "like the nest 
of a golden Oriole." He writes that four other nests were obtained in late 
August. Hartert describes the egg as "pale blue without gloss, spotted over and 
over with rufous, more so on the thicker end, and measures about 20:15 mm." 

Molt. — Specimens taken in July, August, and September are molting. 

Remarks. — Oustalet (1895:202) writes that Marche obtained 25 
specimens of the Golden Honey-eater at Saipan in May, June, and 
July, 1887. Little is known regarding its habits; Moran (1946:262) 
writes that the bird "reminds one of the prothonotary warbler, with 
a long, curved, black bill." Stott (1947:527) writes that "it appears 
to be restricted to a single habitat, that of dense forest." He found 
the bird in forest on the north shore of Magicienne Bay. Coultas 
obtained only one specimen on his visit to Saipan in 1931. Marshall 
(1949:216) records some interesting observations of this bird made 
in 1945. He notes {op. cit. p. 219) that the bird breeds in January, 
February and April. 



304 University of Kansas Publs., Mus. Nat. Hist. 

Not only is it remarkable that the Golden Honey-eater has be- 
come established on a single island in a rather closely associated 
chain of islands, but it is also difficult to determine from where the 
bird came. It seemingly has no close relatives in the Micronesian 
area. Oustalet (1895:202) points out that one has to go to New 
Guinea, Moluccas, Australia, Fiji, Samoa, and Tonga in order to 
find related forms. In looking through the large collections of 
Meliphagidae in the American Museum of Natural History, I found 
only a few genera to which the Saipan Golden Honey-eater seems 
to be closely related. Tijneliopsis of New Guinea has some re- 
semblances to Cleptornis, although the coloration is different. Time- 
liopsis has a similar bill, but has a longer tail and longer wing; the 
shortness of the wing in Cleptornis is not unusual since other insular 
forms also exhibit this characteristic. 

Perhaps Cleptornis is closer to the genus Meliphaga of New 
Guinea and Australia, which has become differentiated into a 
number of diverse species and subspecies. Cleptornis compares 
rather favorably with M. pencillata carteri of Australia, but differs 
by the softness of its feathers and the shorter wing and shorter tail. 
It shows also some affinities with M. flava of Australia, particularly 
in shape of bill; the coloration of the feathers is light olive-green 
in M. flava. The bird at Saipan seemingly has no relationships 
with the Hawaiian honey-eaters. 

Myzomela cardinalis rubratra (Lesson) 
Cardinal Honey-eater 

Cinnyris rubrater Lesson, Diet. Sci. Nat., ed. Levrault, 50, 1827. p. 30. (Type 
locality, OualanrrKusaie.) 

Cinnyris rubrater Lesson (part), Voy. "La Coquille," Zool., 2, 1828, pp. 433, 678 
(Oualan): idem (part), Man. d'Ornith., 2, 1828, p. 55 (Oualan) ; idem (part), 
Traite d'Ornith., 1831, p. 299 (Oualan); Kittlitz (part), Kupfertaf. Naturgesch. 
Vogel, 1832, p. 6, pi. 8, fig. 1 (Ualan); idem (part), Denkw. Reise russ. Amer. 
Micron, und Kamchat., 1, 1858, pp. 364, 381; 2, 1858, pp. 39, 49 (Ualan). 

Certhia Cardinalis Kittlitz, Mem. Acad. Imp. Sci. St. Petersbourg, 2, 1835, p. 4 
(Ualan). 

Cinnyris cardinalis Kittlitz, Obser. Zool., in Lutke, Voy. "Le Seniavine," 3, 1836, 
p. 285 (Ualan). 

Myzomela sanguinolenta Bonaparte, Consp. Avium, 1, 1850, p. 394 (no loc. ^ 
Kusaie?). 

Myzomela rubrater Hartlaub (part), Archiv. f. Naturgesch., 18, 1852, pp. 109, 
131 (Ualan); Finsch and Hartlaub, Fauna Centralpolynesiens, 18()7, p. 57 (Ualan). 

Myzomela lubratra Hartlaub (part), Journ. f. Ornith., 1854, p. 168 (Carolinen = 
Kusaie); idem (part), Proc. Zool. Soc. London, 1867 (186S), p. 829 (Carolines =; 
Kusaie); Hartlaub and Finsch (part), Proc. Zool. Soc. London, 1872, p. 95 (Ualan); 
Giebel (part), Thes. Ornith., 2, 1875, p. 681 (Carolinae = Kusaie) ; Finsch (part), 
Journ. Mus. Godeffroy, 12, 1876, p. 26 Ualan); Forbes (part), Proc. Zool. Soc. 
London, 1879, p. 271 (Ualan); Finsch (part), Journ. f. Ornith., 1880, pp. 285, 298 
(Kuschai); idevi (part). Ibis, 1881, pp. 103, 108, 111 (Kuschai) ; idem (part), Mitth. 
Ornith. Ver. Wien, 1884, p. 48 (Ualan); Hartert, Kat. Vogelsanmil. Senckenb., 1891, 



Baker: The Avifauna of Micronesia 



305 



p. 31 (Ualan); Wiglesworth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 
1890-1891 (1891), p. 31 (Ualan); Oustalet (part), Nouv. Arch. Mus. Hist. Nat. 
Paris, (3), 7, 1895, pp. 201, 202 (Kushai) ; Hartert (part), Novit, Zool., 5, 1898, p. 56 
(Ualan); Dubois (part), Syn. Avium, 1, 1902, p. 716 (Carolines = Kusaie). 

Certhia sanguinolenta Kittlitz, Denkw. Reise russ. Amer. Micron, und Kamchat., 
1, 1858, p. 364 (Ualan). 

Myzomela major Gray, Cat. Birds Trop. Is. Pacific Ocean, 1859, p. 11 (Oualan?). 

Myzomela rubrata Matschie (part), Joum. f. Ornith., 1901, p. 112 (Ualan). 

Myzomela ruhratra ruhratra Wetmore, Proc. Biol. Soc. Washington, 30, 1917, p. 
117 (Kusaie); Wetmore (part), in Townsend and Wetmore, Bull. Mus. Comp. Zool., 
63, 1919, p. 219 (Kusaie); Momiyama, Birds Micronesia, 1922, pp. 15, 20, 21, 22, 
(Kusaie); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 72 (Kusaie); Mathews, 
Syst. Avium Australasianarum, 2, 1930, p. 743 (Oualan) ; Hand-list Japanese Birds 
(part), rev., 1932, p. 172 (Kusaie); Hand-list Japanese Birds (part), 3d ed., 1942, 
p. 191 (Kusaie). 

Myzomela cardinalis rubratra Mayr, Birds Southwest Pacific, 1945, p. 299 (Kusaie). 

Geographic range. — Micronesia : Caroline Islands — Kusaie. 

Characters. — Adult male: Head (except lores), neck back, rump, upper 
tail-coverts, chin, throat, breast, and upper abdomen black with feathers 
tipped with coloring between "scarlet" and *'scarlet-red"; rest of feathering 
black; bill long and curved and black; feet black; iris dark brown. 

Adult female: Resembles adult male, but smaller; red coloring duller; 

Table 48. Me.asxjrements of Myzomela cardinalis of Micronesia 



Subspecies 


Number and sex 


Wing 


Tail 


Full culmen 


Tarsus 


M. 


c. ruhratra .... 


21 adult males 


79 

(76-81) 


55 
(53-56) 


19.5 
(18.5-20.5) 


22 

(21-22) 






20 adult females 


71 
(69-74) 


49 

(45-51) 


18.5 
(17.5-19.5) 


20 

(19-21) 


M. 


c. dichromata . . 


24 adult males 


78 
(76-80) 


53 

(51-56) 


21.5 
(20.0-23.0) 


22 

(21-23) 






22 adult females 


69 
(66-72) 


47 
(45-49) 


19.0 
(17.5-20.5) 


20 

(19-21) 


M. 


c. major 


9 adult males 


77 
(75-78) 


55 
(54-59) 


20.0 
(19.5-20.5) 


22 

(21-22) 






2 adult females 


70 


50 


19.0, 20.5 


21.5 


M. 


c. saffordi 


47 adult males 


73 

(69-77) 


55 

(51-56) 


20.0 
(19.0-20.5) 


22 

(21-24) 






14 adult females 


65 
(63-71) 


49 
(46-51) 


18.5 
(17.5-19.5) 


21 
(19-21) 


M. 


c. kurodai 


2 adult males 


74, 75 


52 


20.0, 20.5 


20, 21 


M. 


c. kobayashii. . 


17 adult males 


74 

(71-76) 


54 

(51-57) 


20.5 
(19.0-22.0) 


21 

(20-22) 






8 adult females 


67 
(65-68) 


48 
(45-50) 


18.0 
(17.5-19.0) 


20 

(19-21) 



20—8131 



306 University of Kansas Publs., Mus. Nat. Hist. 

wings and tail more brownish and less blackish; abdomen and under tail- 
coverts dark gray. 

Immature: Resembles adult, but duller and less blackish and more grayish 
with less red coloring on feathers and an olivaceous-brown tinge to plumage. 

Measurements. — Measurements are listed in table 48. 

Specimens examined. — Total number, 62 (35 males, 27 females), as follows: Caroline 
Islands, USNM— Kusaie, 3 (Feb. 9); AMNH— Kusaie, 59 (Jan., Feb., March). 

Nesting — Finsch records the taking of eggs of the honey-eater at Kusaie 
on February 26 and March 10, 1880. 

Molt. — Evidence of molt was observed in a few specimens taken in January 
and in larger number of birds taken in March. In addition, some skins ob- 
tained in March showed fresh plumage. Although there is little evidence avail- 
able, I suppose that nesting activities of M. r. rubratra at Kusaie occur in the 
winter months of December, January, February, and March, and that molt 
begins in January, especially in the males, and possibly reaches a peak in 
March. 

Remarks. — M. r. rubratra was first ciescribeci by Lesson, who re- 
ferred to it under the name Cinnyris rubrater. The bird was found 
by Lesson at Kusaie, when he visited the island in June, 1924, as a 
member of the expedition from the ship "La Coquille." In his de- 
scription he also stated that the bird was found in the Philippines by 
Dussumier. The report of the bird's occurrence in the Philippines 
proved to be erroneous, as was pointed out by Wetmore (in Town- 
send and Wetmore, 1919:220). Oustalet (1895:200) contended that 
Lesson's description was based on the specimens taken by Quoy and 
Gaimard in the Marianas; he stated that none of the birds which 
Lesson mentions from Kusaie was preserved. Bonaparte also con- 
sidered Cinnyris rubrater to be from the Marianas, and he gave the 
name Myzomela major to the honey-eater of the Caroline Islands 
(apparently including Kusaie) on the basis of specimens taken by 
Hombron and Jacquinot at Truk. Wetmore (in Townsend and Wet- 
more, 1919:220) settles the argument and assigns Lesson's name 
rubratra to the honey-eater at Kusaie; apparently this treatment is 
the correct one inasmuch as Lesson used his own field notes and 
records of the occurrence of this honey-eater at Kusaie in preparing 
his description, even if the actual specimens were not preserved. This 
arrangement makes Bonaparte's name major available for the popu- 
lation at Truk and makes Wetmore's name saffordi available for the 
population in the Marianas. The placing of the honey-eaters of 
Micronesia within the species Myzomela cardinalis by Mayr 
(1932:19) is, I think, justified. 

Little information is available concerning the habits of the honey- 
eater at Kusaie. In 1931, Coultas (field notes) regarded the bird as 



Baker: The Avifauna of Micronesia 307 

common in the lowlands, especially in the coconut groves. He did 
not find the bird at high elevations on the island. 

Myzomela cardinalis dichromata Wetmore 
Cardinal Honey-eater 

Myzomela rubratra dichromata Wetmore, in Townsend and Wetmore, Bull. Mus. 
Comp. Zool., 63, 1919, p. 220. (T>-pe locality, Ponape.) 

Myzomela rubratra Pelzeln, Reise "Novara," Vogel, 1865, pp. 55, 162 (Puynipet = 
Ponape); Finsch (part), Joum. Mus. Godeffroy, 12, 1876, pp. 17, 26 (Ponape); idem, 
Proc. Zool. See. London, 1877 (1878), p. 778 (Ponape); Forbes (part), Proc. Zool., 
Soc. London, 1879, p. 271 (Ponape); Finsch (part), Journ. f. Ornith., 1880, p. 285 
(Ponape) ; idem (part), Proc. Zool. Soc. London, 1880, p. 575 (Ponape) ; idem (part), 
Ibis, 1881, pp. Ill, 115 (Ponape); idem (part), Mitth. Omith. Ver. Wien, 1884, p. 48 
(Ponap6); Wiglesworth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 
(1891), p, 31 (Ponape); Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 
1895, p. 202 (Ponapi). 

Myzomela rubrata Nehrkorn (part), Journ. f. Ornith., 1879, p. 397 (Ponape); 
Christian, The Caroline Islands, 1899, p. 358 (Ponape) ; Matschie (part), Joum. f. 
Ornith., 1901, p. 112 (Ponape); Takatsukasa and Kuroda (part). Tori, 1, 1915, p. 55 
Ponape); Mayr, Proc. 6th Pacific Sci. Congr., 4, 1941, p. 204 (Ponape). 

Myzomela chermesina Gadow, Cat. Birds British Mus., 9, 1884, p. 137 (Ponape); 
Takatsukasa and Kuroda, Tori, 1, 1915, p. 64 (Ponape). 

Myzomela rubratra dichromata Momiyama, Birds Micronesia, 1922, pp. 15, 20, 21, 
22 (Ponape); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 73 (Ponape); Mathews, 
Syst. Avium Australasianarum, 2, 1930, p. 743 (Ponape). 

Myzomela rubratra rubratra Yamashina, Tori, 7, 1932, p. 395 (Ponapi); Hand-list 
Japanese Birds (part), rev., 1932, p. 172 (Ponape); Hand-list Japanese Birds (part), 
3d ed., 1942, p. 191 (Ponapi). 

Myzomela cardinalis dichromata Mayr, Birds Southwest Pacific, 1945, p. 299 
Ponape). 

Geographic range. — Micronesia : Caroline Islands — Ponape. 

Characters. — Adult male: Resembles adult males of M. c. rubratra, but with 
more e.xtensive black markings on lores and below eye; tips of feathers lighter 
"scarlet." 

Adult female : Resembles adult female of M. c. rubratra, but duller and with 
red coloring much reduced; head, neck, shoulder, ear-coverts, and sides of neck 
sooty brownish-gray; rest of upper parts dark brownish-gray with plumage of 
middle of back, rump, and upper tail-coverts tipped with scarlet; wings and 
tail dark brown with outer edges olivaceous-gray; chin and throat reddish; 
breast light brownish-gray, may be washed with reddish; axillaries, abdomen, 
and under tail-coverts grayish. 

Immature male : Resembles adult male, but scarlet coloring less brilliant and 
thinner on forehead, middle of back, rump, upper tail-coverts, and underparts 
and absent, or nearly absent, on crown and neck. 

Immature female : Resembles adult female, but scarlet coloring thinner and 
present only on underparts, back, rump, and upper tail-coverts; abdomen and 
under tail-coverts washed with buff. 

Measurements. — Measurements are listed in table 48. 

Specimens examined. — Total number, 52 (26 males, 24 females, 2 unsexed), as follows: 
Caroline Islands, USNM— Ponape, 3 (Feb. 11, 12); AMNH— Ponape, 49 (Nov., Dec). 

Nesting. — Yamashina (1932a :395) records a large collection of eggs of the 
honey-eater, taken at Ponape in 1931. Of 13 sets of eggs listed, 10 include two 
eggs per set and 3 include one egg per set. These were obtained from July 20 to 



308 University of Kansas Publs., Mus. Nat. Hist. 

September 2. Coultas (field notes) found one nest with young in a tree-fern in 
the period of November and December, 1930. The nest was cup-shaped and 
made of fern and fine grasses and fined with Hchens. Coultas writes that 
only the female feeds the young. He suspects that the honey-eater nests at 
all times of the year. 

Molt. — Most of the birds taken by Coultas in November and December are 
in molting plumage. 

Remarks. — The Cardinal Honey-eater at Ponape is, according to 
Coultas, found in most habitats of the island. He found it to be 
an aggressive bird, often chasing the white-eye Zosterops cinerea. 
The committee (Hachisuka et al.) which prepared the Hand-list 
of Japanese Birds in both the revised edition (1932) and the third 
edition (1942) does not recognize the Ponape honey-eater as sepa- 
rable from the bird at Kusaie. I see no reason for this action 
and find the bird at Ponape to be a well-marked subspecies. 

Myzomela cardinalis major Bonaparte 
Cardinal Honey-eater 

Myzomela major Bonaparte, Comptes Rendus Acaa. Sci. Paris, 38, 18.')4, p. 2G4. 
(Type locality, "ex Ins. Carolinis ab Homb. et Jacq." r: Truk.) 

Myzomela major Gray, Hand-list Birds, 1, 1809, p. 153 (Caroline = Truk); Giebel, 
Thes. Ornith., 1875, p. 681 (Carolinae = Truk?) ; Takatsukasa and Kuroda, Tori, 1, 
1915, p. 64 (Ruk); Kuroda, Dobutsu. Zasshi, 27, 1915, p. 28 (Ruk) ; idem, Dobutsu. 
Zasshi, 28, 1916, p. 71 (Ruk). 

Myzomela rubratra Finsch (part), Proc. Zool. Soc. London, 1880, p. 575 (Ruk); 
Sehmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, p. 253 (Ruk); Wigles- 
worth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 31 
(Ruk); Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, p. 202 (Ruk); 
Hartert (part), Novit. Zool., 5, 1898, p. 56 (Ruk); idevi (part), Novit. Zool., 7, 
1900, p. 2 (Ruk); Dubois (part), Syn Avium, 1, 1902, p. 714 (Carolines = Truk?). 

Myzomela rubrata Matschie (part), Journ. f. Ornith., 1901, p. 112 (Ruck); Takat- 
sukasa and Kuroda (part). Tori, 1, 1915, p. 55 (Ruk). 

Myzomela rubratra rubrata Wetmore (part), in Townsend and Wetmore, Bull. 
Mus. Comp. Zool., 63, 1919, p. 221 (Uala). 

Myzomela rubrata wetmorei Momiyania, Birds Micronesia, 1922, p. 15 (Type lo- 
cality, Ruk); Kuroda, in Momiyania, Birds Micronesia, 1922, p. 73 (Ruk); Mathews, 
Syst. Avium Australasianarum, 2, 1930, p. 743 (Ruk); Hand-list Japanese Birds, rev., 
1932, p. 172 (Ti\ik); Hand-list Japanese Birds, 3d ed., 1942, p. 190 (Truk). 

Myzomela cardinalis major Mayr, Birds Southwest Pacific, 1945, p. 299 (Truk); 
Baker, Srnithson. Misc. Coll., vol. 107, no. 15, 1948, p. 72 (Truk). 

Geographic range. — Micronesia: Caroline Islands — Truk. 

Characters. — Adult male: Resembles adult male of M. c. rubrata, but tips 
of plumage lighter "scarlet," 

Adult female: Resembles adult female of M. c. rubrata, but underparts 
more heavily tipped with scarlet; abdomen and under tail-coverts black; tail 
slightly darker. Differs from M. c. dichromata by presence of scarlet tips on 
feathers of head. 

Immature male: Resembles adult female, but scarlet coloring of tips of 
feathers of head and neck narrower. 



Baker: The Avifauna of Micronesia 309 

Immature female: Resembles immature female of M. c. ruhrata, but upper 
parts grayer; underparts darker. 

Measurements. — Measurements are listed in table 48. 

Specimens examined. — Total number, 19 (13 males, 6 females), as follows: Caroline Is- 
lands, USNM— Truk, 2 (Feb. 16, Dec. 13); AMNH— Truk, 17 (Feb., March, Nov., Dec). 

Nesting. — Concerning the honey-eater at Truk, Hartert (1900:2) writes 
"many nests were found from end of May to July, and one in March." Mc- 
Elroy examined three males in December, which had swollen testes. As 
seems to be the case with other races of thi.? species, the Cardinal Honey- 
eater at Truk may nest at all times of the year. 

Molt. — Specimens examined that were taken in November, December and 
February are in fresh or in molting plumages. 

Remarks. — Bonaparte described his Myzomela major as "Similis 
praecedenti, sed major et percoccinea." He compares it here with 
Myzomela ruhrata, which he considered as a resident of the Mari- 
ana Islands. According to Oustalet (1895:202) Hombron and Jac- 
quinot obtained one specimen of the honey-eater at Truk in 1841. 
This subspecies, as well as most of the others of M. cardinalis in 
Micronesia, is best distinguished by the characteristics of the female. 
The male of the different subspecies shows much less geographic 
variation, 

Myzomela cardinalis saffordi Wetmore 
Cardinal Honey-eater 

Myzomela rubratra saffordi Wetmore, Proc. Biol. Soc. Washington, 30, 1917, p. 117. 
(Type locality, Guam.) 

