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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.
0 f.
■/ i
MARIANA : ISLANDS;
CAROLINE ISLANDS
0 125 250 500 MILES
III I
•WAKE
-%,
■^
^-^o
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V GILBERT ■..-ISLANDS
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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'
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7°00'
BABELTHUAP
7'W
KOROR 'ARAKABESAN
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10
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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 |
|
|
0 50 OO 200 MILES |
* Kopingomofongi |
||
|
0 |
1 1 1 |
, , |
0 |
|
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 0 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, |
0 4 1 2 3 1 0 0 2 1 0 2 2 |
1 |
|
PuMnus |
1 |
|
|
Pterodroma |
1 |
|
|
Phaethon . |
1 |
|
|
Sula |
0 |
|
|
Fregata |
1 |
|
|
Larus |
I |
|
|
Chlidonias |
1 |
|
|
Sterno, |
4 |
|
|
Thalasseiis |
0 |
|
|
Procelsterna |
1 |
|
|
Atious |
0 |
|
|
GvQZS |
0 |
|
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 0 |
|
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
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|
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 |
0 12 7 4 5 |
3 |
|
Melanesian Moluccan-Celebean . Philippine |
0 0 1 |
||||
|
Palearctic |
0 |
||||
|
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 |
0 0 0 0 0 1* 0 0 0 0 0 0 0 0 1 1 0 0 1 1 0 |
0 0 1 0 1 2 4 1 1 0 1 1 0 1 2 6 0 3t 1 6 0 |
0 1 0 0 2 4 11 0 1 1 5 7 3 0 4 9 1 7 6 10 4 |
0 1 |
|
Anatidae |
1 |
||||
|
Accipitridae Megapodidae Rallidae Columbidae Psittacidae Strigidae Caprimulgidae Apodidae A cedinidae Campephagidae. . . . Corvidae |
0 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
0
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 0 |
||
|
Wake Islands |
12.0-14.0 13 0 |
||||||
|
Hawaiian Islands |
13.0-14.0 13 0 |
||||||
|
Marshall Islands |
12.0-14.0 |
||||||
|
Phoenix, Howland, Hull, Canton Islands |
8.5 |
14 0 |
|||||
|
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 0 |
|
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.
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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.
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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
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22-8131
(Continued from inside of front cover.)
16. A mew extinct emydid turtle from the Lower Pliocene of Oklahoma. By Edwin C. Galbreath. Pp. 266-280, 1 plate. August 16, 1948.
17. Pliocene and Pleistocene records of fossil turtles irom western Kan- sas and Oklahoma. By Edwin C. Galbreath. Pp. 281-284, 1 figure in text. August 16, 1948. n *•
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20. Three new beavers from Utah. By Stephen D. Durrant and Harold S. Crane. Pp. 407-417, 7 figures in text. December 24, 194cS.
21. Two new meadow mice from Michoacan Mexico. By E. Raj^mond Hall. Pp. 423-427, 6 figures in text. December 24, 1948.
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24. Geographic range of the hooded skunk. Mephitis macroura, with description of a new subspecies from Mexico. By E. Raymond Hall and Walter W. Dalquest. Pp. 575-580, 1 figure in text. Jan- uary 20, 1950.
25. Pipistrellus cinnamomeus Miller 1902 referred to the genus Myotis. By E. Raymond Hall and Walter W. Dalquest. Pp. 581-590, 5 fig- ures in text. January 20, 1950.
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Raymond Hall and Walter W. Dalquest. Pp. 591-602, 1 figure in
text. January 20, 1950.
Index, Pp. 605-638. "Vol. 2, (Complete) Mammals of Washington. By Walter W. Dalquest. Pp.
1-444, 140 figures in text. April 9, 1948.
"Vol. 3. 1. The avifauna of Micronesia, its origin, evolution, and distribution.
By RoUin H. Baker. Pp. 1-359, 16 figures in text. June 12, 1951.
A Quantitative Study of the Nocturnal Migration of Birds
BY
GEORGE H. LOWERY, JR.
University of Kansas Publications Museum of Natural History
Volume 3, No. 2, pp. 361-472, 47 figures in text June 29, 1951
University of Kansas
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1951
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1951. Vol. 2. (Complete) Mammals of Washington. By Walter W, Dalquest. Pp.
1-444, 140 figures in text. April 9, 1948.
Vol. 3. 1. The avifauna of Micronesia, its origin, evolution, and distribution.
By RoUin 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.
A Quantitative Study of the Nocturnal Migration of Birds
BY
GEORGE H. LOWERY, JR.
University of Kansas Publications Museum of Natural History
Volume 3, No. 2, pp. 361-472, 47 figures in text June 29, 1951
m%. zm?. zooL
LIBRARY
MAR 1 7 1952
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1951
University of Kansas Pubucations, Museum of Natural History
Editors: E. Raymond Hall, Chairman; A. Byron Leonard, Edward H. Taylor, Robert W. Wilson
University of Kansas Lawrence, Kansas
PRINTED BY
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23-10'20
A Quantitative Study of the Nocturnal Migration of Birds
By
GEORGE H. LOWERY, JR.
CONTENTS
PAGE
Introduction 365
Acknowledgments 367
Part i. Flight Densities and Their Determination 370
Lunar Observations of Birds and the Flight Density Concept, 370
Observational Procedure and the Processing of Data 390
Part ii. The Nature of Nocturnal Migration 408
Horizontal Distribution of Birds on Narrow Fronts 409
Density as a Function of the Hour of the Night 413
Migration in Relation to Topography 424
Geographical Factors and the Continental Density Pattern . . 432
Migration and Meteorological Conditions 453
Conclusions 469
Literature Cited 470
(363)
LIST OF FIGURES
Figure page
1. The field of observation as it appears to the observer 374
2. Determination of diameter of cone at any point 375
3. Temporal change in size of the field of observation 376
4. Migration at Ottumwa, Iowa 377
5. Geographic variation in size of cone of observation 378
6. The problem of sampHng migrating birds 380
7. The sampling effect of a square 381
8. Rectangular samples of square areas ..;...... 382
9. The effect of vertical components in bird flight 383
10. The interceptory potential of slanting lines 384
11. Theoretical possibilities of vertical distribution 388
12. Facsimile of form used to record data in the field 391
13. The identification of co-ordinates 392
14. The apparent pathways of birds seen in one hour 393
15. Standard form for plotting the apparent paths of flight 395
16. Standard sectors for designating flight trends 398
17. The meaning of symbols used in the direction formula 399
18. Form used to compute zenith distance and azimuth of the moon 400
19. Plotting sector boundaries on diagrammatic plots 402
20. Form to compute sector densities : . . . 403
21. Detennination of the angle cc 404
22. Facsimile of form summarizing sector densities 405
23. Detennination of net trend density 406
24. Nightly station density curve at Progreso, Yucatan 407
25. Positions of the cone of observation at Tampico, Tamps 411
26. Average hourly station densities in spring of 1948 414
27. Hourly station densities plotted as a percentage of peak 415
28. Incidence of maximum peak at the various hours of the night in 1948, 416
29. Various types of density-time curves 418
30. Density-time curves on various nights at Baton Rouge 422
31. Directional components in the flight at Tampico, Tamps 428
32. Hourly station density curve at Tampico, Tamps , 429
33. The nightly net trend of migrations at three stations in 1948 431
34. Stations at which telescopic observations were made in 1948 437
35. Positions of the cone of observation at Progreso, Yucatan 443
36. Hourly station density curve at Progreso, Yucatan 444
37. Sector density representation on two nights at Rosedale, Miss 451
38. Over-all sector vectors at major stations in spring of 1948 455
39. Over-all net trend of flight directions shown in Figure 38 456
40. Comparison of fhght trends and surface weather conditions on April 22-23, 1948 460
41. Winds aloft at 10:00 P. M. on April 22 (GST) 461
42. Gomparison of flight trends and surface weather conditions on April 23-24, 1948 462
43. Winds aloft at 10:00 P. M. on April 23 (GST) 463
44. Gomparison of flight trends and surface weather conditions on April 24-25 1948 464
45. Winds aloft at 10:00 P. M. on April 24 (GST) '.'.'.'. '. .' .' .' .' ." .' .' ' . 465
46. Gomparison of flight trends and surface weather conditions on May 21-22, 1948 466
47. Winds aloft at 10:00 P. M. on May 21 (GST) 467
Lowery: Noctuenal Migration of Birds 365
INTRODUCTION
The nocturnal migration of birds is a phenomenon that long has intrigued zoologists the world over. Yet, despite this universal in- terest, most of the fundamental aspects of the problem remain shrouded in uncertainty and conjecture.
Bird migration for the most part, whether it be by day or by night, is an unseen movement. That night migrations occur at all is a con- clusion derived from evidence that is more often circumstantial than it is direct. During one day in the field we may discover hundreds of transients, whereas, on the succeeding day, in the same situation, we may find few or none of the same species present. On cloudy nights we hear the call notes of birds, presumably passing overhead in the seasonal direction of migration. And on stormy nights birds strike lighthouses, towers, and other tall obstructions. Facts such as these are indisputable evidences that migration is taking place, but they provide little basis for evaluating the flights in terms of magnitude or direction.
Many of the resulting uncertainties surrounding the nocturnal migration of birds have a quantitative aspect; their resolution hinges on how many birds do one thing and how many do another. If we knew, for instance, how many birds are usually flying between 2 and 3 a. m. and how this number compares with other one-hour intervals in the night, we would be in a position to judge to what extent night flight is sustained from dusk to da^^vn. If we could measure the number of birds passing selected points of observation, we could find out whether such migration in general proceeds more or less uniformly on a broad front or whether it follows certain favored channels or flyways. This in turn might give us a clearer insight into the nature of the orienting mechanism and the extent to which it depends on visual clues. And, if we had some valid way of estimating the number of birds on the wing under varying weather conditions, we might be able to understand better the nature and development of migration waves so familiar to field ornithologists. These are just random examples suggesting some of the results that may be achieved in a broad field of inquiry that is still virtually untouched — the quantitative study of migratory flights.
This paper is a venture into that field. It seeks to evaluate on a more factual basis the traditional ideas regarding these and similar problems, that have been developed largely from circumstantial
366 University of Kansas Publs., Mus. Nat. Hist.
criteria. It is primarily, therefore, a study of comparative quantities or volumes of migration — or what may be conveniently called flight densities, if this term be understood to mean simply the number of birds passing through a given space in a given interval of time.
In the present study, the basic data permitting the numerical ex- pression of such migration rates from many localities under many different sets of circumstances were obtained by a simple method. When a small telescope, mounted on a tripod, is focused on the moon, the birds that pass before the moon's disc may be seen and counted, and their apparent pathways recorded in terms of coordinates. In bare outline, this approach to the problem is by no means new. Ornithologists and astronomers alike have recorded the numbers of birds seen against the moon in stated periods of time (Scott, 1881a and 1881b; Chapman, 1888; Libby, 1889; West, 1896; Very, 1897; Winkenwerder, 1902a and 1902b; Stebbins, 1906; Carpenter, 1906). Unfortunately, as interesting as these observations are, they furnish almost no basis for important generalizations. Most of them lack entirely the standardization of method and the continuity that would make meaningful comparisons possible. Of all these men, Winken- werder appears to have been the only one to follow up an initial one or two nights of observation with anything approaching an organized program, capable of leading to broad conclusions. And even he was content merely to reproduce most of his original data without corre- lation or comment and without making clear whether he fully grasped the technical difficulties that must be overcome in order to estimate the important flight direction factor accurately.
The present study was begun in 1945, and early results obtained were used briefly in a paper dealing with the trans-Gulf migration of birds (Lowery, 1946). Since that time the volume of field data, as well as the methods by which they can be analyzed, has been greatly expanded. In the spring of 1948, through the cooperation and collaboration of a large number of ornithologists and astron- omers, the work was placed on a continent-wide basis. At more than thirty stations (Figure 34, page 437) on the North American continent, from Yucatan to Ontario, and from California to South Carolina, observers trained telescopes simultaneously on the moon and counted the birds they saw passing before its disc.
Most of the stations were in operation for several nights in the full moon periods of March, April, and May, keeping the moon under constant watch from twilight to dawn when conditions permitted. They have provided counts representing more than one thousand
Lowery: Nocturnal Migration of Birds 367
hours of observation, at many places in an area of more than a million square miles. But, as impressive as the figures on the record sheets are, they, like the published observations referred to above, have dubious meaning as they stand. Were we to compare them directly, station for station, or hour for hour, we would be almost certain to fall into serious errors. The reasons for this are not simple, and the measures that must be taken to obtain true com- parisons are even less so. When I first presented this problem to my colleague, Professor William A. Reuse, of the Department of Physics and Astronomy at Louisiana State University, I was told that mathe- matical means exist for reducing the data and for ascertaining the desired facts. Rense's scholarly insight into the mathematics of the problem resulted in his derivation of formulae that have enabled me to analyze on a comparable basis data obtained from different stations on the same night, and from the same station at different hours and on different nights. Astronomical and technical aspects of the problem are covered by Rense in his paper (1946), but the underlying principles are discussed at somewhat greater length in this paper.
Part I of the present paper, dealing with the means by which the data were obtained and processed, will explore the general nature of the problem and show by specific example how a set of observations is prepared for analysis. Part II will deal with the results obtained and their interpretation.
ACKNOWLEDGMENTS
In the pursuit of this research I have received a tremendcus amount of help from my colleagues, students, and other friends. In the first place, in order to obtain much of the data on which the study was based, it was neecssary to enlist the aid of many persons in various parts of the country and to draw heavily on their time and patience to get all-night telescopic counts of mi- grating birds. Secondly, the processing of the primary data and its subsequent analysis demanded that I delve into the fields of astronomy and mathematics. Here, from the outset, I have enjoyed the constant and untiring help of Pro- fessor W. A. Rense of the Department of Physics and Astronomy at Louisiana State University. Without his collaboration, I would not have been able to do this work, for he not only supplied formulae whereby I was able to make de- sired computations, but time and again he maneuvered me through my diffi- culties in the mathematical procedures. Moreover, Professor Rense has mani- fested a great interest in the ornithological aspect of the problem, and his trenchant advice has been of inestimable value to me. No less am I indebted to my associate, Robert J. Newman, with whom I have spent untold hours discussing the various aspects of the pioblem. Indeed, most of the concepts that have evolved in the course of this study have grown out of discussions
368 University of Kansas Publs., Mus. Nat. Hist.
over a four-year period with both Rense and Newman. Whatever merit this work may have may be attributable in no small part to the help these two men have given me. In the preparation of many of the illustrations, I am further obligated to Newman for his excellent creative ideas as well as draftsmanship, and to Miss Helen Behmes and A. Lowell Wood for their assistance.
The mathematical computations required in this study have been laborious and time-consuming. It is estimated that more than two thousand man-hours have gone into this phase of the work alone. Whereas I have necessarily done most of this work, I have received a tremendous amount of help from A. Lowell Wood. Further assistance in this regard came from Herman Fox, Donald Norwood, and Lewis Kelly.
The recording of the original field data in the spring of 1948 from the thirty- odd stations in North America involved the participation of more than 200 ornithologists and astronomers. This collaboration attests to the splendid co- operative spirit that exists among scientists. Many of these persons stayed at the telescope, either as observer or as recorder, hours on end in order to get sets of data extending through a whole night.
The following were responsible for much of the field data herein used: J. R. Andrews, S. A. Arny, M. Dale Arvey, H. V. Autrey, Charles C. Ayres, Mr. and Mrs. Roy Bailey, Irwin L. Baird, Maurice F. Baker, Rollin H. Baker, Bedortha and Edna Baldwin, Mrs. A. Marguerite Baumgartner, T. A. Becket, Paul Bellington, Donald Bird, Carl Black, Jr., Lea Black, Lytle Blankenship, Mr. and Mrs. J. Stewart Boswell, Bruce Boudreaux, Frank Bray, Mr. and Mrs. Leonard Brecher, Homer Brewer, Mrs. Harvey Broome, Hey ward Brown, Floyd Browning, Cyril Broussard, Paul Buress, Ralph M. Burress, Robert Cain, Don Carlos, Mrs. Reba Campbell, Mr. and Mrs. E. Burnham Chamber- lain, Laura Chaney, Van B. Chaney, Jr., Edward Clebsch, Mr. and Mrs. Ben B. Coffey, Wilham Cook, Dr. Jack Craven, Hugh C. and William Davis, Katherine Davis, Richard Davis, Richard DeArment, Robert E. Delphia, J. C. Dickinson, Mr. and Mrs. Otto Dietrich, John Dietrich, Clara Dixon, Nina Driven, John J. Duffy, Mr. and Mrs. R. J. Dunbar, Betty Dupre, Bernard E. Eble, Jr., Robert G. Eble, Dr. and Mrs. William H. Elder, C. C. Emory, Davis Emory, Alice H. Farnsworth, James Fielding, William R. Fish, Mr. and Mrs. Myron Ford, W. G. Fuller, Louis Gainey, Dr. Mary E. Gaulden, Mr. and Mrs. John J. Giudice, Lt. L. E. Goodnight, Earl R. Greene, Max Grilkey, W. W. H. Gunn, Noel Maxwell Hail, Jr., A. J. Hanna, Paul Hansen, Harold W. Harry, Joseph Healy, Dorothy HeJmer, Mr. and Mrs. John H. Helmer, Philip E. Hoberecht, William D. Hogan, Dr. and Mrs. Joseph C. Howell, E. J. Huggins, Mrs. Walter Huxford, Hugh litis, W. S. Jennings, William M. Johnson, William Kasler, Luther F. Keeton, Lawrence C. Kent, W. H. Kiel, L. P. Kindler, Mr. and Mrs. Joseph E. King, Harriet Kirby, E. J. Koestner, Roy Komarek, Ann Knight, Mr. and Mrs. N. B. Langworthy, Mr. and Mrs. C. F. Lard, Prentiss D. Lewis, Ernest Liner, Dr. and Mrs. R. W. Lockwood. Dr. Harvey B. Lovell, William J. Lueck, Don Luethy, James Major, Mr. and Mrs. Russell L. Mannette, Mrs. John B. Mannix, Donald Mary, Dale E. McCollum, Stewart McConnell, Mr. and Mrs. M. L. McCroe, Robert L. McDaniel, Mr. and Mrs. Frank McGill, Thomas Merimer, Mr. and_ Mrs. I. S. H. Metcalf, Ann Michener, John Michener, T. H. Milby, D. S. Miller, Burt Monroe, Jr, Burt Monroe, Sr., Mrs. R. A. Monroe, Gordon Montague, Duryea Morton, James Mosimonn, Don L. Moyle, Grant Murphy,
Lowery: Nocturnal Migration of Birds 369
John T. Murphy, Mrs. H. F. Murphy, Mrs. Hill Myers, Mr. and Mrs. Robert J. Newman, William Nichols, R. A. Norris, Floyd Oaks, Eugene P. Odum, Mrs. E. E. Overton, Lennie E. Pate, Kenneth Patterson, Ralph Paxton, Louis Peiper, Marie Peiper, Mr. and Mrs. Harold S. Peters, Mary Peters, Mr. and Mrs. D. W. Pfitzer, Betty Plice, Max Plice, Lestar Porter, D. R. Power, Kenneth Price, George Rabb, Marge Reese. Wayne L. Reeve, C. L. Riecke, R. D. Ritchie, V. E. Robinson, Beverly J. Rose, Mary Jane Runyon, Roger Rusk, Bernd Safinsley, Mr. and Mrs. Glen C. Sanderson, Lewis L. Sandidge, John Sather, J. Benton Schaub, Evelyn Schneider, Henry W. Setzer, Mr. and Mrs. Walter Shackleton, Mr. and Mrs. Francis P. Shannon, Mr. and Mrs. Charles Shaw, Paul H. Shepard, Jr., Alan C. Sheppard, Mabel Slack, Alice Smith, R. Demett Smith, Jr., Nat Smith, Major and Mrs. Charles H. Snyder, Albert Springs, Dr. and Mrs. Fred W. Stamm, J. S. Steiner, Mrs. Paul Stephen- son, Herbert Stern, Jr., Herbert Stoddard, Mr. and Mrs. F. W. Stomm, Charles Strull, Harold P. Strull, Mrs. Fan B. Tabler, Dr. and Mrs. James T. Tanner, S. M. H. Tate, David Taylor, Hall Tennin, Scott Terry, Mr. and Mrs. S. Charles Thacher, Olive Thomas, G. A. Thompson, Jr., Dr. and Mrs. S. R. Tipton, Robert Tucker, Tom Uzzel, Mr. and Mrs. M. G. Vaiden, Richard Vaught. Edward Violante, Brother I. Vincent. Marilyn L. Walker, Mr. and Mrs. Willis Weaver, Mr. and Mrs. W. L. Webb, Margaret M. L. Wehking, W. A. Welshans, Jr., Mrs. J. F. Wernicke, Francis M. Weston, Miss G. W. Weston. Dr. James W. White, John A. White, A. F. Wicke, Jr., Oren Williams, J. L. Wilson ni, W. B. Wilson, Dr. and Mrs. Leonard Wing, Sherry Woo, Rodney Wuthnow, Grace Wyatt, Mr. and Mrs. Malcom Young, Mr. and Mrs. A. J. Zimmerman. To the scores of other people who assisted in making these observations I extend my hearty thanks.
Drs. E. R. Hall, Edward H. Taylor, and H. B. Hungerford of the University of Kansas have read the manuscript and have made valuable suggestions, as have also Dr. W. H. Gates of Louisiana State University and Dr. Donald S. Famer of the State College of Washington. Dr. Famer has also been of great help, together with Drs. Ernst Mayr, J. Van Tyne, and Ernst Schiiz, in suggesting source material bearing on the subject in foreign literature. Dr. N. Wyaman Storer, of the University of Kansas, pointed out a short-cut in the method for determining the altitude and azimuth of the moon, which resulted in much time being saved. . For supplying climatological data and for guidance in the interpretation thereof, I am grateful to Dr. Richard Joel Russell, Louisiana State University; Commander F. W. Reichelderfer, Chief of the U. S. Weather Bureau, Washington, D. C; Mr. Merrill Bernard, Chief of the CHmatological and Hydrologic Services; and Mr. R-alph Sanders, U. S. Weather Bureau at New Orleans, Louisiana.
Acknowledgment is made to Bausch and Lomb Optical Company for the loan of six telescopes for use in this project. Messrs. G. V. Cutler and George Duff of Smith and Johnson Steamship Company, operators of the Yucatan Line, are to be thanked for granting me free passage on the "S. S. Bertha Br0vig" to Progreso, Yucatan, where I made observations in 1945 and 1948. I am also indebted to the Louisiana State University Committee on Faulty Re- search for a grant-in-aid.
370 University of Kansas Publs., Mus. Nat. Hist.
PART I. FLIGHT DENSITIES AND THEIR DETERMINATION
A. Lunar Observations of Birds and the Flight Density Concept
The subject matter of this paper is wholly ornithological. It is written for the zoologist interested in the activities of birds. But its bases, the principles that make it possible, lie in other fields, in- cluding such rather advanced branches of mathematics as analytical geometry, spherical geometry, and differential calculus. No ex- haustive exposition of the problem is practicable, that does not take for granted some previous knowledge of these disciplines on the part of all readers.
There are, however, several levels of understanding. It is possible to appreciate what is being done without knowing how to do it; and it is possible to learn how to carry out the successive steps of a procedure without entirely comprehending why. Some familiarity with the concepts underlying the method is essential to a full under- standing of the results achieved, and details of procedure must be made generally available if the full possibilities of the telescopic approach are to be realized. Without going into proof of underlying propositions or actual derivation of formulae, I shall accordingly present a discussion of the general nature of the problem, conveyed as much as possible in terms of physical visualization. The develop- ment begins with the impressions of the student when he first at- tempts to investigate the movements of birds by means of the moon.
What the Observer Sees
Watched through a 20-power telescope on a cloudless night, the full moon shines like a giant plaster hemisphere caught in the full glare of a floodlight. Inequalities of surface, the rims of its craters, the tips of its peaks, gleam with an almost incandescent whiteness; and even the darker areas, the so-called lunar seas, pale to a clear, glowing gray.
Against this brilliant background, most birds passing in focus appear as coal-black miniatures, only 1/10 to 1/30 the apparent diameter of the moon. Small as these silhouettes are, details of form are often beautifully defined — the proportions of the body, the shape of the tail, the beat of the wings. Even when the images are so far away that they are pin-pointed as mere flecks of black against the illuminated area, the normal eye can follow their progress easily.
Lowery: Nocturnal Migration of Birds 371
In most cases the birds are invisible until the moment they "enter," or pass opposite, the rim of the moon and vanish the instant they reach the other side. The interval between is likely to be inestim- ably brief. Some birds seem fairly to flash by; others, to drift; yet seldom can their passing be counted in seconds, or even in measure- able fractions of seconds. During these short glimpses, the flight paths tend to lie along straight lines, though occasionally a bird may be seen to undulate or even to veer off course.
Now and again, in contrast to this typical picture, more eerie effects may be noted. Some of them are quite startling — a minute, inani- mate-looking object drifting passively by like a corpuscle seen in the field of a microscope; a gigantic wing brushing across half the moon; a ghost-like suggestion of a bird so transparent it seems scarcely more than a product of the imagination; a bird that pauses in mid-flight to hang suspended in the sky; another that beats its way ineffectually forward while it moves steadily to the side; and flight paths that sweej) across the vision in astonishingly geometric curves. All of these things have an explanation. The "corpuscle" is possibly a physical entity of some sort floating in the fluid of the observer's eye and projected into visibility against the whiteness of the moon. The winged transparency may be an insect uncon- sciously picked up by the unemployed eye and transferred by the camera ludda principle to the field of the telescope. It may be a bird flying very close, so drastically out of focus that the observer sees right through it, as he would through a pencil held against his nose. The same cause, operating less effectively, gives a character- istic gray appearance with hazy edges to silhouettes passing just beneath the limits of sharp focus. Focal distortions doubtless also account for the precise curvature of some flight paths, for this peculiarity is seldom associated with distinct images. Suspended flight and contradictory directions of drift may sometimes be at- tributable to head winds or cross winds but more often are simply illusions growing out of a two-dimensional impression of a three- dimensional reality.
Somewhat more commonplace are the changes that accompany clouds. The moon can be seen through a light haze and at times remains so clearly visible that the overcast appears to be behind, in- stead of in front of, it. Under these circumstances, birds can still be readily discerned. Light reflected from the clouds may cause the sil- houettes to fade somewhat, but they retain sufficient definition to dis- tinguish them from out-of-focus images. On occasion, when white
372 University of Kansas Publs.^ Mus. Nat. Hist.
cloud banks lie at a favorable level, they themselves provide a back- drop against which birds can be followed all the way across the field of the telescope, whether or not they directly traverse the main area of illumination.
Types of Data Obtained
The nature of the observations just described imposes certain limitations on the studies that can be made by means of the moon. The speed of the birds, for instance, is utterly beyond computation in any manner yet devised. Not only is the interval of visibility extremely short, but the rapidity with which the birds go by depends less on their real rate of motion than on their proximity to the observer. The identification of species taking part in the migration might appear to offer more promise, especially since some of the early students of the problem frequently attempted it, but there are so many deceptive elements to contend with that the results cannot be relied upon in any significant number of cases. Shorn of their bills by the diminution of image, foreshortened into un- familiar shape by varying angles of perspective, and glimpsed for an instant only, large species at distant heights may closely resemble small species a few hundred feet away. A sandpiper may appear as large as a duck ; or a hawk, as small as a sparrow. A goatsucker may be confused with a swallow, and a swallow may pass as a tern. Bats, however, can be consistently recognized, if clearly seen, by their tailless appearance and the forward tilt of their wings, as well as by their erratic flight. And separations of nocturnal migrants into broad categories, such as seabirds and passerine birds, are often both useful and feasible.
It would be a wonderful convenience to be able to clock the speed of night-flying birds accurately and to classify them specifically, but neither of these things is indispensable to the general study of nocturnal migration, nor as important as the three kinds of basic data that are provided by telescopes directed at the moon. These concern: — (1) the direction in which the birds are traveling; (2) their altitude above the earth; (3) the number per unit of space passing the observation station.
Unfortunately none of these things can be perceived directly, except in a very haphazard manner. Direction is seen by the observer in terms of the slant of a bird's pathway across the face of the moon, and may be so recorded. But the meaning of every such slant in terms of its corresponding compass direction on the plane of
Lowery: Nocturnal Migration of Birds 373
the earth constantly changes with the position of the moon. Altitude is only vaguely revealed through a single telescope by the size and definition of images whose identity and consequent real dimensions are subject to serious misinterpretation, for reasons already ex- plained. The number of birds per unit of space, seemingly the easiest of all the features of migration to ascertain, is actually the most difficult, requiring a prior loiowledge of both direction and altitude. To understand why this is so, it will be necessary to con- sider carefully the true nature of the field of observation.
The Changing Field of Observation
Most of the observations used in this study were made in the week centering on the time of the full moon. During this period the lunar disc progresses from nearly round to round and back again with little change in essential aspect or apparent size. To the man behind the telescope, the passage of birds looks like a performance in two di- mensions taking place in this area of seemingly constant diameter — not unlike the movement of insects scooting over a circle of paper on the ground. Actually, as an instant's reflection serves to show, the two situations are not at all the same. The insects are all moving in one plane. The birds only appear to do so. They may be flying at elevations of 500, 1000, or 2000 feet; and, though they give the illusion of crossing the same illuminated area, the actual breadth of the visible space is much greater at the higher, than at the lower, level. For this reason, other things being equal, birds nearby cross the moon much more swiftly than distant ones. The field of observa- tion is not an area in the sky but a volume in space, bounded by the diverging field lines of the observer's vision. Specifically, it is an inverted cone with its base at the moon and its vertex at the tele- scope.
Since the distance from the moon to the earth does not vary a great deal, the full dimensions of the Great Cone determined by the diameter of the moon and a point on the earth remain at all times fairly constant. Just what they are does not concern us here, ex- cept as regards the angle of the apex (roughly 1/2°), because ob- viously the effective field of observation is limited to that portion of the Great Cone below the maximum ceiling at which birds fly, a much smaller cone, which I shall refer to as the Cone of Observation (Figure 1).
The problem of expressing the number of passing birds in terms of a definite quantity of space is fundamentally one of finding out
374
University of Kansas Publs., Mus. Nat. Hist.
|
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Fig. 1. The field of ob.-ervation, riiowing its two-dimensional aspect as it appears to the observer and its three-dimensional actuality. The breadth of the cone is greatly exaggerated.
the critical dimensions of this smaller cone. The diameter at any distance from the observer may be determined witii enough accuracy for our purposes simply by multiplying the distance by .009, a con- venient approximation of the diameter of the moon, expressed in
Lowery: Nocturnal Migration of Birds
375
radians (see Figure 2). One hundred feet away, it is approximately 11 inches; 1000 feet away, nine feet; at one mile, 48 feet; at two miles, 95 feet. Estimating the effective length of the field of ob- servation presents more formidable difficulties, aggravated by the fact that the lunar base of the Great Cone does not remain station-
FiG. 2. Method for determining the diameter of the cone at any point. The angular diameter of the moon may be expressed in radi- ans, or, in other words, in terms of lengths of arc equivalent to the radius of a circle. In the diagram, the arc between C and E, being equivalent to the radius CO, represents a radian. If we allow the arc between A and B to be the diameter of the moon, it is by astronomi- cal calculation about .009 radian, or .009 CO. This ratio will hold for any smaller circle inscribed about the center O; that is, the arc be- tween A'B' equals .009 CO. Thus the width of the cone of observa- tion at any point, expressed in degrees of arc, is .009 of the axis of the cone up to that point. The cone is so slender that the arc be- tween A and B is essentially equal to the chord AB. Exactly the same consideration holds true for the smaller circle where the chord A'B' represents part of the flight ceiling.
ary. The moon rises in the general direction of east and sets some- where in the west, the exact points where it appears and disappears on the horizon var\'ing somewhat throughout the year. As it drifts across the sky it carries the cone of observation with it like the slim beam of an immense searchlight slowly probing space. This situa- tion is ideal for the purpose of obtaining a random sample of the number of birds flying out in the darkness, yet it involves great complications ; for the size of the sample is never at two consecutive instants the same. The nearer the ever-moving great cone of the moon moves toward a vertical position, the nearer its intersection with the flight ceiling approaches the observer, shortening, therefore, the cone of observation (Figiu-e 3). The effect on the number of
376 University of Kansas Publs., Mus. Nat. Hist.
MOON 12 PM
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Fig. 3. Temporal change in the effective size of the field of observation. The sample sections, A and B, represent the theoretical densities of flight at 8:20 and 12:00 p.m., respectively. Though twice as many birds are assumed to be in the air at midnight when the moon is on its zenith (Z) as there were at the earlier hour, only half as many are visible because of the decrease in size of the cone of observation.
birds seen is profound. In extreme instances it may completely re- verse the meaning of counts. Under the conditions visualized in Figure 3, the field of observation at midnight is only one-fourth as large as the field of observation earlier in the evening. Thus the twenty-four birds seen from 7 to 8 p. m., represent not twice as many birds actually flying per unit of space as the twelve observed from 11:30 to 12:30 a. m., but only half the amount. Figure 4, based on ob-
Lowery: Nocturnal Migration of Birds
377
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Fig. 4. Migration at Ottumwa, Iowa, on the night of May 22-23, 1948. Curve A is a graphic representation of the actual numbers of birds seen hourly through the telescope. Curve B represents the same figures cor- rected for the variation in the size of the cone of observation. The dis- similarity in the two curves illustrates the deceptive nature of untreated telescopic counts.
servations at Ottumwa, Iowa, on the night of May 22-23, shows a similar effect graphically. Curve A represents the actual numbers of birds per hour seen; Curve B shows the same figures expressed as flight densities, that is, corrected to take into account the changing size of the field of observation. It will be noted that the trends are almost exactly opposite. While A descends, B rises, and vice-versa. In this case, inferences drawn from the unprocessed data lead to a complete misinterpretation of the real situation.
Nor does the moon suit our convenience by behaving night after night in the same way. On one date we may find it high in the sky between 9 and 10 p. m.; on another date, during the same interval of time, it may be near the horizon. Consequently, the size of the cone is different in each case, and the direct comparison of flights in the same hour on different dates is no more dependable than the mis- leading comparisons discussed in the preceding paragraph.
The changes in the size of the cone have been illustrated in Figure 3 as though the moon were traveling in a plane vertical to the earth's surface, as though it reached a point directly over the observer's head. In practice this least complicated condition seldom obtains in the regions concerned in this study. In most of the northern
2—1020
378 University of Kansas Publs., Mus. Nat. Hist.
hemisphere, the path of the moon lies south of the observer so that the cone is tilted away from the vertical plane erected on the parallel of latitude where the observer is standing. In other words it never reaches the zenith, a point directly overhead. The farther north we go, the lower the moon drops toward the horizon and the more, therefore, the cone of observation leans away from us. Hence, at the same moment, stationed on the same meridian, two observers, one in the north and one in the south, will be looking into different effective volumes of space (Figure 5).
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Fig. 5. Geographical variation in the size of the cone of observation. The cones A and B represent the effective fields of observation at two stations situated over 1,200 miles apart. The portions of the great cones included here appear nearly parallel, but if extended far enough would be found to have a common base on the moon. Because of the continental scale of the drawing, the flight ceiling appears as a curved surface, equidistant above each station. The lines to the zenith appear to diverge, but they are both perpendicular to the earth. Although the cones are shown at the same instant in time, and have their origin on the same meridian, the dimensions of B are less than one- half as great as those of A, thus materially decreasing the opportunity to see birds at the former station. This effect results from the different slants at which the zenith distances cause the cones to intersect the flight ceiling. The diagram illustrates the principle that northern stations, on the average, have a better chance to see birds passing in their vicinity than do southern stations.
As a further result of its inclination, the cone of observation, seldom affords an equal opportunity of recording birds that are flying in two different directions. This may be most easily understood by
Lowery: Nocturnal Migration of Birds 379
considering what happens on a single flight level. The plane parallel to the earth representing any such flight level intersects the slanting cone, not in a circle, but in an ellipse. The proportions of this ellipse are very variable. When the moon is high, the intersection on the plane is nearly circular; when the moon is low, the ellipse becomes greatly elongated. Often the long axis may be more than twice the length of the short axis. It follows that, if the long axis happens to lie athwart the northward direction of flight and the short axis across the eastward direction, we will get on the average over twice as large a sample of birds flying toward the north as of birds flying toward the east.
In summary, whether we wish to compare different stations, different hours of the night, or different directions during the same hour of the night, no conclusions regarding even the relative numbers of birds migrating are warranted, unless they take into account the ever-varying dimensions of the field of observation. Otherwise we are attempting to measure migration with a unit that is constantly expanding or contracting. Otherwise we may expect the same kind of meaningless results that we might obtain by combining measure- ments in millimeters with measurements in inches. Some method must be found by which we can reduce all data to a standard basis for comparison.
The Directional Element in Sampling
In seeking this end, we must immediately reject the simple logic of sampling that may be applied to density studies of animals on land. We must not assume that, since the field of observation is a volume in space, the number of birds therein can be directly expressed in terms of some standard volume — a cubic mile, let us say. Four birds counted in a cone of observation computed as 1/500 of a cubic mile are not the equivalent of 500 X 4, or 2000, birds per cubic mile. Nor do four birds flying over a sample 1/100 of a square mile mathe- matically represent 400 birds passing over the square mile. The reason is that we are not dealing with static bodies fixed in space but with moving objects, and the objects that pass through a cubic mile are not the sum of the objects moving through each of its 500 parts. If this fact is not immediately apparent, consider the circumstances in Figures 6 and 7, illustrating the principle as it applies to areas. The relative capacity of the sample and the whole to intercept bodies in motion is more closely expressed by the ratio of their per- imeters in the case of areas and the ratio of their surface areas in the case of volumes. But even these ratios lead to inaccurate results
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unless the objects are moving in all directions equally (see Figure 8) . Since bird migration exhibits strong directional tendencies, I have come to the conclusion that no sampling procedure that can be ap- plied to it is sufficiently reliable short of handling each directional trend separately.