Cinnyris rubrater Lesson (part). Diet. Sci. Nat., ed. LevTault, 50, 1827, p. 30 
(Mariannes); idem, (part), Voy. "La Coquille," Zool., 2, 1828, p. 678 (Mariannes); 
idem (part), Man. d'Ornith., 2, 1828, p. 55 (Mariannes); idem (part), Traite d'Ornith., 
1831, p. 299 (Mariannes): Kittlitz (part), Kupfertaf. Naturgesch. Vogel. 1, 1832, p. 6, 
pi. 8, fig. 1 (Guaham) ; idetn (part), Denkw. Raise russ. Amer. Micron, und Kamchat., 
1, 1858, pp. 364, 381; 2, 1858, pp. 39, 49 (Guaham). 

Certhia cardinalis Kittlitz, Obser. Zool., in Lutke, Voy. "Le Seniavine," 3, 1836, p. 
304 (Guaham). 

Myzomela rubrater Hartlauh (part), Archiv f. Naturgesch., 18, 1852, p. 109 
(Mariannen); Finsch and Hartlaub (part). Fauna Centralpolynesiens, 1867, p. 57 
(Guaham). 

Myzomela rubratra Bonaparte, Comptes Rendus Acad. Sci. Paris, 38, 1854, p. 263 
(Mariannes); Hartlaub (part), Joum. f. Omith., 1854, p. 167 (Mariannen); Gray 
(part). Cat. Birds Trop. Is. Pacific Ocean, 1859, p. 11 (Guam); idem (part). Hand- 
list Birds, 1, 1869, p. 154 (Marian); Finsch (part), Journ. Mus. Godeffroy, 12, 1876, 
pp. 17, 26 (Marianen); Forbes (part), Proc. Zool. Soc. London, 1879, p. 270 (Marianis) ; 
Giebel (part), Thes. Ornith., 2, 1875, p. 681 (Marinae) ; Finsch (part), Mitth. Ornith. 
Ver. Wien, 1884, p. 48 (Guam) ; Wiglesworth (part), Abhandl. und Ber. Zool. Mus. 
Dresden, no. 6, 1890-1891 (1891), p. 31 (Marianne); Oustalet (part), Nouv. Arch. 
Mus. Hist. Nat. Paris, (3), 7, 1895, p. 197 (Guam, Rota, Saypan, Pagan, Agrigan) ; 
Hartert (part), Novit. Zool., 5, 1898, p. 55 (Guam, Saipan, Pagan, Agrigan); idem 
(part), Novit. Zool., 7, 1900, p. 2 (Guam); Wheeler, Report Island of Guam, 1900, 
p. 13 (Guam); Seale (part), Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 55 
(Marianae); Safford, The Plant World, 7, 1904, p. 263 (Guam); idem, Contr. U. S. 
Nat. Herb., 9, 1905, p. 79 (Guam); Mearns, Proc. U. S. Nat. Mus., 36, 1909, p. 477 
(Guam); Reichenow (part). Die Vogel, 2, 1914, p. 482 (Marianen); Takatsukasa and 



310 University of Kansas Publs., Mus. Nat. Hist. 

Kuroda (part), Tori, 1, 1915, p. 64 (Marianas); Cox, Island of Guam, 1917, p. 21 
(Guam). 

Myzomela rubrata Oustalet, Le Nat., 1889, p. 260 (Mariannes); Matschie (part), 
Journ. f. Ornith., 1901, p. 112 (Guam, Saipan); Safford, Osprey, 1902, p. 69 (Guam); 
Prowazek, Die deutschen Marianen, 1913, p. 101 (Saipan). 

Myzomela rubratra saffordi Wetmore, in Townsend and Wetmore, Bull. Mus. Comp. 
Zool., 63, 1919, p. 221 (Guam, Saipan) ; Momiyama, Birds Micronesia, 1922, pp. 17, 
20, 21, 22 (Guam, Rota, Saipan, Pagan, Agrigan) ; Kuroda in Momiyama, Birds Micro- 
nesia, 1922, p. 74 (Guam, Rota, Saipan, Pagan, Agrigan); Mathews, Syst. Avium 
Australasianarum, 2, 1930, p. 744 (Guam); Yamashina, Tori, 7, 1932, p. 395 (Mari- 
anas?); Hand-list Japanese Birds, rev., 1932, p. 171 (Marianas); Bryan, Guam Rec, 
vol. 13, no. 2. 1936, p. 25 (Guam); Hand-list Japanese Birds, 3d ed., 1942, p. 190 
(Guam, Rota, Agiguan, Tinian, Saipan, Almagan, Pagan, Agrigan, Assongsong). 

Mizomela rubrata saffordi Yamashina, Tori, 19, 1940, p. 673 (Assongsong, Agiguan). 

Myzomela cardinalis saffordi Mayr, Birds Southwest Pacific, 1945, p. 299 (Mari- 
anas); Downs, Trans. Kansas Acad. Sci., 49, 1946, p. 103 (Tinian); Borror, Auk, 1947, 
p. 417 (Agrihan); Stott, Auk, 1947, p. 527 (Saipan, Guam); Baker, Smithson. Misc. 
Coll., vol. 107, no. 15, 1948, p. 72 (Guam, Rota). 

Myzomela cardinalis Watson, The Raven, 17, 1946, p. 41 (Guam); Strophlet, Auk, 
1946, p. 540 (Guam); Baker, Condor, 49, 1947, p. 125 (Guam). 

Geographic range. — Micronesia: Mariana Islands — Guam, Rota, Tinian, 
Agiguan, Saipan, Almagan, Pagan, Agrihan, Asuncion. 

Characters. — Adult male: Resembles M. c. rubratra, but smaller with red 
coloring lighter and more orange; edges of wing and tail feathers olivaceous. 
Differs from adult males of M. c. dichromata and M. c. major by smaller size 
and presence of olivaceous edgings on wing and tail feathers. 

Adult female: Resembles adult female of M. c. rubratra, but smaller and 
paler with upper parts dark olivaceous-gray, sparsely mottled with scarlet; 
outer edges of wing and tail feathers greenish-olive; abdomen and under tail- 
coverts buffy-gray. Differs from M . c. dichromata by smaller size and presence 
of scarlet tips of feathers on top of head. Differs from M. c. major by smaller 
size and presence of broad olivaceous edges on tail feathers. 

Immature male : Resembles adult male, but red coloring less brilliant, upper 
parts, lower breast, and abdomen more narrowly edged with the red coloring; 
plumage of breast, abdomen, and under tail-coverts buffy-gray, lighter in very 
young birds. 

Table 49. Measurements op Adult Males of Myzomela cardinalis sajfordi 

FROM THE Mariana Islands 



Island 


No. 


Wing 


Tail 


Full culmen 


Tarsus 


Guam 

Rota 


35 

1 
5 

4 

1 


72 
(69-75) 

73 

73 

(71-74) 

74 
(72-76) 

77 


54 

(51-56) 


20.0 
(19.5-20.5) 

20.0 

19.5 
(19.0-20.0) 

19.5 
(19.0-20.5) 

20.0 


22 

(21-23) 

22 


Tinian 


53 

(52-55) 

54 

(53-55) 

55 


22 


Saipan 


(21-24) 
22 


Agrihan 


(22-23) 
22 







Baker: The Avifauna of Micronesia 311 

Immature female: Resembles adult female, but paler with upper parts 
darker brown; underparts pale buffy -brown; outer edges of wing and tail- 
feathers greenish-olive, more extensive than in adult. 

Measurements. — Measurements of the subspecies of M. cardinalis in Micro- 
nesia are listed in table 48. Measurements of male specimens of M. c. saffordi 
from various islands in the Marianas are listed in table 49. 

Weights. — The author (1948:72) records weights of M. c. sajfordi from Guam 
as: 17 adult males, 12.7-18.0 (15.0), and 5 adult females, 10.4-15.0 (12.7). 

Specimens examined.— Total number, 80 (61 males, 17 females, 2 unsexed), as follows: 
Mariana Islands, USNM — Guam, 43 (Jan. 22, May 26, 30, June 2, 3, 5, 7, 8, 9, 13, 
18, 19, 25, 28, July 6, 10, 12, 17, 19, 20, 21, Sept., Nov. 20, 21)— Rota, 2 (Oct. 10)— Tinian, 
3 (Oct. 23, 25)— Saipan 2 (Sept. 27, 30); AMNH— Guam, 23 (Jan. 22, 23, Feb. 5, 7, 9, 16, 
March 8, 10, 11, 13, 23, June 28, July 8, 21, Aug. 22, Nov. 25, Dec. 4, 11)— Tinian, 2 (Sept. 
7, 14) — Saipan, 3 (July 8, Aug. 5, 22) — Asuncion, 1 (June) — Agrihan, 1 (June). 

Nesting. — Seale (1901 :55) obtained nests and eggs in the period from May to 
July at Guam. He found the nests 8 to 15 feet above the ground. Strophlet 
(1946:540) observed a pair of honey-eaters with two young on October 9 at 
Guam. In 1945 at Guam the NAMRU2 party obtained individuals with en- 
larged gonads on January 22, June 2, 5, July 21 and 23. and found evidence of 
nesting on June 16. Hartert (1898:56) writes that Owston's Japanese collectors 
obtained nests in January, February, and March. Each nest contained two 
eggs; they were placed four to eight feet from the ground. Probably the 
Cardinal Honey-eater in the Marianas nests at most times of the year. 

Molt. — Specimens, with molting plumage, have been examined that were 
taken at most times of the year. I suspect that this bird molts at irregular 
intervals. 

Food habits. — The honey-eater feeds partly on insect life and partly on 
nectar and juices from flowers. At Guam, the honey-eater was frequently 
found at flowers of the ink berry bush, where evidently both nectar and insects 
were obtained. The birds were attracted also to the coconut palms, especially 
when the reproductive parts of the palms were developing. 

Remarks. — The Cardinal Honey-eater is one of the most con- 
spicuous land birds in the Mariana Islands. Its scarlet plumage and 
characteristic fluttering flight cause it to stand out against its habitat 
of forest, scrub, and garden. At Guam, the author (1947b: 124) 
found the honey-eater on 37.6 percent of the 125 roadside birds 
counts made in 1945. The species included 3.9 percent of all of the 
birds observed on these counts. Seale (1901:55) and Strophlet 
(1946:540) also commented on its abundance at Guam; however, 
in 1931, Coultas (field notes) wrote that the bird was rare; he ob- 
tained only one skin at Guam. At Rota, the NAMRU2 party found 
the honey-eater to be abundant. Coultas obtained only a few birds 
at Tinian and Saipan in 1931. In 1945, Downs (1946:103) saw 
only a single pair at Tinian; Gleise (1945:220) estimated the popu- 
lation at Tinian to be 12 in 1945. At Agrihan, Borror (1947:417) 
reported that the honey-eater was a common bird in 1945. 



312 University of Kansas Publs., Mus. Nat. Hist. 

Table 49 lists the measurements of males of M. c. saffordi from 
several islands in the Marianas. Measurements of birds from 
Guam, Rota, Tinian, and Saipan are fairly similar, although the 
birds at Saipan seem to have a slightly longer wing than those at 
Guam. A single skin from Agrigan has larger measurements than 
those of birds obtained in the southern Marianas. Whether the 
birds in the northern Marianas are separable because of larger size 
can only be ascertained by the studying of more material from that 
region. 

Mayr (1945a: 102) writes that males of M. cardinalis seem to 
outnumber the females by approximately four to one. On the basis 
of collections and field observations, the males were found to out- 
number the females in the Micronesian islands; although the ratio 
may not be so great as four to one. At Guam, the NAMRU2 party 
obtained 21 males and 8 females. Although these birds are often 
seen as pairs (male and female), single males are frequently ob- 
served. The females do not appear to have more secretive habits 
than the males. 

Myzomela cardinalis kurodai Momiyama 
Cardinal Honey-eater 

Myzomela rubratra kurodai Momiyama, Birds Micronesia, 1922, p. 17. (Type 
locality. Yap.) 

Myzomela rubratra Hartlaub and Finseli (part), Proc. Soc. London, 1872, pp. 89, 
94 (Uap); Graffe, Journ. Mus. Godeffroy, 2, 1873, p. 122 (Yap); Finsch (part), Journ. 
Mus. Godeffroy, 8, 1875, p. 4 (Yap); Forbes (part), Proc. Zool, Soc. London, 1879, p. 
271 (Yap); Wiglesworth (part), Abhandl. und. Ber. Zool. Mus. Dresden, no. 6, 1890- 
1891 (1891), p.. 31 (Uap); Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 
7, 1895, p. 202 (Yap); Takatsukasa and Kuroda (part). Tori, 1, 1915, p. 64 (Yap); 
Kuroda, Dobutsu. Zasshi, 27, 1915, pp. 331, 332 (Yap). 

Myzomela rubrata Matschie (part), Journ. f. Ornith., 1901, p. 112 (Yap). 

Myzomela rubrata kurodai Kuroda, in Momiyama, Birds Micronesia, 1922, p. 74 
(Yap) ; Mathews, Syst, Avium Australasianarum, 2, 1930, p. 743 (Yap) ; Hand-list 
Japanese Birds (part), rev., 1932, p. 172 (Yap); Hand-list Japanese Birds (part), 3d 
ed., 1942, p. 190 (Yap). 

Myzomela cardinalis kurodai Mayr, Birds Southwest Pacific, 1945, p. 299 (Yap). 

Geographic range. — Micronesia: Caroline Islands — Yap. 

Characteri,. — Adult male: According to Momiyama (1922:17), M. c. kurodai 
is "Similar to M. r. saffordi Wetmore from Southern Marianne islands, but 
the tarsus is decidedly shorter, not e.xceeding 21 mm. (more than 21 mm. in 
M. r. saffordi), and the colour of plumage is not so much tinged with Vermil- 
lion. It differs from M. r. rubratra, M. r. dichromata, and M. r. wetmorei by 
the body measuring much shorter, and by the scarlet colour of plumage being 
less pronounced. The length of bill in M . r. loetniorei and kurodai is nearly 
the same." 

Adult female: According to Momiyama (1922:17), "Upper-parts of body 
dark olivaceous brown ; under-parts, including chin, throat and fore neck like 
upper-parts, but somewhat paler; breast and abdomen yellowish ashy-white; 



Baker: The Avifauna of Micronesia 313 

head, lower back, rump, upper tail-coverts, chin, throat as well as lower breast 
tinged with scarlet (the red colour more distinct on lower back but less so on 
lower breast) ; pale olive margin to the outer web of flight-feathers." 
Measurements. — Measurements are listed in table 48. 

Specimens examined. — Total number, 2 males, from Caroline Islands, AMNH — Yap (Sept.). 

Remarks. — This subspecies is tentatively recognized as distinct 
from M. c. kobayashii of Palau. No female has been examined, 
and the two males seen and the description by Momiyama indicate 
that the population at Yap closely resembles the one at Palau. The 
Hand-list of Japanese Birds (Hachisuka et al., 1932:172) places 
the birds from Yap and Palau in the same subspecies. 

Myzomela cardinalis kobayashii Momiyama 
Cardinal Honey-eater 

Myzomela rubratra kobayashii Momiyama, Birds Micronesia, 1922, p. 19. (Type 
locality, Pelew Islands.) 

Cinnyris rubrater Lesson (part), Diet. Sci. Nat., ed., Levrault, 50, 1827, p. 30 
(Pelew); idem (part), Voy. "La Coquille," Zool., 1, 1828, p. 678 (Pelew); idem (part), 
Man. d'Ornith., 2, 1828, p. 55 (Pelew). 

Myzomela rubratra Gray (part). Cat. Birds Trop. Is. Pacific Ocean, 1859, p. 11 
(Pelew); Hartlaub (part), Proc. Zool. Soc. London, 1867 (1868), p. 829 (Pelew); 
Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 5, 116, 118 (Pelew); Gray 
(part). Hand-list Birds, 1, 1869, p. 154 (Pelew); Hartlaub and Finsch (part), Proc. 
Zool. Soc. London, 1872, pp. 89, 94 (Pelew); Finsch (part), Journ. Mus. Godeffroy, 
8, 1875, pp. 4, 16 (Palau); idem (part), Journ. Mus. Godeffroy, 12, 1876, pp. 17, 
26 (Palau) ; Forbes (part), Proc. Zool. Soc. London, 1879, p. 270 (Pelew) ; Finsch 
(part), Mitth. Ornith. Ver. Wien, 1884, p. 48 (Palau); Gadow, Cat. Birds British 
Mus., 9, 1884, p. 129 (Pelew); Tristram, Cat. Birds, 1889, p. 206 (Pelew); Wigles- 
worth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 31 
(Pelew); Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, p. 202 
(Palaos); Nehrkorn, Kat. Eiers., 1899, p. 79 (Palau-inseln) ; Scale (part), Occ. Papers 
Bernice P. Bishop Mus., 1, 1901, p. 57 (Pelew); Reichenow (part). Die Vogel, 2, 
1914, p. 482 (Palau); Takatsukasa and Kuroda (part). Tori, 1, 1915, p. 64 (Pelew). 

MyzoTnela rubratra Nehrkorn (part), Journ. f. Ornith., 1879, p. 397 (Palau); 
Matschie (part), Journ. f. Ornith., 1901, p. 112 (Palau); Takatsukasa and Kuroda 
(part). Tori, 1, 1915, p. 55 (Pelew); Kuroda, Dobutsu. Zasshi, 28, 1916, p. 71 (Pe- 
lew). 

Myzomela rubratra kobayshii Kuroda, in Momiyama, Birds Micronesia, 1922, p. 
74 (Pelew); Mathews, Syst. Avium Australasianarum, 2, 1930, p. 722 (Pelew). 

Myzomela rubratra kurodai Hand-list Japanese Birds (part), rev., 1932, p. 172 
(Palau); Hand-list Japanese Birds (part), 3d ed., 1942, p. 190 (Babelthuap, Koror, 
Peleliu). 

Mizomela rubratra kurodai Yamashina, Tori, 10, 1940, p. 674 (Palau). 

Myzomela cardinalis kobayashii Mayr, Birds Southwest Pacific, 1945, p. 299 (Pa- 
lau); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 72 (Peleliu). 

Geographic range. — Micronesia: Palau Islands — Babelthuap, Koror, Gara- 
kayo, Peleliu, Angaur. 

Characters. — Adult male: Resembles M. c. rubratra, but smaller and with 
red coloring darker, near "scarlet-red"; margins of wing feathers olivaceous. 
Differs from adult males of other subspecies of M. cardinalis by red coloring 
of feathers being darker. 

Adult female: Resembles adult female of M. c. dichromata but red color- 



314 University of Kansas Publs., Mus. Nat. Hist. 

ing darker, top of head only partly red; abdomen, under tail-coverts, and 
axillaries buff-gray; outer edges of wing and tail feathers light olive. Differs 
from adult females of other subspecies of M. cardinalis by having top of head 
only partly red. 

Immature male: Resembles adult male, but red coloring lighter and thinly 
distributed; wings and tail browish-olive ; abdomen and under tail-coverts 
grayish. 

Immature female : Resembles adult female, but red coloring paler and 
underparts more buffy and less grayish. 

Measurements. — Measurements are listed in table 48. 

Specimens examined. — Total number, 42 (28 males, 11 females, 3 unsexed), as follows: 
Palau Islands, USNM— Koror, 4 (Nov.)— Peleliu, 11 (Aug. 29, 30, 31, Sept. 1, 5); AMNH— 
exact locality not given, 27 (Oct., Nov., Dec). 

Molt. — Many of the specimens taken from late August to December are in 
molt. Of the adult males obtained during this period almost a half had en- 
larged testes. 

Food habits.— Stomachs of specimens obtained by the NAMRU2 party in 
August and September, 1945, contained vegetable matter, seeds and small 
insects. 

Remarks.— Jioney -eaters were found by the NAMRU2 party in 
open woodlands, in coconut groves and about human habitations. 
They were not seen in dense jungle areas, and appeared to prefer 
the plantation areas. 

The Cardinal Honey-eater at Palau is distinguished from other 
subspecies of M. cardinalis in Micronesia by its deeper red coloring. 
In size, it closely resembles the bird at Yap and in the Marianas. 

Evolutionary history of Myzomela cardinalis in Micronesia. — 
The genus Myzomela is found in Australia, northward to Timor, 
Tenimber, Moluccas, Celebes, Melanesia, Polynesia and Micronesia. 
The range of the species M. cardinalis includes the islands from the 
eastern Solomons, New Hebrides, and Loyalty Islands east to central 
Polynesia and north to Micronesia. It appears likely that M. 
cardinalis was derived, probably along with M. nigrita, M. lajargei 
and others, from an ancestral stock in the Melanesian area. Within 
the species M. cardinalis there is one group of subspecies which 
exhibits a marked degree of sexual dimorphism, with the males 
having a much greater amount of red coloration than the females. 
These subspecies occur in the southern part of the range of the 
species (Loyalty, Santa Cruz, New Hebrides, and Samoa islands). 
A second group of subspecies exhibit a lesser amount of sexual 
dimorphism, the females possessing more of the red coloration and 
resembling the males more closely. This second group includes 
subspecies which occur in the more northern part of the range of 
the species (Solomons, Micronesia, and Rotuma islands). The 



Baker: The Avifauna of Micronesia 



315 



males of the various subspecies of M. cardinalis vary one from 
another considerably less than do the females. 

Figure 16 shows the probable routes of colonization used by M. 
cardinalis to attain its present distribution in the Pacific islands. 
The subspecies in the eastern Solomon Islands {M. c. pidchermna 
Ramsey and M. c. sanfordi Mayr) may be representative of the first 
colonization by the supposed ancestral stock. From a focal point in 
this area, M. cardinalis has dispersed by what may be considered as 




120 



140 



160 



180 



160 




'w ./ • 



®*— Q) (5) ••;"'* 




1000 MILES 



20 



c^ 



160. 



_yoLlf 



Fig. 16. Geographic distribution of Myzomela cardinalis and routes of its 
dispersal. (1) Probable center of dispersal of Myzomela; (2) ranges of M. c. 
sanfordi and M. c. pulcherrima in the Solomon Islands; (3) M. c. ruhratra; 
(4) M. c. dichromata ; (5) M. c. major; (6) M. c. kurodai; (7) M. c. kobayashii ; 
(8) M. c. saffordi; (9) M. c. chermesina; (10) range of M. cardinalis in the 

Santa Cruz, New Hebrides, Banks and Loyalty islands; (11) M. c. nigriventris. 

two routes. One route evidently was to the south as far as the 
Loyalty Islands with a side branch extending to the Samoan Islands 
where M. c. nigriventris Peale occurs. The second route extended 
north to the islands of Micronesia. The Caroline Islands were seem- 
ingly inhabited initially, with invasions of the Palaus made via Yap, 
and of the Marianas via Kusaie or Ponape (as indicated by the 
comparison of specimens) . Mayr (in conversation) has pointed out 
the close relationship between the subspecies in Micronesia and M. c. 



316 University of Kansas Publs., Mus. Nat. Hist. 

chermesina Gray of Rotuma Island. This subspecies at Rotiima, 
which is located between Santa Cruz and Samoa, resembles closely 
M. c. dichromata of Ponape, especially in the case of the female. It 
is evident that the honey-eater arrived at Rotuma from Micronesia, 
rather than from the Solomon and Santa Cruz area to the west. 

Zosterops conspicillata conspicillata (Kittlitz) 
Bridled White-eye 

Dicaeum conspicillatum Kittlitz, Kupfertaf. Naturgesch. Vogel, 2, 1833, p. 15, pi. 
19, fig. 1. (Type locality, Guaham.) 

Dicaeum conspicillatum Kittlitz, Mem. Acad. Imp. Sci. St. Petersbourg, 2, 1835, 
p. 3, pi. 4 (Guaham); idem, Obser. Zool., in Lutke, Voy. "Le Seniavine," 3, 1836, p. 305 
(Guaham). 

Zosterops conspicillatum Bonaparte, Consp. Avium, 1, 1850, p. 398 (Mariann. =. 
Guam). 

Zosterops conspicillata Reichenbach. Syn. Avium, 1852, p. 92 (Guaham); Hartlaub, 
Journ. f. Ornith., 1854, p. 187 (Mariannen = Guam) ; Gray, Cat. Birds Trop. Is. 
Pacific Ocean, 1859, p. 16 (Guam); Hartlaub, Journ. f. Ornith., 1865, pp. 5, 17 
(Guaham); Gray, Hand-list Birds, 1, 1869, p. 163 (Ladrone = Guam) ; Hartlaub and 
Finsch, Proc. Zool. Soc. London, 1872, p. 95 (Guaham); Giebel, Thes. Ornith., 3, 
1877, p. 775 (Ladrone = Guam) ; Gadow, Cat. Birds British Mus., 9, 1884, p. 187 
(Guam); Wiglesworth, Abhand. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), 
p. 37 (Guam); Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, p. 205 
(Guam); Hartert (part), Novit. Zool., 5, 1898, p. 57 (Guam); Hartert, Novit. Zool., 
7, 1900, p. 3 (Guam); Matschie (part), Journ. f. Ornith., 1901, pp. 112, 113 (Guam); 
Seale, Occ. Papers Bemice P. Bishop Mus., 1, 1901, p. 58 (Guam); Finsch (part). Das 
Tierreich, no. 15, 1901, p. 37 (Guam); Safford, Osprey, 1902, p. 69 (Guam); Dubois, 
Syn. Avium, 1, 1902, p. 711 (Guam); Safford, The Plant World, 7, 1904, p. 264 
(Guam); idem, Contr. U. S. Nat. Herb., 9, 1905, p. 79 (Guam); Takatsukasa and 
Kuroda (part). Tori, 1, 1901, p. 64 (Marianne = Guam); Cox, Island of Guam, 1917, 
p. 21 (Guam); Mathews, Syst. Avium Australasianarum, 2, 1930, p. 706 (Guam); 
Bryan, Guam. Rec, vol. 13, no. 2, 1936, p. 25 (Guam); Strophlet, Auk, 1948, p. 540 
(Guam). 

Zosterops conspicillatus Kuroda (part), in Momiyama, Birds Micronesia, 1922, p. 
76 (Guam). 

Zosterops conspicillata conspicillata Stresemann, Mitt. Zool. Mus. Berlin, 17, 1931, 
p. 227 (Guam); Hand-list Japanese Birds, rev., 1932, p. 173 (Guam); Hand-list 
Japanese Birds, 3d ed., 1942, p. 192 (Guam); MajT, Amer. Mus. Novit., no. 1269, 
1944, p. 7 (Guam); idem. Birds Southwest Pacific, 1945, p. 299 (Guam); Baker, 
Smithson. Misc. Coll., vol. 107, no. 15, 1948, pp. 72, 73 (Guam). 

Geographic range. — Micronesia: Mariana Islands — Guam. 

Characters. — Adult : A small white-eye with upper parts near "serpentine 
green," becoming slightly lighter on the rump; orbital ring broad and white; 
fronto-loral band light yellowish-white; auriculars grayish-green; chin and 
throat yellowish-white; breast and abdomen dingy yellow; wing and tail 
feathers dark brown with greenish-yellow edges; upper mandible horn colored, 
lower mandible lighter yellow; legs and feet dark olive-gray; iris light umber. 
Adult female may be lighter on underparts. 

Immature: Resembles adult, but underparts paler yellow and upper mandi- 
ble light yellowish-brown. 

Measurements. — Measurements of Z. c. conspicillata are listed in table 50. 
Males and females have measurements which are nearly equal. 



Baker: The Avifauna of Micronesia 



317 



Table 50. Measurements of the Subspecies of Zosterops conspidllata 



Subspecies 



No. 



Wing 


Tail 


56 
(52-59) 


41 
(37-43) 


52 
(50-55) 


38 

(35-40) 


53 

(51-55) 


42 

(42-43) 


55 

(54-57) 


38 
(36-41) 


55 

(52-57) 


36 

(34-38) 


54 

(53-55) 


36 
(34-39) 



Full culmen 



Tarsus 



Z. c. conspicillata 43 



Z. c. saypani. 



Z. c. rotensis. . 



Z. c. semperi. 



Z. c. owstoni. 



Z. c. takatsukasai . 



29 



28 



22 



16 



13.5 
(13.0-14.5) 

12.5 
(12.0-13.5) 

13.0 
(13.0-13.5) 

12.5 
(12.0-13.5) 

12.5 
(12.0-13.0) 

13.0 
(13.0-14.0) 



19 

(18-20) 

18 

(17-19) 

18 
(18-19) 

18 
(17-19) 

19 

(18-20) 

19 
(19-20) 



Weights — The author (1948:73) records the weights of 11 adult males as 
9.5-14.0 (10.5), of 3 adult females as 8.0-10.0 (9.3). 

Specimens examined. — Total number, 61 (33 males, 17 females, 11 unsexed), as follows: 
Mariana Islands, USNM— Guam, 27 (May 24, 2y, 30, June 2, 3, 25, 28, July 12, 18, 19, 20, 
23, 26, Sept., Oct. 8); AMNH — Guam, 34 (Jan., March, July, Aug., Sept., Nov., Dec). 

Nesting. — Seale (1901:58) reports the taking of one nestling and three nests 
with eggs of the bridled white-eye at Ouam in the period from May to July. 
The NAMRU2 party obtained little evidence of nesting in late May to July. 
Three males taken in the period of June and July had enlarged gonads. 
Hartert (1898:57) records several nests taken in February and March at 
Guam. He writes, "The nest is a fairly deep cup, placed in the fork of 
a branch, woven together of fine grasses and roots, and on the outside orna- 
mented with cobwebs, wool and cottonwood, varj'ing in width from 8 to 
5 cm. The clutches consist of 2 or 3 eggs. The eggs are pale blue, like 
all Zosterops eggs. They measure 18:13, 17:13.2, 17:12.2, 15.5:12:5, 17:13.5, 
and between these measurements." Coultas obtained specimens with enlarged 
gonads in August. According to Oustalet (1895:207), Marche found nests and 
young in May or June. 

Remarks. — Kittlitz obtained the Bridled White-eye at Guam, 
when he visited the island, in March, 1828. He found the birds 
common and they reminded him of titmice. Marche obtained a 
series of 21 skins at Guam in August and September, 1887, and in 
February and March, 1888. Seale (1901:58) observed the birds in 
flocks of 10 to 20 in roadside bushes and in vi^aste areas. He men- 
tions that their principal foods are insects. The NAMRU2 party 
found the birds to be restricted to certain areas on Guam, where 
they were found in small flocks moving about in low trees. They 



318 University of Kansas Publs., Mrs. Nat. Hist. 

were taken at only five localities, two of these being at the northern 
end of the island in vegetation along the high, coastal cliffs. The 
other localities were in the central part of the island in low trees in 
the uplands. Strophlet (1946:540) found them in grasslands on the 
foothills. Arvey (field notes) saw a flock of 12 white-eyes at Mount 
Tenjo in July, 1946. 

The white-eye is a very active bird, always moving rapidly 
through the vegetation or flying across open areas to disappear into 
scrub foliage. As they move about they make a twittering sound, 
which is considered to be a flocking call. 

Zosterops conspicillata saypani Dubois 

Bridled White-eye 

Zosterops conspicillata Saypani Dubois, Syn. Avium, 1, 1902, p. 711. (Type locality, 
Say pan.) 

Zosterops conspicillata Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 
1895, p. 205 (Saypan); Hartert (part), Novit. Zool., 5, 1898, p. 57 (Saipan) ; Finsch 
(part), Das Tierreich, no. 15, 1901, p. 37 (Saipan); Matschie (part), Journ. f. Omith., 
1901, pp. 112, 113 (Saipan); Prowazek, Die deutschen Marianen, 1913, p. 101 (Saipan); 
Takatsukasa and Kuroda (part). Tori, 1, 1915, p. 64 (Marianne ^ Saipan). 

Zosterops conspicillata var. saypani Snouckaert, Alauda, (2), 3, 1931, p. 22 (Saypan). 

Zosterops conspicillatus Kuroda (part), in Momiyama, Birds Micronesia, 1922 ; p. 
76 (Saipan). 

Zosterops saipani Mathews, Syst. Avium Australasianarum, 2, 1930, p. 706 (Saipan). 

Zosterops conspicillata saipani Stresemann, Mitt. Zool. Mus. Berlin, 17, 1931, p. 
227 (Saipan); Hand-list Japanese Birds, rev., 1932, p. 173 (Saipan, Tinian); Hand-list 
Japanese Birds, 3d ed., 1942, p. 192 (Saipan, Tinian); Mayr, Amer. Mus. Novit., no. 
1269, 1944, p. 7 (Tinian, Saipan); idem. Birds Southwest Pacific, 1945, p. 299 (Saipan, 
Tinian); Downs, Trans. Kansas Acad. Sci., 49, 1946, p. 104 (Tinian); Stott, Auk, 64, 
1947, p. 527 (Saipan); Baker, Smithson, Misc. Coll., vol. 107, no. 15, 1948, p. 73 
(Saipan, Tinian). 

Zosterops conspicillatus saipani Yamashina, Tori, 7, 1932, p. 398 (Tinian). 

Geographic range. — Micronesia: Mariana Islands — Tinian, Saipan. 

Characters. — Adult: Resembles Z. c. conspicillata, but slightly smaller with 
fronto-loral band more greenish yellow; auriculars olivaceous; orbital ring 
narrower ; upper parts brighter olive ; underparts pale yellowish-white ; bill 
darker. Birds from Saipan resemble closely birds from Tinian, but upper parts 
may be slightly brighter and underparts slightly more yellowish; iris chestnut. 

Measurements. — Measurements are listed in table 50. Twenty-three birds 
from Tinian measure: wing, 51 (50-53) ; tail, 38 (35-41) ; full culmen, 12.0 (12.0- 
13.0) ; tarsus, 18 (17-18) ; six birds from Saipan measure: wing, 54 (52-55) ; tail, 
37 (35-39); full culmen, 13.0 (13.0-15.0); tarsus, 18 (17-19). Birds from Saipan 
are slightly larger than birds from Tinian. 

Specimens examined.- — Total number, 33 (18 males, 13 females, 2 unsexed), as follows: 
Mariana Islands, USNM— 7 (Oct. 7, 8, 9, 10, 23); AMNH— 26 (July, Aug., Sept.). 

Nesting. — Yamashina (1932a :398) records the taking of three nests of the 
Bridled White-eye at Tinian on January 8, 1932. The nests contained one, two, 
and three eggs, respectively. The color of the eggs is uniformly pale blue; the 
nests were situated two to four meters from the ground. Oustalet (1895:207) 
writes that Marche obtained records of nesting at Saipan in the period from 



Baker: The Avifauna of Micronesia 319 

May to July. Of 18 birds taken by Coultas at Tinian in September, 1931, one- 
half of them had enlarged gonads. 

Molt. — Specimens examined that were taken in July, August, September, 
and October have molting plumage. 

Remarks. — Marche obtained the first skins of this white-eye at 
Saipan; he got 23 specimens in May, June, and July, 1887. The 
population at Saipan was initially considered similar to that at 
Guam; it was later given subspecific separation by Dubois. The 
birds at Tinian exhibit some differences from the birds at Saipan, 
and it is possible that these two populations should be regarded as 
subspecifically distinct from one another. 

In 1931, Coultas (field notes) found this white-eye common at 
Saipan and Tinian. He writes "The little fellow has adjusted him- 
self to the gardens and shrubs in the villages. He is a seed eater 
and makes himself at home now around human habitation. I have 
seen him climbing over potted plants on the window ledges of 
dwellings. His cheerful little sibilation uttered continuously while 
at work or while on the wing makes him friends wherever he goes. 
He is no longer a bird of the forest as he has none here to go to." 
Several observers in the late war have published notes on this 
white-eye. Stott (1947:527) writes that he was reminded of the 
bush-tit {Psaltriparus) when he observed the behavior of this white- 
eye; Moran (1946:262) writes that it is "Similar in size and be- 
havior to our vireos." Gleise (1945:220) estimated the population 
of white-eyes at Tinian at 500 plus in 1945. Downs (1946:104-105) 
found the birds to be abundant at Tinian; he found them in small 
flocks in low brush or trees and at edges of open fields as well as 
elsewhere. He saw a white-eye eating "a large green fuzzy cater- 
pillar." 

Zosterops conspicillata rotensis Takatsukasa and Yamashina 

Bridled White-eye 

Zosterops semperi rotensis Takatsukasa and Yamashina, Dobutsu. Zasshi, 43, 1931, 
p. 486. (Type locality, Rota.) 

Zosterops semperi Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, 
p. 207 (Rota); Hartert, (part), Novit. Zool., 5, 1898, p. 57 (Rota); Finsch (part), 
Das Tierreich, no. 15, 1901, p. 30 (Rota); Scale (part), Dec. Papers Bernice P. Bishop 
Mus., 1, 1901, p. 58 (Rota); Dubois (part), Syn. Avium, 1, 1902, p. 710 (Rota); 
Takatsukasa and Kuroda (part). Tori, 1, 1915, p. 64 (Marianne=Rota). 

Zosterops semperi semperi Momiyama (part). Birds Micronesia, 1922, p. 23 (Rota) ; 
Kuroda, (part) in Momiyama, Birds Micronesia, 1922, p. 75 (Rota). 

Zosterops semperi rotensis Snouckaert, Alauda. (2), 4, 1932, p. 459 (Rota); Yama- 
shina, Tori, 7, 1932, p. 399 (Rota); Hand-list Japanese Birds, rev., 1932, p. 173 
(Rota). 

Zosterops conspicillata rotensis Hand-list Japanese Birds, 3d ed., 1942, p. 193 
(Rota); Mayr, Amer. Mus. Novit., no. 1269, 1944, p. 7 (Rota); idem. Birds Southwest 
Pacific, 1945, p. 299 (Rota); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 
73 (Rota). 



320 University of Kansas Publs., Mus, Nat. Hist, 

Geographic range. — Micronesia: Mariana Islands — Rota. 

Characters. — Adult: Upper parts and sides of neck between "warbler 
green" and "pyrite yellow" becoming lighter on the rump; auriculars light 
yellowish-green; orbital ring white; fronto-loral band narrowly tinged with 
yellow; underparts dingy yellow; wing and tail feathers dark with light 
greenish-yellow edges; upper mandible light brown; lower mandible light 
yellowish-brown; feet light brown. 

Resembles Z. c. conspicillata, but brighter greenish-yellow above; chin and 
throat yellow like rest of underparts; fronto-loral band tinged with bright 
yellow; auriculars resemble closely the upper parts in color; narrow orbital 
ring. 

Measurements. — Measurements are listed in table 50. 

Specimens examined. — Total number, 5 (3 males, 1 female, 1 unsexed), from Mariana 
Islands, USNM— Rota (Oct. 18, 20, 22). 

Nesting. — Yamashina (1932a :399) records the taking of one nest contain- 
ing two eggs at Rota on March 7, 1931- 

Molt. — Specimens taken in October were in molt. 

Remarks. — Oustalet (1895:207) reported on two specimens of 
white-eye taken at Rota by Marche. He considered them as being 
similar to the birds at Palau. The birds at Rota were named as 
a separate subspecies by Takatsukasa and Yamashina in 1931. The 
NAMRU2 party found the birds to be numerous at Rota in October, 
1945. 

Zosterops conspicillata semperi Hartlaub 
Bridled White-eye 

Zosterops semperi Hartlaub, in Hartlaub and Finsch, Proc. Zool. Soc. London, 
1868, p. 117. (Type locality, Pelew Islands.) 

Zosterops semperi Hartlaub and Finsch, Proc. Zool. Soc. London, 1872, pp. 89, 95 
(Pelew) ; Finsch, Joum. Mus. Godeffroy, 8, 1875, pp. 4, 16, pi. 4, fig. 1 (Palau) ; 
Giebel, Thes. Ornith., 3, 1877, p. 777 (Pelew); Nehrkorn, Journ. f. Ornith., 1879, 
p. 396 (Palau); Finsch (part), Journ. f. Ornith., 1880, p. 286 (Palau); idem (part). 
Ibis, 1881, p. Ill (Pelew); Schmaltz and Krause (part), Ethnogr. Abth. Mus. Godef- 
froy, 1881, p. 407 (Palau); Finsch (part), Mitth. Ornith. Ver. Wien, 1884, p. 48 (Pa- 
lau); Gadow (part). Cat. Birds British Mus., 9, 1884, p. 183 (Pelew); Tristram, 
Cat. Birds, 1889, p. 212 (Pelew) ; Wiglesworth (part), Abhandl. und Ber. Zool. Mus. 
Dresden, no. 6, 1890-1891 (1891), p. 37 (Pelew); Oustalet (part), Nouv. Arch. Mus. 
Hist. Nat. Paris, (3), 7, 1895, p. 208 (Palaos) ; Hartert (part), Novit. Zool., 5, 1898, 
p. 57 (Pelew); Matschie, Journ. f. Ornith., 1901, pp. 112, 113 (Palau); Finsch (part). 
Das Tierreich, no. 15, 1901, p. 30 (Palau); Scale (part), Occ. Papers Bernice P. 
Bishop Mus., 1, 1901, p. 58 (Pelew); Dubois (part), Syn. Avium, 1, 1902, p. 710 
(Palau); Takatsukasa and Kuroda (part). Tori, 1, 1915, pp. 55, 64 (Pelew). 

Zosterops semperi semperi Hartert, Novit. Zool., 7, 1900, p. 2 (Pelew); Momiyama 
(part). Birds Micronesia, 1922, pp. 22, 23 (Pelew); Kuroda (part), in Momiyama, 
Birds Micronesia, 1922, p. 75 (Pelew); Mathews, Syst. Avium Australasianarum, 2, 
1930, p. 705 (Pelew); Takatsukasa and Yamashina, Dobutsu. Zasshi, 43, 1931, p. 486 
(Pelew); Hand-list Japanese Birds, rev., 1932, p. 174 (Palau). 

Zosterops conspicillata semperi Stresemann, Mitt. Zool. Mus. Berlin, 17, 1931, p. 
227 (Palau); Hand-list Japanese Birds, 3d ed., 1942, p. 193 (Babelthuap, Koror, 
Peliliu); Mayr, Amer. Mus. Novit., no. 1269, 1944, p. 7 (Palau); Mayr, Birds South- 
west Pacific, 1945, p. 299 (Palau); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 
1945, p. 73 (Garakayo). 



Baker: The Avifauna of Micronesia 321 

Geographic range. — Micronesia: Palau Islands — Babelthuap, Koror, Gara- 
kayo, Peleliu. 