Fig. 6. The problem of sampling migrating birds. The large square in the diagram may be thought of as a square mile on the earth's surface, divided into four equal smaller squares. Birds are crossing over the area in three directions, equally spaced, so that each of the subdivisions is traversed by three of them. We might be tempted to conclude that 4x3, or 12, would pass over the large square. Actually there are only seven birds involved all told. Obviously, the intercep- tive potential of a small square and a larger square do not stand in the same ratio as their areas.
For this reason, the success of the whole quantitative study of migration depends upon our ability to make directional analyses of primary data. As I have already pointed out, the flight directions of birds may be recorded with convenience and a fair degree of objec- tivity by noting the slant of their apparent pathways across the disc of the moon. But these apparent pathways are seldom the real pathways. Usually they involve the transfer of the flight line from a horizontal plane of flight to a tilted plane represented by the face of the moon, and so take on the nature of a projection. They are
Lowery: Nocturnal Migration of Birds
381
clues to directions, but they are not the directions themselves. For each compass direction of birds flying horizontally above the earth, there is one, and only one, slant of the pathway across the moon at a given time. It is possible, therefore, knowing the path of a bird in relation to the lunar disc and the time of the observation, to com- pute the direction of its path in relation to the earth. The formula employed is not a complicated one, but, since the meaning of the
+
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SAMPLE SQUARE
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EVERY BIRD FLIES OVER 4 SQUARES THE SIZE OF THE SAMPLE
Fig. 7. The sampling effect of a square. In Diagram A eight evenly distributed birds are flying from south to north, and another four are pro- ceeding from east to west. Three appear in each of the smaller squares. Thus, if we were to treat any of these smaller sections as a directly pro- portionate sample of the whole, we would be assuming that 3x16, or 48, birds had traversed the square mile — four times the real total of 12. If we consider the paths separately as in Diagram B, we see quite clearly what is wrong. Every bird crosses four plots the size of the sample and is being computed into the total over and over a corresponding number of times. Patently, just as many south-north birds cross the bottom tier of squares as cross the four tiers comprising the whole area. Just as many west-east birds traverse one side of the large square as cross the whole square. In other words, the inclusion of additional sections athwart the direction of flight involves the inclusion of additional birds proceeding in that direc- tion, while the inclusion of additional sections along the direction does not. The correct ratio of the sample to the whole would seem to be the ratio of their perimeters, in this case the ratio of one to four. When this factor of four is applied to the problem it proves correct : 4 x 3 (the number of birds that have been seen in the sample square) equals 12 (the exact number of birds that could be seen in the square mile).
lunar coordinates in terms of their corresponding flight paths paral- lel to the earth is constantly changing with the position of the moon, the calculation of each bird's flight separately would require a tre- mendous amount of time and effort.
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Whatever we do, computed individual flight directions must be frankly recognized as approximations. Their anticipated inaccura- cies are not the result of defects in the mathematical procedure em- ployed. This is rigorous. The difficulty lies in the impossibility of reading the slants of the pathways on the moon precisely and in the
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Fig. 8. Rectangular samples of square areas. In Diagram A, where as many birds are flying from west to east as are flying from south to north, the perime- ter ratio (three to eight) correctly expresses the number of birds that have traversed the whole area relative to the number that have passed through the sample. But in Diagram B, where all thirty-two birds are flying from south to north, the correct ratio is the ratio of the base of the sample to the base of the total area (one to four), and use of the perimeter ratio would lead to an inaccurate result (forty-three instead of thirty-two birds). Perimeter ratios do not correctly express relative interceptory potential, unless the shape of the sample is the same as the shape of the whole, or unless the birds are flying in all directions equally.
three-dimensional nature of movement through space. The observed coordinates of birds' pathways across the moon are the projected product of two component angles — the compass direction of the flight and its slope off the horizontal, or gradient. These two factors cannot be dissociated by any technique yet developed. All we can do is to compute what a bird's course would be, if it were flying hori- zontal to the earth during the interval it passes before the moon. We cannot reasonably assume, of course, that all nocturnal migra- tion takes place on level planes, even though the local distractions so often associated with sloping flight during the day are minimized in the case of migrating birds proceeding toward a distant destina- tion in darkness. We may more safely suppose, however, that de-
Lowery: Nocturnal Migration of Birds
383
Fig. 9. The effect of vertical components in bird flight. The four diagrams illustrate various effects that might result if a bird with an undulating flight, such as a Goldfinch, flew before a moon 45° above the horizon. In each case the original profile of the pathways, illustrated against the dark background, is flattened considerably as a result of projection. In the situation shown in Diagram A, where the high point of the flight line, GHJ, occurs within the field of the telescope, it is not only obvious that a deviation is involved, but the line GJ drawn between the entry and departure points coincides with the nor- mal coordinates of a bird proceeding on a horizontal plane. In Diagrams B and C, one which catches an upward segment of flight, and the other, a down- ward segment, the nature of the deviation would not be detectable, and an incorrect direction would be computed from the coordinates. Over a series of observations, including many Goldfinches, one would expect a fairly even dis- tributioij of ups and downs. Since the average between the coordinate angles in Diagrams B and C, -|- 19° and -19°, is the angle of the true coordinate, we have here a situation where the errors tend to compensate. In Diagram D, where the bird is so far away that several undulations are encompassed within the diameter of the field of view, the coordinate readings do not differ mate- rially from those of a straight line.
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University of Kansas Publs., Mus. Nat. Hist.
viations from the horizontal are random in nature, that it is mainly a matter of chance whether the observer happens to see an ascending segment of flight or a descending one. Over a series of observations, we may expect a fairly even distribution of ups and downs. It fol- lows that, although departures from the horizontal may distort in- dividual directions, they tend to average out in the computed trend of the mean. The working of this principle applied to the undulating flight of the Goldfinch (Spinus) is illustrated in Figure 9.
Since individually computed directions are not very reliable in any event, little is to be lost by treating the observed pathways in groups. Consequently, the courses of all the birds seen in a one-hour period may be computed according to the position of the moon at the middle of the interval and expressed in terms of their general positions on the compass, rather than their exact headings. For this latter pur- pose, the compass has been divided into twelve fixed sectors, 22i/^ degrees wide. The trends of the flight paths are identified by the mid-direction of the sector into which they fall. The sectoring method is described in detail in the section on procedures.
The problem remains of converting the number of birds involved in each directional trend to a fixed standard of measurement. Figure 7A contains the partial elements of a solution. All of the west-east flight paths that cross the large square also cross one of its mile-long sides and suggest the practicability of expressing the amount of mi-
NUMBER OF BIROS
Fig. 10. The interceptory potential of slanting lines. The diagram deals with one direction of flight and its incidence across lines of six different slants, lines of identical length oriented in six different ways. Obviously, the number of birds that cross a line depends not only on the length of the line, but also on its slant with respect to the flight paths.
Lowery: Nocturnal Migration of Birds 385
gration in any certain direction in terms of the assumed quantity passing over a one-mile line in a given interval of time. However, many lines of that length can be included within the same set of flight paths (Figure 10) ; and the number of birds intercepted depends in part upon the orientation of the line. The 90° line is the only one that fully measures the amount of fight per linear unit of front; and so I have chosen as a standard an imaginary mile on the earth's surface lying at right angles to the direction in which the birds are traveling. Definitions of Flight Density
When the count of birds in the cone of observation is used as a sample to determine the theoretical number in a sector passing over such a mile line, the resulting quantity represents what I shall call a Sector Density. It is one of several expressions of the more general concept of Flight Density, which may be defined as the passage of migration past an observation station stated in terms of the theo- retical number of birds flying over a one-mile line on the earth's surface in a given interval of time. Note that a flight density is primarily a theoretical number, a statistical expression, a rate of passage. It states merely that birds were moving through the effec- tive field of observation at the rate of so many per mile per unit of time. It may or may not closely express the amount of migration occurring over an actual mile or series of miles. The extent to which it does so is to be decided by other general criteria and by the circum- stances surrounding a given instance. Its basic function is to take counts of birds made at different times and at different places, in fields of observation of different sizes, and to put them on the sta- tistically equal footing that is the first requisite of any sound com- parison.
The idea of a one-mile line as a standard spacial measurement is an integral part of the basic concept, as herein propounded. But, within these limitations, flight density may be expressed in many different ways, distinguished chiefly by the directions included and the orientation of the one-mile line with respect to them. Three such kinds of density have been found extremely useful in subsequent analyses and are extensively employed in this paper: Sector, Net Trend, and Station Density, or Station Magnitude.
Sector Density has already been referred to. It may be defined as the flight density within a 22%° directional spread, or sector, measured across a one-mile line lying at right angles to the mid- direction of the sector. It is the basic type of density from the point
386 University of Kansas Publs., Mus. Nat. Hist.
of view of the computer, the others being derived from it. In analysis it provides a means of comparing directional trends at the same station and of studying variation in directional fanning.
Net Trend Density represents the maximum net flow of migration over a one-mile line. It is found by plotting the sector densities directionally as lines of thrust, proportioned according to the density in each sector, and using vector analysis to obtain a vector resultant, representing the density and direction of the net trend. The mile line defining the spacial limits lies at right angles to this vector re- sultant, but the density figure includes all of the birds crossing the line, not just those that do so at a specified angle. Much of the directional spread exhibited by sector densities undoubtedly has no basis in reality but results from inaccuracies in coordinate readings and from practical difficulties inherent in the method of computa- tion. By reducing all directions to one major trend, net trend den- sity has the advantage of balancing errors one against the other and may often give the truer index to the way in which the birds are actually going. On the other hand, if the basic directions are too widely spread or if the major sector vectors are widely separated with little or no representation between, the net trend density may become an abstraction, expressing the idea of a mean direction but pointing down an avenue along which no migrants are traveling. In such instances, little of importance can be learned from it. In others, it gives an idea of general trends indispensable in comparing station with station to test the existence of flyways and in mapping the con- tinental distribution of flight on a given night to study the influence of weather factors.
Station Density, or Station Magnitude, represents all of the mi- gration activity in an hour in the vicinity of the observation point, regardless of direction. It expresses the sum of all sector densities. It includes, therefore, the birds flying at right angles over several one-mile lines. One way of picturing its physical meaning is to imagine a circle one-mile in diameter lying on the earth with the observation point in the center. Then all of the birds that fly over this circle in an hour's time constitute the hourly station density. While its visualization thus suggests the idea of an area, it is derived from linear expressions of density; and, while it involves no limita- tion with respect to direction, it could not be computed without tak- ing every component direction into consideration. Station density is adapted to studies involving the total migration activity at vari- ous stations. So far it has been the most profitable of all the density
Lowery: Nocturnal Migration of Birds 387
concepts, throwing important light on nocturnal rhythm, seasonal increases in migration, and the vexing problem of the distribution of migrating birds in the region of the Gulf of Mexico.
Details of procedure in arriving at these three types of flight den- sity will be explained in Section B of this discussion. For the mo- ment, it will suffice to review and amplify somewhat the general idea involved.
Altitude as a Factor in Flight Density
A flight density, as we have seen, may be defined as the number of birds passing over a line one mile long; and it may fee calculated from the number of birds crossing the segment of that line included in an elliptical cross-section of the cone of observation. It may be thought of with equal correctness, without in any way contradicting the accuracy of the original definition, as the number of birds pass- ing through a vertical plane one mile long whose upper limits are its intersection with the flight ceiling and whose base coincides with the one mile line of the previous visualization. From the second point of view, the sample becomes an area bounded by the triangu- lar projection of the cone of observation on the density plane. The dimensions of two triangles thus determined from any two cones of observation stand in the same ratio as the dimensions of their ellip- tical sections on any one plane; so both approaches lead ultimately to the same result. The advantage of this alternative way of looking at things is that it enables us to consider the vertical aspects of mi- gration— to comprehend the relation of altitude to bird density.
If the field of observation were cylindrical in shape, if it had parallel sides, if its projection were a rectangle or a parallelogram, the height at which birds are flying would not be a factor in finding out their number. Then the sample would be of equal breadth throughout, with an equally wide representation of the flight at all levels. Since the field of observation is actually an inverted cone, triangular in section, with diverging sides, the opportunity to detect birds increases with their distance from the observer. The chances of seeing the birds passing below an elevation midway to the flight ceiling are only one-third as great as of seeing those passing above that elevation, simply because the area of that part of the triangle below the mid-elevation is only one-third as great as the area of that part above the mid-elevation. If we assume that the ratio of the visible number of birds to the number passing through the density plane is the same as the ratio of the triangular section of the cone
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University of Kansas Publs., Mus. Nat. Hist.
to the total area of the plane, we are in effect assuming that the density plane is made up of a series of triangles the size of the sample, each intercepting approximately the same number of birds. We are assuming that the same number of birds pass through the inverted triangular sample as through the erect and uninvestigable
A" H™K
I D"J
Fig. 11. Theoretical possibilities of vertical distribution. Dia- gram I shows the effect of a uniform vertical distribution of birds. The figures indicate the number of birds in the respective areas. Here the sample triangle, ABD, contains the same number of birds as the upright triangle, ACD, adjacent to it; the density plane may be conceived of as a series of such alternating triangles, equal in their content of birds. Diagram II portrays, on an exaggerated scale, the situation when many more birds are flying below the me- dian altitude than above it. In contrast to the 152 birds occurring in the triangle A'C'D', only seventy-two are seen in the triangle A'B'D'. Obviously, the latter triangle does not provide a represen- tative sample of the total number of birds intersecting the density plane. Diagram III illustrates one method by which this difficulty may be overcome. By lowering the line F'G' to the median alti- tude of bird den.sity, F"G" (the elevation above which there are just as many birds as below), we are able to determine a rectangu- lar panel, HIJK, whose content of birds provides a representative sample of the vertical distribution.
triangle beside it (as in Figure 11, Diagram II). In reality, the as- sumption is sound only if the altitudinal distribution of migrants is uniform.
The definite data on this subject are meagre. Nearly half a cen- tury ago, Stebbins worked out a way of measuring the altitude of
Lowery: Nocturnal Migration of Birds 389
migrating birds by the principle of parallax. In this method, the distance of a bird from the observers is calculated from its apparent displacement on the moon as seen through two telescopes. Stebbins and his colleague, Carpenter, published the results of two nights of observation at Urbana, Illinois (Stebbins, 1906; Carpenter, 1906) ; and then the idea was dropped until 1945, when Rense and I briefly applied an adaptation of it to migration studies at Baton Rouge. Results have been inconclusive. This is partly because sufficient work has not been done, partly because of limitations in the method itself. If the two telescopes are widely spaced, few birds are seen by both observers, and hence few parallaxes are obtained. If the instruments are brought close together, the dis- placement of the images is so reduced that extremely fine readings of their positions are required, and the margin of error is greatly increased. Neither alternative can provide an accurate representa- tive sample of the altitudinal distribution of migrants at a station on a single night. New approaches currently under consideration have not yet been perfected.
Meanwhile the idea of uniform vertical distribution of migrants must be dismissed from serious consideration on logical grounds. We know that bird flight cannot extend endlessly upward into the sky, and the notion that there might be a point to which bird density extends in considerable magnitude and then abruptly drops off to nothing is absurd. It is far more likely that the migrants gradually dwindle in number through the upper limits at which they fly, and the parallax observations we have seem to support this view.
Under these conditions, there would be a lighter incidence of birds in the sample triangle than in the upright triangle beside it (Figure 11, Diagram III). Compensation can be made by de- liberately scaling down the computed size of the sample area below its actual size. A procedure for doing this is explained in Figure 11. If it were applied to present altitudinal data, it would place the computational flight ceiling somewhere below 4000 feet. In arriving at the flight densities used in this paper, however, I have used an assumed ceiling of one mile. When the altitude factor is thus as- signed a value of 1, it disappears from the formula, simplifying computations. Until the true situation with respect to the vertical distribution of flight is better understood, it seems hardly worth- while to sacrifice the convenience of this approximation to a rigorous interpretation of scanty data. This particular uncertainty, however, does not necessarily impair the analytical value of the
390 University of Kansas Publs., Mus. Nat. Hist.
computations. Provided that the vertical pattern of migration is more or less constant, flight densities still afford a sound basis for comparisons, wherever we assume the upper flight limits to be. Raising or lowering the flight ceiling merely increases or reduces all sample cones or triangles proportionately.
A more serious possibility is that the altitudinal pattern may vary according to time or place. This might upset comparisons. If the divergencies were severe enough and frequent enough, they could throw the study of flight densities into utter confusion.
This consideration of possible variation in the altitudinal pattern combines with accidents of sampling and the concessions to perfect accuracy, explained on pages 379-385, to give to small quantities of data an equivocal quality. As large-scale as the present survey is from one point of view, it is only a beginning. Years of intensive work and development leading to a vast accumulation of data must elapse before the preliminary indications yet discernible assume the status of proved principles. As a result, much of the discussion in Part II of this paper is speculative in intent, and most of the conclusions suggested are of a provisional nature. Yet, compared with similar procedures in its field, flight density study is a highly objective method, and a relatively reliable one. In no other type of bird census has there ever been so near a certainty of recording all of the individuals in a specified space, so nearly independently of the subjective interpretations of the observer. The best assurance of the essential soundness of the flight density computations lies in the coherent results and the orderly patterns that already emerge from the analyses presented in Part II.
B. Observational Procedure and the Processing of Data
. At least two people are required to operate an observation station — one to observe, the other to record the results. They should ex- change duties every hour to avoid undue eye fatigue. Additional personnel are desirable so that the night can be divided into shifts.
Essential materials and equipment include: (1) a small telescope; (2) a tripod with pan-tilt or turret head and a mounting cradle; (3) data sheets similar to the one illustrated in Figure 12. Bausch and Lomb or Argus spotting scopes (19.5 x) and astronomical tele- scopes up to 30- or 40-power are ideal. Instruments of higher magnification are subject to vibration, unless very firmly mounted, and lead to difficulties in following the progress of the moon, unless powered by clockwork. Cradles usually have to be devised. An
Lowery: Nocturnal Migration of Birds
391
ORIGINAL DATA SHEET
DATE
OBSEPVERS_ WEATHER __
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Fig. 12. Facsimile of form used to record data in the field. One sheet of the actual observations obtained at Progreso, Yucatan, on April 24-25, 1948, is reproduced here. The remainder of this set of data, which is to be used throughout the demonstration of procedures, is shown in Table 1.
adjustable lawn chair is an important factor in comfort in latitudes where the moon reaches a point high overhead.
As much detail as possible should be entered in the space provided at the top of the data sheet. Information on the weather should include temperature, description of cloud cover, if any, and the
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University of Kansas Publs., Mus. Nat. Hist.
direction and apparent speed of surface winds. Care should be taken to specify whether the telescope used has an erect or inverted image. The entry under "Remarks" in the heading should de- scribe the location of the observation station with respect to watercourses, habitations, and prominent terrain features.
The starting time is noted at the top of the "Time" column, and the observer begins the watch for birds. He must keep the disc of the moon under unrelenting scrutiny all the while he is at the telescope. When interruptions do occur as a result of changing positions with the recorder, re-adjustments of the telescope, or the disappearance of the moon behind clouds, the exact duration of the "time out" must be set down.
Whenever a bird is seen, the exact time must be noted, together with its apparent pathway on the moon. These apparent pathways can be designated in a simple manner. The observer envisions the disc of the moon as the face of a clock, with twelve equally spaced points on the circumference marking the hours (Figure 13) . He calls the bottommost point 6 o'clock and the topmost, 12. The intervals in between are numbered accordingly. As this lunar clockface moves across the sky, it remains oriented in such a way that 6 o'clock con-
FiG. 13. The identification of coordinates. These diagrams illustrate how the moon may be envisioned as a clockface, constantly oriented with six o'clock nearest the horizon and completely independent of the rotation of the moon's topographic features.
Lowery: Nocturnal Migration of Birds
393
tinues to be the point nearest the horizon, unless the moon reaches a position directly overhead. Then, all points along the circumference are equidistant from the horizon, and the previous definition of clock values ceases to have meaning. This situation is rarely encountered in the northern hemisphere during the seasons of migration, except
Time 11:00-12:00 No. of Birds 86
Fig. 14. The apparent pathways of the birds seen in one hour. The obser- vations are those recorded in the 11 :00^12:00 p. m. interval on April 24-25, 1948, at Progreso, Yucatan (see Table 1).
3—1020
394 University of Kansas Publs., Mtjs. Nat. Hist.
in extreme southern latitudes. It is one that has never actually been dealt with in the course of this study. But, should the problem arise, it would probably be feasible to orient the clock during this interval with respect to the points of the compass, calling the south point 6 o'clock.
When a bird appears in front of the moon, the observer identifies its entry and departure points along the rim of the moon with respect to the nearest half hour on the imaginary clock and informs the re- corder. In the case of the bird shown in Figure 13, he would simply call out, "5 to 10:30." The recorder would enter "5" in the "In" col- umn on the data sheet (see Figure 12) and 10:30 in the "Out" col- umn. Other comment, offered by the observer and added in the re- marks column, may concern the size of the image, its speed, distinct- ness, and possible identity. Any deviation of the pathway from a straight line should be described. This information has no bearing on subsequent mathematical procedure, except as it helps to elimi- nate objects other than birds from computation.
The first step in processing a set of data so obtained is to blue- pencil all entries that, judged by the accompanying remarks, relate to extraneous objects such as insects or bats. Next, horizontal lines are drawn across the data sheets marking the beginning and the end of each even hour of observation, as 8 p. M.-9 p. m., 9 p. m.-IO p. M., etc. The coordinates of the birds in each one-hour interval may now be plotted on separate diagrammatic clockfaces, just as they appeared on the moon. Tick marks are added to each line to indicate the number of birds occurring along the same coordinate. The slant of the tick marks distinguishes the points of departure from the points of entry. Figure 14 shows the plot for the 11 p. m. — 12 p. M. observations reproduced in Table 1. The standard form, illustrated in Figure 15, includes four such diargrams.
Applying the self-evident principle that all pathways with the same slant represent the same direction, we may further consolidate the plots by shifting all coordinates to the corresponding lines pass- ing through the center of the circle, as in Figure 15. To illustrate, the 6 to 8, 5 to 9, 3 to 11, and 2 to 12 pathways all combine on the 4 to 10 line. Experienced computers eliminate a step by directly plotting the pathways through center, using a transparent plastic straightedge ruled ofif in parallel lines.
We now have a concise picture of the apparent pathways of all the birds recorded in each hour of observation. But the coordinates do not have the same meaning as readings of a horizontal clock on
Lowery: Nocturnal Migration of Birds
395
|
LOCALITY 0^njeQ,'«*-r> |
L uucoLvn, ll IMC. |
DATE ^^•^5 Pi^inJL. i^sa |
||||||
|
12 |
12 |
|||||||
|
11 ^ ' |
< |
> |
„>•- ' —^^^^t |
|||||
|
10 ^ |
N |
V |
10^ |
€ |
V2 |
|||
|
9-1 ^"^^*^ |
^ |
1,3 9 |
1 |
^^ |
J' |
|||
|
* \ |
y |
^ N8 = N6 = |
8> |
V |
V*4 "^ N8 = N6 = |
|||
|
T^*"**?-— ^ |
r^ |
s |
N4 = Nj = |
7*^ |
6 |
N4 = N2= \ Ni -- Z |
||
|
Tin.e S 00- qOO |
N3 = 2 |
Time |
q-oo- \o 00 |
N3=fe |
||||
|
Md. of Eird |
s 2 |
N5 = N7 = S = S r |
No. |
ol Birds Q |
N5 = N7 = S = S - |
|||
|
12 |
12 |
|||||||
|
'\^C\> |
r |
\ |
11 |
^ |
wHl^"*-^' |
|||
|
lo^s^^^Al |
V 2 |
'2 |
^ |
«i |
"v^ |
|||
|
p^^-^-o^^ |
/ |
\ |
f |
^-^.^ |
^^ |
\ |
||
|
^j ^^--^ |
^^ |
hi c |
{ |
J |
Jr' |
|||
|
eX / |
1 |
N6 = |
^ |
\ |
1 |
^4 N8 = N6 = |
||
|
5 |
N4 = |
7 |
^^**~1 |
-Mr^ 5 |
N4 = 3 |
|||
|
1 |
N2=2 Ni =,o |
6 |
N2= 3 NI = 2H |
|||||
|
Time 10 DO |
: MOO |
N3 = 21 |
Time |
II 00-12:00 |
N3= SS |
|||
|
No. ol Eire |
is -35 |
N5= 2. N7 = S = |
No. |
of Birds g^ |
N5= 2 N7=, S = |
|||
|
Computer (y. X.. |
S r |
S = |
Fig. 15. Standard form for plotting the apparent paths of flight. On these diagrams the original coordinates, exemplified by Figure 14, have been moved to center. In practice the sector boundaries are drawn over the circles in red pencil, as shown by the white lines in Figure 19, making it possible to count the number of birds falling within each zone. These numbers are then tallied in the columns at the lower right of each hourly diagram.
the earth's surface, placed in relation to the points of the compass. They are merely projections of the birds' courses. An equation is available for reversing the effect of projection and discovering the
396
University of Kansas Publs., Mus. Nat. Hist.
Table 1. — Continuation of Data in Figure 12, Showing Time and Readings of Observations on 24-25 April 1948, Progreso, Yucatan
|
Time |
In |
Out |
Time |
In |
Out |
|
10:37-10:41 Time out |
11:15 |
8 |
9:30 |
||
|
10:45 |
5:30 |
10 |
11:16 |
4 |
11 |
|
6 |
9 |
5 |
9 |
||
|
5:30 |
10 |
11:17 |
5 |
11:30 |
|
|
10:46 |
6 |
8 |
11:18 |
5 |
12 |
|
3:30 |
11 |
6 |
11:30 |
||
|
5 |
12 |
11:19 |
5:30 |
11:30 |
|
|
10:47 |
3:15 |
1 |
11:20 |
6 |
10 |
|
6 |
8:30 |
3 |
12 |
||
|
5:45 |
11:45 |
5 |
12 |
||
|
5 |
10 |
11:21 |
5:45 |
11 |
|
|
10:48 |
6 |
9:45 |
5 |
11 |
|
|
10:50 |
5:30 |
11 |
11:23 |
5 |
12 |
|
10:51 |
4 |
11 |
11:25 |
5 |
10:30 |
|
10:52 |
4 |
2 |
6 |
11 |
|
|
5:30 |
11 |
6 |
12 |
||
|
10:53 |
5:30 |
11:30 |
11:27 |
6 |
10 |
|
5 |
11 |
11:28 |
6 |
11:30 |
|
|
10:55 |
5 |
12 |
5:30 |
12:30 |
|
|
5 |
11 |
11:29 |
6 |
11:30 |
|
|
10:56 |
6 |
10 |
4 |
12 |
|
|
10:58 |
4:30 |
11:30 |
6:30 |
10:30 |
|
|
5:45 |
11:45 |
6 |
11 |
||
|
10:59 |
6:30 |
10:30 |
11:30 |
3 |
10 |
|
11:00 |
3:30 |
12 |
(2 birds at |
once) |
|
|
6:30 |
11 |
11:31 |
5 |
10:30 |
|
|
(2 birds at once) |
5:30 |
10:30 |
|||
|
11:03 |
6 |
11 |
11:32 |
6 |
11:30 |
|
11:04 |
3 |
12 |
11:33 |
7:30 |
9:30 |
|
5 |
12 |
4 |
10:30 |
||
|
11:05 |
6 |
10 |
6 |
11:30 |
|
|
5 |
11 |
8 |
9:30 |
||
|
11:06 |
6 |
10:30 |
11:35 |
7 |
10 |
|
11:07 |
3 |
10 |
4:30 |
1 |
|
|
11:08 |
6 |
11 |
11:38 |
6:30 |
11 |
|
11:10 |
7 |
9:30 |
11:40 |
5:30 |
12 |
|
11:11 |
5 |
9:15 |
11:42 |
4 |
2 |
|
11:13 |
5 |
12 |
5 |
12 |
|
|
11:14 |
6:30 |
10 |
6 |
10 |
|
|
5:30 |
1 |
4 |
2 |
||
|
4 |
12 |
5 |
12 |
Lowery: Nocturnal Migration of Birds
397
Table 1. — Concluded
|
Time |
In |
Out |
Time |
In |
Out |
|
11:44 |
8 |
9:30 |
8 |
10:15 |
|
|
7 |
11 |
12:16 |
3:30 |
1:30 |
|
|
6 |
10 |
8 |
11 |
||
|
11:45 |
5 |
12 |
12:23 |
7 |
1:30 |
|
6 |
10:30 |
6 |
12:30 |
||
|
5:45 |
11 |
12:36 |
8 |
11 |
|
|
4 |
12 |
12:37 |
7:30 |
1 |
|
|
11:46 |
7 |
11 |
12:38 |
7 |
12:30 |
|
6 |
12 |
12:40 |
8 |
1 |
|
|
11:47 |
8 |
10 |
12:45 |
7:30 |
1 |
|
11:48 |
6 |
10 |
12:47 |
5:30 |
1 |
|
11:49 |
6:30 |
10:30 |
12:48 |
7 |
1 |
|
11:51 |
8 |
10 |
12:52 |
5:30 |
1:30 |
|
8 |
10 |
12:54-12:55 Time out |
|||
|
8 |
10 |
12:56 |
8 |
10:45 |
|
|
8 |
10 |
12:58 |
5:30 |
1:30 |
|
|
6 |
10 |
7 |
1:30 |
||
|
8 |
10 |
7 |
2 |
||
|
6 |
11 |
12:59 |
5 |
3 |
|
|
7 |
12 |
1 :00-l :30 |
Time out |
||
|
11:52 |
5 |
1 |
1:37 |
8 |
12 |
|
11:54 |
7 |
11 |
1:38 |
8 |
12 |
|
6 |
12:30 |
1:48 |
7 |
1 |
|
|
11:55 |
5 |
12 |
7 |
1 |
|
|
11:56 |
7 |
10 |
1:51 |
5:30 |
11 |
|
5 |
12 |
1:57 |
8 |
1 |
|
|
11:58 |
8 |
11 |
2:07 |
7 |
2 |
|
11:59 |
5:30 |
12 |
2:09 |
9 |
12 |
|
12:00-12 |
:03 Time out |
2:10 |
8 |
1 |
|
|
12:03 |
5:30 |
11:30 |
2:17 |
9 |
12 |
|
12:04 |
8 |
11 |
2:21 |
6 |
2 |
|
12:07 |
6 |
12:30 |
2:30 |
5:30 |
3:15 |
|
7:30 |
1 |
2:32 |
8 |
2 |
|
|
12:08 |
5 |
10:30 |
2:46 |
7 |
1 |
|
12:09 |
5:30 |
1 |
3:36 |
9 |
2 |
|
7:30 |
2 |
3:39 |
8:30 |
2 |
|
|
12:10 |
6:30 |
12:45 |
3:45 |
6 |
4 |
|
12:13 |
8 |
11 |
3:55 |
9 |
2 |
|
12:14 |
7 |
1 |
4:00 |
8 |
3 |
|
12:15 |
7 |
12:30 |
4:03 |
9 |
2 |
|
7:15 |
1:30 |
4:30 |
Closed station |
398
University of Kansas Publs., Mus. Nat. Hist.
true directions of flight. This formula, requiring thirty-five sepa- rate computations for the pathways reproduced in Figure 12 alone, is far too-consuming for the handling of large quantities of data. A simpler procedure is to divide the compass into sectors and, with the aid of a reverse equation, to draw in the projected boundaries of these divisions on the circular diagrams of the moon. A standard-
ENE
ESE
WNW
SSE
SSW
Fig. 16. Standard sectors for designating flight trends. Each zone covers a span of 22^/l>°. The No and Ng, the N.^s and N7, and their south complements, where usually few birds are represented, can be combined and identified as' Ne-s and N5-7, etc.
ized set of sectors, each 22i/2° wide and bounded by points of the compass, has been evolved for this purpose. They are identified as shown in Figure 16. The zones north of the east-west line are known as the North, or N, Sectors, as Ni, N2, Ns, etc. Each zone south of the east-west line bears the same number as the sector opposite, but is distinguished by the designation S.
Several methods may be used to find the projection of the sector boundaries on the plot diagrams of Figure 15. Time may be saved by reference to graphic tables, too lengthy for reproduction here, showing the projected reading in degrees for every boundary, at every position of the moon; and a mechanical device, designed by
Lowery: Nocturnal Migration of Birds
399
Fig. 17. The meaning of symbols used in the direction formula.
C. M. Arney, duplicating the conditions of the original projection, speeds up the work even further. Both methods are based on the principle of the following formula:
tan (17 — ip)
tan 6 :=
(1)
cos Zo The symbols have these meanings:
6 is the position angle of the sector boundary on the lunar clock, with positive values measured counterclockwise from 12 o'clock, negative angles clockwise (Figure 17A).
7) is the compass direction of the sector boundary expressed in de- grees reckoned west from the south point (Figure 17B).
Zo is the zenith distance of the moon's center midway through the
400 University of Kansas Publs., Mrs. Nat. Hist.
hour of observation, that is, at the half hour. It represents the number of degrees of arc between the center of the moon and a point directly over the observer's head (Figure 17C).
ij/ is the azimuth of the moon midway through the hour of obser- vation, measured from the south point, positive values to the west, negative values to the east (Figure 17D).
COMPUTATION OF ZENITH DISTANCE AND AZIMUTH OF MOON DATE 2^-35 (3p/L.4«t or AT TTvU^nyiy^^ /"/..rntniTI I. AT JJ'IT'N LOtJG .89'S9' W
Avg. Time 8;30 CST on Z^dfO^i^ = 03-30 GCT on JZ5 ClfvUt
GHA = /8°4-3' Declination of Moon = 5 I9°^8' ,
LHA = GHA — Local Long. = IB'^S''— 89°39' = 70" 56
Altitude = 5'»Jj' .•,90° — Alt. = 90°— 9*^3' = 80" ZT'= z^
Azimuthn =//5«'44' Az„ - 180° = //5''44' - 180° =-64-''(6'= ^l o
Avg. Time 9:30 CST on Z4 Opnii. = 0330 GCT on 25 dp/Ut
GHA = 33"//' Declination of Moon= S ZO'OO'
LHA = GHA — Local Long. = 33°'/' —89°35' = 3^' Z&'
Altitude = Z-COS' .-.90°- Alt. = 90° — ^.l-'OS'. = feS'SS = Zo
Aziinuthn=/2;2''54' Az„- 180° = /22°^4.'.- 180° =~57'0(,' = Vo
I .. . I II I — - - ^ , 1
Avg. Time /0-30 CST on 24^2/ev«-4 = 0430 GCT on ZSdpruJl,
GHA = 47 "40' Declination of Moon = 5 20°/;^ ,
LHA = GHA — Local Long. = 47°40'— 89° 39' - 4/ ° 59
Altitude = 3/*39' .'.90° — Alt. = 90° — 3/''39' = 5B°Z\' = Zo
Azimuthn = /3Z° 30^ Az^- 180° = /32''30'' - 180° =~^7°30' = ^o
Avg. Time //-SOcSTon Z-4 C^f^ = O530 GCT on Z5 (2f>ru£
GHA = (oT/OZ' Declination of Moon = S S-C^a'
LHA = GHA — Local Long. = ifX" OQ— 89''39' = Z7°3I^
Altitudes 4-0°Z5' .•. 90° — Alt. = 90°— 40''Z5' = ■^g'35'' = Zo
Azimuthn =/45<'/a' Az„ - 180° = /f 5°/g' - 180° ^ =-34''4>s' = Vo
;. Time /Z:30CST on Z^(^2fxnjJL = Ob30 GCT on 25 dattjJi A = 7i,''37'' Declination of Moon = SZO''3Z''
Avg.
GHA
LHA = GHA — Local Long. = 76''37 — 89''39' = fi'O-a/
Altitude = 46''3 5' .'. 90° —Alt. = 90° — 4-4,''35' = 43''.25' = Zo
Azimuthn =/6;Z''0O " Az^ - 180° = /62°CO'- 180° ^-l&°Oo' = ypo
Avg. Time ST on = GCT on
GHA = Declination of Mocrn =
LHA = GHA — Local Long. = — =
Altitude = •. 90° — Alt. = 90° — = = Zo
Azimuthn = Az - 180° = —180° = = U'o
n
11 CHA is greater than Local Long, Azimuth ( ^q) is positive
If GHA is less than Local Long., Azimuth is negative Computer
Fig. 18. Form used in the computation of the zenith distance and azimuth of the moon.