Characters. — Adult: Resembles adult of Z. c. rotensis, but fronto-loral band 
lighter yellow, and coloring is usually not continuous above; auriculars paler; 
breast and abdomen paler yellow; maxilla and feet darker; mandible whitish. 
Resembles adult of Z. c. conspicillata, but brighter greenish-yellow above ; 
coloring of chin and throat like that of rest of undcrparts; auriculars colored 
like back; fronto-loral band narrowly tinged with bright yellow and not com- 
pletely connected above; orbital ring narrow; iris grayish-white. 

Measurements. — Measurements are listed in table 50. 

Specimens examined. — Total number, 30 (15 males, 14 females, 1 unsexed), as follows: 
Palau Islands, USNM— Babelthuap, 2 (Nov. 27)— Koror, 4 (Nov. 14, 19)— Garakayo, 4 
(Sept. 18, 19); AMNH — exact locality not given, 20 (Oct., Nov., Dec). 

Molt. — All birds examined (taken in September, October, and November) 
are in molting plumage. 

Food habits. — At Garakayo, birds were observed in small flocks feeding in 
low trees. Two stomachs examined, which were from individuals of these 
flocks, contained very small seeds. 

Remarks. — Oustalet (1895:207) first pointed out the realtionship 
between the Bridled White-eye at Palau and the one at Rota. 
Hartert (1898:57) thought that the occurrence of the same kind of 
bird at Palau and at Rota was "very peculiar." It was not until 1931 
that Takatsukasa and Ynmashina separated the two populations by 
name. 

Coultas (field notes) found the Bridled White-eye to be uncommon 
in the Palaus in 1931. He observed them in the tops of trees, noting 
that they were wary and easily frightened away by the shooting of a 
gun. Coultas writes that he found the birds to be numerous at 
Peleliu ; in 1945, the NA]\IRU2 party did not find the birds at that 
island. The only locality where they were found to occur was on 
the small island of Garakayo where the writer shot four Bridled 
White-eyes on September 18 and 19. He found two or three small 
flocks in low trees near the summit of a hill on the island. Approx- 
imately 25 birds were in this area. 

Zosterops conspicillata owstoni Hartert 
Bridled White-eye 

Zosterops semperi owstoni Hartert, Novit., Zool., 7, 1900, p. 2. (Type locality, 
Ruk.) 

Zosterops semperi semperi Finsch (part), Journ. f. Ornith., 18S0, p. 287 (Ruck); 
idem (part), Proc. Zool. Soc. London, 1880, p. 575 (Ruk); idem (part). Ibis, 1881, p. 
110 (Ruk) ; Schmeltz and Krause (part), Ethnogr. Abth. Mus. Godeffroy, 1881, p. 353 
(Ruk); Gadow (part), Cat. Birds British Mus., 9, 1884, p. 183 (Central Carolines = 
Truk); Wiglesworth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 
(1891), p. 37 (Ruk); Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 
1895, p. 208 (Ruk); Hartert (part), Novit. Zool., 5, 1898, p. 57 (Ruk); Nehrkorn, 
Kat. Eiers, 1899, p. 80 (Ruk). 

21—8131 



322 University of Kansas Publs., Mrs. Nat. Hist. 

Zosierops semperi owstoni Dubois, Syn. Avium, 1, 1902, p. 710 (Ruk); Wetmore, 
in Townsend and Wetmore, Bull. Mus. Comp. Zool., 63, 1919, p. 223 (Tnik) ; Momi- 
yama, Birds Micronesia, 1922, p. 24 (Ruk) ; Kuroda, in Momiyama, Birds Micronesia, 
1922, p. 75 (Ruk) ; Mathews, Syst. Avium Australasianarum, 2, 1930, p. 705 (Ruk) ; 
Takatsukasa and Yaniashina, Dobutsu. Zasshi, 43, 1931, p. 496 (Ruk); Yamashina, 
Tori, 7, 1932, p. 400 (Truk) ; Hand-list Japanese Birds, rev., 1932, p. 174 (Truk). 

Zosterops owstoni Finsch, Das Tierreich, no. 15, 1901, p. 31 (Ruk); Matschie 
(part), Joum. f. Ornith., 1901, pp. 112, 113 (Ruck); Reichenow, Die Vogel, 2, 1914, 
p. 470 (Karolinen = Truk) ; Takatsukasa and Kuroda, Tori, 1, 1915, pp. 55, 64 (Ruk). 

Zosterops conspicillaia owstoni Streseniann, Mitt. Zool. Mus. Berlin, 17, 1931, p. 
277 (Truk); Hand-list Japanese Birds, 3d ed., 1942, p. 193 (Truk); Mayr, Amer. 
Mus. Novit., no. 1269, 1944, p. 7 (Truk); idem, Birds Southwest Pacific, 1945, p. 
299 (Truk); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, pp. 73, 74 (Truk). 

Geographic range. — Micronesia: Caroline Islands — Truk. 

Characters. — Adult : Resembles adult of Z. c. semperi, but upper parts darker 
olive and less yellowish-green; fronto-loral band deeper yellow; auriculars 
slightly darker; black line on lores and under eye more distinct; underparts 
deeper yellow; abdomen with greenish tinges. Resembles adult of Z. c. 
rotensis, but upper parts duller, more green and less yellow; fronto-loral band 
lighter and less distinct, coloring near that of Z. c. semperi; auriculars darker 
green; underparts slightly darker, more olive-green and less yellow. 

Measurements. — Measurements are listed in table 50. 

Specimens examined. — Total number, 23 (12 males, 10 females, 1 unsexed), as follows: 
Caroline Islands, USNM— Truk, 3 (Feb. 16); AMNH— Truk, 20 (Feb., March, May, Nov.). 

Nesting. — Yamashina ( 1932a :400) records the taking of a nest with one egg 
at Truk in May. Hartert (1900:2) records nests containing single eggs taken 
at Truk from May to July. Nests were found in bushes and trees four to 
eight feet above the ground. The eggs are pale blue. He gives measurements 
of seven eggs. 

Remarks. — Kubary obtained the first specimens of the Bridled 
White-eye at Truk. Hartert described the population as a new sub- 
species using material taken by Owston's collectors. The bird was 
named in honor of Alan Owston. McElroy of the NAMRU2 party 
visited Truk in December, 1945. He found this white-eye in the 
mountainous areas at Moen and Udot islands. 

Zosterops conspicillata takatsukasa! Momiyama 
Bridled White-eye 

Zosterops semperi takatsukasai Momiyama, Bird? Micronesia, 1922, p. 22. (Type 
locality, Ponape.) 

Zosterops semperi (part), Finsch, Journ. f. Omith., 1880, p. 286 (Ponape); idem 
(part), Proc. Zool. Soc. London, 1880, p. 575 (Ponape); idem (part). Ibis, 1881, p. 
115 (Ponape) ; Schmeltz and Krause (part), Ethnogr. Abth. Mus. Godeffroy, 1881, p. 
281 (Ponape); Finsch (part), Mitth. Ornith. Ver. Wien, 1884, p. 48 (Ponape); Gadow 
(part), Cat. Birds British Mus., 9, 1884, p. 183 (Central Carolines, Ponape); Wigles- 
worth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 37 
(Ponape); Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, p. 208 
(Ponapi); Finsch, Das Tierreich, no. 15, 1901, p. 30 (Ponape); Dubois (part), Syn, 
Avium, 1, 1902, p. 710 (Ponape); Takatsukasa and Kuroda (part), Tori, 1, 1915, pp. 
65, 64 (Ponape). 

Zosterops owstoni Matschie (part), Journ. f. Omith., 1901, pp. 112, 113 (Ponapi). 

Zosterops semperi takatsukasai. Kuroda, in Momiyama, Birds Micronesia, 1922, p. 



Baker: The Avifauna of Micronesia 323 

76 (Ponape) ; Mathews, Syst. Avium Australasianarum, 2, 1930, p. 705 (Ponape) ; 
Snouchaert, Alauda, (2), 3, 1931, p. 22 (Ponape) ; Takatsukasa and Yamashina, Tori, 
7, 1932, p. 400 (Ponape); Hand-list Japanese Birds, rev., 1932, p. 174 (Ponape). 

Zosterops conspicillata takatsukasai Stresemann, Mitt. Zool. Mus. Berlin, 17, 1931, 
p. 227 (Ponape); Hand-list Japanese Birds, 3d ed., 1942, p. 193 (Ponape); Mayr, 
Amer. Mus. Novit., no. 1269, 1944, p. 7 (Ponape); idem. Birds Southwest Pacific, 
1945, p. 299 (Ponape); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 73 
(Ponape). 

Zosterops conspicillata Mayr, Proe. 6th Pacific Sci. Congr., 4, 1941, p. 204 (Ponape). 

Geographic range. — Micronesia: Caroline Islands — Ponape. 

Characters. — Adult: Resembles adult of Z. c. semperi, but slightly smaller 
with fronto-loral area more sulfur-yellow; underparts brighter, especially the 
coloring of the abdomen and under tail-coverts; iris light chestnut. 

Measurements. — Measurements are listed in table 50. 

Specimens examined. — Total number, 20 (10 males, 9 females, 1 unsexed) from Caroline 
Islands, AMNH — Ponape (Nov., Dec). 

Nesting. — Yamashina (1932a :400) records nests and eggs of Z. c. takatsu- 
kasai. The nests, each containing a single egg, were taken on Julj^ 10 and 20, 
1931. Coultas (field notes) writes that the nest consists of a small, cup-shaped 
structure of grasses and hair. The natives told him that two eggs were laid. 
In birds taken by Coultas in November the gonads were beginning to enlarge; 
specimens taken in December had swollen gonads. From the evidence at hand, 
it would appear that the Bridled White-eye at Ponape breeds at two periods of 
the year, the winter and the summer. 

Molt. — Specimens examined, which were taken by Coultas in November and 
December, are in fre.«h plumage. 

Remarks. — In 1931, Coultas (field notes) found this white-eye to 
be rare at Ponape. He obtained almost every one that he saw to get 
his series of 20 specimens. He found the birds usually in pairs 
around yellow-flowering bushy trees. A specimen taken by Richards 
had "small insects" in its stomach. 

Zosterops conspicillata hypolais Hartlaub and Finsch 
Bridled White-eye 

Zosterops hypolais Hartlaub and Finsch, Proc. Zool. Soc. London, 1872, p. 95. (Type 
locality, Uap.) 

Zosterops hypolais Griiffe, Journ. Mus. Godcffroy, 2, 1873, p. 122 (Yap); Giebel, 
Thes. Omith., 3, 1877, p. 776 (Carolinae = Yap) ; Schmeltz and Krause, Ethnogr. 
Abth. Mus. Godeffroy, 1881, p. 391 (Yap); Gatlow. Cat. Birds Briti.sh Mus., 9, 1884, 
p. 186 (Uap); Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 
(1891), p. 37 (Uap); Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, p. 208 
(Uap); Bolau, Mitteil. Naturhist. Mus. Hamburg, 1898, p. 60 (Yap); Finsch, Das 
Tierreich, no. 15, 1901, p. 24 (Yap); Matschie, Journ. f. Omith., 1901, pp. 112, 113 
(Yap); Dubois, Syn. Avium, 1, 1902, p. 708 (Uap); Reichenow, Die Vogel, 2, 1914, 
p. 469 (Karolinen = Yap); Takatsukasa and Kuroda, Tori, 1, 1915, p. 64 (Mackenzie 
= Yap) ; Kuroda, in Momiyama, Birds Micronesia, 1922, p. 76 (Yap) ; Mathews, Syst. 
Avium Australasianarum, 2, 1930, p. 700 (Yap); Hand-list Japanese Birds, 3d ed., 
1942, p. 192 (Yap). 

Zosterops conspicillata hypolais Stresemann, Mitt. Zool. Mus. Berlin, 17, 1931, p. 
227 (Yap); Mayr, Amer. Mus. Novit., no. 1269, 1944, p. 7 (Yap); idem. Birds South- 
west Pacific, 1945, p. 299 (Yap). 

Zosterops hyolais Hand-list Japanese Birds, rev., 1932, p. 173 (Yap). 



324 University of Kansas Publs., Mus. Nat. Hist. 

Geographic range. — Micronesia: Caroline Islands — Yap. 

Characters. — According to Hartlaub and Finsch (1872:95), "Upper parts of 
a pale greyish green, throat and under tail-coverts a pure but very pale whitish- 
yellow; breast and abdomen of a mixed pale grey and pale yellow; wing- and 
tail-feathers pale blackish, margined with greenish colour of the back; under 
wing-coverts and inner margins of remiges white; eye-ring indistinct; beak 
fuscous, the under mandible paler, except at the tip; feet plumbeous." 

Remarks. — No specimen has been examined by me. I am fol- 
lowing Stresemann (1931:227) in placing the Bridled White-eye at 
Yap as a subspecies of Z. conspicillata. This is one arrangement; 
the committee who prepared the Hand-list of Japanese Birds (1942: 
192) treat this bird as a separate species. The Japanese probably 
have more specimens of this bird than anyone else and may be in 
a better position to judge its taxonomic status. Specimens of this 
white-eye were taken by Fisher in 1946 at Yap. His report (soon 
to be published) may throw additional light on the degree of dis- 
tinctness of Z. c. hypolais. On the basis of published descriptions 
it is evident that Z. c. hypolais has a few characters in common with 
other members of the species. 

Evolutionary history of Zosterops conspicillata. — The small olive- 
green and yellow white-eyes of Micronesia have been considered as 
belonging to several species by authors in the past. As late as 1930, 
Mathews (1930; 700, 706) placed them in four species. Stresemann 
(1931a:227) put them all in the species Z. conspicillata, an arrange- 
ment which is being followed in this report. It is evident, however, 
that these subspecies of Z. conspicillata can be associated into three 
groups. The author (1948:73) states that Z. c. conspicillata and 
Z. c. saypani have pale chins and throats, light fronto-loral bands, 
blackish coloring at the bend of the wings and broad, white orbital 
rings. Another group, Z. c. rotensis, Z. c. semperi, Z. c. owstoni, and 
Z. c. takatsukasai, have bright yellow chins and throats, matching 
the rest of the underparts, obscure fronto-loral bands, which are 
narrowly tinged with yellow, yellowish coloring at the bend of the 
wings, and narrow, white orbital rings. Z. c. hypolais apparently 
falls into a third group by itself, as indicated by the published de- 
scriptions. There is apparently some variation in the color of the 
eyes of these subspecies; they may be either whitish or chestnut in 
color. The data are insufficient to determine the significance of 
this color character. 

Z. conspicillata is restricted to Micronesia and appears to have 
little close relationship to other species of the genus. Z. conspicil- 



Baker: The Avifauna of Micronesia 325 

lata shows little affinity to white-eyes to the north and northwest 
of Micronesia belonging to the species Z. japonica, of which repre- 
sentatives are found in the Bonin and Volcano islands. Z. conspic- 
illata shows greater affinity to species found to the west and to the 
south of Micronesia. 

It may have colonized Micronesia from the south or southeast 
(Polynesia), even though the species is absent at Kusaie; however, 
Z. conspicillata shows more relationships to species now living to 
the westward and the southwestward, and it probably invaded Mi- 
cronesia from some place in that direction. Z. conspicillata differs 
from species found in Melanesia and Malaysia chiefly in color of 
the forehead, lores, fronto-loral band, crown, nape, breast, abdomen, 
orbital ring, and bill. Also there are differences in the breadth of 
the orbital ring. 

Z. conspicillata shows evidence of relationships with Z. nigrorum 
of the Philippines and Z. montanus of the Philippines and other 
parts of Malaysia. Z. nigrorum resembles Z. c. semperi of Palau in 
size, but is brighter yellow-green above with a darker and less 
curved bill and brighter undcrparts. The fronto-loral band and the 
lores are colored the same in Z. nigrorum and Z. c. semperi. Z. mon- 
tanus resembles Z. conspicillata especially in size and in shape of 
the bill. Z. lutea intermedia of the Makassar area shows some 
affinity to Z. conspicillata, although the bill is heavier. The Micro- 
nesia species also bears a close resemblance to Z. griseotincta of the 
Papuan region. This is especially true of Z. c. takatsukasai at Po- 
nape; however, Z. griseotincta has a heavier and larger bill. Z. 
lateralis from southern Melanesia and Australia is not very different 
from Z. conspicillata aside from its grayish and brownish coloring. 

Z. c&rispicillata probably was derived from an ancestral stock 
which came to Micronesia from the Philippine or Moluccan area, 
rather than directly from Melanesia. Z. conspicillata seemingly 
shows the closest resemblance to Z. nigrorum or to some of its rela- 
tives in the Australo-Moluccan area. The subspecies at Palau, Z. c. 
semperi, appears to be the connecting link. Whether the form at 
Yap represents an independent colonization is not known; such 
might also be true in the case of the subspecies at Guam and at 
Saipan and Tinian. If these are considered as separate coloniza- 
tions, then the populations can be regarded as separate species. 
Mayr, (in conversation) has pointed out the affinity of the white- 
eye at Samoa, Z. samoensis, with Z. conspicillata and suggests that 
Z. samoensis is derived from the Micronesian species. 



326 



University of Kansas Publs., Mus. Nat. Hist. 



Zosterops cinerea cinerea (Kittlitz) 
Micronesian Dusky White-eye 

Drepanis cinerea Kittlitz, Kupfertaf. Naturgesch. Vogel, 1, 1832, p. 6, pi. 8, fig. 2. 
(Type locality, Ualan ^ Kusaie.) 

Drepanis cinerea Kittlitz, Mem. Acad. Imp. Sci., St. Petersbourg, 2, 1835, p. 4, pi. 
5 (Ualan); idem, Obser. Zool., in Lutke, Voy. "Le Seniavine," 3, 1836, p. 285 (Ualan); 
Reichenbach, Syn. Avium, 1853, p. 242 (Ualan) ; Kittlitz, Denkw. Reise, russ. Amer. 
Micron, und Kamchat., 1, 1858, p. 367 (Ualan). 

Zosterops cinerea Hartlaub, Archiv f. Naturgesch., 18, 1852, p. 131 (Ualan); Gray, 
Cat. Birds Trop. Is. Pacific Ocean, 1859, p. 16 (Oualan) ; idem. Hand-list Birds, 1, 1869, 
p. 163 (Caroline = Kusaie) ; Hartlaub and Finsch, Proc. Zool. See. London, 1872, p. 96 
(Ualan); Finsch, Journ. Mus. Godeffroy, 12, 1876, p. 27 (Ualan); idem. Ibis, 1881, pp. 
107, 108 (Kuschai); Gadow, Cat. Birds British Mus., 9, 1884, p. 198 (Kushai); Tris- 
tram, Cat. Birds, 1889, p. 210 (Kuschai) ; Wiglesworth, Abhandl. und Bar. Zool. Mus. 
Dresden, no. 6, 1890-1891 (1891), p. 36 (Ualan); Hartert, Kat. Vogelsamml., Senckenb., 
1891, p. 31 (Ualan); Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, p. 208 
(Oualan); Finsch, Das Tierreich, no. 15, 1901, p. 45 (Kusaie); Dubois, Syn. Avium, 1, 
1902, p. 713 (Kusaie); Takatsukasa and Kuroda, Tori, 1, 1915, pp. 55, 64 (Kusaie); 
Wetmore, in Townsend and Wetmore, Bull. Mus. Comp. Zool., 63, 1919, p. 224 
(Kusaie); Stresemann, Mitt. Zool. Mus. Berlin, 17, 1931, p. 230 (Kusaie); Hand-list 
Japanese Birds, rev., 1932, p. 173 (Kusaie); Hand-list Japanese Birds, 3d ed., 1942, 
p. 192 (Kusaie). 

Dicaeum cuiereum Hartlaub, Journ. f. Ornith., 1854, p. 168 (Carolinen r= Kusaie). 

Zosterops cinereus Finsch, Journ. Mus. Godeifroy, 8, 1875, p. 17 (Ualan); idem, 
Journ. f. Ornith., 1880, pp. 286, 297, 300 (Kuschai); idem, Mitth. Ornith. Ver. Wien, 
1884, p. 48 (Kuschai). 

Zosterops Kittlitzi Finsch, Journ. f. Ornith., 1880, p. 300 (Type locality, Kusaie); 
Reichenow and Schalovv, Journ. f. Ornith., 1881, p. 94 (Kusaie?). 

Tephras cinereiis Matschie, Journ. f. Ornith., 1901, pp. Ill, 112, 113 (Ualan). 

Tephras cinerea Kuroda, in Momiyama, Birds Micronesia, 1922, p. 77 (Kusaie); 
Mathews, Syst. Avium Australasianarum, 2, 1930, p. 712 (Oualan). 

Zosterops cinerea cinerea Mayr, Amer. Mus. Novit., no. 1269, 1944, p. 7 (Kusaie?); 
idem. Birds Southwest Pacific, 1945, p. 300 (Kusaie). 

Geographic range. — Micronesia: Caroline Islands — Kusaie. 

Characters. — Adult: A small, dusky white-eye with upper parts smoky 
olivaceous-gray ; lores dingy white ; auriculars brownish ; no white orbital ring ; 
wing and tail feathers dark brownish-gray with paler greenish-gray outer edges; 
underparts pale ashy-gray, chin lighter, flanks darker; bill black; feet light 
brown; iris brown. 

Measurements. — Measurements of Z. cinerea are listed in table 51. Males 
and females have approximately equal measurements. 

Table 51. Measurements of Zosterops cinerea 



Subspecies 



Z. c. cinerea. 



Z. c. ponapensis . 



Z. c. finschii. 



No. 



47 



38 



30 



Wing 



63 
(60-65) 

59 

(57-61) 

65 

(63-67) 



Tail 



37 
(35-39) 

38 

(36-40) 

43 

(40-46) 



Culmen 



15.0 
(14.0-16.5) 

13.5 
(13.0-14.5) 

17.5 
(16.0-18.5) 



Tarsus 



20 

(19-20) 

20 

(18-21) 

21 

(20-23) 



Baker: The Avifauna of Micronesia 327 

Specimens examined.— Total number, 50 (33 males, 17 females), as follows: Caroline 
Islands, USNM— Kusaie, 1 (Feb. 9); AMNH— Kusaie, 49 (Jan., Feb., March). 

Nesting.— Coulias found that approximately one-half of the males which he 
obtained in March, 1931, had swollen gonads. 

Molt.— Many of the birds obtained in January and February were molting, 
and many of those obtained in March were in fresh plumage. 

Remarks.— Coultas obtained a large series of these birds at 
Kusaie in 1931, where he found them to be common. 

Zosterops cinerea ponapensis Finsch 
Micronesian Dusky White-eye 

Zosterops ponapensis Finsch, Proc. Zool. Soc. London, 1875 (1876), p. 643. (Type 
locality, Ponape.) 

Zosterops ponapensis Finsch, Journ. Mus. Godeffroy, 12, 1876, pp. 17, 27, pi. 2, 
fig. 1 (Ponape); idem. Proc. Zool. Soc. London, 1877 (1878), p. 778 (Ponape); 
Nehrkorn, Journ. Mus. Godeffroy, 1879, p. 396 (Ponape?); Finsch, Journ. f. Omith., 
1880, pp. 286, 300 (Ponape); idem, Ibis. 1881, pp. 110, 111, 115 (Ponape); Schmeltz 
and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, p. 281 (Ponape); Finsch, Mitth. 
Ornith. Ver. Wien, 1884, p. 48 (Ponape); Gadow, Cat. Birds British Mus., 9, 1884, 
p. 198 (Ponape); Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890- 
1891 (1891), p. 36 (Ponape); Bolau. Mitfeil. Naturhist. Mus. Hamburg. 1898, p. 60 
(Ponape); Nehrkorn, Kat. Eiers., 1899, p. 80 (Ponape); Finsch, Das Tierreich, no. 
15, 1901, p. 46 (Ponape); Dubois, Syn. Avium, 1, 1902, p. 713 (Ponape); Reichenow, 
Die Vogel, 2, 1914, p. 470 (Ponape); Takatsukasa and Kuroda, Tori, 1, 1915, pp. 55, 
65 (Ponape); Stresemann, Mitt. Zool. Mus. Berlin, 17, 1931, p. 230 (Ponape); Yama- 
shina, Tori, 7, 1932, p. 397 (Ponape); Hand-list Japanese Birds, rev., 1932, p. 173 
(Ponape); Mayr, Proc. Gth Pacific Sci. Congr., 4, 1941, p. 204 (Ponape); Hand-list 
Japanese Birds, 3d ed., 1942, p. 192 (Ponape). 

Tephras ponapensis Matschie, Journ. f. Ornith., 1901, pp. Ill, 112, 113 (Ponape); 
Kuroda, in Momiyama, Birds Micronesia, 1922, p. 77 (Ponape); Mathews, Syst. Avium 
Australasianarum, 2, 1930, p. 712 (Ponape). 

Zosterops ponapenensis Wetmore, in Townsend and Wetmore, Bull. Mus. Comp. 
Zool., 63, 1919, p. 224 (Ponape). 

Zosterops cinerea ponapensis Mayr, Amer. Mus. Novit., no. 1269, 1944, p. 7 
(Ponape?); idem. Birds Southwest Pacific, 1945, p. 300 (Ponape). 

Geographis range. — Micronesia : Caroline Islands Ponape. 

Characters. — Adult: Resembles adult of Z. c. cinerea, but smaller with upper 
parts umber-brown, forehead pale gray; underparts mostly pale gray, sides 
of breast and abdomen brownish-buff;; under tail-coverts pale buffy-gray. 

Measurements. — Measurements are listed in table 51. 

Specimens examined. — Total number, 47 (28 males, 17 females, 2 unsexed), as follows: 
Caroline Islands, USNM — Ponape, 1 (Feb. 11); AMNH — Ponape, 46 (Nov., Dec). 

Nesting. — Yamashina (1931a: 397-398) describes two nests of Z. c. pona- 
pensis, each containing one egg. These were taken at Ponape on August 4 and 
11, 1931. The nests were located 2.5 meters from the ground. The eggs are 
light blue and pale greenish-blue in color; one measures 18.5 by 13.5. He 
writes, "The nest consists of two layers, the inner and the outer. The outer 
layer is made of fine roots, fibers, leaves and petals, interwoven with a large 
quantity of cotton-wool, and the inner layer is made of fibers of fine roots 
only." Coultas found that a large number of birds taken in November had 
enlarged gonads, especially the males; in December, fewer birds with swollen 
gonads were obtained. 



328 University of Kansas Publs., Mus. Nat. Hist. 

Remarks. — Coultas found this white-eye to be common at Ponape, 
when he visited that island in November and December, 1930. 
He observed the birds in flocks and found them noisy and quarrel- 
some. They feed in bushes and small trees on seeds and insects. 
Richards obtained "small large-seeded blackish berries" from the 
stomach of a female from Ponape. He found the birds to frequent 
low altitudes in and about native gardens. 

Zosterops cinerea finschii (Hartlaub) 
Micronesian Dusky White-eye 

Tephras finschii Hartlaub, in Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, 
p. 6, pi. 3. (Type locality, Pelew Islands.) 

Tephras finschii Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 117, 118 
(Pelew Islands). 

Zosterops finschii Gray, Hand-list Birds, 1, 1869, p. 164 (Pelew); Gadow, Cat. 
Birds British Mus., 9, 1884, p. 197 (Pelew); Bolau, Mitteil. Naturhist. Mus. Ham- 
burg, 1898, p. 60 (Palau). 

Zosterops fitischi Hartlaub and Finsch, Proc. Zool. Soc. London, 1872, pp. 89, 96 
(Pelew); Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 4, 17 (Palau); idem, Journ. 
Mus. Godeffroy, 12, 1876, p. 27 (Palau); Giebel, Thes. Ornith., 3, 1877, p. 775 
(Pelew); Finsch, Journ. f. Ornith., 1880, p. 300 (Pelew?); Schmeltz and Krause, 
Ethnogr. A1)th. Mus. Godeffroy, 1881, p. 407 (Palau); Tristram, Cat. Birds, 1889. 
p. 211 (Pelew); Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890- 
1891 (1891), p. 36 (Pelew); Finsch, Das Tierreich, no. 15, 1901, p. 45 (Palau); 
Dubois, Syn. Avium, 1, 1902, p. 713 (Pelew); Reichenow, Die Vogel, 2, 1914, p. 470 
(Palau); Takatsukasa and Kuroda, Tori, 1, 1915, pp. 55, 64 (Pelew); Stresemann, 
Mitt. Zool. Mus. Berlin, 17, 1931, p. 230 (Palau); Hand-list Japanese Birds, rev., 
1932, p. 173 (Palau); Hand-list Japanese Birds, 3d ed., 1942, p. 192 (Babelthuap, 
Koror). 

Tephras finschi Matschie, Journ. f. Ornith., 1901, pp. 112, 113 (Palau); Kuroda, 
in Momiyama, Birds Micronesia, 1922, p. 77 (Pelew) ; Mathews, Syst. Avium Austral- 
asianarum, 2, 1930, p. 712 (Pelew). 

Zosterops cinerea finschi Mayr, Birds Southwest Pacific, 1945, p. 300 (Palau). 

Zosterops cinerea finschii Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 
74 (Peleliu, Garakayo). 

Geographic range. — Micronesia: Palau Islands — Babelthuap, Koror, Gara- 
kayo, Peleliu, Ngabad. 

Characters. — Adult: Resembles adult of Z. c. cinerea, but upper parts 
mostly browner; wing and tail feathers browner; head blacker; rump lighter 
than back ; auriculars grayish-brown ; lores dark ; sides of head and neck 
brownish; underparts mostly dark; chin and throat smoky gray; breast and 
abdomen more brown less gray; sides, flanks and under tail-coverts brown. 
Resembles adult of Z. c. ponapensis, but larger with underparts more buffy; 
upper parts darker. 

Measurements. — Measurements are listed in table 51. 

Specimens examined. — Total number, 37 (15 males, 19 females, 3 unsexed), as follows: 
Palau Islands, USNM— Babelthuap, 1 (Nov. 27)— Koror, 3 (Nov. 4, 5)— Garakayo, 6 (Sept. 
18)— Peleliu, 5 (Aug. 27, Sept. 10); AMNH— exact locality not given, 22 (Oct. Nov., Dee.). 

Molt. — Many of the specimens of Z. c. finschii taken in the period from 
August to December show evidences of molt. Some of the birds taken in 
November and in December appear to be in fresh plumage. All three sub- 
species of Z. cinera evidently undergo a period of molt in the late summer 
and fall. 



Baker: The Avifauna of Micronesia 329 

Remarks. — The Micronesian Dusky White-eye of Palaii was 
found on several of the islands of the southern Palaus by the 
NAMRU2 party in 1945. The bird was observed in flocks of five or 
more individuals moving rapidly through the foliage of trees and 
shrubs. It was not found in the dense, undisturbed jungle areas, but 
rather in second growth vegetation and along the margins of wood- 
lands. At Peleliu, birds were noted in trees and shrubs along the 
roadways; at Garakayo, birds were seen in low trees near the sum- 
mits of hills. At Garakayo, Z. cinerea and Z. conspicillata were 
found in the same areas near the tops of the hills. Both species ap- 
peared to be feeding on seeds of the same trees (unidentified but 
resembling the hibiscus). Z. cinerea was more numerous than Z. 
conspicillata and appeared (from observations made on September 
18, 1945) to be the dominant species and was seen to chase the 
smaller Z. corispicillat a a,w ay . Coultas (field notes) found Z. cmerea 
"fairly common" in 1931 at Palau. 

Evolutionary history of Zosterops cinerea. — The dusky white-eyes 
of Micronesia were considered as separate species until 1944, when 
Mayr (1944b:7) treated them as conspecific, stating that the bird at 
Ponape has characters intermediate between those at Kusaie and 
Palau. Earlier, Hartert (1900:3) suggested a close association be- 
tween Z. cinerea and the species at Truk (now Rukia ruki). Mayr 
concludes that Z. cinerea and R. ruki are not closely related, and 
points out that the absence of a white orbital ring in Z. cinerea does 
not necessarily mean that the bird should be considered as belonging 
to a genus other than Zosterops. 

The pathway of colonization and the ancestral stock of Z. cinerea 
are not certainly known. Among the white-eyes of the Polynesian, 
Melanesian and Malayan areas, there are few kinds which Z. cinerea 
resembles closely. Mayr (1941b: 204) writes that the Z. cinerea at 
Ponape was derived from either Polynesia or Papua. I find little in 
common between Z. cinerea and the species in these areas, and in my 
opinion Z. cinerea is closest to Z. atriceps of the Moluccas. Z. atriceps 
has plumage which is part grayish and part brownish. Its under- 
parts resemble those of Z. c. cinerea but are paler gray; crown, neck, 
and shoulder much like that of Z. c. ponapensis and Z. c. finschii; and 
bill resembling that of Z. c. cinerea. Z. atriceps differs by having 
olive-green coloring on back and wings and yellowish coloring on 
under side of tail. Thus, it is possible that Z. cinerea invaded Micro- 
nesia from the Moluccan region, reaching either Palau or Ponape 
initially. 



330 



University of Kansas Publs., Mus. Nat. Hist. 



Rukia palauensis (Reichenow) 
Palau Greater White-eye 

Cleptomis palauensis Reichenow, Journ. f. Ornith., 1915, p. 125. (Type locality, 
Babeldzuap := Babel thuap, Palauinseln.) 

Megazosterops palauensis Stresemann, Ornith. Monatsber., 38, 1930, p. 159 (Baobel- 
taob); Snouckaert, Alauda (2), 3, 1931, p. 26 (Palau); Stresemann, Mitt. Zool. Mus. 
Berlin, 17, 1931, p. 235 (Baobel Taob r= Babelthuap) ; Mathews, Ibis, 1931, p. 48 
(Palau); Hand-list Japanese Birds, rev., 1932, p. 172 (Palau); Yamashina, Tori, 10, 
1940, p. 674 (Palau); Hand-list Japanese Birds, 3d ed., 1942, p. 191 (Babelthuap, 
Peleliu). 

Rukia palauensis Mayr, Amer. Novit., no. 1269, 1944, p. 7 (Palau); idem, Birds 
Southwest Pacific, 1945, pp. 294, 300 (Peliliu) ; Baker, Smithson. Misc. Coll., vol. 
107, no. 15, 1948, pp. 67, 74 (Peleliu). 

Geographic range. — ^Micronesia : Palau Islands-Babelthuap, Peleliu. 

Characters. — Adult: A large white-eye with upper parts near "Saccardo's 
olive" (some individuals darker brown), head and neck more olivacious, rump 
browner; auriculars blackish with pale yellow streaks; narrow supra-orbital 
stripe pale olive; orbital ring indistinct; underparts near "olive lake," chin 
lighter, under tail-coverts light yellowish-brown; wing and tail feathers dark 
brown, except for tawny outer edges and whitish inner edges; maxilla horn- 
color; mandible yellowish to tawny; feet tawny; iris grayish-brown. 

Measurements. — Measurements of Rukia are listed in table 52. Measure- 
ments of males and females are comparable within the same species. 

Table 52. Mei4surements of Rukia 



Species 


No. 


Wing 


Tail 


Culmen 


Tarsus 


R. 'palauensis 


19 

8 
18 


80 

(76-84) 

81 

(76-85) 

70 

(67-71) 


54 

(51-57) 

52 

(51-52) 

44 

(41-47) 


21.5 
(20.0-22.5) 

21.5 
(20.0-23.0) 

23.0 

(22.0-24.0) 


25 


R. ruki 


(24-26) 
23 


R. sanfordi 


(22-24) 

21 

(20-22) 



Specimens examined. — Total number, 21 (12 males, 9 females), as follows: Palau Islands, 
USNM— Peleliu, 11 (Aug. 27, 29, 30, Sept. 4, 5, 6, 7, Dec., 4, 5); AMNH— Peleliu?, 10 
(Dec). 

Molt. — Specimens taken in August and September are in worn plumage, a 
few individuals show evidence of molt. Specimens taken in December are in 
fresh plumage, although two or three individuals are in the final stages of 
molt. This places the period of molt as September, October, and November. 
Nesting evidently occurs in the summer; one male taken on August 27, 1945, 
had enlarged gonads. 

Remarks. — The Palau Greater White-eye was ciescribed under the 
generic name Cleptomis by Reichenow. This generic allocation was 
not followed by subsequent authors; Stresemann proposed the 



Baker: The Avifauna of Micronesia 331 

generic name Megazosterops in 1930, and Mayr (1944b :7) placed 
this white-eye in the genus Rukia along with other large white-eyes 
from Micronesia. In employing this name, Mayr writes, "The ge- 
neric names Rukia (for ruki) and Kubaryum (for oleaginea) were 
published simultaneously in the same publication. As first reviser 
I select the name Rukia, which not only is shorter but is also based 
on a species which I have been able to examine." 

R. palauensis is recorded from Babelthuap and Peleliu of the Pa- 
lau Islands. In 1931, Coultas found the birds only at the island of 
Peleliu, where he obtained nine specimens from a flock. In 1940, 
Yamashina (1940:674) writes that it is a \ery rare species at Palau. 
Marshall (1949:219) found the bird at Peleliu but at no other is- 
lands visited. In 1945, the iS'AMRU2 party obtained eight speci- 
mens at Peleliu from two localities on the eastern side of the island 
in jungle areas relatively undisturbed by war activities. The birds 
were fairly common in the brush and vines of the jungle under- 
growth at these two areas. There were no flocks seen; usually 
singles or pairs were noted. The bird bears a striking resemblance 
to Psamathia annae, which lives in the same environment and has 
a somewhat similar coloration, shape and posture. These two birds 
probably have undergone a parallel development. Competition be- 
tween the two was not noted. Psamathia is evidently less re- 
stricted in its distribution. 

R. palauensis has a restricted distribution in the Palau Islands, 
as indicated by the observations of Coultas, the Japanese and the 
NAMRU2 party. The disturbance resulting from the war activities 
has undoubtedly influenced the population and restricted further 
the preferred habitat of this white-eye, especially at Peleliu. 

Rukia oleaginea (Hartlaub and Finsch) 
Yap Greater-White-eye 

Zosterops oleaginea Hartlaub and Finsch, Proc. Z(X)1. Soc. London, 1872, p. 95. 
(Type locality. Uap.) 

Zosterops oleaginea Graffe, Joum. Mus. Godeffroy, 2, 1873, p. 122 (Yap) ; Gadow, 
Cat. Birds British Mus., 9, 1884, p. 187 (Yap); Finsch, Das Tierreich, no. 15, 1901, 
p. 24 (Yap); Dubois, Syn. Avium, 1, 1902, p. 708 (Uap); Reichenow, Die Vogel, 2, 
1914, p. 469 (Karolinen = Yap) ; Takatsukasa and Kuroda, Tori, 1, 1915, p. 64 (Mack- 
enzie); Stresemann, Mitt. Zool. Mus. Berlin, 17, 1931, p. 230 (Yap); Hand-list Japa- 
nese Birds, rev., 1932, p. 173 (Yap); Hand-list Japanese Birds, 3d ed., 1942, p. 192 
(Yap). 

Zosterops oleagina Giebel, Thes. Omith., 3, 1877, p. 777 (Mackenzie); Schmeltz 
and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, p. 391 ((Yap); Wiglesworth, 
Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 37 (Uap); 
Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, p. 208 (Uap); Bolau, 
Mitteil. Naturhist. Mus. Hamburg, 1898, p. 60 (Yap). 

Tephras oleaginea Matschie, Journ. f. Ornith., 1901, pp. 112, 113 (Yap). 



332 University of Kansas Publs., Mus. Nat. Hist. 

Kubaryum oleaginus Moniiyama, Birds Micronesia, 1922, p. 1 (Yap); Kuroda, in 
Momiyania, Birds Micronesia, 1922, p. 77 (Yap). 

Kubaryum oleagineuTn Mathews, Syst. Avium Australalianarum, 2, 1930, p. 712 
(Yap). 

Rukia oleaginea Mayr, Anier. Mus. Novit., no. 1269, 1944, p. 7 (Yap); idem, 
Birds Southwest Pacific, 1945, p. 300 (Yap). 

Geographic range. — Micronesia: Caroline Islands — Yap. 

Characters. — According to Hartlaub and Finsch (1872:95), "General colour 
a deep oil-green, with a decided fulvous hue ; underparts a little paler, and a 
little more yellowish ; eye-ring satin-white ; ears blackish ; upper and under 
tail coverts with a slight rufous tinge; wing- and tail-feathers blackish, with 
oil-green margins; under wing-coverts whitish-grey; beak fulvous, under man- 
dible, except at the tip, yellowish; feet pale, probably yellow; iris reddish 
white." 

Remarks. — No specimens of R. oleaginea have been examineci by 
me, and I am following Mayr (1944b: 7) in including it with the 
other large white-eyes of Micronesia in the genus Rukia. 

Rukia ruki (Hartert) 
Truk Greater White-eye 

Tephras ruki Hartert, Bull. British Omith. Club, 7, 1897, p. 5. (Type locality, 
Euk.) 

Tephras ritki Hartert, Ibis, 1898, p. 144 (Ruk) ; idem, Novit. Zool., 7, 1900, p. 3 
(Ruk); Matschie, Journ. f. Ornith., 1901, pp. Ill, 112, 113 (Ruck); Mathews, Syst. 
Avium Australasianarum, 2, 1930, p. 712 (Ruk). 

Zosterops ruki Finsch, Das Tierreich, no. 15, 1901, p. 46 (Ruk); Dubois, Syn. 
Avium, 1, 1902, p. 713 (Ruk); Reichenow, Die Vogel, 2, 1914, p. 470 (Ruk); Takatsu- 
kasa and Kuroda, Tori, 1, 1915, p. 64 (Ruk); Stresemann, Mitt. Zool. Mus. Berlin. 
17, 1931, p. 230 (Truk); Hand-list Japanese Birds, rev., 1932, pp. 172 (Truk); Hand- 
list Japanese Birds, 3d ed., 1942, p. 191 (Truk). 

Rukia ruki Momiyama, Birds Micronesia, 1922, p. 2 (Ruk); Kuroda, in Momiyama, 
Birds Micronesia, 1922, p. 78 (Ruk); Mayr, Amer. Mus. Novit., no. 1269, 1944, p. 7 
(Truk); idem. Birds Southwest Pacific, 1945, p. 301 (Truk). 

Geographic range. — Micronesia: Caroline Islands — Truk. 