Lowery: Nocturnal Migration of Birds 401
The angle 7? for any sector boundary can be found immediately by measuring its position in the diagram (Figure 16) . The form (Figure 18) for the "Computation of Zenith Distance and Azimuth of the Moon" illustrates the steps in calculating the values of Zo and »/^o- From the American Air Almanac (Anonymous, 1945-1948), issued annually by the U. S. Naval Observatory in three volumes, each covering four months of the year, the Greenwich Hour Angle (GHA) and the declination of the moon may be obtained for any ten-minute interval of the date in question. The Local Hour Angle (LHA) of the observation station is determined by subtracting the longitude of the station from the GHA. Reference is then made to the "Tables of Computed Altitude and Azimuth," published by the U. S. Navy Department, Hydrographic Office (Anonymous, 1936- 1941), and better known as the "H. 0. 214," to locate the altitude and azimuth of the moon at the particular station for the middle of the hour during which the observations were made. The tables employ three variables — the latitude of the locality measured to the nearest degree, the LHA as determined above, and the decli- nation of the moon measured to the nearest 30 minutes of arc. Interpolations can be made, but this exactness is not required. When the latitude of the observation station is in the northern hemi- sphere, the H. O. 214 tables entitled "Declinations Contrary Name to Latitude" are used with south declinations of the moon, and the tables "Declinations Same Name as Latitude," with north declina- tions. In the sample shown in Figure 15, the declination of the moon at 11:30 p. m., midway through the 11 to 12 o'clock interval, was S 20° 22'. Since the latitude of Frogreso, Yucatan i? N 21° 17', the "Contrary Name" tables apply to this hour
Because the H.O. 214 expresses the vertical position of the moon in terms of its altitude, instead of its zenith distance, a conversion is required. The former is the number of arc degrees from the hori- zon to the moon's center; therefore Zo is readily obtained by sub- tracting the altitude from 90°. Moreover, the azimuth given in the H.O. 214 is measured on a 360° scale from the north point, whereas the azimuth used here (i/^o) is measured 180° in either direction from the south point, negative values to the east, positive values to the west. I have designated the azimuth of the tables as Azn and ob- tained the desired azimuth (i/^o) by subtracting 180° from Azn. The sign of ij/o may be either positive or negative, depending on whether or not the moon has reached its zenith and hence the meridian of the observer. When the GHA is greater than the local longitude
402
University of Kansas Publs., Mus. Nat. Hist.
(that is, the longitude of the observation station), the azimuth is positive. When the GHA is less than the local longitude, the azimuth is negative.
Locating the position of a particular sector boundary now be- comes a mere matter of substituting the values in the equation (1) and reducing. The computation of the north point for 11 to 12 p. m. in the sample set of data will serve as an example. Since the north point reckoned west from the south point is 180°, its ?/ has a value of 180°.
Fia. 19. Method of plotting sector boundaries on the diagrammatic plots. The example employed is the 11:00 to 12:00 p. m. diagram of Figure 15.
Lowery: Nocturnal Migration of Birds
403
tan 6 Npt.
tan (180°— V'o)
cos Zo Substituting values of xpo found on the form (Figure 18)
tan 6 Npt.
tan [180°— (—35°)] tan 215'
.700
= 1.09
cos 50° cos 50° .643
e Npt. = 47°28'
Four angles, one in each quadrant, have the same tangent value.
|
COMPUTATIONS O LOCALITY ,Q .np^^liO.'-^p^^it*^ D 60 _ (220) T (No of Birds) {cos^ Zq |
F SECTOR DENSITIES ijau^ 3 ATF. 2S-25n^iqs8cOMPUTERG.L. »? 8 = ^5'?° 'I I = 169° 16= ^36° 1 J i 146° ^6-8 = ^48° 1 5.7= in° 1 4 =2140 V ^ = 124'^ 1 2 = Hl° 1 7 =101° |
||
|
Vl — sin^ Zo cosset a = 180° — Ti t f |
|||
|
Avg TimP 1110 7onP N4 7.,^ (220) (M) ( 3 ) (.Vl2) p - feO , |
50° |
_3in" Z^ /I |
= .588 ,1- . - 35=" |
|
- lano 0 III ^ i — -IC* 1 |
|||
|
Vl _ (.588) ( .128) ,. . (ZT3 ) - 273 |
a |
■= 69' |
|
|
5M^ |
cosher = \23 |
||
|
Vi-(.753) •■^"^' |
|||
|
Avg Tim* 1 1 ' 30 7 .^n» NZ 7 ,- |
SO-' |
sin^Zo- -SSa ;^^= -35- a. 180° -iqi**^ (-35") |
|
|
Vl - (.588) (.^a^) n = (2.73 ) , 273 |
a |
f = Hfe* |
|
|
323 |
cos'ct = 433 |
||
|
Vi_(.28M) •^'^^ |
|||
|
Avg. TLme M 'iO Zonpf^l. Z^= . (220){±^) ( 2M )(.MI2) |
50* |
_sin^ Z^ |
- .588 li - -35" |
|
= |
> ^ 0 r = 180° — U9*t ( -TS" 1 |
||
|
V « 1 |
|||
|
V>- (.588) (.835) D= (2190) - 2190 |
o |
' = 2H« |
|
|
3070 |
cos'^a- = .8 35 |
||
|
Vi_(.S9i) -''^ |
|||
|
Avg TimP U lO 7r,np (N3 7. = ("0)(-||)(5M X.HIZ) |
50" |
_3ir.^ Zo a |
- .588 ^ . -35" |
|
r = 180° —\nc'* ( -i<;o ) |
|||
|
Vi-(.58a) (.s<<<») n= (M920^ . SqzO |
a |
r- (*• |
|
|
76, CO |
cos^a -- .«1<1<T |
||
|
Vi-(.5aa) •^'^'^ |
|||
Fia. 20. Form for computing sector densities.
404
University of Kansas Publs., Mus. Nat. Hist.
Since, in processing spring data, we are dealing mainly with north sectors, it is convenient to choose the acute angle, in this instance 47°28'. In doubtful cases, the value of the numerator of the equa- tion (here 215°) applied as an angular measure from 6 o'clock will tell in which quadrant the projected boundary must fall. The fact that projection always draws the boundary closer to the 3-9 line serves as a further check on the computation.
W
Fig. 21. Determinaton of the angle cc
In the same manner, the projected position angles of all the perti- nent sector boundaries for a given hour may be calculated and plot- ted in red pencil with a protractor on the circular diagrams of Figure 15. To avoid confusion in lines, the zones are not portrayed in the black and white reproduction of the sample plot form. They are shown, however, in the shaded enlargement (Figure 19) of the 11 to 12 p. M. diagram. The number of birds recorded for each sector may be ascertained by counting the number of tally marks between each pair of boundary lines and the information may be entered in the columns provided in the plot form (Figure 15).
Lowery: Nocturnal Migration of Birds
405
We are now prepared to turn to the form for "Computations of
Sector Densities" (Figure 20), which systematizes the solution of
the following equation:
60 (220) — (No. of Birds) (cos^ Zo)
D = (2)
i
sin- Zj cos- a
SUMMARY OF SECTOR DENSITIES
LOCALITY I) -WgryjUto.Uuuc&fcjn. DATE 2S-25 Atvu]!. I'^HScOMPUTER G.L.
|
Zone |
Average Time of Observation C.S-T |
Total |
||||||||
|
8 30 |
q 30 |
10 30 |
11 iO |
1230 |
1 30 |
2 30 |
3 30 |
M'JO |
||
|
N8 E-F.NE |
||||||||||
|
N6 ENE-NE |
||||||||||
|
N4 NE-NNF. |
5sq |
500 |
||||||||
|
N2 NNE-N |
31 |
IS8 |
323 |
25S |
320 |
1 too |
||||
|
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Fig. 22. Facsimile of form summarizing sector densities. The totals at the bottom of each column give the station densities.
406
University of Kansas Publs., Mus. Nat. Hist.
Some of the symbols and factors, appearing here for the first time, require brief explanation. D stands for Sector Density. The constant, 220, is the reciprocal of the quotient of the angular di- ameter of the moon divided by 2. T is Time In, arrived at by sub-
VECTOR RESULTANT
A
4000 BIRDS
Fig. 23. Determination of Net Trend Density.
tracting the total number of minutes of time out, as noted for each hour on the original data sheets, from 60. "No. of Birds" is the number for the sector and hour in question as just determined on the plot form. The symbol a represents the angle between the mid- line of the sector and the azimuth line of the moon. The quantity is found by the equation:
a ^^ 180 — rj -\- <fo
(3)
Lowery: Nocturnal Migration of Birds
407
The symbol t] here represents the position of the mid-line of the sector expressed in terms of its 360° compass reading. This equation is illustrated in Figure 21. The values of rj for various zones are given in the upper right-hand corner of the form (Figure 20) . The subsequent reductions of the equations, as they appear in the figure for four zones, are self-explanatory. The end result, representing the sector density, is entered in the rectangular box provided.
After all the sector densities have been computed, they are tabu- lated on a form for the "Summary of Sector Densities" (Figure 22). By totaling each vertical column, sums are obtained, expressing the Station Density or Station Magnitude for each hour.
An informative way of depicting the densities in each zone is to plot them as lines of thrust, as in Figure 23. Each sector is repre- sented by the directional slant of its mid-line drawn to a length ex- pressing the flight density per zone on some chosen scale, such as 100 birds per millimeter. Standard methods of vector analysis are then applied to find the vector resultant. This is done by consider- ing the first two thrust lines as two sides of an imaginary parallelo- gram and using a drawing compass to draw intersecting arcs locating the position of the missing corner. In the same way, the third vector
14
12
10
in
b o o - 8
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o i 4
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10 It 12 I 2 3
HOUR OF THE NIGHT ( CST )
Fig. 24. Nightly station density curve at Progreso, Yucatan, on
April 24-25, 1948.
408 University of Kansas Publs., Mus. Nat. Hist.
is combined with the invisible resultant whose distal end is repre- sented by the intersection of the first two arcs. The process is re- peated successively with each vector until all have been, taken into consideration. The final intersection of arcs defines the length and slant of the Vector Resultant, whose magnitude expresses the Net Trend Density in terms of the original scale.
The final step in the processing of a set of observations is to plot on graph paper the nightly station density curve as illustrated by Figure 24.
PART II. THE NATURE OF NOCTURNAL MIGRATION
Present day concepts of the whole broad problem of bird migra- tion are made up of a few facts and many guesses. The evolutionary origin of migration, the modem necessities that preserve its biologic utility, the physiological processes associated with it, the sensory mechanisms that make it possible, the speed at which it is achieved, and the routes followed, all have been the subject of some investiga- tion and much conjecture. All, to a greater or less extent, remain matters of current controversy. All must be considered unknowns in every logical equation into which they enter. Since all aspects of the subject are intimately interrelated, since all have a bearing on the probabilities relating to any one, and since new conjectures must be judged largely in the light of old conjectures rather than against a background of ample facts, the whole field is one in which many alternative explanations of the established phenomena remain equally tenable. Projected into this uncertain atmosphere, any statistical approach such as determinations of flight density will require the accumulation of great masses of data before it is capable of yielding truly definitive answers to those questions that it is suited to solve. Yet, even in their initial applications, density analyses can do much to bring old hypotheses regarding nocturnal migration into sharper definition and to suggest new ones.
The number of birds recorded through the telescope at a par- ticular station at a particular time is the product of many potential variables. Some of these — like the changing size of the field of observation and the elevation of flight — pertain solely to the ca- pacity of the observer to see what is taking place. It is the function of the density and direction formulae to eliminate the influences of these two variables insofar as is possible, so that the realities of the situation take shape in a nearly statistically true form. There re- main to be considered those influences potentially responsible for
Lowery: Nocturnal Migration of Birds 409
variations in the real volume of migration at different times and places — things like the advance of season, geographic location, disposition of terrain features, hourly activity rhythm, wind cur- rents, and other climatological causes. The situation represented by any set of observations probably is the end result of the inter- action of several such factors. It is the task of the discussions that follow to analyze flight densities in the light of the circumstances surrounding them and by statistical insight to isolate the effects of single factors. When this has been done, we shall be brought closer to an understanding of these influences themselves as they apply to the seasonal movements of birds. Out of data that is essentially quantitative, conclusions of a qualitative nature will begin to take form. It should be constantly borne in mind, how- ever, that such conclusions relate to the movement of birds en masse and that caution must be used in applying these conclusions to any one species.
Since the dispersal of migrants in the night sky has a fundamental bearing on the sampling procedure itself, and therefore on the reli- ability of figures on flight density, consideration can well be given first to the horizontal distribution of birds on narrow fronts.
A. Horizontal Distribution of Birds on Narrow Fronts
Bird migration, as we know it in daytime, is characterized by spurts and uneven spatial patterns. Widely separated V's of geese go honking by. Blackbirds pass in dense recurrent clouds, now on one side of the observer, now on the other. Hawks ride along in narrow file down the thermal currents of the ridges. Herons, in companies of five to fifty, beat their way slowly along the line of the surf. And an unending stream of swallows courses low along the levees. Everywhere the impression is one of birds in bunches, with vast spaces of empty sky between.
Such a situation is ill-suited to the sort of sampling procedure on which flight density computations are based. If birds always traveled in widely separated flocks, many such flocks might pass near the cone of observation and still, by simple chance, fail to enter the sliver of space where they could be seen. Chance would be the dominating factor in the number of birds recorded, obscuring the effects of other influences. Birds would seldom be seen, but, when they did appear, a great many would be observed simul- taneously or in rapid succession.
4—1020
410 University of Kansas Publs., Mus. Nat. Hist.
When these telescopic studies were first undertaken at Baton Rouge in 1945, some assurance already existed, however, that night migrants might be so generally dispersed horizontally in the darkness above that the number passing through the small segment of sky where they could be counted would furnish a nearly proportionate sample of the total number passing in the neighborhood of the observation station. This assurance was provided by the very interesting account of Stone (1906: 249-252), who enjoyed the unique experience of viewing a nocturnal flight as a whole. On the night of March 27, 1906, a great conflagration occurred in Phila- delphia, illuminating the sky for a great distance and causing the birds overhead to stand out clearly as their bodies reflected the light. Early in the night few birds were seen in the sky, but thereafter they began to come in numbers, passing steadily from the southwest to the northeast. At ten o'clock the flight was at its height. The observer stated that two hundred birds were in sight at any given moment as he faced the direction from which they came. This un- paralleled observation is of such great importance that I quote it in part, as follows: "They [the birds] flew in a great scattered, wide- spread host, never in clusters, each bird advancing in a somewhat zigzag manner . , . Far off in front of me I could see them coming as mere specks . . . gradually growing larger as they approached . . . Over the illuminated area, and doubtless for great distances beyond, they seemed about evenly distributed . . . I am inclined to think that the migrants were not influ- enced by the fire, so far as their flight was concerned, as those far to the right were not coming toward the blaze but keeping steadily on their way . . . Up to eleven o'clock, when my observations ceased, it [the flight] continued apparently without abatement, and I am informed that it was still in progress at midnight."
Similarly, in rather rare instances in the course of the present study, the combination of special cloud formations and certain at- mospheric conditions has made it possible to see birds across the entire field of the telescope, whether they actually passed before the moon or not. In such cases the area of the sky under observation is greatly increased, and a large segment of the migratory movement can be studied. In my own experience of this sort, I have been forci- bly impressed by the apparent unifonnity and evenness of the pro- cession of migrants passing in review and the infrequence with which birds appeared in close proximity.
As striking as these broader optical views of nocturnal migration are, they have been too few to provide an incontestable basis for
Lowery: Nocturnal Migration of Birds
411
generalizations. A better test of the prevailing horizontal distribu- tion of night migrants lies in the analysis of the telescopic data themselves.
The distribution in time of birds seen by a single obsever may be studied profitably in this connection. Since the cone of observation is in constant motion, swinging across the front of birds migrating from south to north, each interval of time actually represents a dif- ferent position in space. This is evident from the map of the prog- ress of the field of observation across the terrain at Tampico, Tamau- lipas, on April 21-22, 1948 (Figure 25). At this station on this night, a total of 259 birds were counted between 7:45 p. m. and 3:45
Fig. 25. Positions of the cone of observation at Tampico, Tamps., on April 21-22. 1948. Essential features of this diagrammatic map are drawn to scale, the triangular white lines representing the projections of the cone of observation on the actual terrain at the mid-point of each hour of observation. If the distal ends of the position lines were connected, the portion of the map en- compassed would represent the area over which all the birds seen between 8:30 p. M. and 3:30 a. m. must have flown.
412 University of Kansas Publs., Mus. Nat. Hist.
A. M. The number seen in a single hour ranged from three to seventy-three, as the density overhead mounted to a peak and then declined. The number of birds seen per minute was not kept with stop watch accuracy; consequently, analysis of the number of birds that passed before the moon in short intervals of time is not justified. It appears significant, however, that in the ninety minutes of heav- iest flight, birds were counted at a remarkably uniform rate per fif- teen minute interval, notwithstanding the fact that early in the pe- riod the flight rate overhead had reached a peak and had begun to decline. The number of birds seen in successive fifteen-minute peri- ods was twenty-six, twenty-five, nineteen, eighteen, fifteen, and fifteen.
Also, despite the heavy volume of migration at this station on this particular night, the flight was sufficiently dispersed horizontally so that only twice in the course of eight hours of continuous observa- tion did more than one bird simultaneously appear before the moon. These were "a flock of six birds in formation" seen at 12:09 a.m. and "a flock of seven, medium-sized and distant," seen at 2:07 a. m. In the latter instance, as generally is the case when more than one bird is seen at a time, the moon had reached a rather low altitude, and consequently the cone of observation was approaching its maxi- mum dimensions.
The comparative frequency with which two or more birds simul- taneously cross before the moon would appear to indicate whether or not there is a tendency for migrants to fly in flocks. It is signifi- cant, therefore, that in the spring of 1948, when no less than 7,432 observations were made of birds passing before the moon, in only seventy-nine instances, or 1.1 percent of the cases, was more than one seen at a time. In sixty percent of these instances, only two birds were involved. In one instance, however, again when the moon was low and the cone of observation near its maximum size, a flock estimated at twenty-five was recorded.
The soundest approach of all to the study of horizontal distribu- tion at night, and one which may be employed any month, any- where, permitting the accumulation of statistically significant quan- tities of data, is to set up two telescopes in close proximity. Pro- vided the flight overhead is evenly dispersed, each observer should count approximately the same number of birds in a given interval of time. Some data of this type are already available. On May 19- 20, at Urbana, Illinois, while stationed twenty feet apart making parallax studies with two telescopes to determine the height above
Lowery: Nocturnal Migration of Birds 413
the earth of the migratory birds, Carpenter and Stebbins {loci cit.) saw seventy-eight birds in two and one-half hours. Eleven were seen by both observers, thirty-three by Stebbins only, and thirty- four by Carpenter only. On October 10, 1905, at the same place, in two hours, fifty-seven birds were counted, eleven being visible through both telescopes. Of the remainder, Stebbins saw seventeen and Carpenter, twenty-nine. On September 12, 1945, at Baton Rouge, Louisiana, in an interval of one hour and forty minutes, two independent observers each counted six birds. Again, on October 17, 1945, two observers each saw eleven birds in twenty-two minutes. On April 10, 1946, in one hour and five minutes, twenty-four birds were seen through one scope and twenty-six through the other. Like- wise on May 12, 1946, in a single hour, seventy-three birds were counted by each of two observers. The Baton Rouge observations were made with telescopes six to twelve feet apart. These results show a remarkable conformity, though the exceptional October ob- servation of Carpenter and Stebbins indicates the desirability of con- tinuing these studies, particularly in the fall.
On the whole, the available evidence points to the conclusion that night migration differs materially from the kind of daytime migra- tion with which we are generally familiar. Birds are apparently evenly spread throughout the sky, with little tendency to fly in flocks. It must be remembered, however, that only in the case of night migration have objective and truly quantitative studies been made of horizontal distribution. There is a possibility that our im- pressions of diurnal migration are unduly influenced by the fact that the species accustomed to flying in flocks are the ones that at- tract the most attention.
These conclusions relate to the uniformity of migration in terms of short distances only, in the immediate vicinity of an observation station. The extent to which they may be applied to broader fronts is a question that may be more appropriately considered later, in connection with continental aspects of the problem.
B. Density as Function of the Hour of the Night
There are few aspects of nocturnal migration about which there is less understanding than the matter of when the night flight begins, at what rate it progresses, and for what duration it continues. One would think, however, that this aspect of the problem, above most others, would have been thoroughly explored by some means of objective study. Yet, this is not the case. Indeed, I find not a
414 University of Kansas Publs., Mus. Nat. Hist.
single paper in the American literature wherein the subject is dis- cussed, although some attention has been given the matter by- European ornithologists. Siivonen (1936) recorded in Finland the frequency of call notes of night migrating species of Turdus and from these data plotted a time curve showing a peak near midnight. Bergman (1941) and Putkonen (1942), also in Finland, studied the night flights of certain ducks {Clangula hy emails and Oidemia jusca and 0. nigra) and a goose (Branta bernicla) and likewise demon- strated a peak near midnight. However, these studies were made at northern latitudes and in seasons characterized by evenings of long twilight, with complete darkness limited to a period of short duration around midnight. Van Oordt (1943: 34) states that in many cases migration lasts all night; yet, according to him, most European investigators are of the opinion that, in general, only a part of the night is used, that is, the evening and early morning hours. The consensus of American ornithologists seems to be that migratory birds begin their flights in twilight or soon thereafter and that they remain on the wing until dawn. Where this idea has been challenged at all, the implication seems to have been that the flights are sustained even longer, often being a continuation far into the night of movements begun in the daytime. The telescopic method fails to support either of these latter concepts.
The Time Pattern
When the nightly curves of density at the various stations are plotted as a function of time, a salient fact emerges — that the flow
to
II 12 I 2
HOUR OF THE NIGHT
Fig. 26. Average hourly station densities in spring of 1948. This curve represents the arithmetic mean obtained by adding all the station densities ■ for each hour, regardless of date, and dividing the sum by the number of sets of observations at that hour (CST).
Lowery: Noctuknal Migration of Birds
415
of birds is in no instance sustained throughout the night. The majority of the curves rise smoothly from near zero at the time of twilight to a single peak and then decline more or less symmetrically to near the base line before dawn. The high point is reached in or around the eleven to twelve o'clock interval more often than at any other time.
Figure 26, representing the average hourly densities for all stations on all nights of observation, demonstrates the over-all effect of these tendencies. Here the highest density is reached in the hour before midnight with indications of flights of great magnitude also in the hour preceding and the hour following the peak interval. The curve ascends somewhat more rapidly than it declines, which fact may or may not be significant. Since there is a great dispro- portion in the total volume of migration at different localities, the thought might be entertained that a few high magnitude stations, such as Tampico and Progreso, have imposed their own character- istics on the final graph. Fortunately, this idea may be tested by subjecting the data to a second treatment. If hourly densities are expressed as a percentage of the nightly peak, each set of observa- tions, regardless of the number of birds involved, carries an equal weight in determining the character of the over-all curve. Figure 27 shows that percentage analysis produces a curve almost identical with the preceding one. To be sure, all of the individual curves do not conform with the composite, either in shape or incidence of
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Fig. 27. Hourly station densities plotted as a percentage of peak. The curve is based only on those sets of data where observations were continued long enough to include the nightly peak. In each set of data the station density for each hour has been expressed as a percentage of the peak for the night at the station in question. All percentages for the same hour ori all dates have been averaged to obtain the percentile value of the combined station density at each hour (CST). *
416 University of Kansas Publs., Mus. Nat. Hist.
peak. The extent of this departure in the latter respect is evident from Figure 28, showing the number of individual nightly station curves reaching a maximum peak in each hour interval. Even this graph demonstrates that maximum densities near midnight represent the typical condition.
O 12
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Z 4
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HOUR OF THE KiSHT
Fig. 28. Incidence of maximum peak at the various hours of the night in 1948. "Number of stations" represents the total foi- all nights of the numbers of station peaks falling within a given hour.
The remarkable smoothness and consistency of the curves shown in Figures 26 and 27 seem to lead directly to the conclusion that the volume of night migration varies as a function of time. Ad- mittedly other factors are potentially capable of influencing the number of birds passing a given station in a given hour. Among these are weather conditions, ecological patterns, and specific topo- graphical features that might conceivably serve as preferred ave- nues of flight. However, if any of these considerations were alone responsible for changes in the numbers of birds seen in successive intervals, the distribution of the peak in time could be expected to be haphazard. For example, there is no reason to suppose that the cone of observation would come to lie over favored terrain at pre- cisely the hour between eleven and twelve o'clock at so many widely separated stations. Neither could the topographical hypothesis ex- plain the consistently ascending and descending pattern of the ordi- nates in Figure 28. This is not to say that other factors are without effect; they no doubt explain the divergencies in the time pattern exhibited by Figure 28. Nevertheless, the underlying circumstances are such that when many sets of data are merged these other in- fluences are subordinated to the rise and fall of an evident time pattern.
Lowery: Noctubnal Migration of Birds 417
Stated in concrete terms, the time frequencies shown in the graphs suggest the following conclusions: first, nocturnal migrations are not a continuation of daytime flights; second, nearly all night mi- grants come to earth well before dawn; and, third, in each hour of the night up until eleven or twelve o'clock there is typically a pro- gressive increase in the number of birds that have taken wing and in each of the hours thereafter there is a gradual decrease. Taken at its face value, the evidence seems to indicate that birds do not begin their night migrations en masse and remain on the wing until dawn and that in all probability most of them utilize less than half of the night.
Interestingly enough, the fact that the plot points in Figure 26 lie nearly in line tempts one to a further conclusion. The curve be- haves as an arithmetic progression, indicating that approximately the same number of birds are leaving the ground in each hour inter- val up to a point and that afterwards approximately the same num- ber are descending within each hour. However, some of the com- ponents making up this curve, as later shown, are so aberrant in this regard that serious doubt is cast on the validity of this gen- eralization.
Because the results of these time studies are unexpected and star- tling, I have sought to explore other alternative explanations and none appears to be tenable. For example, the notion that the vary- ing flight speeds of birds might operate in some way to produce a cumulative effect as the night progresses must be rejected on close analysis. If birds of varying flight speeds are continuously and evenly distributed in space, a continuous and even flow would result all along their line of flight. If they are haphazardly distributed in space, a correspondingly haphazard density pattern would be ex- pected.
Another explanation might be sought in the purely mathematical effects of the method itself. The computational procedure assumes that the effective area of the sample is extremely large when the moon is low, a condition that usually obtains in the early hours of the evening in the days surrounding the full moon. Actually no tests have yet been conducted to ascertain how far away a silhouette of a small bird can be seen as it passes before the moon. Conse- quently, it is possible that some birds are missed under these condi- tions and that the effective field of visibility is considerably smaller than the computed field of visibility. The tendency, therefore, may be to minimize the densities in such situations more than is justified.
418
University of Kansas Publs., Mus. Nat. Hist.
However, in many, if not most, cases, the plotting of the actual number of birds seen, devoid of any mathematical procedures, re- sults in an ascending and descending curve.
A third hypothesis proposes that all birds take wing at nearly the same time, gradually increase altitude until they reach the mid-point of their night's journey, and then begin a similarly slow descent. Since the field of observation of the telescope is conical, it is assumed that the higher the birds arise into the sky the more they increase their chances of being seen. According to this view, the changes
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Fig. 29. Various types of density-time curves. (A) Near typical, Ottumwa, April 22-23; (B) random fluctuation, Stillwater, April 23-24; (C) bimodal, Knoxville, April 22-23; (D) sustained peak, Ottumwa, April 21-22; (E) early peak. Oak Grove, May 21-22; (F) late peak, Memphis, April 23-24.
Lowery: Nocturnal Migration of Birds 419
in the density curve represent changes in the opportunity to see birds rather than an increase or decrease in the actual number of migrants in the air. Although measurements of flight altitude at various hours of the night have not been made in sufficient number to subject this idea to direct test, it is hardly worthy of serious consideration. The fallacy in the hypothesis is that the cone of observation itself would be rising with the rising birds so that actually the greatest proportion of birds flying would still be seen when the field of observation is in the supine position of early evening.
It cannot be too strongly emphasized that the over-all time curves just discussed have been derived from a series of individual curves, some of which differ radically from the composite pattern. In Figure 29, six dissimilar types are shown. This variation is not surprising in view of the fact that many other causative factors aside from time operate on the flow of birds from hour to liour. Figure 29A illustrates how closely some individual patterns con- form with the average. Figure 29B is an example of a random type of fluctuation with no pronounced time character. It is an effect rarely observed, occurring only in the cases where the number of birds observed is so small that pure chance has a pronounced effect on the computed densities; its vacillations are explicable on that account alone. Errors of sampling may similarly accoimt for some, though not all, of the curves of the bimodal type shown in Figure 29C. Some variation in the curves might be ascribed to the vari- ations in kinds of species comprising the individual flights at different times at different places, provided that it could be demonstrated that different species of birds show dissimilar temporal patterns. The other atypical patterns are not so easily dismissed and will be the subject of inquiry in the discussions that follow. It is significant that in spite of the variety of the curves depicted, which represent every condition encountered, in not a single instance is the density sustained at a high level throughout the night. Morover, these dissident patterns merge into a remarkably harmonious, almost normal, average curve.
When, at some future date, suitable data are available, it would be highly desirable to study the average monthly time patterns to ascertain to what extent they may deviate from the over-all average. At present this is not justifiable because there are not yet enough sets of data in any two months representing the same selection of stations.
420 University of Kansas Publs., Mus. Nat. Hist.
Correlations with Other Data
It is especially interesting to note that the data pertaining to this problem derived from other methods of inquiry fit the conclusions adduced by the telescopic method. Overing (1938), who for several years kept records of birds striking the Washington Monument, stated that the record number of 576 individuals killed on the night of September 12, 1937, all came down between 10:30 p. m. and mid- night. His report of the mortality on other nights fails to mention the time factor, but I am recently informed by Frederick C. Lin- coln {in litt.) that it is typical for birds to strike the monument in greatest numbers between ten and twelve o'clock at night. At the latter time the lights illuminating the shaft are extinguished, thus resulting in few or no casualties after midnight. The recent report by Spofford (1949) of over 300 birds killed or incapacitated at the Nashville airport on the night of September 9-10, 1948, after flying into the light beam from a ceilometer, is of interest in this con- nection even though the cause of the fatality is shrouded in mystery. It may be noted, however, that "most of the birds fell in the first hour," which, according to the account, was between 12:30 a. m. and 1 :30 A. M. Furthermore, birds killed at the Empire State Building in New York on the night of September 10-11, 1948, began to strike the tower "shortly after midnight" (Pough, 1948). Also it will be recalled that the observations of Stone {loc. cit.), already referred to in this paper (page 410) , show a situation where the flight in the early part of the night was negligible but mounted to a peak be- tween ten and eleven o'clock, with continuing activity at least until midnight.
All of these observations are of significance in connection with the conclusions herein advanced, but by far the most striking cor- relation between these present results and other evidences is found in the highly important work of various European investigators studying the activity of caged migratory birds. This work was recently reviewed and extended by Palmgren (1944) in the most comprehensive treatise on the subject yet published. Palmgren re- corded, by an electrically operated apparatus, the seasonal, daily, and hourly activity patterns in caged examples of two typical European migrants, Turdus ericetorum philomelas Brehm and Eri- thacus rubecula (Linnaeus). Four rather distinct seasonal phases in activity of the birds were discerned: winter non-migratory, spring migratory, summer non-migratory, and autumn migratory. The first of these is distinguished by morning and evening maxima
Lowery: Nocturnal Migration of Birds 421
of activity, the latter being better developed but the former being more prolonged. Toward the beginning of migration, these two pe- riods of activity decline somewhat. The second, or spring migratory phase, which is of special interest in connection with the present problem, is characterized by what Palmgren describes as nightly migratory restlessness {Zugunruhe) . The morning maximum, when present, is weaker and the evening maximum often disappears al- together. Although variations are described, the migratory restless- ness begins ordinarily after a period of sleep ("sleeping pause") in the evening and reaches a maximum and declines before midnight. This pattern agrees closely with the rhythm of activity indicated by the time curves emerging from the present research. Combining the two studies, we may postulate that most migrants go to sleep for a period following twilight, thereby accounting for the low densities in the early part of the night. On awakening later, they begin to exhibit migratory restlessness. The first hour finds a certain num- ber of birds sufficiently stimulated so that they rise forthwith into the air. In the next hour still others respond to this urge and they too mount into the air. This continues until the ''restlessness" be- gins to abate, after which fewer and fewer birds take wing. By this time, the birds that began to fly early are commencing to descend, and since their place is not being filled by others leaving the ground, the density curve starts its decline. Farner (1947) has called atten- tion to the basic importance of the work by Palmgren and the many experimental problems it suggests. Of particular interest would be studies comparing the activity of caged American migrant species and the nightly variations in the flight rates.
The Baton Rouge Drop-off
As already stated, the present study was initiated at Baton Rouge, Louisiana, in 1945, and from the outset a very peculiar density time pattern was manifest. I soon found that birds virtually disappeared from the sky after midnight. Within an hour after the termination of twilight, the density would start to ascend toward a peak which was usually reached before ten o'clock, and then would begin, surpris- ingly enough, a rapid decline, reaching a point where the migratory flow was negligible. In Figure 30 the density curves are shown for five nights that demonstrate this characteristically early decline in the volume of migration at this station. Since, in the early stages of the work, coordinates of apparent pathways of all the birds seen were not recorded, I am unable now to ascertain the direction of flight and thereby arrive at a density figure based on the dimension
422 University of Kansas Publs., Mxjs. Nat. Hist.
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5
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Fig. 30. Density-time curves on various nights at Baton Rouge. (A) April 25, 1945; (B) April 15, 1946; (C) May 10, 1946; (D) May 15, 1946; (E) April 22-23, 1948. These curves are plotted on a "plus or minus" basis as described in the text, with the bottom of the curve representing the minimum density and the top of the curve the maximum.
Lowery: Nocturnal Migration of Birds 423
of the cone and the length of the front presented to birds flying in certain directions. It is feasible, nevertheless, to compute what I have termed a "plus or minus" flight density figure stating the rate of passage of birds in terms of the maximum and minimum correc- tions which all possible directions of flight would impose. In other words, density is here computed, first, as if all the birds were flying perpendicular to the long axis of the ellipse, and, secondly, as if all the birds were flying across the short axis of the ellipse. Since the actual directions of flight were somewhere between these two ex- tremes, the "plus or minus" density figure is highly useful.
The well-marked decline before midnight in the migration rates at Baton Rouge may be regarded as one of the outstanding results emerging from this study. Many years of ornithological investiga- tion in this general region failed to suggest even remotely that a situation of this sort obtained. Now, in the light of this new fact, it is possible for the first time to rationalize certain previously in- congruous data. Ornithologists in this area long have noted that local storms and cold-front phenomena at night in spring sometimes precipitate great numbers of birds, whereupon the woods are filled the following day with migrants. On other occasions, sudden storms at night have produced no visible results in terms of bird densities the following day. For every situation such as described by Gates (1933) in which hordes of birds were forced down at night by in- clement weather, there are just as many instances, even at the height of spring migration, when similar weather conditions yielded no birds on the ground. However, the explanation of these facts is simple; for we discover that storms that produced birds occurred before midnight and those that failed to produce birds occurred after that time (the storm described by Gates occurred between 8:30 and 9:00 P.M.).
The early hour decline in density at Baton Rouge at first did not seem surprising in view of the small amount of land area between this station and the Gulf of Mexico. Since the majority of the birds destined to pass Baton Rouge on a certain night come in general from the area to the south of that place, and since the distances to various points on the coast are slight, we inferred that a three-hour flight from even the more remote points would probably take the bulk of the birds northward past Baton Rouge. In short, the coastal plain would be emptied well before midnight of its migrant bird life, or at least that part of the population des- tined to migrate on any particular night in question. Although data
424 University of Kansas Publs., Mus. Nat. Hist.
in quantity are not available from, stations on the coastal plain other than Baton Rouge, it may be pointed out that such observations as we do have, from Lafayette and New Orleans, Louisiana, and from Thomasville, Georgia, are in agreement with this hypothesis.
A hundred and seventy miles northward in the Mississippi Valley, at Oak Grove, Louisiana, a somewhat more normal density pattern is manifested. There, in four nights of careful observation, a pro- nounced early peak resulted on the night of May 21-22 (Figure 29E), but on the other three nights significant densities held up until near twelve o'clock, thereby demonstrating the probable effect of the increased amount of land to the south of the station.
Subsequent studies, revealing the evident existence of an under- lying density time pattern, cast serious doubt on the explanations just advanced of the early decline in the volume of migration at Baton Rouge. It has as yet been impossible to reconcile the early drop-off at this station with the idea that birds are still mounting into the air at eleven o'clock, as is implied by the ideal time curves.
C. Migration in Relation to Topography
To this point we have considered the horizontal distribution of birds in the sky only on a very narrow scale and mainly in terms of the chance element in observations. Various considerations have supported the premise that the spread of nocturnal migration is rather even, at least within restricted spacial limits and short in- tervals of time. This means that in general the flow of birds from hour to hour at a single station exhibits a smooth continuity. It does not mean that it is a uniform flow in the sense that approxi- mately the same numbers of birds are passing at all hours, or at all localities, or even on all one-mile fronts in the same locality. On the contrary, there is evidence of a pronounced but orderly change through the night in the intensity of the flight, corresponding to a basic and definitely timed cycle of activity. Other influences may interfere with the direct expression of this temporal rhythm as it is exhibited by observations at a particular geographical location. Among these, as we have just seen, is the disposition of the areas that offer suitable resting places for transient birds and hence con- tribute directly and immediately to the flight overhead. A second possible geographical effect is linked with the question of the tend- ency of night migrants to follow topographical features.
General Aspects of the Topographical Problem
That many diurnal migrants tend to fly along shorelines, rivers,
Lowery: Nocturnal Migration of Birds 425
and mountain ridges is well known, but this fact provides no assur- ance that night migrants do the same thing. Many of the obvious advantages of specialized routes in daylight, such as feeding oppor- tunities, the lift provided by thermal updrafts, and the possible aid of certain landmarks in navigation, assume less importance after night falls. Therefore, it would not be safe to conclude that all nocturnal migrants operate as do some diurnal migrants. For in- stance, the passage of great numbers of certain species of birds along the Texas coast in daylight hours cannot be regarded as certain proof that the larger part of the nocturnal flight uses the same route. Neither can we assume that birds follow the Mississippi River at night simply because we frequently find migrants concentrated along its course in the day. Fortunately we shall not need to speculate indefinitely on this problem ; for the telescopic method offers a means of study based on what night migrants are doing at night. Two lines of attack may be pursued. First we may compare flight densities obtained when the field of the telescope lies over some out- standing topographical feature, such a a river, with the recorded volume of flight when the cone of observation is directed away from that feature. Secondly, we may inquire how the major flight direc- tions at a certain station are oriented with respect to the terrain. If the flight is concentrated along a river, for instance, the flight density curve should climb upward as the cone of observation swings over the river, reganUess of the hour at which it does so. The effect should be most pronounced if the observer were situated on the river bank, so that the cone would eventually come to a position directly along the watercourse. Though in that event birds coming up the river route would be flying across the short axis of an elliptical sec- tion of the cone, the fact that the whole field of observation would be in their path should insure their being seen in maximum proportions. If, on the other hand, the telescope were set up some distance away from the river so that the cone merely moved across its course, only a section of the observation field would be interposed on the main flight lane.
The interaction of these possibilities with the activity rhythm should have a variety of effects on the flight density curves. If the cone comes to lie over the favored topographical feature in the hour of greatest migrational activity, the results would be a simple sharp peak of doubtful meaning. However, since the moon rises at a dif- ferent time each evening, the cone likewise would reach the im-
5—1020
426 University of Kansas Publs., Mi:s. Nat. Hist.
mediate vicinity of the terrain feature at a different time each night. As a result, the terrain peak would move away from its position of coincidence with the time peak on successive dates, producing first, perhaps, a sustention of peak and later a definitely bimodal curve. Since other hypotheses explain double peaks equally well, their mere existence does not necessarily imply that migrants actually do travel along narrow topographical lanes. Real proof requires that we demonstrate a moving peak, based on properly corrected density computations, corresponding always with the position of the cone over the most favored terrain, and that the flight vectors be consistent with the picture thus engendered.
The Work of Winkenwerder
To date, none of the evidence in favor of the topographical hy- pothesis completely fills these requirements. Winkenwerder (lac. cit.) , in analyzing the results of telescopic counts of birds at Madison and Beloit, Wisconsin, Detroit and Ann Arbor, Michigan, and at Lake Forest, Illinois, between 1898 and 1900, plotted the number of birds seen at fifteen-minute intervals as a function of the time of the night. He believed that the high points in the resulting frequency histo- grams represented intervals when the field of the telescope was moving over certain topographically determined flight lanes, though he did not specify in all cases just what he assumed the critical physiographic features to be. Especially convincing to him were results obtained at Beloit, where the telescope was situated on the east bank of the Roek River, on the south side of the city. Immedi- ately below Beloit the river turns southwestward and continues in this direction about five miles before turning again to flow in a southeastward course for approximately another five miles. In this setting, on two consecutive nights of observation in May, the number of birds observed increased tremendously in the 2 to 3 a. m. interval, when, according to Winkenwerder's interpretation of the data (he did not make the original observations at Beloit himself) , the tele- scope was pointing directly down the course of the river. This con- clusion is weakened, however, by notable inconsistencies. Since the moon rises later each evening, it could not have reached the same position over the Rock River at the same time on both May 12-13 and May 13-14, and therefore, if the peaks in the graph were really due to a greater volume of migration along the watercourse, they should not have so nearly coincided. As a matter of fact the incidence of the peak on May 12-13 should have preceded that of
Lowery: Nocturnal Migration of Birds 427
the peak on May 13-14; whereas his figure shows the reverse to have been true. Singularly enough, Winkenwerder recognized this difficulty in his treatment of the data from Madison, Wisconsin. Unable to correlate the peak period with the Madison terrain by the approach used for Beloit, he plotted the observations in terms of hours after moonrise instead of standard time. This procedure was entirely correct; the moon does reach approximately the same posi- tion at each hour after its rise on successive nights. The surprising thing is that Winkenwerder did not seem to realize the incompat- ibility of his two approaches or to realize that he was simply choosing the method to suit the desired results.
Furthermore, as shown in Part I of this paper, the number of birds seen through the telescope often has only an indirect connection with the actual number of birds passing over. My computations reveal that the highest counts of birds at Beloit on May 12-13 were re- corded when the moon was at an altitude of only 8° to 15° and, that when appropriate allowance is made for the immense size of the field of observation at this time, the partially corrected flight density for the period is not materially greater than at some other intervals in the night when the telescope was not directed over the course of the Rock River. These allowances do not take the direction factor into consideration. Had the birds been flying at right angles to the short axis of an elliptical section of the cone throughout the night, the flight density in the period Winkenwerder considered the peak would have been about twice as high as in any previous interval. On the other hand, if they had been flying across the long axis at all times, the supposed peak would be decidedly inferior to the flight density at 10 to 11:00 p. m., before the cone came near the river.
Admittedly, these considerations contain a tremendous element of uncertainty. They are of value only because they expose the equal uncertainty in Winkenwerder's basic evidence. Since the coordi- nates of the birds' apparent pathways at Beloit were given, I at first entertained the hope of computing the flight densities rigorously, by the method herein employed. Unfortunately, Winkenwerder was apparently dealing with telescopes that gave inverted images, and he used a system for recording coordinates so ambiguously de- scribed that I am not certain I have deciphered its true meaning. When, however, his birds are plotted according to the instructions as he stated them, the prevailing direction of flight indicated by the projection formula falls close to west-northwest, not along the course of the Rock River, but at direct right angles to it.
428
University of Kansas Publs., Mus. Nat. Hist.
Interpretation of Recent Data
I am in a position to establish more exact correlations between flight density and terrain features in the case of current sets of observations. Some of these data seem at first glance to fit the idea of narrow topographically-oriented flight lanes rather nicely. At Tampico, where six excellent sets of observations were made in March and April, 1948, the telescope was set up on the beach within a few yards of the Gulf of Mexico. As can be seen from Figure 25 (ante), the slant of the coastline at this point is definitely west of north, as is also the general trend of the entire coast from southern Veracruz to southern Tamaulipas (see Figure 34, beyond). The
n
N10*W
NirW
Nt2*W
22-23 APRIL
23-24 APRIL
Fig. 31. Directional components in the flght at Tampico on three nights in 1948. The lengths of the sector vectors are determined by their respective densities expressed as a percentage of the station density for that night; the vector resultants are plotted from them by standard piocedure. Thus, the nightly diagrams are not on the same scale with respect to the actual number of birds involved.
Lowery: Noctuenal Migration of Birds
429
over-all vector resultant of all bird flights at this station was N 11° W, and, as will be seen from Figure 31, none of the nightly- vector resultants in April deviates more than one degree from this average. Thus the prevailing direction of flight, as computed, agrees with the trend of the coast at the precise point of the observations, at least to the extent that both are west of north. To be sure, the individual sector vectors indicate that not all birds were following this course; indeed, some appear to have been flying east of north, heading for a landfall in the region of Brownsville, Texas, and a very few to have been traveling northeastward toward the central Gulf coast. But it must be remembered that a certain amount of computational deviation and of localized zigzagging in flight must be anticipated. Perhaps none of these eastward vectors represents an actual extended flight path. The nightly vector resultants, on the other hand, are so consistent that they have the appearance of re- markable accuracy and tempt one to draw close correlations with the terrain. When this is done, it is found that, while the prevailing flight direction is 11° west of north, the exact slant of the coastline at the location of the station is about 30° west of north, a difference of around 19°. It appears, therefore, that the birds were not following the shoreline precisely but cutting a chord about ten miles long across an indentation of the coast. If it be argued that the method of calculation is not accurate enough to make a 19° difference signifl- cant, and that most of the birds might have been traveling along the beach after all, it can be pointed out with equal justification that, if this be so, the 11° divergence from north does not mean anything either and that perhaps the majority of the birds were
10
II 12 I
HOUR OF THE NIGHT
Fig. 32. Hourly station density curve at Tampico, Tamaulipas, on the night of April 21-22, 1948 (CST).
430 University of Kansas Publs., Mus. Nat. Hist.
going due north. We are obliged to conclude either that the main avenue of flight paralleled the disposition of the major topographical features only in a general way or that the angle between the line of the coast and true north is not great enough to warrant any inference at all.
Consideration of the Tampico density curves leads to similarly ambiguous results. On the night of April 21-22, as is evident from a comparison of Figures 25 and 32, the highest flight density oc- curred when the projection of the cone on the terrain was wholly included within the beach. This is very nearly the case on the night of April 23-24 also, the positions of the cone during the peak period of density being only about 16" apart. (On the intervening date, clouds prevented continuous observation during the critical part of the night.) These correlations would seem to be good evidence that most of these night migrants were following the coastline of the Gulf of Mexico. However, the problem is much more complicated. The estimated point of maximum flight density fell at 10:45 p. m. on April 21-22 at 11:00 p. m., on April 23-24, both less than an hour from the peak in the ideal time curve (Figure 26, ante). We cannot be sure, therefore, that the increase in density coinciding with the position of the moon over the beach is not an increase which would have occurred anyway. Observations conducted several nights before or after the second quarter, when the moon is not on or near its zenith at the time of the predictable peak in the density curve, would be of considerable value in the study of this particular problem.
The situation at Tampico has been dealt with at length because, among all the locations for which data are available, it is the one that most strongly supports the topographical hypothesis. In none of the other cases have I been able to find a definite relation between the direction of migration and the features of the terrain. Studies of data from some of these stations disclose directional patterns that vary from night to night only slightly more than does the flight at Tampico. In three nights of observation at Lawrence, Kansas, marked by very high densities, the directional trend was north by north-northeast with a variation of less than 8°, yet Lawrence is so situated that there seems to be no feature of the landscape locally or in the whole of eastern Kansas or of western Missouri that co- incides with this heading. At Mansfield, Louisiana, in twelve nights of observation, the strong east by northeast trend varied less than 15°, but again there appears to be no correlation over a wide area
Lower y: Nocturnal Migration of Birds
431
between this direction and any landmarks. And, at Progreso, Yuca- tan, where the vector resultants were 21° and 27° on successive nights, most of the birds seen had left the land and were beginning their flight northward over the trackless waters of the Gulf of Mex- ico. Furthermore, as I have elsewhere pointed out (1946: 205), the whole northern part of the Yucatan Peninsula itself is a flat terrain, unmarked by rivers, mountains, or any other strong physiographic features that conceivably might be followed by birds.
tOUISVlLU
KNOXVIUX
OTTUMWA
Fig. 33. The nightly net trend of migrations at three stations in 1948. Each arrow is the vector resultant for a particular night, its length expressing the nightly density as a percentage of the total station density for the nights repre- sented. Thus, the various station diagrams are not to the same scale.
In Figure 33 I have shown the directional patterns at certain sta- tions where, unlike the cases noted above, there is considerable change on successive nights. Each vector shown is the vector result- ant for one particular night. The lengths of the vectors have been determined by their respective percentages of the total computed density, or total station magnitude, for all the nights in question. In other words, the lengths of the individual vectors denote the per- centile role that each night played in the total density. From the directional spread at these stations it becomes apparent that if most of the birds were traveling along certain topographic feature on one
432 University of Kansas Publs., Mus. Nat. Hist.
night, they could not have been traveling along the same feature on other nights.
The possibility should be borne in mind, however, that there may be more than one potential topographic feature for birds to follow at some stations. Moreover, it is conceivable that certain species might follow one feature that would lead them in the direction of their ultimate goal, whereas other species, wishing to go in an en- tirely different direction, might follow another feature that would lead them toward their respective destination. It would seem un- likely, however, that the species composition of the nocturnal flights would change materially from night to night, although there is a strong likelihood that it might do so from week to week and cer- tainly from month to month.
By amassing such data as records of flight direction along the same coast from points where the local slant of the shoreline is materially different, and comparisons of the volume of migration at night along specialized routes favored during the day with the flight densities at progressive distances from the critical terrain feature involved, we shall eventually be able to decide definitely the role topography plays in bird migration. We cannot say on the basis of the present ambiguous evidence that it is not a factor in determining which way birds fly, but, if I had to hazard a guess one way or the other, I would be inclined to discount the likelihood of its proving a major factor.
D. Geographical Factors and the Continental Density
Pattern
A study of the total nightly or seasonal densities at the various stations brings forth some extremely interesting factors, many of which, however, cannot be fully interpreted at this time. A com- plete picture of the magnitude of migration at a given station can- not be obtained from the number of birds that pass the station on only a few nights in one spring. Many years of study may be re- quired before hard and fast principles are justifiable. Nevertheless, certain salient features stand out in the continental density pattern in the spring of 1948. (The general results are summarized in Tables 2-5; the location of the stations is shown in Figure 34.) These features will be discussed now on a geographical basis.
Lowery: Noctuenal Migration of Birds
433
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434 University of Kansas Publs., Mus. Nat. Hist. Table 3. — Average Hourly Station Densities in 1948
|
Observation Station |
March |
April |
May |
Season |
|
Canada Pt. Pelee |
400 |
400 |
||
|
Mexico S. L. P. : Ebano |
400 700 |
400 |
||
|
Tamps. : Tampico |
6,300 2,800 0 + 300 400 500 1,700 1,400 2,000 2,000 700 600 700 800 1,400 400 |
3,500 |
||
|
Yuc: Progreso |
2,800 |
|||
|
United States Fla. : Pensacola |
200 200 |
100 |
||
|
Winter Park |
300 |
|||
|
Ga. : Athens |
400 |
|||
|
Thomasville |
100 3,800 |
300 |
||
|
Iowa: Ottumwa |
3 100 |
|||
|
Kans. : Lawrence |
4,000 |
3,400 |
||
|
Ky. : Louisville |
700 |
1 500 |
||
|
Murray |
2 000 |
|||
|
La. : Baton Rouge |
700 |
|||
|
Lafayette |
• |
600 |
||
|
Mansfield New Orleans |
0 60 |
800 |
600 300 |
|
|
Oak Grove |
800 |
1 100 |
||
|
Mich.: Albion |
400 |
|||
|
Minn. : Hopkins |
500 700 1,700 200 1,000 0 + 800 900 1,500 |
500 |
||
|
Miss. : Rosedale |
1,100 400 500 200 200 1,300 800 1,100 1,600 |
900 |
||
|
Mo.: Columbia |
900 |
|||
|
Liberty |
300 |
|||
|
Okla.: Stillwater S. Car. : Charleston Tenn. : Knoxville |
500 200 |
400 100 1,100 |
||
|
Memphis Tex.: College Station |
300 |
700 1 200 |
||
|
Rockport |
1 600 |
|||
Lowery: Nocturnal Migration of Birds
435
Table 4. — Maximum Hourly Station Densities in 1948
Observation Station
Canada Pt. Pelee .
Mexico S. L. P.: Ebano... Tamps. : Tampico . Yuc. : Progreso . . . .
United States
Fla.: Pensacola
Winter Park
Ga. : Athens
Thomasville
Iowa: Ottumwa
Kans. : Lawrence
Ky. : Louisville
Murray
La. : Baton Rouge . . . .
Lafayette
Mansfield
New Orleans
Oak Grove
Mich.: Albion
Minn.: Hopkins
Miss. : Rosedale
Mo.: Columbia
Liberty
Okla. : Stillwater
S. Car. : Charleston . . . Tenn. : Knoxville
Memphis
Tex. : College Station .
Rockport
March
GOO 3,100
14,500
200
900 400
1,200
April
21,200 11,900
100 2,300
900 1,500 3,800 2,200 5,000 3,700 3,400 1,800 2,100 1,100 2,700
700
2,200
800
800
700
600
5,800
3,400
3,400
2,400
May
1,400
700 1,000
200 ' 12,500
1,400
1,600
2,500
1,100 1,400 3,400
800 1,400
200 1,900 2,100 3,100
436
University of Kansas Publs., Mus. Nat. Hist.
Table 5. — Maximum Nightly Densities at Stations with More Than One
Night of Observation
Observation Station
Mexico Tamps.: Tampico. Yuc: Progreso....
United States Fla.: Winter Park ... . Ga.: Athens
Thomasville
Iowa: Ottumwa
Kans. : Lawrence
Ky.: Louisville
Murray
La. : Baton Rouge . . . .
Mansfield
Oak Grove
Miss.: Rosedale
Mo.: Columbia
Okla.: Stillwater
Tenn. : Knoxville
Memphis
Tex. : College Station .
March
5,500
51 ,600
2,700
3,600
April
63,600 31,600
6,200 2,600 3,900
15,300 5,400
17,000
16,400 6,200 4,900
13,600 6,800 1,400 1,900
15,200 7,900 6,200
May
54,600 "8,400'
5,200 5,800 5,800
10,300 3,000 9,000 7,000
13,200
Lowery: Nocturnal Migration of Birds
437
Fig. 34. Stations at which telescopic observations were made in 1948.
438 University of Kansas Publs., Mus. Nat. Hist.
Gulf Migration: A Review of the Problem
In view of the controversy in recent years pertaining to migration routes in the region of the Gulf of Mexico (Williams, 1945 and 1947 ; Lowery, 1945 and 1946), the bearing of the new data on the prob- lem is of especial interest. While recent investigations have lent further support to many of the ideas expressed in my previous papers on the subject, they have suggested alternative explanations in the case of others. In the three years that have elapsed since my last paper dealing with Gulf migration, some confusion seems to have arisen regarding the concepts therein set forth. Therefore, I shall briefly re-state them.
It was my opinion that evidence then available proved conclu- sively that birds traverse the Gulf frequently and intentionally; that the same evidence suggested trans-Gulf flights of sufficient mag- nitude to come within the meaning of migration ; that great numbers of birds move overland around the eastern and western edges of the Gulf; that it was too early to say whether the coastal or trans-Gulf route was the more important, but that enough birds cross the water from Yucatan to account for transient migration in the ex- treme lower Mississippi Valley; and, that, in fair weather, most trans-Gulf migrants continue on inland for some distance before coming to land, creating an area of "hiatus" that is usually devoid of transient species. I tried to make it emphatically clear that I realized that many birds come into Texas from Mexico overland, that I did not think the hordes of migrants normally seen on the Texas coast in spring were by any means all trans-Gulf migrants. I stated (1946: 206): "Proving that birds migrate in numbers across the Gulf does not prove that others do not make the journey by the coastal routes. But that is exactly the point. No one has ever pretended that it does." Although some ornithologists seem to have gained the impression that I endorse only the trans-Gulf route, this is far from the truth. I have long held that the mi- grations overland through eastern Mexico and southern Texas on one hand, and the over-water flights on the other, are each part of the broad movement of transients northward into the United States. There are three avenues of approach by which birds making up the tremendous concentrations on the Texas coast may have reached there: by a continental pathway from a wintering ground in eastern and southern Mexico; by the over- water route from Yucatan and points to the southward; and, finally, by an overland route from Central America via the western edge of the Gulf. As a result of Louisiana State University's four-year study of the avifauna in
Lowery: Nocturnal Migration of Birds
439
eastern Mexico, I know that migrants reach Texas from the first source. As a consequence of my studies in Yucatan of nocturnal flight densities and their directional trends, I strongly believe that migrants reach Texas from this second source. As for the third source, I have never expressed an opinion. I am not prepared to do so now, for the reason that today, as three years ago, there is no de- pendable evidence on which to base a judgment one way or another.
Western Gulf Area
Among the present flight density data bearing on the above issues, are the six sets of observations from the vicinity of Tampico, Tamau- lipas, already referred to. These were secured in the spring of 1948 by a telescope set up on the Gulf beach just north of the Miramar pavilion and only a hundred feet from the surf (see Figure 25, ante). The beach here is approximately 400 feet wide and is backed by scrub-covered dunes, which rapidly give way toward the west to a ratlier dense growth of low shrubs and trees. One might have ex- pected that station densities at Tampico in March would be rather high. Actually, though they are the second highest recorded for the month, they are not impressive and afford a striking contrast with the record flights there in April (Table 6). Unfortunately, only
Table 6. — Computed Hourly Densities at Tampico, Tamps., in Spring of 1948
|
Average hour of observation |
|||||||||
|
Date |
8:30 |
9:30 |
10:30 |
11:30 |
12:30 |
1:30 |
2:30 |
3:30 |
4:30 |
|
22-23 March 23-24 March |
600 0 300 1,100 700 600 |
700 400 700 7,000 2,900 4,700 |
1,000 1,200 800 14,900 7,500 19,100 |
800 3,100 1,600 12,900 |
100 800 1,100 8,100 |
100 |
0 |
100 |
|
|
24-25 March |
|||||||||
|
21-22 April 22-23 April 23-24 April |
3,800 |
3,500 |
200 |
||||||
|
21 ,200 |
5,500 |
5,900 |
4,000 |
2,000 |
200 |
a few stations were operating in March and thus adequate compari- sons are impossible; but the indications are that, in March, migra- tion activity on the western edges of the Gulf is slight. It fails even to approach the volume that may be observed elsewhere at the same time, as for example, in eastern Kansas where, however, the migration is not necessarily correlated with the migration in the
440 University of Kansas Publs., Mus. Nat. Hist.
lower Gulf area. Strangely enough, on the night of March 22-23, at Tampico, approximately 85 per cent of the birds were flying from north of an east-west line to south of it, opposite to the normal trend of spring migration. This phenomenon, inexplicable in the present instance, will be discussed below. On the other two nights in March, the directional trend at Tampico was northward with few or no aberrant components. Observations made approximately thirty-five miles inland from the Gulf, at Ebano, San Luis Potosi, on the night of March 25-26, show lower station densities than the poorest night at Tampico, but since they cover only a three-hour watch, they reveal little or nothing concerning the breadth of the so-called coastal flyway.
April flight densities at Tampico are the highest recorded in the course of this study. The maximum hourly density of 21,200 birds is 46 per cent higher than the maximum hourly density anywhere else. The average hourly density of 6,300 in April is more than twice as great as the next highest average for that month. These figures would seem to satisfy certain hypotheses regarding a coast- wise flight of birds around the western edge of the Gulf. Other aspects of the observations made at that time do not satisfy these hypotheses. Texas ornithologists have found that in periods of heavy spring migration, great numbers of birds are invariably pre- cipitated by rainy weather. On April 23, in the midst of the record- breaking telescopic studies at Tampico, Mr. Robert J. Newman made a daytime census immediately following four hours of rain. He made an intensive search of a small area of brush and low growth back of the beach for traces of North American migrants. In his best hour, only thirteen individual birds out of seventy-five seen were of species that do not breed there. The transient species were the Ruby-throated Hummingbird (1), Scissor- tailed Flycatcher (1), Western Wood Pewee (1), Black-throated Green Warbler (2) Orchard Oriole (7), and Baltimore Oriole (1), all of which winter extensively in southern Mexico. Perhaps, however, the apparent scarcity of transients on this occasion is not surprising in the light of the analysis of flight density in terms of bird density on the ground which I shall develop beyond. My only point here is to demonstrate that rain along the coast does not always produce birds.
As large as the nocturnal flights at Tampico have so far proved to be, they are not commensurate with the idea that nearly all birds follow a narrow coastwise route around the Gulf. To establish the
Lowery: Nocturnal Migration of Birds 441
latter idea, one must be prepared to show that the migrant species returning to the United States pass along two flyways a few miles wide in the immense volume necessary to account for their later abundance on a 1500-mile front extending across eastern North America. One might expect at least ten to twenty fold the number observable at any point in the interior of the United States. In ac- tuality, the highest nightly density of 63,600 birds at Tampico is barely sufficient to account for the highest nightly density of 54,600 at Ottumwa, Iowa, alone.
Of course, there is no way of knowing how closely a ratio of any- where from ten to one through twenty to one, employed in this com- parison, expresses the true situation. It may be too high. It could be too low, particularly considering that preliminary studies of flight density in Florida indicate that the western shores of the Gulf of Mexico must carry the major part of the traffic if migratory flights back to the United States in spring take place only along coastwise routes. Consideration of the data obtained in Florida in 1948 will serve to emphasize the point.
Eastern Gulf Area
At Winter Park, Florida, seventy-seven hours were spent at the telescope in April and May. This was 71 per cent more hours of actual observation than at the next highest station. Nevertheless, the total seasonal density amounted to only 21,700 birds. The aver- age hourly density was only 300 birds, with the maximum for any one hour being 2,300 birds. In contrast, forty-five hours of observa- tion at Tampico, Tamaulipas, in March and April, yielded a total station density of 140,300 birds. At the latter place, on the night of April 23-24, almost as many birds passed in a single hour as passed Winter Park in all of its seventy-seven hours of observation.
Should future telescopic studies at Florida stations fail to produce densities appreciably higher than did Winter Park in 1948, the cur- rently-held ideas that the Florida Peninsula is a major fly way will be seriously shaken. But one consideration must be kept in mind regarding the present picture. No observations were made at Winter Park in March, when it is conceivable that densities may have been materially higher. We know, for instance, that many of the early migrants to the southern United States are species whose winter homes are in the West Indies. Numbers of Vireonidae and Parulidae (notably the genera Vireo, Parula, Protonotaria, Mnio- tilta, Seiurus, Geothlypis, Setophaga, and certain Dendroica and Vermivora) winter extensively in this region and are among the first
6—1020
442 University of Kansas Publs., Mrs. Nat. Hist.
birds to return to the southern states in the spring. Many of them often reach Louisiana and other states on the Gulf coastal plain by mid-March. In the same connection, it may be mentioned that many of the outstanding instances of birds striking lighthouses in southern Florida occurred in March and early April (Howell, 1932).
Yucatan Area
I have long felt that the answers to many of the questions which beset us in our study of Gulf migration are to be found on the open waters of the Gulf of Mexico itself or on the northern tip of the Yucatan Peninsula. Accordingly, in the spring of 1945 I crossed the Gulf by slow freighter for the purpose of determining how many and what kinds of birds might be seen between the mouth of the Mississippi River and the Yucatan Peninsula in fair weather, when it could not be argued that the birds had been blown there by in- clement weather. To my own observations I was able to add those of other ornithologists who likewise had been aboard ship in the Gulf.
The summary of results proved that birds of many species cross the Gulf and do so frequently. It failed to demonstrate beyond all doubt that they do so in large numbers. Nor had I expected it to do so. The concensus of Gulf coast ornithologists seemed to be that transient migration in their respective regions is often per- formed at too high an elevation to be detected unless the birds are forced to earth by bad weather. I saw no reason to anticipate that the results would be otherwise over the waters of the Gulf of Mexico.
The application of the telescopic method held promise of supply- ing definite data on the numbers of trans-Gulf migrants, however high their flight levels. The roll and vibration of the ship had pre- vented me in 1945 from making telescopic observations at sea. Since no immediate solution to the technical difficulties involved presented itself, I undertook to reach one of the small cays in Alacran Reef, lying seventy-five miles north of Yucatan and in line with the coast of southern Louisiana. Because of transportation difficulties, my plans to place a telescopic station in this strategic location failed. Consequently, I returned in 1948 by freighter to Progreso, Yucatan, where telescopic counts were made for three nights, one of which was rendered almost valueless by the cloud cover.
The observation station at Progreso was situated on the northern end of the new wharf which projects northward from the beach to a point one mile over the Gulf. As will be seen from Figure 35, the entire cone of observation lay at nearly all times over the interven-
Lowery: Nocturnal Migration of Birds
443
PREVAILING
DIRECTION OF
FLIGHT
, Of ^^^''°
WHARF
3:53
ij-f^ 5Vc--^ Tfe-^'-^ ?,„__
■ ^ ■' ■/■!% „, <-•> 'i ^ f' I
c
^.
(c.
Z^-,
> 'i Z' ( '■
'•'^
/•! ^^ ^ n rt ^^i
tProgreso ,■: ■% * yJ^^^' >,
<^j
<p
-ilt
rjJi'
M/^~
i?r!stit.~i:V.
Fig. 35. Positions of the cone of observation at Progreso, Yucatan, on the night of April 23-24, 1948, from 8:53 p.m. to 3:53 a.m. Essential features of this map are drawn to scale. The telescope was set up on the end of a one- mile long wharf that extends northward from the shore over the waters of the Gulf of Mexico. The triangular (white) lines represent the projections of the cone of visibility on the earth at the mid-point of each hour of observation. Only briefly, in the first two hours, did the cone lie even in part over the ad- jacent mainland. Hence, nearly all of the birds seen in the course of the night had actually left the land behind.
ing water between the telescope on the end of the wharf and the beach. Therefore, nearly all of the birds seen were actually ob- served leaving the coast and passing out over the open waters of the Gulf. The hourly station densities are shown in Table 7 and Figures 24 and 36. In the seventeen hours of observation on the nights of April 23-24 and April 24-25, a total computed density of 59,200 birds passed within one-half mile of each side of Progreso. This is the third highest density recorded in the course of this study. The
444 University of Kansas Publs., Mus. Nat. Hist.
14
12
b o o '- 8
z
>- to 6
X 4
8 9 iO II 12 I 2 3 4
HOUR OF THE NIGHT (CST)
Fig. 36. Hourly station density curve for night of April 23-24, 1948, at
Progreso, Yucatan.
Table 7. — Computed Hourly Densities at Progreso, Yuc, in Spring of 1948
|
Average hour of observation |
|||||||||
|
Date |
8:30 |
9:30 |
10:30 |
11:30 |
12:30 |
1:30 |
2:30 |
3:30 |
4:30 |
|
23-24 April 24-25 April |
400 0 |
3,000 500 |
5,100 3,700 |
10,000 11,900 |
9,000 7,900 |
2,800 1,900 |
900 1,100 |
400 400 |
200 |
maximum for one hour was a computed density of 11,900 birds. This is the fourth highest hourly density recorded in 1948.
It is not my contention that this many birds leave the northern coast of Yucatan every night in spring. Indeed, further studies may show negligible flight densities on some nights and even greater den- sities on others. As a matter of fact several hours of observation on the night of April 25-26, at Merida, Yucatan, approximately twenty- five miles inland from Progreso, indicated that on this night the density overhead was notably low, a condition possibly accounted for by a north wind of 10 mph blowing at 2,000 feet. I merely sub-
Lowery: Nocturnal Migration of Birds 445
mit that on the nights of April 23-24 and 24-25, birds were leaving the coast of Yucatan at Progreso at the rate indicated. But, as I have emphasized in this paper and elsewhere (1946: 205-206), the northern part of the Yucatan Peninsula is notably unmarked by- streams or any other physiographic features which birds might fol- low. The uniformity of the topography for many miles on either side of Progreso, if not indeed for the entire breadth of the Penin- sula, makes it probable that Progreso is not a particularly favored spot for observing migration, and that it is not the only point along the northern coast of Yucatan where high flight densities can be recorded. This probability must be considered when comparisons are made between Progreso densities and those at Tampico. The argument could be advanced that the present densities from Tam- pico do not sufficiently exceed those at Progreso to establish the coastal route as the main avenue of traffic in spring, since there is every reason to suspect topography of exerting some influence to produce a channeling effect in eastern Mexico. Here the coast par- allels the directional trend of the migratory movement for more than 600 miles. Likewise the Sierra Madre Oriental of eastern Mexico, situated approximately 100 miles inland (sometimes less), lies roughly parallel to the coast. Because of the slant of the Mexican land mass, many winter residents in southern Mexico, by short northward movements, would sooner or later filter into the coastal plain. Once birds are shunted into this lowland area, it would seem unlikely that they would again ascend to the top of the Sierra Madre to the west. In this way the great north-south cordillera of moun- tains may act as a western barrier to the horizontal dispersion of transients bound for eastern North America. Similarly, the Gulf itself may serve as an eastern barrier; for, as long as migrants may progress northward in the seasonal direction of migration and still remain over land, I believe they would do so.
To put the matter in a slightly different way, the idea of a very narrow flight lane is inherent in the idea of coastwise migration. For, as soon as we begin to visualize flights of great volume over fronts extending back more than fifty miles from the shore line, we are approaching, if indeed we have not already passed, the point where the phenomenon is no longer coastwise in essence, but merely overland (as indeed my own unprocessed, telescopic data for 1949 indicate may be the case) . In actuality, those who have reported on the migration along the western edge of the Gulf of Mexico have never estimated the width of the main flight at more than fifty miles
446 University of Kansas Publs., Mus. Nat. Hist.
and have intimated that under some circumstances it may be as narrow as two miles. No evidence of such restrictions can be dis- cerned in the case of the trans-Gulf flights. If it cannot be said that they may be assumed to be as wide as the Gulf itself, they at least have the potential breadth of the whole 260-mile northern coast of the Yucatan Peninsula. On these premises, to be merely equal in total magnitude, the coastwise flights must exhibit, de- pending on the particular situation, from five to 130 times the con- centrations observable among trans-Gulf migrants. This point seems almost too elementary to mention, but I have yet to find any- one who, in comparing the two situations, takes it into consideration.
Judged in this light, the average hourly density of 2,800 birds at Progreso in April would appear to be indicative of many more mi- grants on the entire potential front than the 6.300 birds representing the average hourly density for the same month at Tampico.
That the Progreso birds were actually beginning a trans-Gulf flight seems inevitable. The Yucatan Peninsula projects 200 miles or more northward into the vast open expanses of the Gulf of Mexico and the Caribbean Sea, with wide stretches of water on either side. The great majority of the birds were observed after they had pro- ceeded beyond the northern edge of this land mass. Had they later veered either to the east or the west, they would have been obliged to travel several hundred miles before again reaching land, almost as far as the distance straight across the Gulf. Had they turned southward, some individuals should have been detected flying in that direction. As can be seen from Figures 23, 42, and 44, not one bird observed was heading south of east or south of west on either night. No other single piece of evidence so conclusively demon- strates that birds cross the Gulf of Mexico in spring in considerable numbers as do flight density data recorded from Progreso in 1948.
Northern Gulf Area
Unfortunately only a few data on flight density are available from critical localities on the northern shores of the Gulf in spring. As the density curves in Figure 30 demonstrate, several sets of observa- tion, including some phenomenal flights, have been recorded at Baton Rouge. This locality, however, lies sixty-four miles from the closest point on the Gulf coast, and the point due southward on the coast is eighty-four miles distant. Since all of the birds seen at Baton Rouge on any one night may have come from the heavily forested area between Baton Rouge and the coast of the Gulf, we cannot use data from Baton Rouge as certainly representative of
Lowery: Nocturnal Migration of Birds 447
incoming trans-Gulf flights. Data from repeated observations at stations on the coast itself are needed to judge the degree of trans- Gulf migration northward. On the few nights of observation at such localities (Cameron and Grand Isle, Louisiana, and Pensacola, Florida), flight densities have been zero or negligible. To be sure, negative results have been obtained at stations in the interior of the United States, and flights of low density have been recorded on occasion at stations where the flight densities are otherwise high. Nevertheless, in view of the volume of migration departing from Progreso, Yucatan, it would appear, upon first consideration, that we should at times record on the coast of Louisiana enough birds arriving in a night of continuous observation to yield a high density figure.
Upon further consideration, however, there are factors mitigating against heavy densities of birds in northern flight on the northern coast of the Gulf. In the first place, presuming the main trans-Gulf flight to originate from northern Yucatan, and that there is a direc- tional fanning to the northward, the birds leave on a 260-mile front, and arrive on a front 400 miles or more wide. Consequently, other factors remaining the same, there would be only approxi- mately half the number of birds on the coast of arrival, at a given time and place, as there was on the coast of departure. Secondly, we may now presume on the basis of the telescopic studies at Pro- greso, that most migrants leaving northern Yucatan do so in the few hours centering about midnight. The varying speeds of the birds making the 580-mile flight across the Gulf distribute them still more sparsely on the north coast of the Gulf both in time and in space. Also we can see only that segment of the flight, which arrives in that part of a twenty-four hour period when the moon is up. This circumstance further reduces the interceptive potential because the hours after dark, to which the present telescopic studies have been restricted, comprise the period in which the fewest mi- grants arrive from over the water. To illustrate: it is a mathe- matical certainty that none of the birds leaving Yucatan in the hours of heaviest flight, before 12 p. m., and flying on a straight course at a speed of approximately 33 mph will reach the northern Gulf coast after nightfall; they arrive in the daytime. It will be useful to devise a technique for employing the sun as a background for tele- scopic observation of birds, thereby making observations possible on a twenty-four hour basis, so as to test these inferences by objec- tive data.
448 University of Kansas Publs., Mus. Nat. Hist.
When a whole night's observation (1949 data not yet processed) at Port Aransas, on the southern coast of Texas, on the great over- land route from eastern Mexico, yields in one night in April only seven birds, the recording of no birds at a station near the mouth of the Mississippi River becomes less significant.
As I have previously remarked in this paper, the new data obtained since 1946, when I last wrote on the subject of migration in the re- gion of Gulf of Mexico, requires that I alter materially some of my previously held views. As more and more facts come to light, I may be compelled to alter them still further. For one thing, I have come to doubt seriously the rigidity of the coastal hiatus as 1 envisioned it in 1945. I believe instead that the scarcity of records of transient migrants on the Gulf coastal plain in fair weather is to a very large extent the result of a wide dispersion of birds in the dense cover that characterizes this general region. I now question if appreciable bird densities on the ground ever materialize any- where except when the sparseness of suitable habitat for resting or feeding tends to concentrate birds in one place, or when certain meteorological conditions erect a barrier in the path of an oncoming migratory flight, precipitating many birds in one place.