Characters. — According to Hartert (1897:5), "Entirely sepia-brown, the 
inner webs of the remiges and under wing-coverts lighter, inclining to whitish; 
the primaries darker, the outer webs bordered with the same colour as the 
back. Bill black; iris red; tarsi and feet orange-rufous; claws mouse-brown." 
R. ruki may be distinguished from other species of Rukia by its dark olive- 
brown coloring. 

Measurements. — Measurements are listed in table 52. 

Specimens examined. — Total number, 7 (4 males, 2 females, 1 un.sexed), from Caroline 
Islands, AMNH— Truk (Nov., Dec). 

Remarks. — This white-eye was first obtained by Owston's col- 
lectors in 1895 at Truk. Hartert (1900:3) writes, "It is most pecul- 
iar that the late J. Kubary, who was an excellent collector, and 
who spent more than fourteen months on Ruk, did not obtain this 
bird. It is probably not numerous, and occurs only on a certain 
secluded spot not visited by Kubary." In like manner, R. palauen- 



Baker: The Avifauna of Micronesia 333 

sis was not described from Palau until 1915, although several col- 
lectors had visited the island at previous times. Hartert included 
the Truk Greater White-eye in the genus Tephras of Hartlaub. 
Later, Momiyama (1922:2) made this bird the type for his new 
genus Rukia, in which Mayr has placed all of the large white-eyes 
of Micronesia. 

Rukia sanfordi (Mayr) 

Ponape Greater White-eye 

Rhampozosterops sanfordi Mayr, Ornith. Monatsber., 39, 1931 [mailing date, Nov. 
4, 1931, ex Mayr, 1944b :8], p. 182. (Type locality, Ponape.) 

Cinnyrorhyncha longirostra Takatsukasa and Yamashina, Dobutsu. Zasshi, 43, 1931 
[printed date, Oct. 15, 1931, but mailing date for extra -Japanese recipients, Nov. 23, 
1931, ex Mayr, 1944b :8], p. 599. (Type locality, Ponape); Hand-list Japanese Birds, 
rev., 1932, p. 172 (Ponape); Hand-list Japanese Birds, 3d ed., 1942, p. 191 (Ponape). 

Cinnyrorhy7icha longirostris Mathews, Ibis, 1933, p. 94 (Ponape). 

Rhamphozosterops sanfordi Mayr, Proc. 6th Pacific Sci. Congr., 4, 1941, p. 204 
(Ponape). 

Rukia sanfordi Mayr, Amer. Mus. Novit., no. 1269, 1944, p. 7 (Ponape); idem. 
Birds Southwest Pacific, 1945, p. 301 (Ponape). 

Geographic range. — Micronesia: Caroline Islands — Ponape. 

Characters. — Adult: upper parts buffy-olive, head greenish, rump and upper 
tail-coverts buffy -brown; wing and tail feathers dark brown, outer edges 
yellowish-olive; underparts grayish-buff, chin and throat faintly washed with 
greenish-yellow; under tail-coverts darker; bill long, curved and brownish- 
black, base of mandible paler; feet yellowish; iris chestnut. R. sanjordi is 
distinguished from other species of Rukia by its smaller size, its paler colora- 
tion and its longer and more curved bill. 

Measurements. — Measurements are listed in table 52. 

Specimens examined. — Total number, 18 (12 males, 6 females), from Caroline Islands, 
AMNH— Ponape (Nov., Dec.). 

Remarks. — Coultas obtained this white-eye at Ponape in 1931 ; he 
writes (field notes) that it is "a very rare bird on Ponape. I found 
them at one tree, a sort of a gum-tree, at about 2,000 feet, where they 
were collecting from the flowers of the tree. I was attracted by their 
deep-throated sibilation that is uttered while feeding. They were 
not in the least disturbed by the noise of the gun and remained long 
enough for me to collect a substantial series. One old man, who 
lives not far from the tree, was the only one I could find who knew 
the bird." Six males and one female taken in December had swollen 
gonads. Richards found this bird to be rare at Ponape in 1947-1948. 
He writes (field notes) that the bird was seen twice (he obtained one 
male) , once in deep forest at about 700 feet and once at the summit 
of Jokaj at 900 feet. He observed a group of three birds "wildly and 
loudly chasing one another from tree to tree." The male obtained 
had yellowish sap adhering to its bill. 



334 University of Kansas Publs., Mus. Nat. Hist. ' 

The Ponape Greater White-eye has an appearance very much like 
that of some of the honey-eaters. Takatsukasa and Yamashina 
(1931c: 599) write, "General appearance very much like either 
Cinnijris or Myzomela, but it differs from them by its very small 
first primary, which is far shorter than the primary coverts, and also 
the smooth cutting edge of the bill, though the bill is similarly shaped 
as to that of Cinnyris. These characteristics show that this bird 
belongs to Zosteropidae but not Nectarinidae or Meliphagidae.'' 

Mayr and the Japanese workers, Takatsukasa and Yamashina, 
published descriptions of this white-eye at Ponape almost simulta- 
neously. Mayr (1944b:8) contends that his name, Rharnphozosterops 
sanfordi, is valid because the mailing date of the journal (Ornitholo- 
gische Monatsberichte) in which R. sanfordi was proposed was No- 
vember 4, 1931, while his investigations show that the earliest mail- 
ing date to European and American ornithologists and libraries of 
the issue of Dobutsugaku Zasshi in which the name Cinnyrorhyncha 
longirostra, proposed by Takatsukasa and Yamashina, appeared was 
November 23, 1931. Mayr (1944b:8) points out that Japanese 
friends of the authors of the name C. longirostra assert that they saw 
copies of the description [inferentially printed copies] prior to No- 
vember 23, 1931. These Japanese, as far as is known, have not 
claimed that they saw copies before November 4, 1931, and Mayr's 
conclusion that his name, R. sanfordi, has priority is here accepted. 
If the name C. longirostra Takasukasa and Yamashina appeared in 
printed form and if copies, in requisite number, were distributed to 
specialists or libraries in Japan, or anywhere else, on or before 
November 3, 1931, the name C. longirostris has priority over R. 
sanfordi. 

Evolutionary history of Riikia in Micronesia. — There is little 
known concerning the status of the large white-eyes of Micronesia. 
Most of them were not found by the earlier collectors and are at 
present reported to be rare or restricted in their distribution. Little 
is known concerning the food preferences and nesting activities of 
the birds and also whether they are actually in danger of extermina- 
tion or whether their populations are normally as low as have been 
reported. Originally described under four different generic names, 
they are now considered as belonging in a single genus, Rukia. 

I have compared specimens of Rukia with those of other members 
of the family Zosteropidae found in the Pacific area. Rukia is ap- 
parently not closely related to Z. conspicillata and Z. cinerea of 
Micronesia but has been derived from a different source or sources. 



Baker : The Avifauna of Micronesia 335 

The author has compared Rukia with the genera Zosterops, Wood- 
fordia, Hypocryptadivs, Apoia, Chlorocharis, Pseudozosterops, and 
Tephrozosterops. Results of these comparisons indicate that large 
and well-differentiated white-eyes are found on a number of the 
islands of Oceania. These white-eyes include Woodfordia, Rukia, 
Zosterops inornata, Z. albogularis, Z. tenuirostris, and Z. strenua. 
These birds are all large, have large bills (either longer or stouter or 
both), large and long tarsi, and often short and rounded wings. 
Rukia apparently has undergone a differentiation which parallels 
that which has taken place in these other white-eyes, but there is no 
evidence of a close relationship between these birds and Rukia. 
There are some resemblances between Rukia and Woodfordia super- 
ciliosa of Rennell Island ; W. superciliosa is the same size and has a 
bill somewhat similar to that of R. ruki and a coloration not very 
different from that of R. sanfordi. R. ruki and R. sanfordi may have 
been derived originally from a common ancestral stock in INIelanesia, 
with subsequent isolation on small islands for considerable time 
where differentiation took place. Rukia also shows some resem- 
blance to the genus Apoia, especially to A. pinaiae of Ceram. There 
is also a possibility that the large white-eyes of Micronesia are 
merely highly modified species of the genus Zosterops; this has been 
suggested by Mayr (1944b: 7). It is my opinion that Rukia is a 
valid genus and is as much different from the genus Zosterops (or 
more so) than other recognized genera of large white-eyes {Wood- 
jordia and Apoia). There is also the strong possibility that the 
large white-eyes of Micronesia have been derived from more than 
one source (and are falsely united in one genus) ; however, it is my 
feeling that they represent a single colonization, which successfully 
established itself at four islands and evolved into four divergent 
species. Possibly R. oleaginea is the least specialized and is closest 
to the ancestral stock; however, this supposition is based on study 
of the original description and on a colored plate of the bird in a 
paper by Kuroda (1922b: pi. 7, fig. 4). 

In summary, it seems that the large Micronesian white-eyes of the 
genus Rukia came originally from Melanesia. Possibly they came 
from Malaysia. Probably the birds have been derived from a single 
ancestral stock, that became established at four islands of Micro- 
nesia and became differentiated along diverse lines, so much so that 
some ornithologists have considered them as belonging to separate 
endemic genera. 



336 University of Kansas Publs., Mus. Nat. Hist. 

Erythrura trichroa trichroa (Kittlitz) 
Blue-faced Parrot-finch 

Fringilla trichroa Kittlitz, Mem. Acad. Imp. Sci. St. Peter.sbourg, 2, 1835, p. 8, pi. 
10. (Type locality, Ualan =: Kusaie.) 

Fringilla trichroa Kittlitz, Obser. Zool., in Lutke, Voy. "Le Seniavine," 3, 1836, p. 
285 (Ualan); idem, Denk. Reise russ. Anier. Micron, unci Kamchat., 2, 1858, p. 38 
(Ualan). 

Estrelda trichroa Gray, Genera Birds, 2, 1849, p. 369 (Kusaie?); Gray, Cat. Birds 
Trop. Is. Pacific Ocean, 1859, p. 27 (Oualan). 

Erythrura trichroa Bonaparte, Consp. Avium, 1, 1850, p. 457 (Ualan); Hartlaub, 
Archiv f. Naturgesch., 18, 1852, p. 133 (Carolinen = Kusaie); idem, Joum. f. Ornith., 
1854, p. 168 (Carolinen = Kusaie) ; Gray, Hand-list Birds, 2, 1870, p. 58 (Ualan); 
Giebel, Thes. Ornith., 2, 1875, p. 118 (Carolinen r= Ualan); Finsch, Joum. Mus. Godef- 
froy, 12, 1876, p. 36 (Ualan); idem (part), Journ. f. Ornith., 1880, pp. 290, 297, 302 
(Kusaie); idem (part). Ibis, 1881, pp. 104, 108 (Kuschai) ; Salvadori (part), Ornith. 
Papuasia, 2, 1881, p. 442 (Carolinis = Kusaie?); Schmeltz and Krause (part), Ethnogr. 
Abth. Mus. Godeffroy, 1881, p. 281 (Kusaie); Sclater (part). Ibis, 1881, p. 545 (Ualan); 
Sharpe (part). Cat. Birds British Mus., 13, 1890, p. 385 (Carolines = Kusaie) ; Wigles- 
worth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 42 
(Ualan); Matschie (part), Journ. f. Ornith., 1901, p. 112 (Ualan); Dubois (part), 
Syn. Avium, 1, 1902, pp. 583 (Carolines = Kusaie) ; Takatsukasa and Kuroda (part), 
Tori, 1, 1915, p. 64 (Kusaie). 

Erythrura kittlitzi Bonaparte, Consp. Avium, 1, 1850, p. 457 ( ex Bonaparte MSS.) 
(Type locality, Ualan); Gray, Hand-list Birds, 2, 1870, p. 58 (Caroline Islands = 
Kusaie). 

Erythrura trichros trichros Hartert (part), Novit. Zool., 7, 1900, p. 6 (Kusaie); 
Kuroda (part), in Momiyama, Birds Micronesia, 1922, pp. 27, 29, 78 (Kusaie) ; Mayr 
(part), Amer. Mus. Novit., no. 489, 1931, p. 4 (Kusaie); Takatsukasa and Yamashina, 
Tori, 7, 1931, p. 110 (Kusaie); Hand-list Japanese Birds, rev., 1932, p. 170 (Kusaie); 
Hand-list Japanese Birds, 3d ed., 1942, p. 189 (Kusaie); Mayr, Birds Southwest Pacific, 
1945, p. 302 (Kusaie). 

Chloromunia trichroa trichroa Mathews (part), Syst. Avium Australasianarum, 2, 
1930, p. 840 (Ualan). 

Geographic range. — Micronesia: Caroline Islands — Kusaie. 

Characters. — Adult: A small finch with thick, stout bill; head, neck, back, 
and scapulars between "parrot green" and "grass green"; forehead, orbital area, 
auriculars. and malar area bluish ; sides of neck green tinged with yellowish ; 
edge of forehead and lores blackish ; wing-coverts and outer margins of wings 
yellowish-green; underparts like back but paler green; rump, upper tail-coverts 
and outer edges of tail feathers near "Pompeian red"; wing and tail feathers 
mostly brownish; bend of wing greenish; under wing-coverts brownish; axil- 
laries buffy tinged with greenish; bill black; feet light yellowish-brown; iris 
brown. Adult female duller than male. 

Immature : Resembles adult, but lacks bluish coloring on sides of head and 
on forehead; underparts washed with buffy brown; rump and tail duller 
carmine. 

Measurements. — Measurements are listed in table 53. 

Specimens examined. — Total number, 14 (12 males, 2 females), from Caroline Islands, 
AMNH — Kusaie (Feb., March, April). 

Molt. — Specimens taken in February and March have mostly new feathers, 
molt having been almost completed when obtained. 

Remarks. — Kittlitz was the first person to describe the Blue-faced 

Parrot-finch ; he found it at Kusaie when he visited the island in the 

winter of 1827-28. Later, it was found to have an extensive range in 



Baker: The Avifauna of Micronesia 



337 



Table 53. Measurements of Erythrura tnchroa in Micronesia 



Subspecies 



E. t. trichroa. 



E. t. clara. 



E. t. ■pelewensis* 



No. 



29 



Wing 



58 
(57-59) 

59 

(57-62) 

61.5 



Tail 



46 

(43-48) 

45 

(41-50) 

51 



Culmen 



13.0 

(12.5-13.5) 

13.5 
(13.0-14.5) 

13.5 



Tarsus 



17 

(16-17) 

17 
(17-18) 

18 



*Kuroda (1922:28). 

Micronesia, Melanesia, northern Australia, Celebes, and the Moluc- 
cas. This small finch may be kept as a pet in a cage by native peo- 
ples, but as far as I know there is no evidence that the bird has been 
introduced to island areas as a result of this practice. 

Coultas observed the finch at Kusaic in 1931; he wrote (field 
notes) that it is a common bird but difficult to obtain. He found it 
in most parts of the island and at all elevations; the bird appeared to 
prefer dense underbrush of the jungle or marginal vegetation. He 
found no evidence of breeding activity in February, March or April. 

Erythrura trichroa clara Takatsukasa and Yamashina 
Blue-faced Parrot-finch 

Erythrura trichroa clara Takatsukasa and Yamashina, Tori, 7, 1931, p. 110. (Type 
locality, Ruk Island.) 

Erythrura trichroa Finsch (part), Journ. f. Ornith., 1880, p. 290 (Ponape, Hugeln = 
Truk); idem, Proc. Zool. Soc. London, 1880, p. 576 (Ruk); idem (part), Ibis, 1881, 
pp. 104, 110, 112, 115 (Ponape); Schmeltz and Krause (part), Ethnogr. Abth. Mus. 
Godeffroy, 1881, p. 353 (Ruk); Salvadori (part), Ornith. Papuasia, 2, 1881, p. 442 
(Ponape); Sclater (part), Ibis, 1881, p. 545 (Ponape, Ruk); Sharpe (part). Cat. Birds 
British Mus., 13, 1890, p. 385 (Carolines = Truk, Ponape); Wiglesworth (part), 
Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 42 (Ponape, Ruk); 
Nehrkorn, Kat. Eiers, 1899, p. 122 (Ruk); Matschie (part), Journ. f. Omith., 1901, 
p. 112 (Ruk, Ponape) ; Dubois (part), Syn. Avium, 1, 1902, p. 583 (Carolines = 
Ponape); Takatsukasa and Kuroda (part), Tori, 1, 1915, pp. 55, 64 (Ponape); Mayr, 
Proc. 6th Pacific Sci. Congr., 4, 1941, p. 204 (Ponape). 

Erythrura trichroa trichroa Hartert (part), Novit. Zool., 7, 1900, p. 6 (Ruk, Po- 
nape); Kuroda (part), in Momiyama, Birds Micronesia, 1922, pp. 27, 28, 29, 78 
(Ponape, Ruk); Mayr (part), Amer. Mus., Novit., no. 489, 1931, p. 4 (Ponape, Ruk). 

Chloromunia trichroa Mathews, Birds Australia, 12, 1925, p. 208 (Ruk). 

Chloromunia trichroa trichroa Mathews (part), Syst. Avium Australasianarum, 2, 
1930, p. 840 (Carolines = Truk, Ponape). 

Erythrura trichroa clara Hand-list Japanese Birds, rev., 1932, p. 170 (Truk, Po- 
nape); Hand-list Japanese Birds, 3d ed., 1942, p. 189 (Truk, Ponape); Mayr, Birds 
Southwest Pacific, 1945, p. 302 (Truk, Ponape); Baker, Smithson. Misc. Coll., vol. 107, 
no. 15, 1948, p. 74 (Truk). 

Lobospingus trichroa clara Mathews, Ibis, 1933, p. 96 (Ruk, Ponape). 

Geographic range.— Micronesia: Caroline Islands — Truk, Ponape, Lukunor? 
Characters.— Adult: Resembles adult of E. t. trichroa, but slightly larger 
with underparts more yellowish and less greenish; blue on head slightly paler; 
22—8131 



338 University of Kansas Publs., Mus. Nat. Hist. 

sides of neck tinged more strongly with yellowish. Birds from Ponape are 
shghtly paler than those from Truk. 

Measuremejits. — Measurements are listed in table 53. Birds from Ponape 
and Truk differ but little in measurements. 

Specimens examined. — Total number, 39 (22 males, 16 females, 1 unsexed), as follows: 
Caroline Islands, USNM— Truk, 2 (May 5, Dec); AMNH— Truk, 15 (March, June, Nov.) 
— Ponape, 22 (Dec). 

Molt. — Birds taken in March and June are not in molt. Some of the 
specimens obtained in November and December are in molt. 

Remarks. — The differences between E. t. trichroa at Kusaie anci 
E. t. clara at Ponape anci Truk are slight. Takatsiikasa an(i Yama- 
shina (1931d:110) separate E. t. clara from E. t. trichroa of Kusaie 
on the basis of a paler blue coloring on head, body more yellowish 
green and sides of neck more distinctly golden-yellow. 

Coultas obtained specimens at Ponape in 1930 and reports (field 
notes) that the bird occurs in the extensive grassland areas of the 
island but that the numbers are small. He estimates the population 
to be less than 100 individuals. He learned that the Japanese had 
trapped them for shipment to Japan as caged birds. Coultas writes 
that the finch at Ponape "is very shy and flies readily when he is 
disturbed. As soon as a call of alarm is uttered the whole flock 
flies up from the ground and heads for the true forest where they 
will hide. They will also work along in the grass, and make a 
getaway. The bird has a little hissing sybilation that it utters 
when on the wing." He found the bird in flocks of 3 to 20; imma- 
tures were frequently found alone. 

McElroy of the NAMRU2 party obtained a female at Moen Is- 
land in the Truk Atoll in December, 1945. He found small flocks 
of these birds in dense vegetation along streams. 

Erythrura trichroa pelewensis Kuroda 
Blue-faced Parrot-finch 

Erythrura trichroa pelewensis Kuroda, in Momiyama, Birds Micronesia, 1922, p. 27. 
(Type locality, Pelew Islands). 

Erythrura trichroa pelewensis Kuroda, Ibis, 1927, p. 692 (Pelew); Mayr. Amer. Mus. 
Novit., no. 489, 1931, p. 4 (Pelew); Hand-list Japanese Birds, rev., 1932, p. 171 
(Palau); Hand-list Japanese Birds, 3d ed., 1942, p. 189 (Babelthuap) ; Mayr, Birds 
Southwest Pacific, 1945, p. 301 (Palau). 

Chlorumunia trichroa pelewensis Mathews, Syst. Avium Australasianarum, 2, 1930, 
p. 840 (Pelew). 

Geographic range. — Micronesia: Palau Islands — Babelthuap. 

Characters. — Kuroda (1922a: 27) describes the bird as follows, "Resembles 
E. trichroa (Kittlit^) from Carolines (the type from Kusaie), but distinguish- 
able from it by the bill being much, thicker and stouter, by the chin being 
tinged with blue, by the under-parts being paler throughout and somewhat 



Baker: The Avifauna of Micronesia 339 

tinged with bluish, by the rump and upper tail-coverts being bright crimson 
instead of dull crimson, by the central tail-feathers brownish red instead of 
dull crimson, by the distinct shafts of central tail-feathers and by longer wing 
and tail." 

Measurements. — The measurements by Kuroda of a single specimen are 
listed in table 53. 

Remarks. — Only one specimen of this subspecies is known. The 
NAMRU2 party did not obtain any record of it in the southern 
Palaus in 1945. If still present in the islands, it may be confined 
to the higher forested areas of Babelthuap. 