This retrenchment of ideas is a direct consequence of the present study, for time and again, as discussed in the case of Tampico densi- ties, maximal nightly flights have failed to produce a visible abun- dance of transients on land the following day. A simple example may serve to illustrate why. The highest one-hour density recorded in the course of this study is 21,200 birds. That means that this many birds crossed a line one mile long on the earth's surface and at right angles to the direction of flight. Let us further assume that the average flight speed of all birds comprising this flight was 30 mph. Had the entire flight descended simultaneously, it would have been dispersed over an area one mile wide and thirty miles long, and the precipitated density on the ground would have been only 1.1 birds per acre. Moreover, if as many as ten species had been involved in the flight, this would have meant an average per species of less than one bird per nine acres. This would have failed, of course, to show appreciable concentrations to the observer in the field the following day. If, however, on the other hand, the same flight of 21,200 birds had encountered at one point a weather barrier, such as a cold-front storm, all 21,200 birds might have been precipi- tated in one place and the field observer would have recorded an "inundation of migrants." This would be especially true if the
Lowery: Nocturnal Migration of Birds 449
locality were one with a high percentage of open fields or prairies and if the flight were mainly of woodland dwelling species, or con- versely, if the locality were densely forested with few open situations and the flight consisted mainly of open-country birds. As explained on page 389, the density formula may be too conservative in its expression of actual bird densities. Even if the densities computed for birds in the air are only half as high as the actual densities in the air, the corresponding ground density of 2.2 birds per acre that results if all the birds descended simultaneously would hardly be any more impressive than the 1.1 bird per acre.
This consideration is doubtless highly modified by local circum- stances, but, in general, it seems to suggest a working hypothesis that provides an explanation for many of the facts that we now have. For example, on the coast of Texas there are great expanses of terrain unattractive to such birds as warblers, vireos, tanagers, and thrushes. The precipitation there by bad weather of even a medi- ocre nightly flight composed of birds of the kinds mentioned would surely produce an overwhelming concentration of birds in the scattered woods and shrubs.
In spite of all that has been written about the great concentrations of transient migrants on the coast of Texas in spring, I am not con- vinced that they are of a different order of magnitude than those con- centrations that sometimes occur along the cheniers and coastal is- lands of Louisiana and Mississippi. I have read over and over the highly informative accounts of Professor Williams {loci cit.) and the seasonal summaries by Davis (1936-1940) and Williams (1941- 1945). I have conversed at length with Mrs. Jack Hagar, whom I regard as one of the leading authorities on the bird life of the Texas coast, and she has even permitted me access to her voluminous records covering a period of fifteen years residence at Rockport. Finally, I have spent a limited amount of time myself on the Texas coast studying first-hand the situation that obtains there in order that I might be in a position to compare it with what I have learned from observations elsewhere in the region of the Gulf of Mexico, Louisiana, Florida, Yucatan, and eastern Mexico.
Although the concentrations of birds on some days near the mouth of the Mississippi River are almost incalculable, the fact remains that in Texas the densities of transient species on the ground are more consistently high from day to day. The reason for this may be simple. As birds move up daily from Mexico overland, a certain percentage would be destined to come down at all points along the
450 University of Kansas Publs., Mus. Nat. Hist.
route but so dispersed in the inland forest that they might pass un- noticed. However, that part of the same fiight settling down in coastal areas, where trees are scarce, would produce visible con- centrations of woodland species. With the advent of a cold-front storm, two diametrically opposite effects of the same meteorological phenomenon would tend to pile up great concentrations of migrants of two classes — the overland and the trans-Gulf flights. During the prepolar-front weather the strong southerly (from the south) and southeasterly winds would tend to displace much of the trans-Gulf segment to the western part of the Gulf. With the shift of the winds to the north and northwest, which always occurs as the front passes, the overland flight still in the air would tend to be banked up against the coast, and the incoming trans-Gulf fiight would be confronted with a barrier, resulting in the precipitation of birds on the first available land.
These postulated conditions are duplicated in part in autumn along the Atlantic coast of the eastern United States. There, as a result of the excellent work of Allen and Peterson (1936) and Stone (1937), a similar effect has been demonstrated when northwest winds shove the south-bound flights up against the coast of New Jersey and concentrate large aggregations of migrants there.
Interior of the United States
Attention has been drawn already to the nature of the nightly flights at stations immediately inland from the Gulf coast, where densities decline abruptly well before midnight. I have suggested that this early drop-off is mainly a result of the small amount of terrain south of these stations from which birds may be contributed to a night's flight. At Oak Grove, Louisiana, the fiight exhibited a strong directional trend with no significant aberrant components. Therefore, one may infer that a considerable part of the flight was derived from regions to the south of the station.
At Mansfield, Louisiana, thirty-eight hours of observation in April and May resulted in flight densities that are surprisingly low — much lower, in fact, than at Oak Grove. In eleven of the hours of observation no birds at all were seen. A possible explanation for these low densities lies in the fact that eastern Texas and western Louisiana, where, probably, the Mansfield flights originated, is not an especially attractive region to migrants because of the great amount of deforested and second growth pine land. Oak Grove, in contrast, is in the great Tensas-Mississippi River flood plain, char- acterized by an almost solid stand of deciduous forest extending over thousands of square miles in the lower Mississippi valley.
Lowery: Nocturnal Migration of Birds
451
In further contrast to the considerable flight densities and pro- nounced directional trend at Oak Grove, we have the results from Rosedale, Mississippi, only seventy miles to the north and slightly to the east. At Rosedale the densities were mediocre and the flight
21-22 APRIL
20-21 MAY
Fig. 37. Sector density representation on two nights at Rosedale, Mississippi, in 1948. The white lines are the vector resultants.
directions were extremely divergent. Many of the nights of observa- tion at this locality were seriously interrupted by clouds, but such counts as were made on those dates indicated little migration taking place. On two nights, however, April 21-22 and May 20-21, visi- bility was almost continuous and densities were moderately high. In Figure 37 I have shown the flight directions for these two nights. The lengths of the individual sector vectors are plotted as a per- centage of the total station density for each of the two nights (5,800 and 6,800 birds, respectively). Although the vector resultants show a net movement of birds to the northeast, there are important di- vergent components of the flights. This "round-the-compass" pat- tern is characteristic of stations on the edge of meteorological disturbances, as was Rosedale on April 21-22, but not on the night of May 20-21. If bats are presumed to have played a role in these latter observations, their random flights would tend to cancel
452 University of Kansas Publs., Mus. Nat. Hist.
out and the vector resultant would emerge as a graphic representa- tion of the actual net trend density of the birds and its prevailing direction of flow. Although I do not believe that bats are the real reason for the diverse directional patterns at Rosedale, I can offer no alternative explanation consistent with data from other stations.
Moving northward in the valley of the Mississippi and its tribu- taries, we find a number of stations that yielded significantly high densities on most nights when weather conditions were favorable for migration. Louisville and Murray, Kentucky, and Knoxville, Ten- nessee, each show several nights with many birds flying, but only Lawrence, Kansas, and Ottumwa, Iowa, had migrations that ap- proach in magnitude the record station densities at Tampico. In- deed, these were the only two stations in the United States that pro- duced flights exceeding the densities at Progreso, Yucatan. The densities at Lawrence are unique in one respect, in that they were extremely high in the month of March. Since there were very few stations in operation then, these high densities would be of little significance were it not for the fact that at no time in the course of this study from 1945 to the present have comparable densities been obtained this early in the migration period. Examination of the "Remarks" section of the original data sheets from Lawrence show frequent mention of "duck-like" birds passing before the moon. We may infer from these notations that a considerable part of the overhead flight was composed of ducks and other aquatic birds that normally leave the southern United States before the main body of transient species reach there. The heavy flight densities at Law- rence may likewise have contained certain Fringillidae, Motacillidae, Sylviidae, and other passerine birds that winter mainly in the southern United States and which are known to begin their return northward in March or even earlier. Observations in 1948 at Law- rence in April were hindered by clouds, and in May no studies were attempted. However, we do have at hand two excellent sets of data recorded at Lawrence on the nights of May 3-4 and May 5-6, 1947, when the density was also extremely high.
At Ottumwa, Iowa, where a splendid cooperative effort on the part of the local ornithologists resulted in forty-four hours of observation in April and May, densities were near the maximum for all stations. Considering this fact along with results at Law- rence and other mid-western stations where cloud cover did not interfere at the critical periods of observation, we have here evi- dence supporting the generally held thesis that eastern Kansas, Mis- souri, and Iowa lie on a principal migratory flyway.
Lowery: Nocturnal Migration of Birds 453
Stations in Minnesota, Illinois, Michigan, Massachusetts, and Ontario were either operated for only parts of one or two nights, or else clouds seriously interfered with observations, resulting in dis- continuous counts. It may be hoped that future studies will include an adequate representation of stations in these states and that observations will be extensive enough to permit conclusions re- garding the density and direction of migration.
Charleston, South Carolina, which does not conveniently fall in any of the geographic regions so far discussed, had, to me, a sur- prisingly low flight density; twenty-two hours of observation there in March, April, and May yielded a total flight density of only 3,000 birds. This is less, for example, than the number of birds computed to have passed Lawrence, Kansas, in one hour, or to have passed Progreso, Yucatan, in one twenty-minute interval! Possibly observations at Charleston merely chanced to fall on nights of in- explicably low densities; further observations will be required to clear up this uncertainty.
E. Migration and Meteorological Conditions
The belief that winds affect the migration of birds is an old one. The extent to which winds do so, and the precise manner in which they operate, have not until rather recently been the subject of real investigation. With modern advances in aerodynamics and the de- velopment of the pressure-pattern system of flying in aviation, at- tention of ornithologists has been directed anew to the part that air currents may play in the normal migrations of birds. In America, a brief article by Bagg (1948), correlating the observed abundance of migrants in New England with the pressure pattern obtaining at the time, has been supplemented by the unpublished work of Winni- fred Smith. Also Landsberg (1948) has pointed out the close cor- respondence between the routes of certain long-distance migrants and prevailing wind trajectories. All of this is basis for the hypoth- esis that most birds travel along definite air currents, riding with the wind. Since the flow of the air moves clockwise around a high pres- sure area and counterclockwise around a low pressure area, the birds are directed away from the "high" and toward the center of the "low." The arrival of birds in a particular area can be predicted from a study of the surrounding meteorological conditions, and the evidence in support of the hypothesis rests mainly upon the success of these predictions in terms of observations in the field.
From some points of view, this hypothesis is an attractive one. It explains how long distances involved in many migrations may be
454 University of Kansas Publs., Mus. Nat. Hist.
accomplished with a minimum of effort. But the ways in which winds affect migration need analysis on a broader scale than can be made from purely local vantage points. Studies of the problem must be implemented by data accumulated from a study of the proc- ess in action, not merely from evidence inferred from the visible results that follow it. Although several hundred stations operating simultaneously would surely yield more definite results, the tele- scopic observations in 1948 offer a splendid opportunity to test the theory on a continental scale.
The approach employed has been to plot on maps sector vectors and vector resultants that express the directional trends of migra- tion in the eastern United States and the Gulf region, and to com- pare the data on these maps with data supplied by the U. S. Weather Bureau regarding the directions and velocities of the winds, the loca- tion of high and low pressure areas, the movement of cold and warm fronts, and the disposition of isobars or lines of equal pressure. It should be borne in mind when interpreting these vectors that they are intended to represent the directions of flight only at the proximal ends, or junction points, of the arrows. The tendency of the eye to follow a vector to its distal extremity should not be allowed to create the misapprehension that the actual flight is supposed to have con- tinued on in a straight line to the map location occupied by the arrowhead.
A fundamental difficulty in the pressure-pattern theory of migra- tion has no doubt already suggested itself to the reader. The diffi- culty to which I refer is made clear by asking two questions. How can the birds ever get where they are going if they are dependent upon the whim of the winds? How can pressure-pattern flying be reconciled with the precision birds are supposed to show in return- ing year after year to the same nesting area? The answer is, in part, that, if the wind is a major controlling influence on the routes birds follow, there must be a rather stable pattern of air currents prevailing from year to year. Such a situation does in fact exist. There are maps showing wind roses at 750 and 1,500 meters above mean sea level during April and May (Stevens, 1933, figs. 13-14, 17-18). Similarly, the "Airway Meteorological Atlas for the United States" (Anonymous, 1941) gives surface wind roses for April (Chart 6) and upper wind roses at 500 and 1,000 meters above mean sea level for the combined months of March, April, and May (Charts 81 and 82). The same publication shows wind resultants at 500 and 1,000 meters above mean sea level (Charts 108 and 109).
Loweky: Nocturnal Migration of Birds
455
Fig. 38. Over-all sector vectors at major stations in the spring 1948. See text for explanation of s>'stem used in determining the length of vectors. For identification of stations, see Figure 34.
456
University of Kansas Publs.. Mus. Nat. Hist.
Fig. 39. Over-all net trend of flight directions at stations shown in Figure 38. The arrows indicate direction only and their slants were obtained by vector analysis of the over-all sector densities.
Lowery: Nocturnal Migration of Birds 457
Further information permitting a description in general terms of conditions prevailing in April and May is found in the "Monthly Weather Review" covering these months (c/. Anonymous, 1948 a, Charts 6 and 8; 1948 b, Charts 6 and 8).
First, however, it is helpful as a starting point to consider the over-all picture created by the flight trends computed from this study. In Figure 38, the individual sector vectors are mapped for the season for all stations with sufficient data. The length of each sector vector is determined as follows: the over-all seasonal density for the station is regarded as 100 per cent, and the total for the season of the densities in each individual sector is then expressed as a percentage. The results show the directional spread at each sta- tion. In Figure 39, the direction of the over-all vector resultant, obtained from the sector vectors on the preceding map, is plotted to show the net trend at each station.
As is evident from the latter figure, the direction of the net trend at Progreso, Yucatan, is decidedly west of north (N 26° W). At Tampico this trend is west of north (N 11° W) , but not nearly so much so as at Progreso. In Texas, Louisiana, Georgia, Tennessee, and Kentucky, it is decidedly east of north. In the upper Missis- sippi Valley and in the eastern part of the Great Plains, the flow appears to be northward or slightly west of north. At Winter Park, Florida, migration follows in general the slant of the Florida Peninsula, but, the meager data from Thomasville, Georgia, do not indicate a continuation of this trend.
It might appear, on the basis of the foregoing data, that birds migrate along or parallel to the southeast-northwest extension of the land masses of Central America and southern Mexico. This would carry many of them west of the meridian of their ultimate goal, obliging them to turn back eastward along the lines of net trend in the Gulf states and beyond. This curved trajectory is un- doubtedly one of the factors — but certainly not the only factor — contributing to the effect known as the "coastal hiatus." The ques- tion arises as to whether this northwestward trend in the southern part of the hemisphere is a consequence of birds following the land masses or whether instead it is the result of some other natural cause such as a response to prevailing winds. I am inclined to the opinion that both factors are important. Facts pertinent to this opinion are given below.
In April and May a high pressure area prevails over the region of the Gulf of Mexico. As the season progresses, fewer and fewer
7— 1C20
458 University of Kansas Publs., Mus. Nat. Hist.
cold-front storms reach the Gulf area, and as a result the high pressure area over the Gulf is more stable. Since the winds move clockwise around a "high," this gives a general northwesterly trajec- tory to the air currents in the vicinity of the Yucatan Peninsula. In the western area of the Gulf, the movement of the air mass is in general only slightly west of north, but in the central Gulf states and lower Mississippi Valley the trend is on the average northeast- erly. In the eastern part of the Great Plains, however, the average circulation veers again slightly west of north. The over-all vector resultants of bird migration at stations in 1948, as mapped in Figure 39, correspond closely to this general pattern.
Meteorological data are available for drawing a visual com- parison between the weather pattern and the fight pattern on indi- vidual nights. I have plotted the directional results of four nights of observation on the Daily Weather Maps for those dates, showing surface conditions (Figures 40, 42, 44 and 46). Each sector vector is drawn in proportion to its percentage of the corresponding nightly station density; hence the vectors at each station are on an inde- pendent scale. The vector resultants, distinguished by the large arrowheads, are all assigned the same length, but the nightly and average hourly station densities are tabulated in the legends under each figure. For each map showing the directions of flight, there is on the facing page another map showing the directions of winds aloft at 2,000 and 4,000 feet above mean sea level on the same date (see Figures 41-47). The maps of the wind direction show also the velocities.
Unfortunately, since there is no way of analyzing the sector trends in terms of the elevations of the birds involved, we have no certain way of deciding whether to compare a given trend with the winds at 2,000, 1,000, or 0 feet. Nor do we know exactly what wind corre- sponds to the average or median flight level, which would otherwise be a good altitude at which to study the net trend or vector resultant. Furthermore, the Daily Weather Map illustrates conditions that obtained at 12:30 a. m. (CST) ; the winds aloft are based on obser- vations made at 10:00 p. m. (CST) ; and the data on birds covers in most cases the better part of the whole night. Add to all this the fact that the flight vectors, their resultants, and the wind representations themselves are all approximations, and it becomes apparent that only the roughest sort of correlations are to be expected.
However, as will be seen from a study of the accompanying maps (Figures 40-47), the shifts in wind direction from the surface up to 4,000 feet above sea level are not pronounced in most of the in-
Lowery: Nocturnal Migration of Birds 459
stances at issue, and such variations as do occur are usually in a clockwise direction. All in all, except for regions where frontal activity is occurring, the weather maps give a workable approxi- mation to the average meteorological conditions on a given night.
The maps (Figures 40-47) permit, first, study of the number of instances in which the main trend of flight, as shown by the vector resultant, parallels the direction of wind at a reasonable potential mean flight elevation, and, second, comparison of the larger indi- vidual sector vectors and the wind currents at any elevation below the tenable flight ceiling — one mile.
On the whole, inspection of the trend of bird-flight and wind di- rection on specific nights supports the principle that the flow of migration is in general coincident with the flow^ of air. It might be argued that when the flow of air is toward the north, and when birds in spring are proceeding normally in that direction, no signifi- cance can be attached to the agreement of the two trends. However, tlie same coincidence of wind directions and bird flights seems to be maintained when the wind currents deviate markedly from a north- ward trajectory. Figures 46 and 47, particularly in regard to the unusual slants of the flight vectors at Ottumwa, Knoxville, and Memphis, illustrate that this coincidence holds even when the wind is proceeding obliquely eastward or westward. On the night of May 22-23, when a high pressure area prevailed from southern Iowa to the Atlantic coast, and the trajectory of the winds was northward, migration activity at Knoxville and Ottumwa was greatly increased and the flow of birds was again northward in the normal seasonal direction of migration.
Further study of the data shows fairly conclusively that maxi- mum migration activity occurs in the regions of high barometric pressure and that the volume of migration is either low or negligible in regions of low pressure. The passage of a cold-front storm may almost halt migration in spring. This was demonstrated first to me by the telescopic method at Baton Rouge, on April 12, 1946, following a strong cold front that pushed southeastward across the Gulf coastal plain and over the eastern Gulf of Mexico. The winds, as usual, shifted and became strong northerly. On this night, following the shift of the wind, only three birds were seen in seven hours of continuous observation. Three nights later, however, on April 15, when the warm air of the Gulf was again flowing from the south, I saw 104 birds through the telescope in two hours. Apropos of this consideration in the 1948 data are the nights of May 21-22 and 22-23.
460
University of Kansas Publs., Mus. Nat. Hist.
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Fid. 40. Comparison of flight trends and surface weather conditions on April 22-23, 1948. The meteorological data were taken from the U. S. Weather Bureau Daily Weather Map for 12:30 a.m. (CST) on April 23. The nightly station densities and the average hourly station density (shown in parentheses) are as follows:
5. Louisville: 9,100 (1,100) 16. College Station: 13,300 (1.900)
6. Murray: 16,300 (2,700) 17. Baton Rouge: 6,200 (1.000)
8. Stillwater: 1,900 (500) 19. Lafayette: 2,800 (600)
9. Knoxville: 15.200 (1.700) 21. Winter Park: 6,200 (700) 13. Oak Grove: 13,600 (1.700) 23. Tampico: 11,100(3,700)
Lowery: Nocturnal Migration of Birds
461
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Fig. 41. Winds aloft at 10:00 p. m. on April 22 (CST). Winds at 2,000 feet above mean sea level are shown in black; those at 4.000 feet, in white. Veloc- ities are indicated by standard Beaufort S'cale of Wind Force. The numbers in circles refer to the stations shown in Figure 40.
Correction: Figures 41 and 45 were inadvertently transposed.
462 Univ-ersity of Kansas Publs., Mus. Nat. Hist.
Fig. 42. Comparison of flight trends and surface weather conditions on April 23-24, 1948. The meteorological data were taken from the U. S. Weather Bureau Daily Weather Map for 12:30 .'\. m. (CST) on April 24. The nightly station densities and the average hourly station density (shown in parentheses) are as follows:
1. Albion: 1,100 (300) 14. Mansfield: 4.900 (1,200)
2. Ottumwa: 5,500 (90O) 16. College Station: 700 (100)
4. Lawrence: 5,400 (1,400) 17. Baton Rouge: 1,700 (400)
5. Louisville: 13,300 (2,700) 18. Pensacola: migraKon negligible
6. Murray: 9,800 (1,400) 20. New Orleans: 1,600 (800)
8. Stillwater: 800 (100) 21. Winter Park: 2,700 (300)
9. Kno.xville: 8,000 (900) 23. Tampico: 63,600 (6,300) 10. Memphis: 7,900 (1,000) 24. Progreso: 31,300 (3,900)
Lowery: Nocturnal Migration of Birds
463
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Fig. 43. Winds aloft at 10:00 p. m. on April 23 (CST). Winds at 2,000 feet above mean sea level are shown in black; those at 4,000 feet, in white. Veloc- ities are indicated by standard Beaufort Scale of Wind Force. The numbers in circles refer to the stations shown in Figure 42.
464
University of Kansas Publs., Mrs. Nat. Hist.
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Fig. 44. Comparison of flight trends and surface weather conditions on April 24-25. 1948. The meteorological data were taken from the U. S. Weather Bureau Daily Weather Map for 12:30 a.m. (CST) on April 25. The nightly station densities and the average hourly station density (shown in parentheses) are as follows:
1. Albion: migration negligible 12. Rosedale : 1.100 (100)
2. Ottumwa: 4.600 (1,500) 14. Mansfield: 1.700 (400)
3. Cohimbia: 1.400 (400) 18. Pensacola: migration negligible 5. Louisville: 1,700 (200) 21. Winter Park : 600 (100)
10. Memphis: 6,600 (900) 24. Progreso: 27,300 (3.000)
Lowery: Nocturnal Migration of Birds
465
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Fig. 45. Winds aloft at 10:00 p. m. on April 24 (CST). Winds at 2,000 feet above mean sea level are shown in black; those at 4,000 feet, in white. Veloc- ities are indicated by standard Beaufort Scale of Wind Force. The numbers in circles refer to the stations shown in Figure 44.
Correction: Figures 41 and 45 were inadvertently transposed.
466 University of Kansas Publs., Mus. Nat. Hist.
Fig. 46. Comparison of flight trends and surface weather conditions on May 21-22, 1948. The meteorological data were taken from the U. S. Weather Bureau Daily Weather Map for 12:30 a.m. (CST) on May 22. The nightly station densities and the average hourly station density (shown in parentheses) are as follows: 2. Ottumwa: 6,900 (1,400) 13. Oak Grove: 5,800 (800)
5. Louisville: 1,500 (200) 14. Mansfield: 2,500 (800)
9. Knoxville: 3,200 (500) 18. Pensacola: migration negligible
10. Memphis: 7,000 (1,200) 21. Winter Park: 1,200 (200)
Lowery: Nocturnal Migration of Birds
467
Fig. 47. Winds aloft at 10:00 p. m. on May 21 (CST). Winds at 2,000 feet above mean sea level are shown. Velocities are indicated by standard Beaufort Scale of Wind Force. The numbers in circles refer to the stations shown in Figure 46.
468 University of Kansas Publs., Mus. Nat. Hist.
On the first night, following the passage of a cold front, migration at Ottumwa was comparatively low (6,900 birds in five hours) . On the following night, when the trajectory of the winds was toward the north, the volume of migration was roughly twice as high (22,300 birds in eight hours). At Louisville, on May 21-22, the nightly station density was only 1,500 birds in seven hours, whereas on the following night, it was 8,400 birds in the same length of time, or about six times greater.
The evidence adduced from the present study gives support to the hypothesis that the continental pattern of spring migration in eastern North America is regulated by the movement of air masses. The clockwise circulation of warm air around an area of high pres- sure provides, on its western edge, tail winds which are apparently favorable to northward migration. High pressure areas exhibit a centrifugal force outward from the center, which may tend to dis- perse the migratory flight originating at any given point. In con- trast, the circulation of air in the vicinity of a low pressure area is counterclockwise with the force tending to be directed inward toward the center. Since the general movement of the air is from the high pressure area toward a low pressure area, birds starting their mi- grations with favorable tail winds, are often ultimately carried to a region where conditions are decidedly less favorable. In the vicinity of an area of low pressure the greater turbulence and high wind velocities, combined with the possibly slightly less buoyant property of the air, cause birds to descend. Since low pressure areas in spring generally precede cold fronts, with an attending shift of the wind to the north, an additional barrier to the northward mi- gration of birds is imposed. The extreme manifestation of low pressure conditions and the manner in which they operate against bird flight, are associated with tropical hurricanes. There, the cen- tripetal force of the wind is so great that it appears to draw birds into the "eye" of the hurricane. A classic example of this effect is seen in the case of the birds that came aboard the "West Quechee" when this vessel passed through the "eye" of a hurricane in the Gulf of Mexico in August, 1927. I have already discussed the details of this incident in a previous paper (1946:192). There is also the interesting observation of Mayhew (1949), in which a similar observation was made of large numbers of birds aboard a ship passing through one of these intense low-pressure areas.
Although the forces associated with an ordinary low-pressure area are by no means as intense as those associated with a tropical hurri-
Lowery: Nocturnal Migration of Birds 469
cane, the forces operating are much the same. Consequently birds conceivably might tend to be drawn toward a focal point near the center of the low, where the other factors already mentioned would tend to precipitate the entire overhead flight. Visible evidence of migration would then manifest itself to the field ornithologists.
CONCLUSIONS
1. Telescopic counts of birds passing before the moon may be used to determine reliable statistical expressions of the volume of migration in terms of direction and of definite units of time and space.
2. Night migrants fly singly more often than in flocks, creating a remarkably uniform dispersion on a local scale throughout the sky, quite unlike the scattered distributions observable in the daytime.
3. The nocturnal migration of birds is apparently preceded by a resting or feeding pause during which there are few migrants in the air. It is not to an important degree a non-stop continu- ation of flights begun in the daylight.
4. Nightly migrational activity in North America varies from hour to hour according to a definite temporal pattern, corre- sponding to the Zugunruhe of caged European birds, and ex- pressed by increasingly heavy flights up until the hour before midnight, followed by a pronounced decline.
5. The visible effects of the time pattern are subject to modifica- tion at a particular station by its location with respect to the resting areas from which the night's flight originates.
6. Quantitative and directional studies have so far failed to prove that nocturnal migrants favor narrow, topographically-deter- mined flight lanes to an important degree.
7. Flight densities on the east coast of Mexico, though of first magnitude, have not yet been demonstrated in the volume de- manded by the premise that almost all migrants returning to the United States from regions to the south do so by coastal routes.
8. Heavy flights have been recorded from the northern coast of Yucatan under circumstances leading inevitably to the con- clusion that birds migrate across the Gulf of Mexico in consid- erable numbers.
9. There is reason to believe that the importance of the Florida Peninsula as an April and May flyway has been over-estimated, as regards the numbers of birds using it in comparison with the numbers of birds using the Mexican and Gulf routes.
470 University of Kansas Publs., Mus. Nat. Hist.
10. The amount of migration is apparently seldom sufficient to pro- duce heavy densities of transient species on the ground without the operation of concentrative factors such as ecological pat- terns and meteorological forces.
11. The absence or scarcity of transients in some areas in fine weather may be explained by this consideration.
12. A striking correlation exists between air currents and the direc- tional flight trends of birds, suggesting that most night migrants travel by a system of pressure-pattern flying.
LITERATURE CITED
Allen, R. P., and R. T. Peterson
1936. The hawk migrations at Cape May Point, New Jersey. Auk, 53: 393-404.
Anonymous
1936-1941. Tables of computed altitude and azimuth. U. S. Navy De- partment Hydrographic OflBce. U. S. Govt. Printing Office, Wash- ington, D. C, vols. 3-5.
1941. Airway meteorological atlas for the United States. Weather Bureau Publ. 1314. U. S. Dept. Commerce, Washington, D. C.
1945-1948. The American air almanac. U. S. Naval Observatory. U. S.
Govt. Printing Office, Washington, D. C, 3 vols., issued annually. 1948a. Meteorological and climatological data for April 1948. Monthly
Weather Review, April 1948, 76:65-84, 10 charts. 19486. Meteorological and climatological data for May 1948. Monthly
Weather Review, May 1948, 76:85-103, 11 charts.
Bagg, a. M.
1948. Barometric pressure-patterns and spring migration. Auk, 65:147.
Bergman, G.
1941. Der Fruhlingszug von Clangula hyemalis (L.) und Oidemia nigra (L.) bei Helsingfors. Eine Studie iiber Zugverlauf und Witterung sowie Tagesrhythmus und Flughohe. Ornis Fennica, 18:1-26.
Brat, R. A.
1895. A remarkable flight of birds. Nature (London), 52:415.
Carpenter, F. W.
1906. An astronomical determination of the height of birds during noc- turnal migration. Auk, 23:210-217.
Chapman, F. M.
1888. Observations on the nocturnal migration of birds. Auk, 5:37-39.
Davis, L. I.
1936-1940. The season : lower Rio Grande Valley region. Bird-Lore (now Audubon Mag.), 38-42.
Lowery: Nocturnal Migration of Birds 471
F. [arner], D. [onald] S.
1947. Studies on daily rhythm of caged migrant birds (review of Palmgren article). Bird-Banding, 18:83-84.
Gates, W. H.
1933. Hailstone damage to birds. Science, 78:263-264.
Howell, A. H.
1932. Florida bird life. Florida Department Game and Fresh Water Fish, Tallahaseee, 1-579 -|- 14 pp., 58 pis., 72 text figs.
Lansberg, H.
1948. Bird migration and pressure patterns. Science, 108:708-709.
LiBBY, O. G.
1899. The nocturnal flight of migratory birds. Auk, 16:140-146.
LowERY, G. H., Jr.
1945. Trans-Gulf spring migration of birds and the coastal hiatus. Wilson Bull., 57:92-121.
1946. Evidence of trans-Gulf migration. Auk, 63:175-211.
Mayhew, D. F.
1949. Atmospheric pressure and bird flight. Science, 109:403.
OVERING, R.
1938. High mortality at the Washington Monument. Auk, 55:679.
Palmgren, P.
1944. Studien iiber die Tagesrhythmik gekafigter Zugvogel. Zeitschrift fi'r Tierpsychologie, 6:44-86.
POUGH, R. H.
1948. Out of the night sky. Audubon Mag., 50:354-355.
PUTKPNEN, T. A.
1942. Kevatmuutosta Viipurinlakdella. Omis Fennica, 19:33-44.
Rense, W. a.
1946. Astronomy and ornithology. Popular Astronomy, 54:55-73.
Scott, W. E. D.
ISSla. Some observations on the migration of birds. Bull. Nuttall Orni.
Club, 6:97-100. 18816. Migration of birds at night. Bull. Nuttall Orni. Club, 6:188.
SlIVONEN, L.
1936. Die Stiirkevariation des Nachtlichen Zuges bei Turdiis ph. philomelos Brehn und T. musicus L. auf Grund der Zuglaute geschatz und mit der Zugunruhe einer gekafigten Singdrossel Verglichen. Omis Fen- nica, 13:59-63.
Spofford, W. R.
1949. Mortality of birds at the ceilometer of the Nashville airport. Wilson Bull., 61:86-90.
Stebbins, J.
1906. A method of determining height of migrating birds. Popular As- tronomy, 14:65-70.
472 University of Kansas Publs., Mus. Nat. Hist.
Stephens, Loyd A.
1933. Upper-air wind roses and resultant winds for the eastern United States. Monthly Weather Review, Supplement No. 35, November 13, pp. 1-3, 65 figs.
Stone, W.
1906. Some light on night migration. Auk, 23:249-252. 1937. Bird studies at Old Cape May. Delaware Valley Omi. Club, Phila- delphia, Vol. 1, 1-520 + 15 pp., pis. 1-46 and frontis.
Thomson, A. L.
1926. Problems of bird migration. Houghton Mifflin Company, Boston.
Van Oordt, G.
1943. Vogeltrek. E. J. Brill, Leiden, xii + 145 pp.
Very, F. W.
1897. Observations of the passage of migrating birds across the lunar disc on the nights of September 23 and 24, 1896. Science, 6:409-411.
Walters, W.
1927. Migration and the telescope. Emu, 26:220-222.
West, R. H.
1896. Flight of birds acro&s the moon's di.sc. Nature (London), 53:131.
Williams, G. G.
1941-1948. The season: Texas coastal region. Audubon Mag., 43-50. 1945. Do birds cross the Gulf of Mexico in spring? Auk, 62:98-111. 1947. Lowery on trans-Gulf migration. Auk, 64:217-238.
Winkenwerder, H. a.
1902a. The migration of birds with special reference to nocturnal flight. Bull. Wisconsin Nat. Hist. Soc, 2:177-263.
1902ti. Some recent observations on the migration of birds. Bull. Wis- consin Nat. Hist. Soc, 2:97-107.
Transmitted June 1, 1949.
□
23-1020
ogeny of the Waxwings and Allied Birds
BT
M. DALE ARVEY
University of Kansas Publications Museum of Natural History
Volume 3, No. 3, pp. 473-530, 49 figures in text, 13 tables October 10, 1951
Mire. Zm?. ZCOL LIBRAinf
MAR 1 7 1952
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The University of Kansas Publications, Museum 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, Univeksity of Kansas Li- brary, Lawrence, Kansas, U. S. A.
Museum op Natural History. — E. Raymond Hall, Chairman, 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. , Durrant. Pp. 1-82, 1 figure in text. August 15, 1946.
2. The systematic status of Eumeces pluvialis Cope, and noteworthy records of other a,mphibians 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 two 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. Kjrphosis 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 of two broods of great horned owls (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. Lowery, 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.
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14. A new pocket gopher (Thomomys) and a new spiny pocket mouse (Liomys) from Michoacan, Mexico. By E. Ra3anond Hall and Bernardo Villa-R. Pp. 249-256, 6 figures in text. July 26, 1948.
(Continued on inside of back cover.)
O'
Phylogeny of the Waxwings and Allied Birds
BY
M. DALE ARVEY
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MAR 1 7 1952
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University of Kansas Publications Museum of Natural History
Volume 3, No. 3, pp. 473-530, 49 figures in text, 13 tables October 10, 1951
University of Kansas
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1950
23-1019
Phylogeny of the Waxwings and Allied Birds
by M. DALE ARVEY
CONTENTS
PAGE
Introduction 476
Acknowledgments 476
nomenclatural history 477
Materials 478
Diagnoses 478
Coloration 485
Courtship 489
Nest Building 491
Food 493
Skeleton 494
Skull 494
Humerus 499
Pygostyle 502
Sternum 505
Relative Lengths of Bones 505
Leg-Trunk Percentages 509
Arm-Trunk Percentages 511
Musculature 514
Caudal Muscles 514
Pectoral Muscles 517
Hind Limb Musculature 517
Digestive Tract 517
Origin of the Species 519
Conclusions 521
Summary 524
Bibliography 525
(475)
476 University of Kansas Publs., Mus. Nat. Hist.
INTRODUCTION
A small family of passerine birds, the Bombycillidae, has been selected for analysis in the present paper. By comparative study of coloration, nesting, food habits, skeleton and soft parts, an attempt is made to determine which of the differences and similarities be- tween species are the result of habits within relatively recent geolog- ical time, and which differences are the result of inheritance from ancient ancestral stocks, which were in the distant past morphologi- cally different. On the basis of this information, an attempt is made to ascertain the natural relationships of these birds. Previous workers have assigned waxwings alone to the family Bombycillidae, and a question to be determined in the present study is whether or not additional kinds of birds should be included in the family.
It has generally been assumed that the nomadic waxwings origi- nated under boreal conditions, in their present breeding range, and that they did not undergo much adaptive radiation but remained genetically homogeneous. Also it is assumed that the species were wide ranging and thus did not become isolated geographically to the extent that, say, the Fringillidae did. The assumption that wax- wings originated in the northern part of North America or Eurasia may be correct, but it is more probable that the origin was more southerly, perhaps, in northern Mexico, of North America (see p. 519. Subsequent to the differentiation of this stock in the south, there was a northerly movement, while certain populations remained behind and underwent an evolution different from the northern group. Since the fossil record does not permit us to say when in geological time the family originated, we must rely on anatomical evidence and the distributional evidence of present-day species to estimate when the family stock had diverged from some unknown group sufficiently to merit the status of a separate family.
ACKNOWLEDGMENTS
It is witli pleasure that I acknowledge the guidance received in this study from Professor E. Raymond Hall of the University of Kansas. I am indebted also to Dr. Herbert Friedmann of the United States National Museum for the loan of certain skins, skeletons, and alcoholic material; to Mr. Alexander Skutch, for notes on cer- tain Central American birds; and to Dr. Henry W. Setzer, Mr. George H. Lowery, Jr., Mr. Victor E. Jones, Mr. Victor Housholder,
Arvey: Waxwings and Allied Birds 477
Mr. Alvaro Wille-Trejos, and Mr. Morton F. Davis, for gifts of specimens that have been used in this work. Suggestions and criti- cal comments from Professors Worthie H. Horr, Charles G. Sibley and Edward H. Taylor are gratefully acknowledged. I wish also to thank Mrs. Virginia Unruh for the preparation of the drawings used in this work,
NOMENCLATURAL HISTORY
The oldest name available for any species of the waxwings is Lanius garrulus Linnaeus (1758). Lanius garrulus and Lanius gar- rulus variety B carolinensis were described as conspecific. The de- scription has been associated with the first of the two names. The latter name is a nomen nudum since it was not accompanied by a separate description. The generic name Lanius was originally ap- plied to both shrikes and waxwings by Linnaeus. Since that name is applied to the shrikes only, the next available generic name that may be applied to the generically different waxwings must be used. This is Bo?nby cilia, a name originally proposed by Brisson (1760) for the Cedar Waxwing. In the 12th Edition of the Systemae Na- turae (1766) Gmelin proposed the generic name Ampelis for the Bohemian Waxwing, and combined it with the specific name gar- rulus, the Cedar Waxwing being termed variety B. Vieillot (1807) l^roposed the generic name Bomby cilia and combined it with a new specific name, cedrorum, for the Cedar Waxwing. Vieillot has been cited as the author of Bombycilla since that time, although Brisson used Bombycilla 33 years before. Oberholser (1917) did not cite Brisson's work in his discussion of the proper generic name for the waxwings, and Bombycilla should be ascribed to Brisson and not Vieillot, since Opinion 37, rendered by the International Zoological Committee on Nomenclature, states that generic names used by Brisson (1760) are valid under the Code. In consequence, the spe- cific name available for the Cedar Waxwing, since Brisson is ruled not to be a binomialist, is Bombycilla cedrorum Vieillot (1807).
Most workers prior to 1900 utilized the family name Ampelidae to include waxwings, silky flycatchers, and palm-chats. Ridgway (1904:113) elevated the silky flycatchers to family rank under the name Ptilogonatidae, and assigned the palm-chats to a separate family, the Dulidae.
478 University of Kansas Publs., Mus. Nat. Hist.
MATERIALS
The following specimens, numbering 238, and representing each currently recognized species and subspecies, were used in the study, and were supplemented by observation in 1947 on specimens in the United States National Museum.
Species or Subspecies
Skin
Skeleton
Alcoholic
Phainoptila melanoxantha melanoxantha .
Phainoptila melanoxantha minor
Ptilogonys cinereus cinereus
Ptilogonys cinereus molybdophanes
Ptilogonys caudatus
Phainopepla nitens nitens
Phainopepla nitens lepida
Boynhycilla cedrorum
Bombycilla garrula garrula
Bombycilla garrula centralasiae
Bombycilla garrula pallidiceps
Bombycilla japonica
Dulus dominicus dominicxis
Dulus dominicus oviedo
Totals.
8 2
13 6
16
12 53 4 9 7 10 9 4
153
1 3'
3 1 5
27 3 2 3
5 1
54
2
'4'
4 5 4
8
2
'2'
31
DIAGNOSES Family Bombycillidae
Diagnosis. — Bill short, flat, somewhat obtuse, minutely notched near tip of each maxilla, flared at base; gape wide and deeply cleft; culmen convex; nasal fossa broad, exposed, or filled with short, erect or antrorse, close-set velvety feathers; nostril narrowly elliptical; rictal vibrissae long, short, or absent; lacrimal bone free, articulating at two points; wings long and pointed, or short and rounded; primaries ten, tenth reduced in some species; tail short, narrow, even, two thirds or less length of wing, or much longer and forked or rounded; feet weak (except in Dulus and Phainoptila) ; tarsus generally shorter than middle toe and claw, distinctly scutellate with five or six divisions, the lateral plate subdivided (except in Phainoptila) ; lateral toes of nearly equal length; hallux approximately as long as inner lateral toe, or shorter; basal phalanx of middle toe more or less united to that of outer and inner toes; body stout; head generally conspicuously crested; plumage soft, smooth and silky (except in Dulu^) ; eggs spotted; nest in trees; three subfamilies, five genera, eight species.
Subfamily Ptilogonatinae
Diagnosis. — Rictus with conspicuous bristles ; nasal fossa almost entirely exposed; tail long and rounded, graduated, or square; caudal muscles and pygostyle well developed; wings rounded and short, first primary a half to a third as long as second; second primary shorter than third; humerus long.
Arvey: Waxwings and Allied Birds 479
with small external condyle; plumage soft and silky, less so in Phainoptila; sexes dissimilar, young like adult female; three genera, four species.
Genus Phainoptila Salvin
Phainoptila Salvin, Proc. Zool. Soc. London, 1877 :367, April 17, 1877. Type Phainoptila melanoxantha Salvin.
Diagnosis. — Without crest; tarsus longer than middle toe and claw, and booted or very slightly reticulate; tail shorter than wing, rounded; nostril exposed, ovate; rictal bristles distinct; first primary well developed; plumage normal, bill flared slightly at base.
Range. — Costa Rica and Panama.
Phainoptila melanoxantha melanoxantha Salvin Phainoptila
Phainoptila melanoxantha melanoxantha Salvin, Proc. Zool. Soc. London, 1877:367; April 17, 1877.
Diagnosis. — Coloration of adult males: Pileum, hindneck, back, scapulars, and upper tail coverts Black (capitalized color terms after Ridgway, Color Standards and Color Nomenclature, Washington, D. C, 1912), with Bluish Gray-Green gloss; rump Lemon Yellow tinged with Olive; lower breast and abdomen Gull Gray or Slate Gray; sides and flanks clear Lemon Yellow; lower chest, upper breast, and under tail coverts Yellowish Olive-Green, extending to patch on sides and flanks of same color; bill and feet Black or Blackish Brown. Coloration of adult females: Most of upper parts Olive-Green, with Yellowish Olive on rump; thighs Olive-Gray, as are sides of head; rest of coloration as in male. Coloration of young: As in adult female, but duller throughout.
Measurements. — Wing 99.0, tail 88.5, culmen 15.2, tarsus 28.4.
Range. — Highlands of Costa Rica and extreme western Panama (Volcan de Chiriqui).
Phainoptila melanoxantha minor Griscom
Phainoptila
Phainoptila melanoxantha minor Griscom, Amer. Mus. Novitates, 141 :7, 1924.
Diagnosis. — Coloration as in P. m. melanoxantha, but female with hindneck more extensively gray and of slightly darker shade; rump, upper tail coverts, and edgings to tail feathers slightly greener, less yellow; average size smaller than in P. m. melanoxantha.
Range. — Highlands of westeran Panama (Cerro Flores and eastern Chiriqui).
Genus Ptilogonys Swainson
Ptilogonys Swainson, Cat. Bullock's Mex. Mus., App. 4, 1824. Type Ptilogonys cinereus Swainson.
Diagnosis. — Tail much longer than wing, even or graduated; head with bushy crest; nostril large, rounded and fully exposed, bordered by membrane; rictal bristles well developed; tarsus shorter than middle toe with claw; plumage soft, blended.
Range. — Southwestern United States to Costa Rica.
480 University of Kansas Publs., Mrs. Nat. Hist.
Ptilogonys cinereus cinereus Swainson Ashy Ptilogonj^s
Ptilogonys cinereus cinereus Swainson, Cat. Bullock's Mex. Mus., App. 4, 1824.
Diagnosis.— Colovsition of adult male: Frontals, supralorals, malars, and chin White; orbital ring White; auriculars and nape grayish brown; rest of head smoke gray; back, scapulars, wing coverts, rump, and upper tail coverts plain Bluish Black; rectrices (except middle pair) with large patch of White midway between base and tip, rest plain Bluish Black; chest, breast, and anterior parts of sides plain Bluish Gray-Green, much lighter than back, and fading into paler Gray on throat; abdomen and thighs White; flanks and posterior part of sides Olive-Yellow or Yellowish Olive; under tail coverts Lemon Yellow; bill, legs and feet Black. Coloration of adult females: Head plain Smoke Gray, passing into White on frontals, malars, and chin; back, scapulars, wing coverts, and rump Hair Brown; upper tail coverts Dark Gull Gray; remiges and rectrices Black with faint Dusky Green gloss, edged with Gull Gray; chest Dark Grayish Brown lightening to Wood Brown on sides and flanks; abdomen White; under tail coverts Yellow Ocher. Coloration of young : As in adult female, but paler throughout.
Meas^lrements.—Jn adult male, wing 94.0. and tail 104.2: in adult female, wing 93.3, and tail 94.8; both sexes, culmen 11.1, and tarsus 18.7.
Range. — Mountainous districts of central and southern Mexico, in states of Durango, Zacatecas, Hidalgo, Mexico, Oaxaca, Colima, Morelos, Veracruz, San Luis Potosi, Guerrero and Michoacan.
Ptilogonys cinereus molybdophanes Ridgway Ashy Ptilogonys
Ptilogonys cinereus molybdophanes Ridgway, Man. N. American Birds, 464 (footnote), 1887.
Diag7iosis. — Coloration of adult male: Upper parts darker bluish than in P. c. cinereus; venter paler; flanks Olive-Green rather than Olive as in P. c. cinereus. Coloration of adult female : Like female of P. c. cinereus but colors darker throughout; dorsum more olivaceous.
Measurements. — In adult male, wing 89.4, and tail 97.1 ; in adult female, wing 89.4, and tail 93.3; both sexes, culmen 11.7, and tarsus 17.3.
Range. — Western Guatemala, in subtropical and temperate zones.
Ptilogonys caudatus Cabanis Costa Rican Ptilogonys
Ptilogonys cuudatus Cabanis, Jour, fiir Orn., 1866:402, Nov. 1866.
Diagnosis. — Coloration of adult male : Forehead and crown Pale Grayish Blue, slightly paler anteriorly; orbital ring Lemon Yellow; rest of head and neck, including crest, Olive-Yellow; throat paler and tinged with Light Gull Gray; back, scapulars, rump, upper tail coverts and wing coverts uniform Bluish Slate-Black; chest and breast similar but paler; sides and flanks Yellowish Olive-Green; thighs, lower abdomen, and under tail coverts Lemon Yellow; remiges, primary coverts, and tail Black, glossed with Bluish Black and edged with Gull Gray; inner webs of rectrices (except two middle pair)
Arvey: Waxwings and Allied Birds 481
with large middle patch of White; bill, legs, and feet Black. Coloration of adult female : Forehead and crown Pale Gull Gray, becoming paler anteriorly ; rest of head, together with neck, back, scapulars, rump, and wing coverts plain Yellowish Olive Green; chest and breast similar but more grayish; lower abdomen and flanks White tinged with Yellowish Olive; under tail coverts Olive-Gray ; remiges, primary coverts, and rectrices Black with Gull Gray edges. Coloration of young: Dorsum plain Light Grayish Olive; upper tail coverts Brownish Olive ; underparts Grayish Olive anteriorly, becoming more Yellowish Olive on abdomen; under tail coverts pale Yellowish Olive with Grayish Olive base; bill and feet Brownish Drab.
Measureemnts. — In adult male, wing 96.2, and tail 135.7; in adult female, wing 93.9, and tail 113.7; both sexes, culmen 12.6, and tarsus 19.1.
Range. — Highlands of Costa Rica and extreme western Panama.
Genus Phainopepla Sclater
Phainopepla Sclater, Proc. Zool. Soc. London, 26:543, 1858. Type Phainopepla nitena (Swainson).
Diagnosis. — Tail almost as long as wing ; head with pointed crest of narrow, separated feathers ; rectrices without white ; bill narrow, compressed terminally ; conspicuous white patch under wing; nostril small, exposed; rictal bristles dis- tinct; tail slightly rounded.
Phainopepla nitens nitens (Swainson) Phainopepla
Phainopepla nitena nitens (Swainson), Anim. in Menag., 1838:285, Dec. 31, 1837.
Diagnosis. — Coloration of adult male: Uniform glossy Bluish Black; inner webs of primaries except innermost pair with middle portion White; bill, legs, and feet Black. Coloration of adult female : Plain Olivaceous Black, longer feathers of crest Black, edged with Gull Gray; remiges and rectrices Dusky Drab to Black; rectrices and coverts margined by White; bill and feet Brown- ish Drab to Dusky Brown. Coloration of young : Like adult female but more Brownish Drab.
Measurements. — No specimens examined; larger than P. n. lepida (Van Tyne, 1925).
Range. — Central and southern Me.xico, in states of Coahuila, San Luis Po- tosi, Durango, Guanajuato, Mexico, Puebia, and Veracruz.
Phainopepla nitens lepida Van Tyne Phainopepla
Phainopepla nitens lepida Van Tyne, Occ. Pap. Bost. Soc. Nat. Hist., 5 :149, 1925.
Diagnosis. — Coloration same as P. n. nitens; separated by smaller size.
Measurements. — Wing 91.0, tail 90.3, culmen 11.5, tarsus 17.6.
Range. — Southwestern United States, from central California, southern Utah, and central western Texas southward to Cape San Lucas in Baja California, and into northwestern Mexico (Sonora and Chihuahua).
482 University of Kansas Publs., Mus. Nat. Hist.
Subfamily Bombycillinae
Diagnosis. — Wings long and pointed, reaching almost to tip of tail; first pri- mary spurious; second primary longest; tail short and even; rictal vibrissae few and short; secondaries generally, and sometimes also rectrices, tipped with red, corneous appendages; nasal fossa partly filled with short, antrorse, close-set velvety feathers; plumage soft, silky; tail tipped with yellow band (red in B. japonica) ; sexes alike; humerus short with large external condyle; caudal muscles and pygostyle not well developed ; bill flared widely at base ; one genus, three species.
Range of subfamily. — Holarctic breeding area; wanders nomadically south in winter to Central America and West Indies, southern Europe and Asia.
Genus Bombycilla Brisson
Bombycilla Brisson, Orn. ii, 1760:337. Type Bombycilla garrula (Linnaeus).
Diagnosis. — As described for the subfamily.
Bombycilla cedrorum Vieillot Cedar Waxwing
Bombycilla cedrorum Vieillot, Hist. Nat. Amer., 1 :88, Sept. 1, 1807
Diagnosis. — Coloration of adults: Shading from Saccardo's Umber on dorsum to Bister on top of head; upper tail coverts and proximal rectrices Gull Gray; underparts shade through pale Lemon Yellow wash on belly into White on under tail coverts; forehead, lores, and eye-stripe Black; chin same, soon shading into Blackish Mouse Gray and into color of breast; side of under jaw with sharp White line; narrow line bordering forehead, and lores. White; lower eyelid White; quills of remiges Dark Mouse Gray, darkening at tips; inner quills tipped with red horny wax appendages; tail feathers like primaries, but tipped with Lemon Yellow, and occasionally showing also red homy wax appendages; bill and feet Black. Coloration of young: Dorsum as in adult, but lightly streaked with White ; head concolor with dorsum ; forehead White ; lores Black; eye stripe Black anterior to eye and White posterior to eye; throat Light Buff; belly with alternate streaks of Dresden Brown and light Ochraceous Buff but posteriorly White; tail tipped with Lemon Yellow bar; bill black at tip, shading to Sepia at base.
Measurements. — Wing 92.9, tail 55.5, culmen 10.9, tarsus 16.8.
Range. — Breeds from central British Columbia, central Alberta and Mani- toba, northern Ontario, southern Quebec and Cape Breton Island south to northwestern California, northern New Mexico, Kansas, northern Arkansas, North Carolina, and northern Georgia. Winters south to Louisiana, Missis- sippi, Texas, Arizona, Colorado, Florida, Honduras, Costa Rica, Jamaica, Little Cayman Island, Haiti, and Panama.
Bombycilla garrula (Linnaeus) Bohemian Waxwing Bombycilla garrula (Linnaeus), Syst. Nat., 10th Ed., 1758:55.
Diagnosis. — Coloration of adults: General color Olive-Brown, shading in- sensibly from clear Smoke Gray of upper tail covert.s and nimp to Cinnamon- Drab anteriorly, heightening on head and forehead to Hazel; narrow frontal line, lores, broader mask through eye, chin, and upper throat. Sooty Black; under tail-coverts Cinnamon-Brown; tail Smoke Gray, deepening to Blackish
Arvey: Waxwings and Allied Birds 483
Mouse Gray distally, and tipped with Lemon Yellow; wings Blackish Mouse Gray; primaries tipped with sharp spaces of Lemon Yellow or White, or both; secondaries with White spaces at ends of outer web, shafts usually ending with enlarged, horny red appendages; primary coverts tipped with White; bill Blackish Slate and paler at base; feet Black. Coloration of young: Much like adult, but general color duller; some streaking on venter and back; chin, throat, and malar region dull White. Three subspecies.
Bombycilla garrula garrula (Linnaeus) "
Bohemian Waxwing
Bombycilla garrula garrula (Linnaeus), Syst. Nat., 10th Ed., 1758:55.
Diagnosis. — Coloration: As described for the species, but darkest of the three subspecies; tending to be more Vinaceous dorsally than either pallidiceps or centralasiae.
Measurements. — Wing 113.5, tail 63.1, culmen 12.5, tarsus 20.7.
Range. — Europe; breeds north to northern Russia and Norway, south to about 65° N latitude ; winters south to England and Ireland, southern France, northern Italy, and Turkey.
Bombycilla garrula centralasiae Poljakov Bohemian Waxwing
Bombycilla garrula centralasiae Poljakov, Mess. Om. vi :137, 1915.
Diagnosis. — Coloration: As described for the subspecies garrula, but less Vinaceous dorsally, and more Cinnamon; venter lighter gray than garrula, and much paler than pallidiceps.
Measurements. — Wing 114.7, tail 63.0, culmen 12.2, tarsus 21.0.
Range. — Asia; breeds northern Siberia south to Vladivostok; winters to Turkestan and central eastern China and Japan.
Bombycilla garrula pallidiceps Reichenow Bohemian Waxwing
Bombycilla garrula pallidiceps Reichenow, Orn. Monats. 16 :191, 1908.
Diagnosis. — Coloration: As described for the species, but more grayish above and below than B. g. garrula; darker gray than in centralasiae.
Measurements. — Wing 115.1, tail 71.7, culmen 12.6, tarsus 21.1.
Range. — Breeds from western Alaska to northern Mackenzie and north- western Manitoba south to southern British Columbia, southern Alberta, northern Idaho, and possibly Colorado (Bergtold 1924) and Montana (Bur- leigh 1929) ; winters east to Nova Scotia and irregularly over much of Canada, and south irregularly to Penns^-lvania, Ohio, Michigan, Indiana, Kansas, Colo- rado, California, Arizona, and Texas.
Bombycilla japonica (Siebold) Japanese Waxwing
Bombycilla japonica (Siebold), Nat. Hist. Jap., St. No. 2:87, 1824.
Diagnosis. — Coloration: Dorsum generally Brownish Drab shading to Light Brownish Drab on lower back, rump, and upper tail coverts; secondary and tertiary coverts Pale Brownish Drab, washed on outer web with Carmine;
484 University of Kansas Publs., Mus. Nat. Hist.
primary coverts Blackish Slate, with White edging; tail feathers Slate-Gray, broadly tipped with Carmine, bordered anteriorly by subterminal Black bar; head crested, forehead Chestnut; lores, frontals, and stripe extending around eye and nape, Black; throat Black, narrowing on lower throat; breast, sides of flanks Light Drab; venter pale Sulphur Yellow; thighs Brownish Drab; under tail coverts Carmine; bill, legs, and feet Black.
Measurements. — Wing 108.3, tail 53.6, culmen 11.2, tarsus 19.4.
Range. — Breeds eastern Siberia, northern China; winters south in China, and to Japan (Hokkaido, Kyushu), Taiwan, and Korea.
Subfamily Dulinae
Diagnosis. — Bill deep and compressed, culmen strongly depressed; nostrils circular, wholly exposed; tail even, and shorter than wing; tenth primary less than half length of ninth; under parts streaked; plumage hard and harsh; rictal bristles minute; wing rounded; humerus long and with small external condyle; pygostyle and caudal muscles not well developed; one genus, one species. ' Range of subjamily. — Islands of Haiti and Gonave, Greater Antilles.
Genus Dulus Vieillot
Dulus Vieillot, Analyse, 1816:42.
Diagnosis. — Like the subfamily.
Dulus dominicus dominicus (Linnaeus) Palm-chat
Dulus dommicus dominicus (Linnaeus), Syst. Nat., 12th Ed., 1766 :316.
Diagnosis. — Coloration: Dorsum Olive, back, scapulars, and wing coverts more Brownish Olive; lower rump and upper tail coverts Olive-Green; pileum and hindneck with indistinct streaks of Brownish Olive; tail Brownish Drab, edged with Light Olive Gray; lores, suborbital region, and auricular regions Dusky Brown; malars Dusky Brown and streaked with Sooty Black, streaka narrower on abdomen, broader and paler on under tail coverts, bill Light Brownish Drab; legs and feet Brownish Drab.
Measurements. — Wing 85.0, tail 68.8, culmen 15.0, tarsus 24.7.
Range. — Island of Haiti, Greater Antilles.
Dulus dominicus oviedo Wetmore Palm-chat
Dulus dominicus oviedo Wetmore, Proc. Biol. Soc. Wash., 42:117, 1929.
Diagnosis. — Coloration: Like D. d. dominicus, but averaging more Grayish Olive ; rump and tail coverts with less greenish wash.
Measurements. — Wing 90.1, tail 71.3, culmen 16.2, tarsus 25.1. Range. — Gonave Island, off Haiti, Greater Antilles.
Arvey: Waxwings and Allied Birds 485
COLORATION
The general coloration of waxwings is cryptic, that is to say, con- cealing or blending. The lighter color of the venter, especially of the belly, contrasts with the duller, darker vinaceous color of the dorsum. Several ruptive marks tend to obliterate the outline of the body. The crest of the head, when elevated, tends to elongate the body, making the outline less like that of a normal bird. The facial mask effectively breaks up the outline of the head, and con- ceals the bright eye, which would otherwise be strikingly distinct. The white spots on the distal ends of the secondaries of B. garrula and the yellow color on the distal ends of the rectrices (red in B. japonica) are also ruptive. These ruptive marks on an otherwise blending type of plumage might be important to waxwings, and probably are more effective when the birds remain motionless in either a well-lighted area or in one that is partly in shadow, rather than in one that is wholly in shadow.
The red wax tips on the secondaries of the flight feathers, and sometimes found on the ends of the rectrices in Bombycilla, are puzzling and no wholly convincing reason has been suggested for their occurrence. Two instances are known of yellow instead of red-colored wax tips in B. cedrorum (Farley, 1924). It is well known that many individuals, especially of B. cedrorum, do not possess these tips; they are absent in a smaller proportion of indi- viduals of B. garrula. Of the 53 skins of B. cedrorum available in the University of Kansas Museum of Natural History, which might be taken as a sampling at random of the general population of this species, only 17 possess wax tips. A few specimens are unilateral, and the tips are of varying sizes in different individuals. Of these 17 birds, 6 are female and 7 male, the others being unsexed at the time of skinning. This proportion is, roughly, half and half. Of the seven skins of B. garnda pallidiceps in the same Museum, five possess the tips, and two that are females have no trace of the red tips at all. Of the five which do have the tips, two are males, two are females, and one is unsexed. In a series of 13 specimens of the three subspecies of B. garnda, loaned by the United States Na- tional Museum, all but two individuals possess the tips on the sec- ondaries, and, in addition, four specimens, equally divided between the two sexes, have color on the rachis of some rectrices, and small appendages of pigment extend beyond the feathers. Stevenson (1882) found that among 144 specimens of B. garrula garrula killed by storms in England in the winter of 1866-67, 69 individuals had
486 University of Kansas Publs., Mrs. Nat. Hist.
wax tips. Of these, 41 were males and 27 were females; the remain- ing one was of uncertain sex. Among 38 definitely sexed B. garrula pallidiceps in the California Museum of Vertebrate Zoology, Swarth (1922:276) lists tips in 22 males and 16 females. These data indi- cate that the proportion of birds with the wax tips is higher in B. garrula than in B. cedrorum. The potentiality for wax tips is pos- sibly inherited according to Mendelian ratio.
Bombycilla japonica is of interest in that the adults, at least, sel- dom have the waxy appendages. Nevertheless, in the specimens observed, the entire distal ends of the feathers normally possessing the tips in other species are suffused with red color. This may be the original condition of all waxwings, or perhaps, instead, this spe- cies is in a transitional stage in the development of the tips. Swarth (1922:277) says concerning the probable derivation of the wax tips in B. garrula (and in B. cedrorum) : ''the ornamentation, in fact, may well have begun with the coloring of the shaft, spreading later over adjoining feather barbs. The last stage would have been the coalescing of the barbs, forming the waxlike scale as is now seen. Various steps of this hypothetical development are supplied in the wing and tail feathers of different birds of this series." Bombycilla japonica thus may be close to the ancestral condition in the waxwing stock in the development of the waxy appendage.
The rectrices of all three species of waxwings seldom possess the wax tips, unless the secondaries have the maximum number of tips. In these individuals, the pigment seems to "spill over" onto the tail feathers. Eight is the maximum number of tips found on the second- aries. Rectrices with wax tips are more frequently found in B. gar- rula, and only occasionally in B. cedorum. The pigment in the tip of the tail of B. japonica is red rather than yellow as it is in the other two species, and some individuals of the Japanese Waxwing show a slight amount of coalescence of wax in the tail feathers as well as in the secondaries.
If the tips were present in all members of the two species, it could be postulated, in line with recent investigational work by Tinbergen (1947), that the tips are in the nature of species "releasers," facili- tating species recognition. Such recognition is now regarded as of prime importance in the formation of species. It is improbable that sex recognition may be aided, as there is no evidence to indicate that the tips are found predominantly in either sex.
The wax tips are not limited to the adult birds in the species B. garrula. Swarth {op. cit.) mentions the capture of several youn^^
Arvey: Waxwings and Allied Birds 487
Bohemian Waxwings, and describes them as "possessing all the dis- tinctive markings of the most highly developed adult." This in- cludes wax appendages, and several citations are given (Wolley 1857, Gould 1862) to indicate that this is the rule rather than the exception, not only for the American subspecies pallidiceps, but at least for the European subspecies garrula as well. On the other hand, the young of B. cedrorum lack the wax tips, at least as far as available data show.
Some characteristics of living animals are of the "relict" type; that is to say, they were developed in ancient times when some unknown ecological factor was operative which is no longer demon- strable, and the characteristic is now neutral or at least not detrimental, although of no positive value to the organism. Possibly the wax tips of waxwings are thus to be explained. I am more in- clined to the opinion that the wax tips are adaptations to present-day ecological conditions for the birds.
The wax tips are ruptive in effect, bince the birds, especially in winter, are habitues of bushes and trees that have berries, and the tips, on the otherwise dull body, suggest berries. The red tips tend further to disrupt the body outline at the midline, or slightly pos- terior to this. Perhaps the wax tips on the rectrices emphasize the end of the tail, the region of the body that is the least vital and that may be expendable in times of pursuit by an enemy.
Any characteristic is of survival value to an organism if in any way the characteristic enhances the chances of survival up to the time when the organism can successfully raise even a few young to maturity. If that character, as for example, the red wax tips on the secondaries, helps to maintain the individual until it can raise to independence a greater number than merely a few young, such a character can be said to be of greater survival value. The character may be effective for a brief period of time and may be uncommon ; it might be effective for a split second in time, and only at a particular stage in the life history.
The winter period probably is the most hazardous for waxwings, in that they then depend at times upon long flights to find food. The food is vegetable, and thus is comparatively low in food value; the birds must ingest large quantities of berries or dried fruits to maintain themselves. In winter, in northern latitudes at least, predators are more apt to prey upon those species which, like wax- wings, do not migrate south. The winter months are those in which waxwings frequent berry bushes, and it may well be that in these
488 University of Kansas Publs., Mus. Nat. Hist.
months, the wax tips that appear like berries, are especially valuable to the birds, and operate selectively.
It is suggested, therefore, that the wax tips are of positive value to waxwings, rather than being relict characters. Coalescence of pig- ment has taken place in the formation of the wax tips. B. japonica is closer to the ancestral stock insofar as wax tips are concerned, and generally lacks the tips. B. cedrorum has the tips in approxi- mately half of the adults, and not at all in the young. B. garrula has the tips in almost all the adults, and in a like proportion of the young, and probably has evolved further in the development and retention of the wax tips than has either of the other two species.
The streaked plumage of Dulus is decidedly generalized, and is probably more nearly like the color of the ancestral stock. In this connection it is notable that young Cedar Waxwings are streaked, and young Bohemian Waxwings are streaked to a lesser degree. This streaking is apparently a recapitulation of the feather color of the stock. Perhaps the color of Dulus has not changed, as the streaking would not be a disadvantage to the birds in their environment of light and shadow. In joining together in groups and in the construction of large communal nests, Dulus has evidently gained sufficient pro- tection against predators; other birds solve this problem by modify- ing their coloration.
Ptilogonys is ruptively colored, but in a different fashion than Bomhycilla. The tail markings, the distinct yellow on the under tail coverts, the sharply marked pileum, are all examples of ruptive coloration. The generally lighter venter (especially under tail coverts), the crest that may be elevated, and the generally drab bluish dorsum, are cryptic and serve to hide the animal insofar as is possible considering its habits. The very conspicuous coloration of the male, in contrast to the more drab color of the female, how- ever, would lead one to believe that in Ptilogonys, following the pat- tern of many passerine birds, the male leads a predator from the nest, leaving the drab female to incubate the eggs, and thus pre- serve the young.
It is difficult to suggest reasons for the brilliant coloration of the male Phainopepla, unless it is for decoying predators away from the nest. Possibly some birds survive not because of, but in spite of, their coloration, and Phainopepla may be a case of this sort. Anyone who has observed Phainopepla in life will agree, certainly, that the male makes no attempt at concealment, and flaunts his color to all comers.
Arvey: Waxwings and Allied Birds 489
The coloration of Phainoptila, in contrast to Phainopepla, is much more plain, and is suited to its habits of brush dwelling ; in a brush habitat the drab coloration is difficult to detect. The Yellowish Olive under tail-coverts and the Olivaceous dorsum are all evidences of cryptic coloration, and undoubtedly, this bird depends upon hiding for escape from its enemies, since it is a bird of the dense forest cover.
Coloration, which varies relatively rapidly in response to differing ecological conditions, has become more different in the species of Bombycillidae than is true in many other families of passerine birds. The explanation lies in early geographical isolation of the three subfamilies, with consequent radiation in three directions. Wax- wings have become adapted by possessing a thick protective layer of feathers and drab coloration broken by ruptive marks. They still retain the streaked plumage, which is probably ancestral, in the juveniles; this is lost at the first molt in the fall. In its evolution. Dulus has developed large feet, heav\' dccurved beak, and the large communal nest that affords protection from enemies; as a conse- quence, perhaps Dulus did not need a plumage different from the primitive and streaked one. The survival of Dulus may not have depended on either ruptive marks or on brilliant and outstanding plumage. The large feet and large bill seem to be responses to par- ticular ecological requirements, as will be shown later.
The Ptilogonatinae, with habits paralleling those of the flycatch- ers, probably are considerably modified from the ancestral stock; the coloration probably is more brilliant and conspicuous. Perhaps this type of coloration and the habit of capturing insects from a perch are correlated. Some amount of territoriality is characteristic of this subfamily and dimorphism in color — the plumage of the male is outstandingly conspicuous — possibly is of selective value to the race. In a tropical forest community, a duller pattern possibly would be more visible and thus would be selectively disadvantageous.
COURTSHIP
Waxwings are gregarious birds and individuals establish no well- defined territories as do many birds. The nest itself is the only defended territory, and as Crouch (1936) has shown, the Cedar Waxwing will nest in close proximity to others of the same species. Swarth (1932:275) mentions that the Bohemian Waxwing is toler- ant of the nests of other pairs near by. The extreme condition is that found in Dulus, in which the territory is not limited even to
2—1019
490 University of Kansas Publs., Mus. Nat. Hist.
the nest, but to the individual compartment of the community nest. Phainopepla, a less gregarious bird than Dulus and waxwings, has a much more definite territory, although individuals of Phainopepla are tolerant of others of the same species; no feeding territory is established, and small flocks of birds feed together at any time of the year.
In birds whose territories lack well-defined boundaries, it would be expected that elaborate song would not have evolved, and that most of the recognition of kind and sex would be dependent upon the behavior of the birds. This is the fact; song, as such, is lacking in the three subfamilies Bombycillinae, Ptilogonatinae, and Dulinae. Waxwings utter (1) notes that serve to keep the flock together, (2) calls used by the young in begging for food, and (3) some low notes that Crouch [op. cit.:2) considered as possibly concerned with court- ship. Phainopepla has various call notes, and in addition, a suc- cession of notes which are run together. Ptilogonijs utters a note which Skutch (MS) characterizes as a loud, not unmusical "tu- whip" that is used as the birds "fly in straggling parties which keep in contact by their constant chatter." Dulus is described by Wet- more and Swales (1931:349) as having only a variety of rather harsh chattering notes in chorus.
The most notable behavior pattern associated with courtship in Waxwings, in the absence of song, is the so-called "mating dance" described by Crouch (1936), and observed by me in Lawrence, Kan- sas, in the spring of 1948. This consists of one bird of a pair (pre- sumably the male) hopping along a branch toward the other bird (the female), then away again, repeating the procedure for some little time. The female remains motionless until, as the male ap- proaches, mutual fondling of the head and neck feathers takes place, or the birds may peck at each other's bill. A berry may be passed from bill to bill, although generally the berry is not utilized for food, and this can be interpreted as a nervous reaction of the birds. It may be an instance of "false feeding" as is seen in many birds, in which the female begs for food, as a nestling would beg, as a preliminary to the sexual act. I am of the opinion that these re- actions are in the nature of behavioristic patterns that bring the birds into the emotional balance for copulation, as copulation fol- lows the "dance." Sometimes, however, copulation is preceded by a "nuptial flight" around the nesting area, at which time the birds utter loud calls. Armstrong (1924:183) is of the same opinion, citing numerous instances in which nuptial flights and elaborate
Arvey: Waxwings and Allied Birds 491
displays have evolved for just this purpose. The birds are then in the proper physiological balance to initiate the complicated sequence of copulation, nesting, incubation, feeding, and brooding of the young.
It would be valuable to know more concerning the life histories of the other birds considered in this paper, since behavior is inherent, and probably can be cited as evidence of close relationship or the opposite. All that I have been able to learn is that Phainopepla has a nuptial flight in which the male chases the female, and that Dulus (Wetmore and Swales, 1931:347) seeks the company of others of its kind at all times, and that two birds, presumably paired, will sidle up to one another when they are perched.
NEST BUILDING
There are numerous papers concerning the nesting of waxwings. B. garrula, owing to its nesting in the far north, where observers are few, has received less attention than B. cedrorum. There is, on the other hand, no literature that deals with the nesting habits of the majority of the Ptilogonatines, with the exception of Phaino- pepla, on which there is considerable literature (Merriam, 1896; Myers, 1907, 1908). No detailed study of the nesting of Dulus has been reported, although Wetmore and Swales (1931) have described carefully the large communal nest of this genus.
In Bomhycilla, both members of a pair apparently aid in the construction of the nest (Crouch, 1936; Swarth, 1932). Although the sexes are alike in plumage and general appearance, most stu- dents of the nesting of waxwings agree that one bird, assumed to be the female, does most of the arranging of the material, and does the shaping of the nest, whereas both birds carry materials to the nest site. As is characteristic of many passerine birds, both members of the pair gather materials and fly back to the nest site, where the female takes the more active part in the construction of the nest itself.
Both species of American waxwings build bulky nests, with the base or platform composed of a large amount of twigs and sticks, from which there often trails a mass of sticks and moss or string. Softer materials such as moss, plant fibers, and string, are placed inside the platform; moss is readily available to, and preferred by, B. garrula according to Swarth [op. cit. :211), and various plant fibers and string are used by B. cedrorum. The inner lining consists of soft plant fibers or down, dry grasses, and feathers. The nest is usually unconcealed in a tree either adjacent to a trunk or on a main
492 University of Kansas Publs., Mus. Nat. Hist,
side branch, but sometimes in a fork. Nest building by both Cedar and Bohemian waxwings is rapid, taking from three to five days, and is followed immediately by egg laying.
Nesting by waxwings is late in the season; June is the month in which the nest is usually started. This is readily explainable in Bohemian Waxwings, since adverse weather would prohibit earlier nesting in the area in which they spend the summer. Crouch (op. cit.:l) remarks that B. cedronim possibly evolved in the far north where it was impossible for it to start nesting earlier, and that the habit has been retained. Perhaps, on the other hand, nesting is de- layed until the berry crop is ripe, to insure sufficient food for the young.
Desertion of the nest is not uncommon in waxwings, despite the tolerance to other animals that is shown by the birds. A new nest may suddenly be begun before the first one is finished, and all the materials from the first nest may be removed, or the nest may be abandoned before it is completed. The eggs may be left at any time up to hatching, and the young may be deserted, especially in the earlier stages of development.
The very large and bulky communal nest of Dulus is not radically different from the nest of waxwings. In the absence of sufficient nesting sites, a pair of gregarious birds such as Dulus could com- bine their nest with those of other pairs, retaining for their own territory only the nest cavity, and in this way communal nests might have evolved. The nest of Dulus is communal probably because of the lack of suitable trees for nesting sites, and only inci- dentally does this type of nest afford better protection from natural marauders. Large numbers of Palm-chats work together in the construction of the nest platform, and both sexes probably take part in the work.
In Phainopepla the nest is built mostly by the male (Merriam, 1896; Myers, 1908), although the female does some of the work, especially in the shaping and lining of the nest. In this genus, the nest is usually a compact structure, but exceptional nests are of con- siderable bulk. The nest is commonly placed in a fork near the main trunk of a tree, in a conspicuous location, and generally is 10 to 20 feet from the ground. In shape and location, the nest closely cor- responds to that of Bombycilla, but the materials used for a base are stems of annual plants, whereas Bombycilla uses more woody twigs. The finer materials used by Phainopepla are more readily obtainable in the ecological association inhabited by Phainopepla than would be heavier twigs such as Bombycilla uses.