Evolutionary history of Erythrura irichroa in Micronesia. — The 
Blue-faced Parrot-finch has been recorded from Kusaie, Ponape, 
Truk and Palau, which are all "high" islands of southern Micro- 
nesia. This bird belongs to a species which occurs in Melanesia, 
northern Australia, Celebes, and the Moluccas. Stresemann (1940: 
40) points out the interesting observation that this species ranges 
only east of Wallace's Line. Mayr (1931c: 1-10) has reviewed the 
parrot-finches of the genus Erythrura and places E. trichroa in the 
subgenus Erythrura, noting that E. t. cyaneifrons from Banks and 
the New Hebrides is similar to the subspecies found in Micronesia. 
As a group the subspecies of E. trichroa are very similar, but the 
populations in Micronesia appear closest to subspecies from the 
Solomons, Admiralty Islands and possibly to E. t. modesta from the 
Moluccas, which appears to indicate that Micronesia was invaded 
from the south or from the southwest via the Moluccas. Whether 
the little known subspecies at Palau represents an independent in- 
vader from the Moluccas is uncertain. 

Lonchura nigerrima minor (Yamashina) 
Black-breasted Weaver-finch 

Munia (Donacola) hunsteini minor Yamashina, in Takatsukasa and Yamashina, 
Dobutsu. Zasshi, 43, 1931, p. 600. (Type locality, Ponape.) 

Lonchura hunsteini minor Hand-list Japanese Birds, rev., 1932, p. 171 (Ponape, 
Truk); Hand-list Japanese Birds, 3d ed., 1942, p. 190 (Ponape. Truk). 

Donacola hunsteini minor Mathews, Ibis, 1933, p. 95 (Ponape). 

Lonchura nigerrima minor Mayr, Birds Southwest Pacific, 1945, p. 301 (Ponap^, 
?Truk). 

Geographic range. — Micronesia: Caroline Islands — Ponape, Truk? 

Characters. — Yamashina in Takatsukasa and Yamashina ( 1931c :600) char- 
acterizes this subspecies as similar to M. hunsteini from New Ireland, but 
smaller; the wing of the adult of the bird from Ponape is from 46 to 49 mm, 
instead of 50-51 mm. as in the New Ireland bird. Moreover the crown and 
nape are white instead of pearl gray. 

Remarks. — Little is known concerning this subspecies named by 



340 University of Kansas Publs., Mus. Nat. Hist. 

Yamashina at Ponape. No specimens have been seen by me. 
Richards obtained one male at Ponape in 1947-1948. He found 
the birds in large flocks. 

Lonchura punctulata cabanisi (Sharpe) 
Philippine Nutmeg Marmikin 

Munia cabanisi Sharpe, Cat. Birds British Mus., 13, 1890, p. 3.53. (Type locahty, 
Luzon.) 

Munia punctulata cabanisi Kuroda, in Momiyama, Birda Micronesia, 1922, p. 7S 
(Yap). 

Lonchura punictulata cabanisi Yamashina, Tori, 7, 1932, p. 395 (Yap); Hand-list 
Japanese Birds, rev., 1932, p. 171 (Yap); Hand-list Japanese Birds, 3d ed., 1942, p. 
189 (Yap). 

Geographic range. — Philippine Islands and Micronesia. In Micronesia; 
Palau Islands; Caroline Islands — Yap. 

Characters. — A small finch with upper parts light grayish-brown, feathers 
with white shafts producing a streaked appearance; lores, anterior part of 
auriculars, malar region, and feathers of chin and throat chocolate-brown with 
faint white shafts;; breast and sides mottled white and dark brown, middle 
of abdomen and under tail-coverts pale buffy-white, wings brown with lighter 
edges, under wing dark with lighter coverts; upper tail-coverts and middle tail 
feathers dark olive, outer tail feathers colored like wings; bill heavy and black; 
feet dark brown. 

Re77iarks.— The Philippine Nutmeg Mannikin is a resident on the 
island of Yap. Yamashina (1932a: 395) records a nest containing 
one egg taken there on May 15, 1932. Marshall (1949:221) records 
this bird at Palau on November 6 and December 2, 1945. Whether 
this bird was introduced to Yap and Palau by man or whether it 
reached there by independent invasion is unknown. 

SUMMARY AND CONCLUSION 

The avifauna of Micronesia consists of 206 kinds of birds belong- 
ing to 37 families and 91 genera. Of these, 30 kinds are sea birds, 29 
kinds are migratory shore birds, and 146 kinds are land and fresh- 
water birds. Of the 30 sea birds, 18 kinds are resident; of the 147 
land and fresh-water birds, 104 kinds are resident and 6 kinds have 
been introduced by man. There are no resident shore birds in Micro- 
nesia. The following conclusions can be drawn from this study: 

1. The islands of Micronesia are oceanic islands and were seem- 
ingly formed independently of any present day continental land 
mass. Terrestrial organisms have reached these islands by "over- 
water dispersal." The avifauna of Micronesia has been received 
from the following sources: Polynesia, Melanesia, the Moluccas, 
Celebes, Phillipines, and Palearctica (see figure 8). 

2. Oceanic birds are among the oldest forms of bird life inhabiting 



Baker : The Avifauna of Micronesia 341 

Micronesia. The presence of elevated islands containing phosphate, 
resulting from the deposition of guano by oceanic birds, is some indi- 
cation of the length of time during which these birds have been pres- 
ent. In number of individuals, the oceanic birds inhabiting the in- 
shore zone are more numerous than those inhabiting the offshore and 
pelagic zones, although twelve of the eighteen resident kinds of 
oceanic birds prefer the offshore and pelagic zones. Most of the 
species of oceanic birds resident in Micronesia are circumtropical in 
distribution; no residents are known in Micronesia which have been 
derived from Palearctica or the North Pacific. Micronesia has no 
endemic oceanic birds. 

3. On the migratory flights, shore birds reach Micronesia along 
three distinct flyways, which in this report are named the Asiatic- 
Palauan Flyway, the Japanese-Marianan Flyway, and the Nearctic- 
Hawaiian Flyway (see figure 7). The shore birds began to utilize 
the Pacific islands as wintering grounds by gradually spreading from 
the Eastern Hemisphere rather than from the AVestern Hemisphere. 

4. More than half (52 percent) of the land birds and fresh-water 
birds in Micronesia were derived directly from ancestral stocks in 
Melanesia. The areas of the Moluccas and of Celebes (Malaysia) 
supplied 21 percent of the birds; the Philippines, 10 percent; Poly- 
nesia, 9 percent; and Palearctica, 8 percent. Results of this study 
show that there may have been only 46 actual colonizations of 
Micronesia by birds from other areas, and that many of the large 
number of endemics present have been the result of secondary 
colonizations within the islands of Micronesia. It is concluded that 
Micronesia, except for the Marshall Islands, has a much closer 
affinity to Melanesia than to any other area as regards avifauna. The 
Marshall Islands may be regarded as a part of the Polynesian Sub- 
region from the viewpoint of avian zoogeography. 

5. Endemism in tlic land birds and fresh-water birds of Micro- 
nesia is extreme. Of 104 native, resident birds, 97 (93.5 percent) 
have become differentiated and can be separated taxonomically from 
related forms. In Micronesia, there are 5 endemic genera, 31 en- 
demic species, and 76 endemic subspecies. The families containing 
the greatest number of endemic forms are Muscicapidae (14) , Zos- 
teropidae (14), Columbidae (13), and Sturnidae (9). 

6. It is concluded that some of the more important factors con- 
trolling the dispersal of the bird life to Micronesia are the direction 
and the intensity of the winds, the small size of the islands, the isola- 
tion of the islands (especially those "high" islands), and the insular 



342 University of Kansas Publs., Mus. Nat. Hist. 

climates, which appear to favor colonists from tropical homes rather 
than those from Palearctic homes. 

7. The factors most important in the process of differentiation of 
birds in the islands of Micronesia are isolation, paucity in numbers 
of individuals, freedom from predation, absence (and presence) of 
interspecific and intraspecific strife, and nutrition. The importance 
of the "dilution" factor is discussed, and the possibility of cross- 
breeding between different kinds of birds is considered. It is con- 
cluded that genetic change altering the phenotypic expression of 
avian characteristics is no more apt to occur in insular populations 
than in continental populations, but such changes have a greater 
chance of being perpetuated in insular populations. 



Baker: The Avifauna of Micronesia 343 



BIBLIOGRAPHY 

Amadon, D. 

1942. Notes on some non-passerine genera, 1. Amer. Mus. no. 1175:1-11, 
1 fig. 

1943. Notes on some non-passerine genera, 3. Amer. Mus. Novit., no. 
1237:1-22. 

Austin, 0. L., Jr. 

1948a. Japanese ornithology and mammalogy during World War II. 
Natural Resources Section, General Headquarters, Supreme Com- 
mander Allied Powers. Report No. 102:1-47. 

1948b. The birds of Korea. Bull, Mus. Comp. Zool., 101, no. 1:1-301, 1 pi. 

Baker, R. H. 

1946a. A new race of Rhipidura rufifrons from Rota Island, Mariana Is- 
lands. Proc. Biol. Soc. Washington, 55:77-78. 

1946b. Some effects of the war on the wildlife of Micronesia. Trans. 11th 
N. Amer. Wildlife Conf.:205-213, 3 figs. 

1946c. A study of rodent populations on Guam, Mariana Islands. Ecol. 
Monogr., 16:393-408, 11 figs. 

1947a. Observations on the birds of the North Atlantic. Auk, 64:245-259, 
5 figs. 

1947b. Size of bird populations at Guam, Mariana Islands. Condor, 49:124- 
125. 

1948. Report on collections of birds made by United States Naval Medical 
Research Unit No. 2 in the Pacific war area. Smithsonian Misc. 
Coll., 107, no. 13:ii-i-74 pp., 6 pis. 

Bequaert, J. C. 

1939. Hippoboscidae of the Caroline Islands (including the Palau Group). 
Mushi, 12:81-82. 

1941. The Hippoboscidae of Oceania (Diptera). Occ. Papers Bernice P. 
Bishop Mus., 16:247-292, 5 figs. 

Berlioz, M. J. 

1929. Catalogue systematique des type de la collection d'oiseaux du mu- 
seum. Bull. Mus. Hist. Nat., Paris, 2nd Ser., 1 :58-69. 

BOGERT, C. 

1937. The distribution and the migration of the long-tailed cuckoo 
iUrodynamis taitensis Sparrman). Amer. Mus. Novit., no. 933:1-12, 
1 map. 

BOLAU, H. 

1898. Die typen der Vogelsammlung des Naturhistorischen Museums zu 
Hamburg. Mitteil. Naturhist. Mus. Hamburg, 15:47-71. 

Bonaparte, C. R. 

1850-'57. Conspectus generum avium. 1850, 1:1-543; 1855-57, 2:1-232. 

BORROR, D. J. 

1947. Birds of Agrihan, Auk, 64:415-417. 

Brigham, W. T. 

1900. An index to the islands of the Pacific Ocean. Mem. Bernice P. 
Bishop Mus., 1, no. 2:1-170, maps. 



344 University of Kansas Publs., Mus. Nat. Hist. 

Bryan, E. H., Jr. 

1936. Birds of Guam. Guam Recorder [newspaper]. 13, no. 2:14-15, 24- 
25; no. 3:10; no. 4:14, 30; no. 5:15-16. 30-31; no. 6:14-15, 34; no. 
7:18-20, 34-35. 

1941. A summary of the Hawaiian birds. Proc. 6th Pacific Sci. Cong., 
1939, 4:185-189. 

Bryan, E. H. Jr., and Greenway, J. C., Jr. 

1944. Contribution to the ornithology of the Hawaiian Islands. Bull. 
Mus. Comp. Zool., 94:80-142. 

Bryan, W. A. 

1903. A monograph of Marcus Island. Occ. Papers Bemice P. Bishop 
Mus., 2:77-139. 

BiJTTIKOFER, J. 

1893. A review of the genus Rhipidura. Notes Leyden Mus., 15:65-98. 

Buxton, P. A. 

1938. The formation of species among insects in Samoa and other oceanic 
islands. Proc. Zool. Soc. London: 264-267. 

Cabanis, J. 

1859-'60. Museum Heineanum. Verzeichniss der Ornithologoschen Samm- 
lung. pt. 2:1-175. 

Cassin, J. 

1858. United States Exploring Expedition during the years 1838, 1839, 
1840, 1841, 1842 under the command of Charles Wiles, U. S. N. 
Mammalogy and ornithology. Philadelphia, viii -|- 466 pp. 

Chamisso, a. von 

1821. Remarks and opinions of the naturalist of the expedition, in Kotze- 
bue's Voyage "Rurick," London, 2:351-433; 3:1-318. 

Christian, F. W. 

1899. The Caroline Islands, travels in the sea of little islands. Charles 
Scribner's Sons, New York :xiii -f 412 pp., illus. 

1926. The megapode bird and the story it tells. Journ. Polynesian Soc, 
35:260. 

Churchill, W. 

1911. The peopling of Yap. Bull. Amer. Geogr. Soc, 43:510-518. 

Cox, L. M. et at. 

1917. The island of Guam. U. S. Govt. Print. Office, Washington, 95 pp., 
illus., map. 

Cram, E. B. 

1927. Bird parasites of the nematode suborders Strongylata. Ascaridata 
and Spirurata. Bull. U. S. Nat. Mus., 140:xvii -[- 465 pp. 

Crampton, H. E. 

1925. Studies on the variation, distribution and evolution of the genus 
Partula. The species of the Mariana Islands, Guam and Saipan. 
Publ. Carnegie Inst., Washington, 228A:vi + 116 pp., 14 pis. 

Darlington, P. J. 

1938. The origin of the fauna of the Greater Antilles, with discussion of 
of dispersal of animals over water and through the air. Quart. Rev 
Biol.. 13:274-300, 5 figs. 



Baker: The Avifauna of Micronesia 345 

Delacour, J. ^ , n 7 • 1 

1942- '43. The bush-warblers of the genera Cettia and Bradypterus, with 

notes on allied genera and species. Ibis, 84 (1942) : 509-5 19; 85 
(1943) : 27-40. 

1943. A revision of the subfamily Estrildinae of the family Ploceidae. 
Zoologica, 28:69-86. 

1946. Notes of the taxonomy of the birds of Malaysia. Zoologica, 31:1-8. 

1947. Birds of Malaysia. The Macmillan Co., New York, xvi + 382 pp., 

84 figs. 

Delacour, J., and Mayr, E. 

1945a. The family Anatidae. Wilson Bull., 57:3-55, 24 figs. 

1945b. Notes on the taxonomy of the birds of the Philippmes. Zoologica, 

30:105-117. 
1946. Birds of the Philippines. New York, xv + 309 pp., 69 figs., 1 map. 

Desmarest, a. G. 

1826. Columba oceanica. Diet. Sci. Nat., ed. Levrault, Paris, 40:316. 

DOBZHANSKY, T. G. 

1941. Genetics and the origin of species. Columbia Univ. Press, New 
York, 2d ed., rev., ix + 466 pp., 24 figs. 

Downs, T. j ci - 

1946. Birds on Tinian in the Marianas. Trans. Kansas Acad. Sci., 49:8/- 
106, 3 figs., 1 map. 

Dubois, A. 

1902-'04. Svnopsis avium. Nouveau manuel d'ornithologie. Bruxelles, 1 
(1899-1902) :xv -h 1-729, pis. i-xii; 2 (1902-1904) :ix-f 730-1399, pis. 
xii-xvi. 

Eliott, D. G. 

1878a. The genus Porphyria and its species. Stray Feathers, 7:6-25. 

1878b. On the fruit-pigeons of the genus Piilopus. Proc. Zool. Soc. London: 
500-575. 

FiNSCH, O. 

1867-'68. Die Papageien. Leiden, 1 (1867) :xii -f 561 pp.; 2 (1868) :vii 

4- 996 pp. 
1870. Zur ornithologie der Tonga-Inseln. Joum. f. Ornith.. 18:119-140. 
1852. Zur ornithologie der Samoa-Inseln. Journ. f. Ornith., 20:30-58. 

1875. Zur ornithologie der Siidsee-Inseln. 1. Die Vogel der Palau-Gruppe. 
Journ. Mus. GodefTroy, 8:133-183 (in reprint: 1-51), 5 pis. 

1876a. Characters of six new Polynesian birds in the Museum Godeffroy at 
Hamburg. Proc. Zool. Soc. London for 1875:643-644. 

1876b. Zur ornithologie der Siidsee-Inseln. II. LTeber neue und weniger 
grekannte Vogel von Viti-, Samoa- und Carolinen-Inseln. Journ. 
Mus. Godeffroy, 12:1-41. 

1876c. Uber den Schwalbenwurger {Artamus) der Palau Inseln. Journ. Mus. 
Godeffroy, 12:41-42. 

1878. On the birds of the island of Ponape, eastern Carolines. Proc. Zool. 
Soc. London for 1877:777-782. 

1880a. Beobachtungen iiber die Vogel der Inseln Ponape (Carolinen). 
Journ. f. Ornith.. 28:283-295. 



346 University of Kansas Publs., Mus. Nat. Hist. 

1880b. Beobachtungen ijber die Vogel der Insela Kuschai (Carolinen). 
Journ. f. Ornith., 28:296-310. 

1880c. Ornithological letters from the Pacific, no. II. Taluit (Bonham) 
Island. Ibis, 4th Ser., 4:218-220. 

1880d. Ornithological letters from the Pacific, no. III. Taluit (Bonham) 
Island. Ibis, 4th Ser., 4:329-333. 

1880e. List of the birds of the islands of Ruk in the central Carolines. 
Proc. Zool. Soc. London: 574-577. 

1881a. On two species of pigeons from the Caroline Islands. Proc. Zool. 
Soc. London for 1880:577-578. 

1881b. Ornithological letters from the Pacific. No. V. Kushai; no. VI. 
Ponape. Ibis, 4th Ser., 5:102-115. 

1881c. Ornithological letters from the Pacific, no. VII. Nawodo (Pleasant 
Island). Ibis, 4th Ser., 5:245-249. 

1883. On a new reed-warbler from Nawado or Pleasant Island, in the 
western Pacific. Ibis, 5th Ser., 1:142-144. 

1884. xjber Vogel der Siidsee. Auf Grund eigener beobachtungen and 
sammlungen. Mitt. Ornith. Ver. Wien, V. Micronesia :46-56. 

1893a. Ueber die Monasaralle von Kuschai [Kittlitzia Monasa (Kittlitz) 
und die bisher mit ihr verwechselten Arten]. Mitth. Ornith. Ver, 
Wien, "Die Schwalbe," 17:1-5. 

1893b. Einiges iiber Siidsee-Rallen. Anzeigeblatt Ornith. Monats. des Deut. 
Ver. z. Schutze der Vogelwelt, 18:457-463. 

1900a. Zur catalogisirung der ornithologischen Abtheilung. Notes Leyden 
Mus, 22:75-125. 

1900b. Meine Beobachtunger uber Fregattvogel (Fregata aquila L., F. ariel 
Gould). Anzeigeblatt Ornith. Monats. des Deut. Ver z. Schutze der 
Vogelwelt, 25 :446-452. 

1901. Das Tierreich. no. 15. Zosteropidae. Berlin, xiv -}- 54 pp., 31 figs. 

FiNSCH, 0., and Hartlaub, G. 

1867. Beitrag zur Fauna Centralpolynesiens. Ornithologie der Viti-, Samoa- 
und Tonga-Inseln. Halle, xl + 290 pp., 14 pi. 

Fisher, H. I. 

1946. The type localities of Pufjinus pacificus cuneatus Salvin and Ptero- 
droma leucoptera hypoleuca (Salvin). Auk, 63:587-588. 

1947. Utinomi's Bibliographica Micronesica: Chordate Sections. Pacific 
Science, 1, no. 3:129-150. 

Forbes, W. A. 

1879. A synopsis of the Meliphagine genus Myzomela, with descriptions of 
two new species. Proc. Zool. Soc. London: 256-279. 

Gadow, H. 

1884. Catalogue of the Passeriformes or perching birds in the collections 
of the British Museum. Cinnyrimorphae. 9:xii-j-310 pp., 7 pis. 

Gaimard, p. J. 

1823. Memoire sur un nouveau genre de Gallinaces, establi sous le nom de 
Megapode. Bull. Gen. et Univ. des Annonces et des Nouvelles Sci., 
2:450-451. 



Baker: The Avifauna of Micronesia 347 

GlEBEL, C. G. 

1872-77. Thesaurua ornithologiae. Reportorium der gesammten omitho- 
logischen Literatur und Nomenclator sammtlicher Gattungen und 
Arten der Vogel nebst Svnonymem und geographischer Verbreitung. 
F. A. Brockhaus, Leipzig, 1 (1872) :vii + 868 pp.; 2 (1875) :vi + 
787 pp.; 3 (1877) :vi + 861 pp. 

Gleize, D. a. 

1945. Birds of Tinian. Bull. Mass. Audubon Soc. 29:220. 

Gleize D. A., and Genelly, D. 

1945. in With the colors. Bull. Mass. Audubon Soc, 29:221-222. 

Godman, F. 

1907-1910. A monograph of the Petrels. Witherby, London, Iv -1- 381 pp., 
pis. 

Graffe, E. 