Arvey: Waxwings and Allied Birds 493
FOOD
Waxwings are typically frugivorous; berries are the staple food. The birds are known to catch insects, especially in the spring and summer, and their insect gathering technique has been likened to that of Tyrannid flycatchers. Nice (1941) experimented with a young captive Cedar Waxwing and found that it had a decided preference for red or blue berries, and that meal worms were utilized as food only when the birds became educated by other captive birds of other species as to the food value of the worms. Post (1916) indicates that the food given to the nestlings of Cedar Waxwings is entirely animal for the first three days, and that a mixed diet of berries and insects is subsequently offered.
In feeding of the young, regurgitation of partly digested food does not take place, according to Wheelock (1905). Rather, the adults "store" food in the form of berries in the expanded esophagus or crop, feeding them whole to the young. Digestion is an unusually rapid process, involving merely minutes for the passage of berries and cherries. This is correlated with a short intestinal tract, which is unusual for a frugivorous bird. Nice's (1940) experiments with Cedar Waxwings revealed that cherries would pass through the digestive tract in 20 minutes, blueberries in 28 minutes, and choke- cherries in 40 minutes. Heinroth (1924) states that berries pass through the digestive tract of Bohemian Waxwings in the space of a "few minutes." This rapid digestion is obviously adaptive, since the value of the food is slight and therefore large quantities of it must be ingested; the large seeds would hamper further ingestion until they were eliminated, since they seem not to be regurgitated.
Members of the subfamily Ptilogonatinae are both insectivorous and frugivorous insofar as available data show, although again there is relatively little information available concerning them. Skutch (MS) has found that the Guatemalan Ptilogonys cinereus catches insects by repeated sallies into the air from a perch, after the man- ner of flycatchers. He notes also that the birds feed on berries of Eurya theoides and Monnina xalapensis. It is well known that Phainopepla catches insects when these are available, and its liking for berries is so apparent that in parts of its range, it is known as the "pepper bird," since it frequents pepper trees {Schinus molle) and feeds on the small red berries. The preserved specimens of Ptilogonys and Phainoptila available for this study contain only berries in the digestive tract. Didus feeds mostly, if not wholly, on plant food. According to Wetmore and Swales (1931:349), berries, fruits, and parts of flowers are eaten.
494 University of Kansas Publs., Mus. Nat. Hist.
SKELETON
A critical analysis of the skeletons provides evidence that aids the student in estimating which differences are merely the result of habits developed in relatively recent geological time as opposed to those which owe their existence to more ancient heritage. Stresses caused by the action of different sets of muscles can apparently stimulate changes in bones to meet new needs, and the evidence from genetics is that such mutations in wild birds are minute and cumu- lative, rather than of large degree and of sudden appearance. Once adaptive mutations have occurred, if genetic isolation from one source or another accompanies it, a new population different from the parental stock may become established. Study of the skeleton of any species of living bird may indicate those characters identifi- able as modifications fitting it to a particular environment. If no distinguishing characters are discovered that may be attributed to environmental factors, such a species can be spoken of as general- ized; the inference then is that such a species is not modified for a single, particular ecological niche.
Some parts of the skeleton, obviously, are more adaptable or plastic than others. The beak seems to be the most adaptable part. Probably this results from its frequent use; it is the part of the bird to capture the food. The long bones, meeting the environment as legs which serve as landing mechanisms or as locomotory ap- pendages, and as wings which provide considerable locomotion for most birds, probably come next in order as regards plasticity. In these parts, then, one may look for the most change in birds, which, within relatively recent geologic times, have been modified to fit a particular set of conditions. From the beak and long bones of a species in which habits are unknown, one can infer the habits and habitat from a comparison with the skeletal features of species of known habits.
Skull. — The skulls in all three subfamilies have essentially the same general appearance and structure, the most marked differences being, as would be expected, in the bills and associated bones.
The most specialized bill is to be found in Dulus; its bill is de- curved, and the associated bones are correspondingly changed for support of the bill. For example, the palatines and "vomer" are much wider, the palatines are more concave from below and have longer posterior processes than the corresponding bones in Bombycilla. Moreover, the "vomer" in Dulus and in Phainoptila is larger and heavier than in Bombycilla, and the quadrate and pterygoid bones are relatively large for support of the beak. The palatines, however.
Arvey: Waxwings and Allied Birds
495
are weak in Phainoptila. In the Ptilogonatinae, with the exception of Phainoptila, the wings of the palatines flare more than in Bomby- cilla, but not to the extent that they do in Dulus, nor does the pala- tine bone present a concave appearance in the Ptilogonatinae. The premaxilla is a relatively weak bone in Bombycilla and Phainopepla, stronger in Ptilogonys, and is notably heavy in Phainoptila and Dulus, and in these latter two genera shows a sharply-ridged to- mium. The maxillae connect to somewhat widened nasal and naso- lateral processes in all the genera, and the premaxillae narrow abruptly from this point forward. In the family, Phainopepla and Phainoptila show the least flaring in this region.
1-
Skulls in lateral view of five genera of Bombycillidae. Natural
Figs. size.
1. Phainoptila m. melanoxantha, sex?, MNH no. 26493, 15 mi. SE Cartage, Costa Rica.
2. Ptilogonys caudatus, male, MNH no. 24492, 15 mi. SE Cartago, Costa Rica.
3. Phainopepla nitens, male, MNH no. 24752, Pima Co., Arizona.
4. Ptilogonys cincreus, female, Louisiana State University no. 297, Xilitla Region, San Luis Potosi, Mexico.
5. Duhu^ dominicus, female, USNM no. 292652, Don Don, Haiti.
6. Bombycilla cedrorum, male, MNH no. 15331, Bexar Co., Texas.
7. Bombycilla garrula, sex?, USNM no. 223S95, Bozeman, Montana.
496 University of Kansas Publs., Mus. Nat. Hist.
Figs. 8-14. Skulls in ventral view of five genera of Bombycillidae. Natural size.
8. Phainoptila m. melanoxantha, sex?, MNH no. 26492, 15 mi. SE Cartago, Costa Rica.
9. Ptilogonys caudattts, male, MNH no. 24492, 15 mi. SE Cartago, Costa Rica.
10. Phainopepla nitens, male, MNH no. 24754, Pima Co., Arizona.
11. Ptilogonys cinereus, female, Louisiana State University no 297, Xilitla Region, San Luis Potosi, Mexico.
12. Dulus dominicus, female, USNM no. 292652, Don Don, Haiti.
13. Bombycilla cedrorum, male, MNH no. 15331, Bexar Co., Texas.
14. Bombycilla garrula, sex?, USNM no. 223895, Bozeman, Montana.
Arvey: Waxwings and Allied Birds
497
Figs. 15-21. Skulls in dorsal view of five genera of Bombycillidae. Natural
size.
15. Phainoptila m. melanoxantha, sex?, MNH no. 26493, 15 mi. SE Cartago, Costa Rica.
16. Ptilogonys caudatus, male, MNH no. 24492, 15 mi. SE Cartago, Costa Rica.
17. Phainopepla nitens, male, MNH no. 24752, Pima Co., Arizona.
18. Ptilogonys cincreus, female, Louisiana State University no. 297, Xilitla Region, San Luis Potosi, Mexico.
19. Duhis dominicus, female, USNM no. 292642, Don Don, Haiti.
20. Bomhycilla cedrorum, male, MNH no. 15331, Bexar Co., Texas.
21. Bomhycilla garrula, sex?, USNM no. 223895, Bozeman, Montana.
498 University of Kansas Publs., Mus. Nat. Hist.
This flaring, immediately lateral to the antorbital plate, is com- mon to all Bombycillids and constitutes a major skeletal character- istic useful for recognition of the members of the family, since the swelling is easily discernible both externally and on the cleaned skulls. In Phainopepla there is much variability in this character; some specimens have a narrower antorbital bridge than others. Only one skeleton of Phainopepla n. nitens was available. The flaring in the skull of this specimen is identical with that in Ptilagonys. Among the skulls of P. n. lepida in the University of Kansas Mu- seum of Natural History, is No. 19228, a juvenile, taken 5 miles south of Tucson, Arizona. In this specimen, the flaring in the antorbital region is clearly evident and equal in amount to that in skulls of P. n. nitens, but the bird had not attained full skeletal growth. However, the flaring of the antorbital region appears to be common in the nestlings of many species of passerine birds. Other specimens of the subspecies lepida show a varying amount of flaring, the least (in the series available) being in No. 24754, MNH, in which the proportion of the skull (length divided by width) closely corresponds to that in Phain<yptila; the skull of No. 24754 is long and thin, and the base of the bill is only slightly swollen. The skull of Phainopepla nitens lepida is more generalized than that of Phainopepla n. nitens, having a longer and narrower bill like the generalized Phainoptila. In Phainopepla n. nitens and in members of the genus Ptilogonys, more flaring occurs in the antorbital region.
Phainoptila, as noted above, has no great amount of flaring in the antorbital region. When more specimens of Phainoptila are examined, the base of the bill probably will be found to flare more in some individuals than in others; this would be expected if we may judge by the data on Phainopepla. The premaxilla and maxilla of Phainoptila are similar to the same bones in Dulus, and there is a well-marked ridge on the tomium (possibly for cutting flower parts) . In Phainoptila, the palatines are narrower than in any other genus of the family and abut the lacrimals. The entire skull ap- pears to be modified along different lines from those of the skull of Dulus; the skull of Phainoptila seems to be modified for a frugiv- orous rather than an insectivorous diet. The skull of Phainoptila probably is more nearly similar to the ancestral skull than is that of any other living species in the family. The wide gape character- istic of some members of the family is undoubtedly a modification for aiding in the capture of insects, and Phainoptila has progressed less in this direction than have other species in the family.
Akvey: Waxwings and Allied Birds 499
The mandibles vary somewhat in the shape and proportionate size of the bones. The mandible is proportionately, as well as ac- tually, highest in Dulus. The medial condyle varies to some extent, being slightly flattened mediad in Bomhycilla, and less so in the other genera. The mandible of Bomhycilla narrows to the sym- physis much more gradually than it does in the other genera.
The antorbital plate is large and divides the orbital chamber from the nasal chamber. The small lacrimal bone anterior to the plate articulates with the maxilla and the premaxilla. Shufeldt (1889) states that the free lacrimal ossicle might be of some taxo- nomic importance in the passerines, since it is found in the general- ized Corvids and in nestling Turdids. I find it well developed and identical, with a double articulation and free ends, in all the Bomby- cillids. There is no significant variability in the family, and this is more evidence of close taxonomic relationship between the mem- bers of the family.
The size of the crania is somewhat variable, although the differ- ences seem to be primarily those of proportion. Ptilogonatinae have long crania, whereas the crania of the Bombycillinae and Dulinae are shorter but deeper. I regard the longer cranium as primitive, and it is longest in Phainoptila. In order of decreasing relative length of the cranium, Phainoptila is followed by Ptilogonys cau- datus, P. cinereus, and Phainopepla. Bomhycilla garrula has the deepest cranium in the family.
The measurements of the lengths and widths of the skulls are given in Table 9. The relative length of the bill and relative width of the skull are given in Table 10. These relative measurements are calcu- lated by using the actual measurements in Table 9 as numerators, the length of the skull from the lacrimal bone to the posteriormost end of the skull being used as the denominator. The data indicate that Phainoptila has a slightly narrower cranium.
Humerus. — Certain families of passerine birds have a noticeable variation in the characteristics of the humerus; the bone varies in length, in diameter, and in the complexity of the processes at either end. In the Bombycillids, however, the amount of variation is relatively small, and the diaphysis of the bone is somewhat twisted, especially so in Dulus. The deltoid tuberosity is variable, being shorter but more elevated in Bomhycilla than it is in the Ptilogo- natinae and in the Dulinae. The tendon from the pectoralis major muscle, which inserts on this process, probably finds better insertion on a higher process than on a lower but longer one.
500
University of Kansas Publs., Mus. Nat. Hist.
Distally, the two major condyles and the intercondylar groove or olecranon fossa that make efficient articulation with the ulnar proc- ess, are not variable. The external condyle, however, is significantly variable in the family. This condyle is longest and most pronounced in birds in which the humerus is short in relation to the trunk, as for example in Tachycineta. In the Bombycillidae the condyle is smallest in Phainoptila, where it is a mere suggestion of a process. In the remainder of the Ptilogonatinae, the condyle is larger but rounded, and shows a double process in Ptilogonys caudatus, and a slightly pointed process in P. cinereus. The external condyle in Duhis is not specialized, being low and rounded, but in Bombycilla, it is noticeably elongated, indicating a better attachment distally for
(J^2Z
li^
24
6=^
26
Figs. 22-28. Humeri of five genera of Bombycillidae. Natural size.
22. Phainoptila m. melanoxantha, sex?, MNH no. 26493, 15 mi. SE Cartage, Costa Rica.
23. Ptilogonys caudatxis, male, MNH no. 24492, 15 mi. SE Cartago, Costa Rica.
24. Phainopepla nitens, male, MNH no. 24754, Pima Co., Arizona.
25. Ptilogonys cinereus, female, Louisiana State University no. 297, Xilitla Region, San Luis Potosi, Mexico.
26. Dulus dominicus, female, USNM no. 292652, Don Don, Haiti.
27. Bombycilla cedrorum, male, MNH no. 15331, Bexar Co., Texas.
28. Bombycilla garrula, sex?, USNM no. 223895, Bozeman, Montana.
Arvey: Waxwings and Allied Birds
501
the deltoid muscle. (No measurements are tabulated for this condyle, as the percentage of error in measuring this small structure is great.) Table 1 gives lengths of humeri, and Table 2 gives lengths of the humeri expressed as percentages of the length of the trunk, a standard measurement.
The area of insertion of the deltoid muscle is elongated in those birds with shortened humeri; these birds have also greater flight power than do birds with longer humeri and therefore a shorter external condyle.
Table 1. Lengths of Arm Bones in cm.
Species
|
Humerus |
Radius |
Ulna |
|
2.39 |
2.57 |
2.79 |
|
2.24 |
2.48 |
2.78 |
|
2.21 |
2.59 |
2.82 |
|
2.40 |
2.51 |
2.70 |
|
2.23 |
2.38 |
2.63 |
|
2.35 |
2.58 |
2.88 |
|
2.06 |
2.34 |
2.60 |
Manus
Ptilogonys caudatus
Ptilogonys cinereus
Phainopepla nitens
Phainoptila melanoxantha .
Dulus dominions
Bombycilla garrula
Bombycilla cedrorum
2.25 2.38 2.39 2.25 2.31 2.67 2.38
Table 2. Arm-trunk Ratios (in percent)
|
Species |
Humerus |
Radius |
Uhia |
Manus |
Total |
|
Ptilogonys caudatus Ptilogonys cinereus Phainopepla nitens |
85 84 84 73 78 69 67 |
92 90 98 77 83 75 76 |
93 103 107 82 92 87 85 |
80 89 91 69 81 78 77 |
2.58 2.76 2.82 |
|
Phainoptila melanoxantha Dulus dominicus |
2.31 2.51 |
||||
|
Bombycilla garrula Bombycilla cedrorum |
2.34 2.29 |
||||
502 University of Kansas Publs., Mus. Nat. Hist. Table 3. Arm-trunk Ratios (in percent)
Species
|
Humerus |
Radius |
Ulna |
Manus |
|
90 |
101 |
111 |
106 |
|
68 |
82 |
90 |
77 |
|
69 |
82 |
91 |
75 |
|
71 |
84 |
96 |
81 |
|
72 |
84 |
98 |
86 |
|
75 |
81 |
92 |
80 |
|
85 |
90 |
106 |
81 |
|
71 |
95 |
107 |
128 |
|
83 |
105 |
115 |
92 |
|
76 |
75 |
89 |
78 |
|
85 |
93 |
105 |
71 |
|
91 |
99 |
110 |
95 |
|
85 |
111 |
120 |
108 |
Total
Corvus brachyrynchos . Dendroica audubonii . . Setophaga ruticilla. . . . Myadestes townsendi . .
Sialia sialis
Hylocichla mustelina. .
Parus atricapillus
Tachycineta thalassina Myiarchus crinitus . . . . Dumetella carolinensis . Polioptila caerulea . . . . Eremophila alpestris . . Muscivora forficata . . .
307 237 235 248 256 247 272 306 290 243 261 296 313
Pygostyle. — This part of the skeletal system is variable in the species dealt with, not so much in size as in complexity. It reflects, of course, the character of the caudal muscles and their size, as well as the length of the rectrices and the corresponding force necessary to hold these feathers upright and in a useful position. Firm attach- ment is important even in flight, because the tail is used as a rudder, and in the Ptilogonatinae as a brake. The pygostyle is most modi- fied in this subfamily.
In lateral aspect, the pygostyles of the species of the Ptilogona- tinae are similar. The crest of the bone is flattened dorsally, and has a broad anterior surface that is thin and bladelike. This is widest in Ptilogonys caudatus, and narrowest in Phainoptila, in which genus, however, the entire bone is of small size. The centrum is widest in Ptilogonys caudatus, and is progressively narrower in P. cinereus, Phainopepla, and Phainoptila. Greater width provides a larger area of attachment for the larger rectrices and also more area for insertion of the lateralis caudae muscle, the size of which varies more than that of the other caudal muscles in the different species of the Bombycillidae.
Arvey: Waxwings and Allied Birds
503
33
34
Figs. 29-35. Pygostyles in posterior view of five genera of Bombycillidae. X 2.
29. Phainoptila m. melanoxantha, sex?, MNH no. 26493, 15 mi. SE Cartago, Costa Rica.
30. Ptilogonys caudatus, male, MNH no. 24492. 15 mi. SE Cartago, Costa Rica.
31. Phninopepla nitens, male, MNH no. 24754, Pima Co., Arizona.
32. Ptilogonys cinereus, female, Louisiana State University no. 297, Xilitla Region, San Luis Potosi, Mexico.
33. Dulus dominicm, female, USNM no. 292652, Don Don, Haiti.
34. Bombycilla ccdrorum, male, MNH no. 15331, Bexar Co., Texas.
35. Bombycilla garrula, sex?, USNM no. 223S95, Bozeman, Montana.
In proportionate size (see Table 7), the pygostyle of Bombycilla is the smallest in the family. The dorsal spinous portion is acutely pointed instead of flattened as in the Ptilogonatinae. In Dulus, the spinous portion is extremely thin, and shows a decided curve dorsad from the centrum, and there is no flattened area anterior to the spinous portion as is seen in Ptilogonys.
The centrum in cross section varies considerably. In Bombycilla the walls are indented, with definite terminal knobs; both knobs and indentations are more pronounced in B. garrula than in cedrorum, however. The spinous portion is enlarged in both species, and the rest of the neck region is constricted (Figs. 29-35) .
The centrum of Dulus in posterior aspect presents the appearance of a simple shield; little of the indentation seen in Bombycilla is
504 University of Kansas Publs., Mrs. Nat. Hist.
Figs. 36-42. Pygostyles in lateral view of five genera of Bombycillidae. X 2.
36. Phainoptila m. melanoxantha, sex?, MNH no. 26493, 15 mi. SE Cartago, Costa JRica.
37. Ptilogonys caudatus, male, MNH no. 24492, 15 mi. SE Cartago, Costa Rica.
38. Phainoptila nitens, male, MNH no. 24754, Pima Co., Arizona.
39. Ptilogonys cinereiis, female, Louisiana State University no. 297, Xilitla Region, San Luis Potosi, Mexico.
40. Dulus dominicus, female, USNM no. 292652, Don Don, Haiti.
41. Bombycilla ccdrorum, male, MNH no. 15331, Bexar Co., Texas.
42. Bombycilla garrula, sex?, USNM no. 223895, Bozeman, Montana.
present. The spinous portion is plain, with no constriction nor terminal enlargement in the neck. The centrum in Phainopepla is similar to that in Dulus, but has a small expansion at the base of the spine, the entire centrum being wider in proportion to its over-all size than in any of the other species mentioned previously. The centrum in Ptilogonys shows great width, and the spine is in a large expanded tip as in Bombycilla. The lateral edges of the cen- trum in P. cinereus are "winged" and in two separate halves,- whereas the centrum of P. caudatus is fairly plain, its specialization being reflected primarily in breadth and flatness. In cross section of the centrum, Phainoptila is similar to Phainopepla, although, in the former, the bone is smaller in proportion to the size of the animal, and the lateral wings are more angular than in Phainopepla.
In specialization for muscle attachment, the centra of the pygo- styles of the Ptilogonatinae have more area for muscle attachment
Arvey: Waxwings and Allied Birds 505
than do the centra in the Bombycillinae and Dulinae; the centrum is wide, the spinous portion is long, and the bone is flattened ante- riorly. The most generalized pygostyle is in Phainoptila, and that of Dulus differs only slightly. In Bombycilla the pygostyle is pro- portionately small, but is complex in shape; there is seemingly not the need for greatly expanded areas since the caudal muscles are less specialized in this genus.
Sternum. — The sternum in Bombycillids is typically passerine in general shape and in having a long and deep carina or sternal crest. The caudal process of the bone is broad, with the terminal ends flattened, forming dorsally a graceful V-shaped outline, whereas the outline of the posterior end of the sternum is broad and convex.
In lateral aspect, the carina is deeper in Bombycilla than in other genera of the family, and is deepest in B. garrula. In this species, the manubrium is more extended and comparatively larger than in the other species of the family. The anterior edge of the keel forms the sharpest angle in B. cedrorum. In Dulus, the keel is moderately deep, the manubrium short, and there is a distinct indented curve between the manubrium and the anterior angle of the keel.
In ventral aspect the lateral processes of the sternum tend to flare outwards in adult Ptilogonatines on almost the same plane as the rest of the bone, whereas in Bombycilla and Dulus the same process is closer to the body of the sternum. In Bombycilla the xiphoid process is more dorsal in position than in other species in the family, and in Dulus an upward curve is very noticeable. The proc- ess in these two genera is narrower than in the Ptilogonatinae, and lacks the heavy distal terminal enlargement which is apparent in Ptilogonys.
Relative Lengths of Bones. — In instances where the animals being compared are obviously different in over-all size, it is useful to express the size of a given part in relation to some other part of the same individual organism if the aim is to obtain clues as to differences in functions of the parts being compared. Differences in actual lengths of corresponding bones in two kinds of animals often, of course, reflect only the difference in over-all size of the animals. Consequently, the relative size of the part is expressed as a percentage in this paper. In computing a percentage it is well, of course, to select some relatively stable part of the animal to use as a denominator in the mathematical expression that yields the per- centage. The thoracic region of the vertebral column is thought to
3—1019
506
University of Kansas Publs., Mus. Nat. Hist.
be such a part. For example, the length of the humerus divided by the length of the thoracic region yields, in Phainopepla and Ptilo- gonys, respective percentages of .84 and .85. These are roughly the same, whereas the actual lengths of the humeri are 2.21 and 2.39 cm.
Table 4. Lengths of Leg Bones in cm.
Species
Femur
Tibio- tarsus
Tarsomet- atarsus
Ptilogonys caudatus
Ptilogonys cinereus
Phainopepla nitens
Phainoptila melanoxantha
Dulus dominicus
Bombycilla garrula
Bombycllla cedrorum
2.04
1.89 1.76 2.43 2.09 2.32 1.92
3.10 2.90 2.78 3.77 3.34 3.46 2.95
1.94 1.77 1.72 2.58 2.09 1.99 1.64
Table 5. Leg-trunk Ratios (in percent)
Species
Ptilogonys caudatus
Ptilogonys cinereus
Phainopepla nitens
Phainoptila melanoxantha
Dulus dominicus
Bombycilla garrula
Bombycilla cedrorum ....
Femur
73 71 69 74 73 68 63
Tibio- tarsus
110 109 106 115 119 101 96
Tarso- meta-
tarsus
69 66 65 60 73 59 53
Total
252 246 240 249 265 228 212
Arvey: Waxwings and Allied Birds
507
Table 6. Leg-trutik Ratios (in percent)
Species
Femur
Tibio- tarsus
Tarso-
meta- tarsus
Total
Corvus brachyrynchos .
Corvus corax
Dendroica audubonii . . Setophaga ruticilla . . . . Myadestes townsendi . ,
Sialia sialis
Hylocichla mustelina . .
Parus atricapillus
Tachycineta thalassina Myiarchus crinitus . . . Dumetella carolinensis Polioptila caerulea .... Eremophila alpestris. . Muscivora forficata. . .
71
73
62
66
61
66
75
78
61
68
73
75
73
62
120 139 109 127
99 111 133 138
97 106 136 144 113
98
77 78 81 94 60 72 97 99 56 74 94 113 115 61
268 290 252 287 220 249 305 315 214 248 303 332 301 221
Table 7. Actual Length and Width in mm. of Pygostyle and Proportionate Length and Width of Pygostyle in percent of Lacrimal Length
Species
Ptilogonys caudatus
Ptilogonys cinereus
Phainopepla nitens
Phainoptila melanoxantha
Dulus dominicus
Bombycilla garrula
Bombycilla cedrorum ....
Length
9.8 8.8 8.4 8.5 8.5 7.0 7.1
Width
3.9 4.1 3.9 3.5 2.9 3.5 2.9
Length, percent
45 41 41 35 38 31 35
Width, percent
18 19 19 14 13 15 14
508
University of Kansas Publs., Mus. Nat. Hist.
Table 8. Length of Sternum and Depth of Carina expressed as percentages
of the Length of the Trunk
Species
Sternum
Carina
Ptilogonys caudatus
Ptilogonys cinereus
Phainopepla nitens
Phainoptila melanoxantha .
Dulus dominicus
Bombycilla garrula
Bomby cilia cedrorum
|
85 |
28 |
|
91 |
32 |
|
81 |
26 |
|
76 |
25 |
|
07 |
28 |
|
88 |
33 |
|
82 |
31 |
Table 9. Skull and Sternum, Length and Width in mm.
|
Species |
Length of Skull |
Width of Skull |
Length of Sternum |
Width of Sternum |
|
Ptilogonys caudatus |
34.9 33.4 33.3 39 7 36.4 37.0 34.0 |
15.6 14.7 15.1 16.0 16.6 16.8 15.5 |
23.9 24.3 21.3 24.8 30.5 30.0 25.3 |
7.8 |
|
Ptilogonys cinereus |
8.5 |
|||
|
Phainooepla nitens |
6.9 |
|||
|
Phainoptila melanoxantha Dulus dominicus |
8.2 8.0 |
|||
|
Bombycilla garrula Bombycilla cedrorum |
11.2 9.6 |
The length of the trunk was taken as the distance from the anterior tip of the neural crest of the last cervical vertebra to the anterior edge of an acetabulum. The number of free thoracic vertebra was five in each specimen ; consequently, there was no error from this source. In the cranium, a measurement was taken from the anterior edge of the lacrimal bone to the posteriormost end of the cranium, and the resultant figure was employed for a constant in cases in which small bones were compared.
Arvey: Waxwings and Allied Birds
509
Table 10. Relative Length and Width of Skull (in percent)
Species
|
Length of Skull |
Width of Skull |
|
160 |
72 |
|
158 |
69 |
|
162 |
73 |
|
161 |
65 |
|
164 |
75 |
|
164 |
74 |
|
162 |
74 |
Ptilogonys caudatus
Ptilogonys cinereus
Phainopepla nitens
Phainoptila melanoxantha
Dulus dominicus
Bombycilla garrula
Bombycilla cedrorum ....
Trunk Measure
Fig. 43. Part of skeleton of Bombycilla cedrorum showing method of measuring the length of the trunk. Natural size.
Leg-trunk Percentages. — Table 4 shows the relative lengths of the legs and of the separate bones in the legs of the different species of the Bombycillids. Table 5 shows corresponding lengths for other passerine birds. The total length of the leg was computed by adding the figures obtained for the lengths of the femur, tibiotarsus and
510 University of Kansas Publs., Mus. Nat. Hist.
tarsometatarsus. The lengths of the toes were disregarded. Length of leg was recorded in this same way by Richardson (1942:333), who thought that only in swimming and running birds do the toes con- tribute to the functional length of the hind limb.
Table 4 shows that of the birds compared in this paper, Dulus has the longest legs. In order of decreasing length the others are the Ptilogonatinae, and finally the Bombycillinae, which have the shortest legs of all. In Waxwings the length of the legs, expressed as percentages of the body-lengths, are identical with those birds that are similar in habits, that is to say, birds which do not use the hind limb except in perching. It can be noted by reference to Table 5 that Tachydneta and Myadestes fall into this category. This shortness of limb is obviously adaptive, and each of the segments of the limb has been correspondingly shortened, with no element re- duced at the expense of the other two. The short leg can be more easily folded against the body while the bird is in flight, than can a long leg which is more unwieldy. It may be noted from tables 4 and 5 that birds which spend much time on the ground, or that hop a great deal in the underbrush, have longer legs than do birds which spend much time in flight. Two birds with noticeably long legs are Hylocichla mustelina, a typical ground dweller, and Parus atric- apillus, which hops about in the trees and underbrush.
Insofar as the lengths of the legs show, Dulus and Phainoptila are the most generalized of the Bombycillidae, since the relative length of leg is approximately the same as that of more generalized birds such as warblers, crows and thrushes of similar locomotory habits. In other words, Dulus and Phainoptila have remained unspecialized, in contrast to the waxwings in which adaptive changes fitting them for a perching habit have taken place. Ptilogonys and Phainopepla are intermediate in length of leg between Phainoptila and Bomby- cilla, and Ptilogonys and Phainopepla have progressed from life on the ground toward the perching habit. Bombydlla cedrotrum is more specialized than is B. garrula in shortness of leg, and the re- duction is comparable, as is noted above, to that in the legs of Tachydneta.
In birds which have the legs much modified for walking or for hop- ping in the brush, such as Polioptila and Eremophila, it is noteworthy that the distal segment, the tarsometatarsus, is the longest, whereas in birds such as Myiarchus and Tachydneta, that do not utilize the limbs in this manner, the tibiotarsus, the middle segment, is the longest. Mammals much modified for walking or hopping likewise
Arvey: Waxwings and Allied Birds
511
250-
200
abed abed A B
abed abed abed abed abed C D E F G
Fig. 44. Graph showing relative lengths of bones of the leg. The percentage values are shown on the axis of the ordinates. A. Bombycilla cedrorum; B. Bomhycilla garrula; C. Dulus dominicus; D.
Phainoplila melanoxantha ; E. Fhainopepla nitens; F. Ptilogonys cinereus;
G. Ptilogonys caudatus. a. femur; b. tibiotarsus; c. tarsometatarsus;
d. total.
have the proximal segment, the femur, short, and the distal segment long (Howell, 1944). The waxwings have all of the segments short; these birds are modified for strong and sustained flight. Their hind limbs are used principally for landing devices and for perching. No one element of the leg has been shortened much, if any, more than any other.
Arm-trunk Percentages. — Tables 1 and 2 show the total length of the arm, and lengths of the separate arm elements, relative to the trunk. Table 3 gives the corresponding lengths for birds other than the Bombycillidae. Total length of arm was obtained by adding together the lengths of the humerus, ulna, and manus, and by di- viding the figure thus obtained by the length of the trunk as was done for leg lengths in tables 4 and 5. The method of adding to- gether the component parts does not give the entire length of the
512 University of Kansas Publs., Mus. Nat. Hist.
wing, since the length of the feathers, which add effectively to the total length, as well as do the lengths of the small carpal elements, is lacking.
It may be noted that Phainoptila and Bombycilla have the short- est arm in the family Bombycillidae. The humerus, radius and ulna
Figs. 45-46. Outlines of wings. X ¥2
45. Ptilogonys caudatus, showing relation of outline of wing to bones of arm.
46. Bombycilla cedrorum, showing relation of outline of wing to bones of arm.
are comparable to the same elements in thrushes and the catbird, and it is only the extremely short manus in Phainoptila that affects the total. The manus in Phainoptila is comparatively smaller than in any other genus of the family Bombycillidae, and this indicates poor flight power. Bombycilla has a total length corresponding
Arvey: Waxwings and Allied Birds
513
closely to that in warblers, but the lengths of the distal elements correspond closely to those in the catbird and thrushes. Of the three segments, the humerus is, relatively, the most shortened. Next in order of increasing length of arm is Dulus; measurements for it are roughly the same as those of Myadestes. The wing bones of the Ptilogonatinae, other than Phainoptila, are the longest in this series, and they most nearly resemble the same bones in flycatchers, Parids, and gnatcatchers.
300
250
200
150
abode abcde abcde abcde abcde Gbcde abcde A B C D E F G
Fig. 47. Graph showing relative lengths of bones of the arm. The percent- age values are shown on the axis of the ordinates. A. Bomhycilla cedrorum; B. Bomhycilla garrula; C. Dulus dominicus; D.
Phainoptila melanoxantha ; E. Phainopcpla nitens; F. Ptilogonys cinereus;
G. Ptilogonys caudatus. a. humerus; b. radius; c. ulna; d. manus;
e. total.
It is notable that, in general, birds with long and narrow wings appear to have relatively the shortest humeri, with the distal bones, especially the manus, variable in length and seemingly correlated with the manner of feather attachment. Those birds with rounded and short wings have the longest humeri. In swallows, for example, the humerus is short, whereas the other arm bones are long, and the manus is unusually large and heavy. A short humerus gives better lever action in the flight stroke than a long humerus does.
514 University of Kansas Publs., Mus. Nat. Hist.
MUSCULATURE
Dissections showed the same muscles to be present in all genera of the Bombycillidae. There are, nevertheless, differences in the size of the muscles in the various species, and these differences have been investigated primarily as a check on differences noted in the structure of the bones. Even slight differences in mass can be im- portant functionally, but the difficulty in accurately measuring the mass prevents wholly reliable conclusions. The method first used in the attempt to determine the mass of a given muscle was that of immersing the muscle in a liquid-filled graduated tube, and then measuring the amount of liquid displaced. This method, although adequate for large muscles, was subject to a great amount of error in the case of small muscles, and consequently was abandoned. The technique eventually used was that previously employed by Rich- ardson (1942). It consisted of dissecting out the muscle, placing it in embalming solution, leaving it there until a later period, and finally, weighing the muscle on scales, accurate to a milligram, after the muscle had been out of the liquid for a period of one minute. After being weighed, the muscle was measured by the dis- placement method in a graduated tube, as a check. The results in- dicate that, although the two methods give the same general results, weighing is accurate to one-hundredth of a gram, whereas the dis- placement method was accurate to only a tenth of a gram.
In determining the percentage of the weight of a muscle in relation to the total weight of the bird, the weight of the muscle was used as the numerator, and the weight of the preserved specimen was used as the denominator. Before weights were taken, all specimens were plucked in identical fashion.
Caudal Muscles. — The muscles of the caudal area that were used for comparison were the levator caudae and the lateralis caudae. These muscles are used by the living bird to maintain the position of the pygostyle and therefore the rectrices; these muscles are es- pecially important to those birds that utilize the tail as a rudder in flight and as a brake. As may be seen by reference to Table 11, the two muscles are largest in proportion to body weight in the Ptilogonatinae, in which subfamily the species have long rectrices and must have correspondingly well-developed muscles in order to utilize the rectrices to best advantage in flight. The lateralis caudae differs more according to species than does the levator caudae, showing that rudder action of the tail is of primary importance in the adaptation for capturing insects. It will be remembered that the
Arvey: Waxwings and Allied Birds
515
pygostyle in this subfamily has a flattened lateral surface for attach- ment of the levator caudae muscle, and it is therefore to be expected that this muscle will be larger in the Ptilogonatinae than it is in either the Bombycillinae or the Dulinae. The levator coccygis, to- gether with the two muscles mentioned above, is responsible for elevation of the tail. The levator coccygis is less altered in different
Fig. 48. Caudal musculature, of Phainopepla nitens lepida, in dorsal view. X 2. a. Levator coccygis; b. Levator caudae; c. Lateralis caudae; d. Lateralis coccygis; e. oil gland; f. dorsal tip of pygostyle.
species of the family than is the lateralis caudae. It may be noted that the caudal muscles of Dulns and Bombycilla constitute a smaller percentage of the total weight of the bird than in any of the genera in the subfamily Ptilogonatinae.
Table 11. Caudal Muscles (Actual and Relative Weights)
Species
Ptilogonys caudatus .
Ptilogonys cinereus .
Phainopepla nitens .
Phainoptila melanoxantha .
Dulus dominicus .
Bombycilla garrula.
Bombycilla cedrorum.