1873. Die Carolineninsel Yap oder Guap. Journ Mus. Godeffroy, 2:84- 
130 (in reprint 1-58). 

Gray, G. R. 

1859. Catalogue of the birds of the tropical islands of the Pacific Ocean 
in the collection of the British Museum. Taylor and Francis, Lon- 
don: 1-72. 

1866. A synopsis of the species of the genus Collocalia, with descriptions 
of new species. Ann. and Mag. Nat. Hist., 3d ser., 17:118-128. 

Gray, J. E. 

1866. Naturalization of Zosterops dorsalis in New Zealand. Ann, and 
Mag. Nat. Hist., 3d ser., 17:237. 

GuucK, A. 

1932. Biological peculiarities of oceanic islands. Quat. Rev. Biol., 7:405- 
427. 

Uachtsuka ]VI XT 

1931-'35. The birds of the Philippine Islands. Witherby, London, 1 (1931- 
'32) :xx + 439 pp.; 2 (1934-'35) :xxxi + 469 pp. 

1939a. New races of a rail and a fruit pigeon from Micronesia and Pala- 
wan. Bull. British Ornith. Club, 59:151-153. 

1939b. The red jungle fowl from the Pacific islands. Tori, 10:596-601. 

Hachisuka, M. U., Kuroda, N., Takatsukasa, N., Uchida.S.. and Yamashina, Y. 
1932. A hand-list of the Japanese birds, revised. The Ornith. Soc. of 
Japan, Tokyo, iv-)-211 pp. 

1942. A hand-list of Japanese Birds, third revised edition. The Ornith. 
Soc. of Japan, Tokyo, vii -|- 238 pp. 

Harrison, T. H., and M.-vrshall, A. J. 

1937. Aplonis santovestris, sp. nov. Bull. British Ornith. Club, 57:148-150. 

Hartert, E. 

1891. Katalog der Vogelsammlung in Museum der Senckenbergischen 
Naturforschenden Gesselschaft in Frankfurt am Main. Frankfurt 
am Main, xxii + 259 pp. 

1892. in Salvin and Hartert, Catalogue of the Picariae in the collection 
of the British Museum. Coraciae. 16:434-654, pis. 10-14. 

1897. On a new species of Tephras (Zosteropidae) from the Caroline Is- 
lands. Bull. British Ornith. Club, 7:5. 



348 University of Kansas Publs., Mus. Nat. Hist. 

1898. On the birds of the Marianne Islands. Novit. Zool., 5:51-69. 

1900. The birds of Ruk in the central CaroHnes. Novit. Zool., 7:1-11. 

1903-'22. Die Vogel der palaarktischen Fauna. Berlin, 1 (1910) :xlix + 1- 
832, figs. 1-134; 2 (1912-21) :xxiv + 833-1764, figs. 135-256; 3 (1921- 
22) : xii -|- 1765-2328, figs. 257-268. 

1917. On some Rallidae. Novit. Zool. 24:265-274. 

1926. Types of birds in the Tring Museum. B. Types in the general col- 
lection, VII. Tubinares. Novit. Zool., 33:344-357. 

1927. Types of birds in the Tring Museum. B. Types in the general 
collection. VIII. Columbae-Struthionidae. Novit. Zool., 34:1-38. 

1930. List of the birds collected by Ernst Mayr. Novit. Zool., 36:27-128. 

Hartlaub, G. 

1852. R. Titian Peale's Vogel der "United States Exploring Expedition." 
Archiv f. Naturg., 18:93-138. 

1854. Zur Ornithologie Oceanien's. Journ. f. Ornith., 2:160-170. 

1865. Monographischer Versuch iiber die Gattung Zosterops. Journ. f. 
Ornith., 13:1-30. 

1868. On a collection of birds from some less-known localities in the west- 
ern Pacific. Proc. Zool. Soc. London for 1867:828-832, pi. 38. 

1892. Vier .«elten Rallen. Abh. Nat. Ver. Bremen, 12:389-402. 

Hartlaxjb, G., and Finsch, 0. 

1868a. On a collection of birds from the Pelcw Islands. Proc. Zool. Soc. 
London: 4-9, pis. 2-3. 

1868b. Additional notes on the ornithology of the Pelew Islands. Proc. 
Zool. Soc. London: 116-118. 

1872. On a fourth collection of birds from the Pelew and Mackenzie islands. 
Proc. Zool. Soc. London: 87-114. 

Hedley, C. 

1899. A zoogeographic scheme for the mid-Pacific. Proc. Linnean Soc. 
New South Wales, 24:391-417. 

Hesse, R., Allee, N. C.. and Schiviidt, K. P. 

1937. Ecological Animal Geography. An authorized, rewritten edition 
ba.sed on Tiergeographie auf oekologischer Grundlage by R. Hesse, 
prepared by W. C. Alee and K. P. Schmidt. John Wiley & Sons, 
New York, xiv -1- 597 pp., 135 figs. 

HOBBS. W. H. 

1945. Fortress islands of the Pacific. J. W. Edwards, Ann Arbor, xiii + 
186 pp., 107 figs. 

HoMBRON, J. B., and Jacquinot, C. H. 

1841. Description de plusieurs oiseaux nouveaux ou peu connus, provenant 
de I'expedition autour du monde faite sur les corvettes I'Astrolabe 
and La Zelee. Ann. Sci. Nat., Paris, 2d ser., 16:312-320. 

Htjxley, J. S. 

1938. Species formation and geographical isolation. Proc. Zool. Soc. Lon- 
don : 253-264. 

Kelso, L. 

1938. An addition to the range of the short-eared owl. Oologist, 60:138. 

KiRBT. H., Jr. 

1925. The Birds of Fanning Island, Central Pacific Ocean. Condor, 27:185- 
196, figs. 47-54. 



Baker: The Avifauna of Micronesia. 349 

KiTTLITZ, F. H. VON 

1832-'33. Kupfertafein zur Naturgeschichte der Vogel. Frankfurt am Main, 
parts 1-3:1-28, 36 pis. 

1835. uber einige noch unbeschriebene Vogel von der Insel Luzon, den 
Carolinen und den Marianen. Mem. Acad. Imp. Sci. St. Petersbourg, 
2:1-9, pis. 1-10. 

1836. Observations zoologiques faites pendant I'expedition de la corvette 
Le Seniavine, par F. H. de Kittlitz. in Lutke, Voyage "Le Senia- 
vine," 3:237-330. 

1858. Denkwiirdigkeiten einer Reise nach dem russischen Amerika, nach 
Micronesien und durch Kamtschatka. Gotha, 1:383 pp.; 2:463 pp. 

Krieger, H. W. 

1943. Island peoples of the western Pacific. Micronesia and Melanesia. 
Smithson. Inst. Publ. 3737 (War Background Ser. No. 16) : iv + 104 
pp., 21 pis., 2 maps. 

KUROD.A, N. 

1922a. Descriptions of two new forms of birds from Pelew Islands, Micro- 
nesia, in Momiyama, Birds of Micronesia, pt. 1:25-30. 

1922b. A list of the birds of Micronesia Group, exclusive of Magalhaes, 
Gilbert and Ellice islands, in Momivama, Birds of Micronesia, pt. 
1:31-78. 

1925. A contribution to the knowledge of the avifauna of the Riu Kiu 
Islands and vicinity, publ. by the author, Tokyo, vi + 293 pp., 
illus. 

1932. A revision of the types of birds described by Japanese authors dur- 
ing the years 1923 "to 1931. Novit. Zool., 37:384-405. 

1941-'42. A study of the Marianas Duck, Anas oustaleti. Tori, pt. 1, 11: 
99-119, 6 pis; pt. 2, 11:443-448, 3 photos. [In Japanese.] 

L.ACK, D. L. 

1947. Darwin's finches. Cambridge Univ. Press, Cambridge, x -f 209 pp., 
8 pis., 27 figs. 

Lesson, R. P. 

1828. Manuel d'ornithologie, ou description des genres et des principales 
especes d'oiseaux. Paris, 1: iv -|- 421 pp.; 2:448 pp. 

1829. Voyage autour du monde, execute par ordre du Roi, sur la corvette 
de la Majeste La Coquille. pendant les annees 1822, 1823, 1824, et 
1825. Zoologie. Paris, 1 : iv -|- 1-360 ; 2:361-743; Atlas:157 col. pis. 

1830-'31. Traite d'ornithologie, ou description des oiseaux reunis dans les 
principaies collection de France. Paris, xxxii -f 659 pp., 119 col. pis. 

LiNSLEY, L. N. 

1935. Curious things about Guam: the mountain chicken. Guam Re- 
corder [newspaper], 12:249-250. 

Lister, J. J. 

1911. The distribution of the avian genus Megapodius in the Pacific is- 
lands. Proc. Zool. Soc. London: 749-759, 1 fig. 

Lutke, F. 

1835-'36. Voyage autour de monde, execute par ordre de sa majeste I'em- 
pereur Nicolas 1, sur la corvette Le Seniaville, dans los annees 1826, 
1827, 1828, et 1829. Partie historique avec un atlas, lithographie 
d'apres les dessins originaux d'Alexandre Postels et du Baron Kitt- 
litz. Didot, Paris, 1 (1835) :xxiv -|- 410 pp.; 2(1835) :387 pp.; 3 
(1836):xiv-h352 pp. 



350 University of Kansas Publs., Mus. Nat. Hist. 

McAtee, W. L. (editor) 

1945. The ring-necked pheasant and its management in North America. 
Amer. Wildlife Inst., Washington : xi + 320 pp., 31 pis., 12 figs. 

Marche, a. 

1891. Rapport general sur une mission aux iles Mariannes. Nov. Arch 
Missions Sci. et Litt., 1 :241-280. 

Marshall, J. T., Jr. 

1949. The endemic avifauna of Saipan, Tinian, Guam and Palau. Condor, 
51:200-221, figs. 35-40. 

Mathews, G. M. 

1925. Monarcharses, gen. nov. Bull. British Ornith. Club, 45:95. 

1926. On some changes in names. Nycticorax caledonicus pelewensis subsp. 
nov. Bull. Brit. Ornith. Club, 46:60. 

1927-30. Systema Avium Australasianarum. British Ornith. Union, pt. 1 
(1927):l-426; pt. 2 (1930) : 427-1048. 

1931. Additions and corrections to the 'Systema Avium Australasianarum'. 
Ibis, 13th ser., 1 : 44-57. 

1932. Additions and corrections to the 'Systema Avium Australasianarum'. 
pt. II. Ibis, 13th ser., 2:132-161. 

1933. Additions and corrections to the 'Systema Avium Australasianarum'. 
pt. III. Ibis, 13th ser., 3:87-97. 

1934. A check-list of the order Procellariiformes. Novit. Zool., 39:151-206. 
1938. Aplornis versus Aplonis. Ibis, 14th ser., 2:342. 

Matschie, p. 

1901. Bermerkungen zur Zoogeographie des westlichen Mikronensiens. 
Journ. f. Ornith., 49:109-114. 

Mayr, E. 

1931a. Notes on Halcyon chloris and some of its subspecies. Amer. Mus. 
Novit., no. 469:1-10. 

1931b. A systematic list of the birds of Rennell Islands with descriptions of 
new species and subspecies. Amer. Mus. Novit., no. 486:1-29. 

1931c. The parrot finches (genus Erythrura). Amer. Mus. Novit., no. 
489:1-10. 

1931d. Notes on fantails of the genus Rhipidura. Amer. Mus. Novit., no. 
502:1-21. 

1931e. Rhamphozosterops nov. gen. Ornith. Monatsber., 39:182. 

1932. Notes on Meliphagidae from Polynesia and the Solomon Islands. 
Amer. Mus. Novit., no. 516:1-30. 

l93Za. Rhamphozosterops versus Cinnyrhyncha. Ibis, 13th ser., 3:389-390. 
1933b. Three new genera from Polynesia and Melanesia. Amer. Mus. 
Novit., no. 590:1-6, 1 fig. 

1933c. Two new birds from Micronesia. Amer. Mus. Novit., no. 609:1-4. 

1933d. Notes on Polynesian flycatchers and a revision of the genus 
Clythorhynchus Elliott. Amer. Mus. Novit., no. 628:1-21. 

1933e. Notes on the genera Myiagra and Mayrornis. Amer. Mus. Novit., 
no. 651:1-20. 

1933f. Notes on the variation of immature and adult plumages in birds and 
a physiological explanation of abnormal plumages. Amer. Mus. 
Novit., no. 666:1-10. 



Baker: The Avifauna of Micronesia 351 

1933g.Die vogelwelt Polynesiens. Mitt. Zool., Mus. Berlin, 19:306-323. 

1935. Descriptions of twenty-five new species and subspecies. Amer. Mus. 
Novit., no. 820:1-6. 

1936. Descriptions of twenty-five species and subspecies. Amer. Mus. 
Novit., no. 828:1-19. 

1937. Notes on New Guinea birds I. Amer. Mus. Novit., no. 915:1-19. 

1938. Notes on New Guinea birds IV. Amer. Mus. Novit., no. 1006:1-16. 

1940. Speciation phenomena in birds. Amer. Nat., 74:249-278, 7 figs. 

1941a. Borders and subdivisions of the Polynesian Region based on our 
knowledge of the distribution of the birds. Proc. 6th Pacific Sci. 
Cong. 1939, 4:191-195. 

1941b. The origin and the historv of the bird fauna of Polynesia. Proc. 
6th Pacific Sci. Cong., 1939, 4:197-216. 

1942a. Notes on the Polynesian species of Aplonis. Amer. Mus. Novit., 
no. 1166:1-6. 

1942b. Systematics and the origin of species from the viewpoint of a zo- 
ologist. Columbia Univ. Press, New York, xiv -f 334 pp., 29 figs. 

1943. The zoogeographical position of the Hawaiian Islands. Condor, 
45:45-48. 

1944a. The birds of Timor and Sumba. Bull. Amer. Mus. Nat. Hist., 83: 
127-194, 4 figs. 

1944b. Notes on some genera from the Southwest Pacific. Amer. Mus. 
Novit, no. 1269:1-8. 

1945a. Birds of the Southwest Pacific. The Macmillan Co., New York, xix 
-f 316 pp., 3 pis., 16 figs., 1 map. 

1945b. Bird habitats in the Southwest Pacific. Audubon Mag., 47:207-221, 
5 figs. 

1945c. Bird conservation problems in the Southwest Pacific. Audubon 
Mag., 47:279-282, 2 figs. 

Mayr, E., and Amadon, D. 

1941. Geographical variation in Demigretta sacra (Gmelin). Amer. Mus 
Novit., no. 1144:1-11, 1 map. 

Mayr, E., and Moynihan, M. 

1946. Evolution in the Rhipidura rufijrons group. Amer. Mus. Novit., 
no. 1321:1-21, 6 figs. 

Mearns, E. a. 

1909. A list of birds collected by Dr. Paul Bartsch in the Philippine Is- 
lands, Borneo, Guam, and Midway Island, with descriptions of three 
new forms. Proc. U. S. Nat. Mus., 36:463-478. 

Meinertzhagen, R. 

1926. Introduction to a review of the genus Corvus. Novit. Zool., 33:57- 
121, 12 pis, 

Merrill, E. D. 

1945. Plant life of the Pacific world. Fighting Forces Ed., The Infantry 
Journal, Washington, xviii -f 298 pp., 256 figs. 

1947. A botanical bibliographv of the islands of the Pacific. Contr. U. S. 
Nat. Herb., 30, pt. 1 :i-v, 1-322. 



352 University of Kansas Publs., Mus. Nat. Hist. 

momiyama, t. t. 

1920. Description of a new subspecies of Aplonis from the islands of the 
western Micronesia. Tori, 2:1-3. 

1922. Birds of Micronesia. Descriptions of two new genera and nine new 
subspecies of birds from Micronesia. The Ornith. Soc. Japan, pt. 
1:1-24. 

1930. On the birds of Bonin and Iwo Islands. Annot. Ornith. Orientalisi, 
2:1-186. 

MORAN. J. V. 

1946. Birds of Saipan and Tinian. Bull. Mass. Audubon Soc, 30:261-262 

Murphy, R. C. 

1927. On certain forms of Puffinus assimilis and its allies. Amer. Mus. 
Novit., no. 276:1-15, 1 fig. 

1929. On Pterodroma cooki and its allies. Amer. Mus. Novit., no. 370:1-17. 
1 fig. 

1936. Oceanic birds of South America. Amer. Mus. Nat. Hist., New York, 
1 :i-xxii -|- 1-640, 5 col. pis. (unnumbered), numbered pis. 1-38, 
figs. 1-61; 2:641-1245, 10 col. pis. (unnumbered), numbered pis. 
39-42, figs. 62-80. 

1938. The need of insular exploration as illustrated by birds. Science, 
88:533-539, 2 figs. 

Murphy, R. C, and Mathews, G. M. 

1929. Birds collected during the Whitney South Sea Expedition. VI. 
Amer. Mus. Novit., no. 350:1-21, 1 fig. 

Nehrkorn, a. 

1879. Mittheilungen liber Nester und Eier des Museums Godeffroy zu 
Hamburg. Journ. f. Ornith., 27:393-410. 

1899. Katalog der Eiersammlung. Braunschweig, vii -f 256 pp. 

Nelson, E. W. 

1921. The importance of systematic exploration of the land fauna of 
islands in the Pacific. Proc. 1st Pan-Pacific Sci. Conf., Honolulu, 
1920:265-276. 

Neumann, O. 

1922. Neue Formen aus dem papuanischen und polynesischen Inselreich. 
Verh. Ornith. Ges. Bayem, 25:234-238. 

Oberholser, H. C. 

1906. A monograph of the genus Collocalia. Proc. Acad. Nat. Sci. Phila., 
58:177-212. 

1912. A revision of the forms of the edible-nest swiftlet, Collocalia juct- 
phaga (Thumberg). Proc. U. S. Nat. Mus., 42:11-20. 

1915. A synopsis of the races of the crested tern, Thalasseus bergii (Licht- 
en.?tein). Proc. U. S. Nat. Mus., 49:515-526, pi. 66. 

1917. The birds of the Anamba Islands. Bull. U. S. Nat. Mus., 98 :v -f 75 
pp., 2 pis. 

1919. A revision of the subspecies of the white-collared kingfisher, Saw- 
opatis Moris (Boddaert). Proc. U. S. Nat. Mus., 55:351-395. 

Ogilvie-Grant, W. R. 

1893. Catalogue of the game birds in the collection of the British Museum. 
22:xvi-t-585, 8 pis. 



Baker: The Avifauna of Micronesia 353 

1898. In Sharpe and Ogilvie-Grant, Catalogue of the Plataleae, Herodiones. 
Steganopodes, Pygopodes. Alcae, and Impennes in the collection of 
the British Museum. 26:329-653, pis. 5-8. 

Oliver, W. R. B. 

1930. New Zealand birds. Fine Arts Ltd., Wellington, viii -f 541 pp., illus. 

OUSTALET, M. E. 

1881a. Observations sur divers oiseaux de I'Asie et de la Nouvelle-Guinee. 
Bull. Soc. Philom. de Paris, ser. 7, 5:71-80. 

1881b. Monographie des oiseaux de la familie des Megapodiides. III. 
Description des genres et spece.s. Ann. Sci. Nat., Zool., ser. 6, 9:1- 
182. 

1889. Sur la faune ornithologique des lies Mariannes. Le Naturaliste, ser. 
2, 3:260-261. 

1891. Note sur las megapodc de la Perouse. Ann Sci. Nat. Zool., Paris, 
17th ser., 11:196. 

1895-'96. Les mammiferes et les oiseaux des lies Mariannes. Nouv. Arch. 
Mus. Hist. Nat. Paris, ser. 3, 7 (1895) :141-228, 7 pis.; 8 (1896) :24-74. 

PeLZELN, A. VON 

1865. Reise de ostereichischen Fregatte Novara um die Erde in de Jahren 
1857, 1858, 1859, unter dm Befehlen des Commodure B. von Wiiller- 
storf-Urbair, 1, pt. 2, Vogel. Vienna, iv -f 176 pp., 6 pis. 

Peters, J. L. 

1930. Notes on some night herons. Proc. Boston Soc. Nat. Hist., 39:26S 

277. 

1931-'45. Check-list of the birds of the World. Harvard Univ. Press. Cam- 
bridge, 1 (1931) :xviii + 345 pp.; 2 (1934) :xvii -f- 401 pp.; 3 (1937): 
xiii + 311 pp.; 4 (1940) :xii + 291 ; 5 (1945) :xi + 306. 

Peters, J. L., and Griscom, L. 

1928. A new rail and a new dove from Micronesia. Proc. New England 
Zool. Club, 10:99-106. 

Phillips, J. C. 

1923. A natural histoiy of the ducks. Houghton Mifflin Co., Boston and 
New York, 2:xii + 409 pp., 26 pis., 38 maps. 

Price, W., 

1936a. The South Sea adventure. The Hokuseido Press, Tokyo, xiv -1- 
313, illus. 

1936b. Mysterious Micronesia, Yap, Map and other islands under the Japa- 
nese Mandate, etc. Nat. Geogr. Mag., 69:481-510. 

Prowazek, S. Von 

1913. Die deutschen Marianen. Leipzig, 125 pp. 

PUCHERAN, J. 

1853. Voyage au Pole Sud et dans I'Oceanie, sur les corvettes I'Astro