Lateralis
.022g. .045%
.OlOg. .026%
.008g. .029%
.015g. .014%
.006g. .014%
.OlOg. .014%
.OOSg. .014%
516 University of Kansas Publs., Mus. Nat. Hist.
Table 12. Weights of Muscles (These percentages expressed in terms of
weights of the body)
|
Species |
P. major |
P. minor |
Deltoid |
|
Ptilogonys caudatus |
2.42g. 4.94% 2.19g. 5.57% 1.30g. 4.99% 3.93g. 6.18% 2.09g. 4.81% 3.85g. 5.31% 2.58g. 5.00% |
.29g. .59% .28g. .71% .20g. .77% .44g. .69% .22g. .50% .45g. .62% .35g. .68% |
.55g. |
|
PtiloEonvs cinereus |
1.12% .53g. |
||
|
Phainopepla nitens Phainoptila melanoxantha Dulus dominicus |
1.35% .30g. 1.15% 1.45% .50g. |
||
|
Bombycilla garrula |
1.15% .55g. |
||
|
Bombycilla cedroruni |
.76% .50k. |
||
|
.97% |
|||
|
Thigh |
Peroneus |
Gatrocnemius |
|
|
Ptilogonys caudatus Ptilogonys cinereus |
.43g. .88% .30g. .71% .28g. 1.12% 1.09g. 1.61% • 73g. 1.68% .50g. .69% .37g. .73% |
.15g. .31% .08g. .21% ■ lOg. .40% .48g. .75% .18g. .41% .15g. .18% .lOg. .19% |
.96% |
|
PhainoDCola nitens |
1.02% |
||
|
Phainoptila melanoxantha Dulus dominicus |
1.42% 2.97% |
||
|
Bombycilla earrula |
1.01% |
||
|
Bombycilla cedrorum |
.59% |
||
|
.83% |
Arvey: Waxwings and Allied Birds 517
Pectoral Muscles. — The pectoral set of muscles varies but little in the family; flight power is seemingly not dependent upon size of either the pectoralis major or pectoralis minor. The data indicate that the insertion on the humerus, with consequent changes in the relative length of that bone, is more significant in type of flight and over-all flight power than is the actual size of the muscle mass. The deltoid muscle, for example, is smaller in Bombycilla than in mem- bers of the other two subfamilies. The humerus in Bombycilla is shortened, and the muscle therefore does not need to be large to accomplish the same powerful stroke that would be accomplished by a longer humerus and a larger, more powerful deltoid muscle. In the case of the deltoid, the shortening of the humerus and the more complex arrangement of the points of insertion have obviated the necessity of enlarging the muscle.
Leg Musculature. — The muscles of the thigh are noticeably larger in birds that have long leg bones. (See Table 12 for size of mus- cles.) On the tibiotarsus, the peroneus and gastrocnemius muscles were measured. When expressed as a percentage of the weight of the bird, the peroneus has much the same relative weight in all but one of the species, whereas the gastrocnemius varies much. The peroneus is proportionately large only in Phainoptila, in which genus all the leg muscles are well developed, but the gastrocnemius is larger in all the Ptilogonatinae and in Dulus than it is in the specialized Bombycilla, in which it has probably been reduced as the leg bones and other muscles have been reduced.
The volume of the muscles of the hind limb changes more readily in response to saltation and running than do the muscles of the forelimb to flying.
DIGESTIVE TRACT
The digestive tract is relatively uniform in all genera of the fam- ily; there are only slight differences between the species. The degree of compactness of the visceral mass varies, Phainoptila and Ptilogonys caudatus having the folds of the digestive tract loosely arranged, whereas Ptilogonys cinereus and Phainopepla have folds which adhere more tightly to the ventriculus and liver. In Dulus and Bombycilla, as compared with the Ptilogonatinae, the visceral mass (primarily liver and ventriculus) is situated more posteriorly in the body cavity, and is more compact, and the intestine is more tightly coiled.
The coiling of the intestine, if its degree of compactness is dis- regarded, is nearly identical in the birds of the family; there are
518 University of Kansas Publs., Mus. Nat. Hist.
four major loops between the ventriculus and the anus. The length of this section of the tract is, however, somewhat variable, as can be seen by reference to Table 13, in which the actual and relative lengths of the intestine are given. It may be seen that in Bomby- cilla and in Phainopepla, the tracts are much shortened. This is notable, since these are frugivorous birds, and in many frugivorous birds, the tract is lengthened for better extraction of edible portions of the food. Possibly the action of the digestive juices is corre- spondingly more rapid in Bombycilla and Phainopepla, thereby per- mitting the necessary nutriment to be extracted by a short digestive tract.
In a migratory bird, or one that depends on flight power to find food and escape capture by predators, as in the case of the wax- wings, the compacted and shortened visceral mass would seem to be advantageous, because of the consequent reduction in weight. I consider the longer intestine to be the ancestral condition, and that the intestine has become shorter to meet new environmental condi- tions.
Beddard (1898:30) states that caecae in the tract may be highly variable in a single family of birds. The Bombycillidae is no exception in this regard. At the junction of the cloaca and the large intestine, there are two small caecae, the function of which is un- known to me. The caecae are largest in the Ptilogonatinae, smaller in the Bombycillinae, and smallest in the Dulinae. There may be a
Table 13. Digestive Tract: Actual Length, and Length Relative to
Thoracic Length
Species
Relative
length
(in percent)
Ptilogonys caudatus
Ptilogonys cinereus
Phainopepla nitens
Phainoptila melanoxantha
Dulus dominicus
Bombycilla garrula
Bombycilla cedrorum
476.9 415.6 357.5 457.1 451.0 298.2 309.5
Arvey: Waxwings and Allied Birds 519
correlation between large caecae and more insectivorous diet and small caecae and frugivorous diet; however, the data are not con- clusive in this regard.
ORIGIN OF THE SPECIES
It is here postulated that the center of origin for the ancestral stock of the Bombycillidae was in a region of North America, which at the time concerned was temperate or possibly even semi-tropical in climate. Probably Northern Mexico was the place and probably the climate was temperate. It is reasonably certain, because of the distribution of the species of the family, that they originated in the Americas. In the absence of paleontological data {Bombycilla alone is reported, in essentially its modern form, from the late Pleistocene — Wetmore, 1940a), the place and time of origin cannot certainly be determined.
The distribution of the family is such that the more primitive groups are in the south. These are the Ptilogonatinae in Central America and Mexico, and the isolated Dulinae in Haiti and the Dominican Republic. This distribution would support the view that the origin was in the south. However, the Holarctic Bomby- cillinae are so typically birds of northern latitudes that, were it not for such close relatives south of their range, it would appear logical to infer a northerly origin with a subsequent shifting of populations both southward and northward. The phyletic age of the family is probably great, however, as evidenced by the spotty dis- tribution of the birds.
In the evolution of this family, population pressure possibly played the initial role in forcing members of the primitive, southern stock to seek habitable areas on the periphery of the range. Some birds also, being possessed of the ^'adventuresome spirit", aided the northerly movement, thus effecting an extension of the breeding ranges to the north. So far as is now known, this family did not seek living space in South America. By extending its range, a species might find more abundant food and nesting sites. This process of extending the range probably would be costly to the species con- cerned, because only those individuals best able to adapt themselves to the new environmental conditions would be able to survive long enough to reproduce their kind.
The return flight to the south could, in time, be dispensed with, except in the coldest weather or when the local berry- and fruit-crop failed. Birds such as waxwings are, of course, able to subsist on
520
University of Kansas Fuels., Mus. Nat. Hist.
dried fruits and berries in the critical winter season when strictly- insectivorous birds, not so catholic in their food habits, must return south. It appears that waxwings are descendants of migratory birds that have adjusted themselves to a life in the north; and they are judged not to have evolved from year-round residents of the north.
Even a short migratory journey in spring by part of a population of birds, while the other part remained in the original range, would quickly isolate one breeding population from the other, resulting in the formation of different genetic strains that lead to subspecies, species, and finally to genera and families. Any variation away from the ancestral, "sedentary" stock would become established more quickly because of such isolation at the breeding period. By the same token, the parental stock can, and no doubt does, become modified to suit its environment more perfectly, thus accelerating the tempo of this type of divergent evolution.
The original "split" of the Bombycillines is thought then to have been the result of migration on the part of some of the ancestral stock, with subsequent loss of regular migration because the need to return south was lost. Early in development, and before the mi- grational tendency was entirely lost, an isolated population, which later became sedentary, as it was an island population, diverged to give rise to the Dulinae. The Dulinae are a homogeneous group since on the islands now inhabited by the birds, they have not been isolated sufficiently long to produce even w^ell-marked subspecies.
The present day Phainoptila is most nearly like the ancestral group, and the remainder of the Ptilogonatinae have diverged to
RECENT TIME
B japonica B gorrula B cedrorum Dulus Phainoptila Phoinopepla P cinereus P caudatus
PAST TIME
ANCESTRAL STOCK
Fig. 49. Hypothetical family tree of the Bombycilhdae.
Arvey: Waxwings and Allied Birds 521
fit conditions similar to those to which the Tyrannid flycatchers, which parallel them, are also fitted.
In comparatively recent geological time, two basic lines developed from the Bombycilline stock, the future B. garrula and B. cedrorum. Possibly garrula originally was isolated in Europe and Asia, and later came into contact with B. cedrorum, following the time at which the two species were genetically well differentiated. It appears certain that B. japonica was an offshoot of the Bombycilline stock at an early time, since it has characteristics that seem rela- tively unspecialized. It possibly was isolated in the Orient.
Structural affinities of Dulus and Bomb y cilia are more pronounced than are those of Dulus and Ptilogonys, for example. Many of the structural features of Dulus parallel those of Phainoptila, and it seems likely that the Dulinae were separated early in the history of the family, perhaps as an isolated offshoot of the early migratory Bombycillinae.
CONCLUSIONS
Nomenclature, as used by a taxonomist, should of course indicate affinities as well as apply a name, and the rank of the family should be applied to a structural unit based on common anatomical char- acters that are more fundamental than, in my opinion, are those used by Ridgway (1904) in proposing family status for the silky flycatchers and the palm-chats. The characters in the diagnosis (page 478) of the family Bombycillidae are common features re- garded as warranting a single family unit for the waxwings, silky flycatchers, and palm-chats. The differences in morphology used by previous workers to characterize each of these groups: (1) the silky flycatchers; (2) waxwings and; (3) palm-chats are regarded as more properly characters of only subfamily rank.
The existing coloration of the species of the Bombycillidae ap- pears to have been acquired relatively late, geologically speaking. The three subfamilies responded to ecological stimuli in three dif- ferent ways, and the resulting color patterns are unlike in the three groups. Dulinae to this day have a color pattern that is most like the ancestral color pattern, and this is recapitulated in the juvenal plumage of the Bombycillinae before they attain their adult plum- age.
Consideration of the geographic distribution of the species of the family indicates that the center of origin of the family Bombycil-
4—1019
522 University of Kansas Publs., Mus. Nat. Hist.
lidae was south of the present range of the waxwings (subfamily Bombycillinae) . Waxwings probably are the descendants of a mi- gratory population that diverged from, the primitive population at an early time in the history of the family. Owing to their adapta- tions to survive in the north, waxwings no longer return south in the autumn. Palm-chats (subfamily Dulinae) are descendants of an isolated population of the family stock that developed communal living habits as one specialization. Silky Flycatchers (subfamily Ptilogonatinae) became modified to catch insects, and have special- izations that roughly parallel those of the Tyrannid flycatchers.
Osteologically, the various species of the Bombycillidae are re- markably similar. Small variations do exist, but these are pri- marily differences in relative size. The modifications of the beak enable palm-chats to feed on parts of plants, and the beak of Phainoptila shows some similarity in this respect. Rounded wings, which cause a bird to fly by means of short, relatively weak strokes, are correlated with a comparatively long humerus, whereas long and pointed wings, which enable a bird to fly with more powerful strokes of the wing, are correlated with a relatively short humerus. There is a positive correlation between a short humerus and a long ex- ternal condyle, and between a long humerus and the absence or smallness of the external condyle.
In the Bombycillidae short bones of the leg are adaptive, and long bones of the leg are the generalized condition. Although all passerine birds were differentiated relatively late in geologic time, long hind limbs still could have been present in the immediate an- cestors of passerine birds. As adaptive radiation took place in the class Aves, some birds, the Bombycillidae included, becaiiie more and more adapted for an arboreal, and eventually an aerial habitat, with consequent loss of saltatorial and running ability.
Birds, like mammals, have a short femur, the most proximal ele- ment in the leg, if the species is adapted to run fast. If the species is not adapted to run fast, birds, unlike mammals, have the tibio- tarsus longer than any of the other elements; in mammals that are not adapted to run fast, the femur and tibia are approximately the same length. In non-running birds as compared with running birds, the leg element distal to the tibiotarsus, and the one proximal to it, are considerably shortened. In waxwings, all three elements of the hind limb are shortened, indicating that the reduction in length has been, evolutionarily speaking, a rapid process, in order to reduce the limbs to a convenient size as soon as possible.
Akvey: Waxwings and Allied Birds 523
The shape of the pygostyle varies in the Bombyeillidae, but the simple shieldhke bone of Phainoptila is judged to resemble closely the ancestral type. In Ptilogonys there is a tall dorsal spine, coupled with a wide and heavy centrum and flattened lateral areas, for support of the long rectrices. In Bombycilla the bone is small with knobs on the centrum that have been developed for muscle attachment.
The muscles were carefully dissected in each genus and in most of the species. The same homologous muscles are present in all species. Significant differences were found only in the relative size of certain muscles. No satisfactorily accurate method of measuring these differences was found. Consequently, less use was made of the results of the dissections than was originally planned.
The set of pectoral muscles varies but slightly in relative mass, and the variation is not considered significant. The deltoid muscle was selected for measurement since its point of insertion is unusually variable, while the mass of the muscle varies little. We can con- clude that the extent of the area of insertion of the tendon of a muscle can determine that muscle's relative efficiency, while the nuiscle itself remains the same in bulk.
The muscles of the hind limb are notably larger in species that have long legs, and a good index of the hopping ability may be gained by study of certain of these muscles. In the Bombyeillidae, and in those Ptilogonatinae that do not use the hind limbs for hopping, the bones are shortened, and the associated muscles are correspondingly smaller.
The gross anatomy of the digestive tract is practically identical in the members of the family. The variability noted is mainly in the degree of compactness of the visceral mass in Bo^nbycilla and in Phainopepla. Also there is a tendency for the Bombycillinae and the Dulinae to have the mass situated more posteriorly than it is in the Ptilogonatinae. Moreover, Bombycilla has a shorter in- testine than do the other genera. All of this indicates that the wax- wings (Bombycillinae) have the center of gravity situated more advantageously for flight than do the birds of the two other sub- families.
524 University of Kansas Publs., Mus. Nat. Hist.
SUMMARY
1. The silky flycatchers, waxwings, and palm-chats are included in the family Bombycillidae; the Ptilogonatidae and Dulidae are reduced to subfamily rank.
2. The coloration of the birds of each subfamily is different be- cause the ecological needs are different.
3. Waxwings were at one time regularly migratory, but are now nomadic, since they are adapted to live in northern latitudes for the entire year.
4. The corresponding bones in different members of the family closely resemble one another, and the differences which do exist are the results of responses within relatively recent times to changes in habits.
5. In the Bombycillidae a rounded wing is judged to be the primi- tive condition. As the wing becomes more pointed, the humerus becomes shorter and its external condyle longer.
6. The hind limbs are short in birds that depend most on flight power, but are longer and the distal elements are disproportion- ately longer in birds that depend on saltation or on running.
7. The pygostyle varies in shape and size between genera and even between some species.
8. The pectoral muscles differ in size only slightly in the different members of the family, but the insertions are more extensive for these muscles in birds that fly a great deal.
9. The muscles of the hind limb vary in mass, but not in kind, in the members of the family Bombycillidae.
10. In the Bombycillidae that depend on flight power, rather than on saltation or on running power, there is a tendency for the digestive tract to become shorter and for the whole visceral mass to become more compact.
Arvey: Waxwings and Allied Birds 525
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Transmitted July 29, 1949. 5—1019
530 University of Kansas Publs., Mus. Nat. Hist.
Mention should be made here of an important paper by Jean Delacour and Dean Amadon (1949). The Relationships of Hypocolius. Ibis, 91:427-429, plates 19 and 20) which appeared after the present paper by Arvey was written. Delacour and Amadon stated that Hypocolius, a monotypic Persian genus, should be assigned to the Bombycillidae. Their conclusions (op. cit.: 429) were as follows: "It might be advisable to set up three subfamilies in the Bombycillidae, one for Bombycilla, one for Hypocolius, and a third for the silky flycatchers, Ptilogonys, Phainopepla and Phainoptila. Further study may show that Dulus can be added as a fourth subfamily.
"Previously the BombyciUidae appeared to be an American group of which one genus (Bombycilla) had reached the Old World. Inclusion of Hypocolius in the family makes this theory uncertain. Without obvious affinities to other families, and consisting of a small number of scattered and rather divergent genera, the Bombycillidae would seem to be a declining group whose origin cannot safely be deduced from the distribution of the few existing species."
—Eds.
□
23-1019
(Coatinued from inside of front cover.)
15. A new hylid frog from eastern Mexico. By Edward H. Taylor. Pp. 257-264, 1 figure in text. August 16, 1948.
16. A new extinct emydid turtle from the Lower Pliocene of Okla- homa. By Edwin C. Galbreath. Pp. 265-280, 1 plate. August 16, 1948.
17. Pliocene and Pleistocene records of fossil turtles from western Kan- - saa and Oklahoma. By Edwin C. Galbreath. Pp. 281-284, 1
figure in text. August 16, 1948.
18. A new species of heteromyid rodent from the Middle OUgocene of northeastern Colorado with remarks on the skull. By Edwin C. Galbreath. Pp. 285-300, 2 plates. August 16, 1948.
19. Speciation in the Brazilian spiny rats (genus Proechimys, Family Echimyidae). By Joao Moojen. Pp. 301-406, 140 figures in text. December 10, 1948.
20. Three new beavers from Utah. By Stephen D. Durrant and Harold S. Crane. Pp. 407-417, 7 figures in text. December 24, 1948.
21. Two new meadow mice from Michoacan, Mexico. By E. Ray- mond Hall. Pp. 423-427, 6 figures in text. December 24, 1948.
22. An annotated check list of the mammals of Michoacan, Mexico. By E. Raymond Hall and Bernardo Villa R. Pp. 431-472, 5 figures in text. December 27, 1949.
23. Subspeciation in the kangaroo rat, Dipodomys ordii. By Hemy W. Setzer. / Pp. 473-573, 27 figures in text, 7 tables. December 27, 1949.
24. Geographic range of the hooded skunk. Mephitis macroura, with description of a new subspecies from Mexico. By E. Raymond Hall and Walter W. Dalquest. Pp. 575-580, 1 figure in text. January 20, 1950.
25. Pipistrellus cinnamomeus Miller 1902 referred to the genus Myotis. By E. Raymond Hall and Walter W. Dalquest. Pp. 581-590, 5 figures in text. January 20, 1950.
26. A synopsis of the American bats of the genus Pipistrellus. By E. Raymond Hall and Walter W. Dalquest. Pp. 591-602, 1 figure in text. January 20, 1950.
Index. Pp. 605-638.
(Complete) Mammals of Washington. By Walter W. Dalquest. Pp.
1-444, 140 figures in text. April 9, 1948.
1. The Avifauna of Micronesia, its origin, evolution, and distribution. By RoUin 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, 46 figures in text. June 29, 1951.
3. Phylogeny of the waxwings and allied species. By M. Dale Arvey, Pp. 473-530, 49 figures in t€xt, 13 tables. October 10, 1951.
*"*-*^fei?ds from the State of Veracruz, Mexico
BY
GEORGE H. LOWERY, JR., and WALTER W. DALQUEST
University of Kansas Publications Museum of Natural History
Volume 3, No. 4, pp. 531-649, 7 figures in text, 2 table* October 10, 1951
MAR 1 7 1952
UNIVERSITY OF KANSAS
LAWRENCE
1»61
^'felrSs^om the State of Veracruz, Mexico
BY
GEORGE H. LOWERY, JR., and WALTER W. DALQUEST
University of Kansas Publications Museum of Natural History
Volume 3, No. 4, pp. 531-649, 7 Bgures in text, 2 table* October 10, 1951
mi. tn?. zooL imm
MAR 1 7 1952
UNIVERSITY OF KANSAS
TAWKENCE 1961
UNIVERSITY OF KANSAS PUBLICATIONS
The University of Kansas Publications, Museum 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 op Natural History. — E. Raymond Hall, Chairman, 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. Durrant. Pp. 1-82, 1 figure in text. August 15, 1946.
2. The systematic status of Eumeces 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 two 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 Micro tus). By E. W. Jameson, jr.. Pp. 125-151, 4 figures in text. October 6, 1947.
8. The postnatal development of two broods of great homed owls (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. Lowery, 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. Raymond Hall and Bernardo Villa-R. Pp. 249-256, 6 figures in text. July 26, 1948.
(Continued on inside of back eover)
Birds from the State of Veracruz, Mexico
BY
GEORGE H. LOWERY, JR., and WALTER W. DALQUEST
T^
?' ^
Mi. CO^P. Z60L LiEPtARt
MAR I r ^'^^
%
University of Kansas Publications Museum of Natural History
Volume 3, No. 4, pp. 531-649, 7 figures in text, 2 tables October 10, 1951
UNIVERSITY OF KANSAS
LAWRENCE
1951
University of Kansas Publications, Museum of Natural History
Editors: E. Raymond Hall, Chairman, A. Byron Leonard, Edward H. Taylor, Robert W. Wilson
Volume 3, No. 4, pp. 531-649, 7 figures in text, 2 tables October 10, 1951
University of Kansas Lawrence, Kansas
PRINTED BY
FERD VOILAND. JR. . STATE PRINTER
TOPEKA. KANSAS
1951
23-3247
Birds from the State of Veracruz, Mexico
By GEORGE H. LOWERY. JR., AND WALTER W. DALQUEST
From the arid tropics in the north to the lush, steaming jungles in the south, and from the sand dunes and mangrove swamps of its Gulf shores to the top of its snow-capped, volcanic peak, Orizaba, the 400-mile length of Veracruz embraces a wide variety of habitats and diversity of life conditions. As a result no other state in the Republic of Mexico exceeds Veracruz in the richness of its avifauna. But, despite this fact, the only comprehensive treatment of the birds of this state is still an unpublished manuscript, "Ornithology of the Mexican State of Veracruz with an Annotated List of Birds," by Frederick W. Loetscher, Jr. (Thesis, Cornell University, 1941). In this voluminous manuscript, the author brings together records pub- lished up to 1940, as well as references to specimens in the major museums. Supplementing these data are Loetscher's notes on five months of intensive personal observation of the birds of the state in which time he covered much ground and visited many localities not studied ornithologically since the days of Salle, Sumichrast, de Oca, and others. Loetscher listed 586 species as definitely occur- ring in Veracruz, and this was a treatment based on an ultra-con- servative point of view as to what constitutes a valid state record. A number of species recorded in the literature as occurring in Veracruz were rejected from his faunal list.
Loetscher's manuscript, which we have been permitted to examine through the courtesy of the author and the Laboratory of Ornithol- ogy at Cornell University, has provided a very helpful background against which to evaluate certain of our own records. Reference has been made also to several excellent papers published since the completion of Loetscher's manuscript. These include publications by Brodkorb (1943 and 1948), Davis (1945), Traylor (1949), and Wetmore (1943).
The large number of discoveries and important distributional data brought to light by these papers show that Veracruz still is one of the richest fields in North America for avifaunal discoveries. The present work is primarily a report on three collections coming from widely scattered localities and represents neither a comprehensive survey of the avifauna of the state nor an exhaustive study of any
(533)
534 University of Kansas Publs., Mus. Nat. Hist.
one locality. It contains, however, a considerable amount of new distributional data.
In late 1945 Dalquest entered Mexico to collect zoological speci- mens for the University of Kansas Museum of Natural History. Headquarters were established at Potrero Viejo, Veracruz, in Feb- ruary, 1946, and work continued from that date, with interruptions of two to three months each year in the rainy season, until July, 1949. Although the time was spent principally in studying the mammals, collections were made also of fishes, amphibians, reptiles, and birds. A total of 1007 specimens of birds, including a large number of skeletons, was secured in the four seasons of field work. Also, in addition to the collection made by Dalquest, which is deposited at the University of Kansas Museum of Natural History, we have had at our disposal for study certain material from Vera- cruz that reposes in the Louisiana State University Museum of Zoology. This includes approximately 200 specimens collected in Veracruz in 1937 and 1938 by Mr. Dyfrig McHattie Forbes of Potrero Viejo, and 87 specimens taken by Lowery and Robert J. Newman, in 1937 and 1949, respectively.
These combined collections and accompanying field notes have provided distributional data relating to 297 species or 312 species and subspecies. For eighteen of these kinds of birds we have been able to find no previously published records for Veracruz. Other speci- mens were of species reported from Veracruz only many years ago, and the validity of some of these old records was in question. The specimens listed in the following accounts of species and subspecies, unless otherwise stated, are in the collection of the University of Kansas Museum of Natural History.
The nomenclature employed, except where explanations are made to the contrary, follows the 4th edition of the American Ornitholo- gists' Union "Check-list" (1931) or its various Supplements (1944- 1949), Peters' "Birds of the World" (1931-1948), or Cory, Hellmayr, and Conover's "Birds of the Americas" (1918-1949).
In this paper, Lowery is responsible for the identification of the specimens, the general organization of the material, and the nomen- clatural and taxonomic comments. Dalquest is responsible for the information dealing with the actual field work, such as natural history notes and most of the distributional facts, including the dis- cussion of life-zones and their application in the individual species accounts.
LowERY AND Dalquest: Birds From Veracruz, Mexico 535
ACKNOWLEDGMENTS
We are grateful to Mr. and Mrs. Dyfrig McHattie Forbes of Potrero Viejo, Veracruz, whose interest in the fauna of this state has continued over many years. Mr.. Forbes personally collected nearly 200 specimens of birds in Veracruz for the Louisiana State University in 1937 and 1938. Lowery was the guest of Mr. Forbes for several days in August, 1937, and Dalquest lived with Mr. Forbes and his family for four seasons of field work in Veracruz. It is a pleasure to name in the present report a new race of owl in honor of Mr. Forbes.
For assistance in various ways and for the loan of comparative material we are indebted to the following: John W. Aldrich, Allen J. Duvall, and the United States Fish and Wildlife Service; Emmet R. Blake, Melvin A. Tray lor, and the Chicago Natural History Museum; John Davis; Frederick W. Loetscher, Jr.; Robert J. New- man; Kenneth C. Parkes and the Laboratory of Ornithology of Cornell University ; James L. Peters and the Museum of Compara- tive Zoology; Charles G. Sibley; George M. Sutton; J. Van Tyne and the University of Michigan Museum of Zoology; Alexander Wetmore, Herbert Friedmann, and the United States National Museum. The University of Kansas Endowment Association spon- sored the major part of the field work. Finally, we are indebted to Professor E. Raymond Hall and Mr. Harrison B. Tordoff for per- mission to report upon the collections from Veracruz in their care, and for suggestions pertaining to the manuscript.
LIFE-ZONES OF VERACRUZ
The life-zone classification employed in this paper represents Dalquest's concepts entirely and is based largely on his observations of the mammals of Veracruz.
Merriam (1892) dealt only superficially with the life-zones as they pertain to areas south of the United States. He did not subdivide the Tropical Region, but other authors applying Merriam's princi- ples have done so. Goldman (1920) separated Panama into the Lower Tropical Life-zone, with arid and humid divisions, the Upper Tropical Life-zone, and the Temperate Life-zone. The latter is an inclusive term to cover the small, isolated areas that are higher than the Tropical life-zones but that are not identifiable in detail with the life-zones of the temperate region farther north. Dickey and van Rossem (1938) divided the Tropical Region of El Salvador into four parts: the Humid Upper Tropical, the Arid Upper Tropical,
536 University of Kansas Publs., Mus. Nat. Hist.
the Humid Lower Tropical, and the Arid Lower Tropical life-zones. Other authors have applied somewhat different terminology to the divisions of the Tropical Region. There is general agreement, however, that the Tropical Region is divisible into at least two major life-zones, and that locally the life-zones in turn may each be divisible into humid and arid parts. Where mountains or high- lands rise above the Tropical Region they present conditions some- what comparable to those found in the Austral and Boreal regions, and they have been grouped together by some authors under the name of the Temperate Zone.
Central Mexico consists of a plateau which, in those parts that adjoin Veracruz, is elevated six to nine thousand feet above sea level. This plateau is an arid region, and was mapped by Merriam, et al. (1910) as lying in the Upper and the Lower Sonoran life- zones. At the eastern edge of the tableland, the lip or rim of the plateau is raised in a series of hills or mountains, of which the Pico de Orizaba and Cofre de Perote are two. From the elevated crest of the rim, the land plunges down abruptly to the tropics. The drop from the Pico de Orizaba to the Lower Tropical Life- zone has an average ratio of one foot in six feet, or a drop of 4,200 meters in twenty-three kilometers. The average distance from the Lower Sonoran Life-zone of the Mexican Plateau to the Lower Tropical Life-zone, in central Veracruz, is approximately nine miles. Where the crest is low, the life conditions are essentially like those of the Transition Life-zone of the United States. Where mountains are encountered, the land may rise to levels where Cana- dian, Hudsonian, or Arctic-Alpine conditions are simulated. But these minor zonal areas are so anomalous, so compressed, so isolated, and of such minor geographic extent that the identification of any one life-zone in any area, save where the land is level for a short distance, is scarcely possible.
West of the main crest of the rim of the Mexican Plateau, the land drops less abruptly on the average, but the slope is still steep. Where the geographic limits of Veracruz extend westward past the western base of the mountains or hills that form the rim of the plateau, the extensive Lower Sonoran Life-zone of the Mexican plateau is reached. This occurs in two small areas only, one of which is north and west of the Cofre de Perote. The other is north of Tulancingo, Hidalgo, where a long arm of Veracruz projects westward across the plateau for some distance.
Approximately ninety per cent of Veracruz lies in the Tropical
LowERY AND Dalquest: Birds From Veracruz, Mexico 537
Region. It is with this region, and its bird fauna, that this paper is primarily concerned. The Tropical Region is clearly divided into two parts. One of these, the Upper Tropical Life-zone, includes the eastern slopes of the mountains forming the rim of the Mexican Plateau from an approximate upper average elevation of 5,500 feet to an approximate lower average elevation of 1,700 feet. The Upper Tropical Life-zone in Veracruz, because it exists principally on the eastern face of the abruptly dropping face of the Mexican Plateau, is a narrow belt separating the extensive Lower Tropical Life-zone from the higher zones of these same mountains. Also included mainly in this life-zone are the Tuxtla Mountains of southern Veracruz. The highlands of the Tuxtla Mountains and the Mexican Plateau both intercept the moisture-laden winds from the Gulf of Mexico, after their passage over the low, comparatively level Lower Tropical Life-zone.
The Upper Tropical Life-zone in Veracruz is not divisible into humid and arid parts. If further study of the Upper Tropical Life- zone of the Americas should show that the recognition of humid and arid divisions is practicable, the Upper Tropical Life-zone of Veracruz would probably be considered as belonging entirely to the humid division. The Upper Tropical Life-zone includes the cloud- hung upland forests and lower mountain slopes, where there is an abundance of water from springs as well as from mountain snows to the west, and where precipitation is great and humidity is high. Streams are clear, cold, and swift. Exposed outcrops of rock are numerous. The forest is high and lush with saprophytes and epi- phytes. The ground is usually well drained and covered with an understory vegetation.
At the lower edge of the Upper Tropical Life-zone, at an approxi- mate average elevation of 1,700 feet, the land levels off and descends gently to sea level at the Gulf of Mexico. This area in Veracruz is the Lower Tropical Life-zone. Division of this extensive lowland into humid and arid divisions is practicable. It should be pointed out that the divisions of the Lower Tropical Life-zone in Veracruz may not correspond to the same divisions of this zone used by other authors for areas farther south. In Veracruz the arid and humid divisions are easily discerned, but in their faunas there is little qualitative difference. The arid division of the Lower Tropical Life- zone consists of an extensive grassy plain, usually spoken of herein as the coastal plain, where the conditions are arid over a long, dry season. The plain is fairly level, with no exposed outcrops of country
538 University of Kansas Publs., Mus. Nat. Hist.
rock. Vegetation on the grass lands consists of thorny thickets and flat-topped trees with lacy foliage. Streams are deep and sluggish with muddy banks. Near the streams and rivers there are jungles, with typical jungle birds and mammals. The arid division of the Lower Tropical Life-zone extends southward (from the arm of mountains that reaches eastward from the Mexican Plateau, north of Jalapa), nearly to the Isthmus of Tehuantepec, skirting about the base of the Tuxtla Mountains. It exists as a nearly continuous unit, except where cut by a network of narrow bands of jungle along the major streams.
The humid division of the Lower Tropical Life-zone consists of the lowland jungles lying, in Veracruz, mostly under 500 feet in ele- vation. A considerable part of the northern third of Veracruz, north of the twenty-degree parallel, lies in this division, as does extreme southern Veracruz, south of the coastal plain and the Tuxtla Mountains. Little collecting of birds was done in the northern part of this area.
GAZETTEER OF LOCALITIES
The following place names and geographical features are those to which reference is made in this paper. The spellings employed are based principally on the American Geographical Society's "Map of Hispanic America on the Scale of 1:1,000,000 (Millionth Map)" and its accompanying Index (1944). A few place names, however, were found only by consulting the "Coordenadas geograficas y alturas en metros sobre el nivel del mar, de las cabeceras municipales de la Republica [de Mexico]," a publication issued by the Secretaria de la Economia Nacional (1944). Numbers in brackets identify the position of a locality on the accompanying map (Fig. 1).
Acultzingo.— Lat. 18° 42'; long. 97° 18'. Village on road from Puebla to
Cordoba [47]. Altotonga.— Lat. 19° 46'; long. 97° 13'. Village on road from Perote to
Tlapacoyan, Puebla [18].
Alvarado.— Lat. 18° 47'; long. 95° 45'. Village on coast [51].
Arroyo de la Piedra. — Lat. 18° 59'; long. 96° 20'. Arro3'o on road from Corboda to Veracruz [37].
Boca del Rio.— Lat. 19° 07'; long. 96° 07'. Village beside Rio Jamapa on
road from Cordoba to Veracruz [36]. Buena Vista.— Lat. 17° 38'; long. 95° 13'. Village on headwaters of Rio San
Juan in southern Veracruz [63]. Canada Blanca. — Same as Ojo de Agua. Cerro Gordo.— Lat. 19° 25'; long. 96° 42'. Village on road from Jalapa to
Veracruz [28]. Coatzacoalcos.— Lat. 18° 09'; long. 94° 25'. Also known as Puerto Mexico.
A city at mouth of the Rio Coatzacoalcos in southern Veracruz [59].
LowERY AND Dalquest: Birds From Veracruz, Mexico 539
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Fig. 1. Map of Veracruz showing the locahties that are numbered in the Gazetteer (pp. 538-543). The numerical sequence of localities is an arrangement whereby north takes precedence over south and west over east. 1. Laguna Cerro Pez. 2. Tampico Alto. 3. El Cepillo. 4. Ozuluama. 5. Potrero Llano. 6. Tuxpan. 7. San Isidro. 8. Tehuatlan. 9. Jacales. 10. Zacualpilla. 11. Papantla. 12. Gutierrez Zamora. 13. San Marcos. 14. Nautla. 15. ElJobo. 16. Tlapacoyan. 17. Jalacingo. 18. Altotonga. 19. Las Vigas. 20. La Joya. 21. Perote. 22. Valsequilla. 23. Limon. 24. Jalapa. 25. Cofre de Perote. 26. Coa- tepec. 27. Jico. 28. Cerro Gordo. 29. Teocelo. 30. Plan del Rio. 31. Puenta Nacional. 32. Mirador. 33. Tlacotepec. 34. Paso de San Juan. 35. Veracruz. 36. Boca del Rio. 37. Arroyo de la Piedra. 38. La Capilla. 39. Fortin. 40. Cordoba. 41. Potrero Viejo. 42. Orizaba. 43. Mecayucan. 44. Yanga. 45. Cuitlahuac. 46. San Juan de la Punta. 47. Acultzingo. 48. Omealco. 49. El Faro. 50. Piedras Negras. 51. Alvarado. 52. El Conejo. 53. Presidio. 54. Motzorongo. 55. Volcan San Martin Tuxtla. 56. San Andres Tuxtla. 57. Cosamaloapan. 58. Otatitlan. 59. Coatzacoalcos. 60. Minatitlan. 61. Jaltipan. 62. Jimba. 63. Buena Vista. 64. Jesus Carranza. For the location of localities mentioned in the text but not shown on the map, see Gazetteer of Localities.
540 University of Kansas Publs., Mus. Nat. Hist.
Coatepec— Lat. 19° 27'; long. 96° 57'. Town south of Jalapa [26].
Cofre de Perote. — Lat. 19° 30'; long. 97^ 09'. A volcanic peak approximately
fifteen miles west of Jalapa. [25]. Cordoba.— Lat. 18° 54'; long. 96° 56'. City in central Veracruz. [40]. Cosamaloapan. — Lat. 18° 22'; long. 95° 48'. A town on Rio Papaloapan [57].
Cuitlahuac— Lat. 18° 49' ; long. 96° 43'. Also known as San Juan de la Punta.
Town on road from Cordoba to Veracruz [45]. El Cepillo.— Lat. 21° 42'; long. 97° 45'. Village approximately forty-eight
miles south of Tampico, Tamaulipas [3]. El Faro. — Lat. 18° 45'; long. 96° 14'. Not on maps. A few houses beside
the Rio Blanco [49]. El Jobo.— Lat. 20° 01'; long. 97° 10'. Village on road from Teziutlan, Puebla,
to the Gulf of Mexico [15]. El Conejo.— Lat. 18° 43'; long. 95° 37'. A Wetmore locality (1943:217) [52]. Fortin.— Lat. 18° 54'; long. 97° 00'. Town on road from Orizaba to Cordoba
[39]. Gutierrez Zamora.— Lat. 20° 28'; long. 97° 06'. Town on road south of
Papantla [12]. Hacienda Mirador. — See Mirador. Hacienda Potrero. — Same as Potrero Viejo [41].
Isla.— Lat. 18° 03'; long. 95° 28'. A Loetscher locality situated on the rail- road between Cordoba and Jesus Carranza, at a point approximately 8.4 miles northwest of Jimba.
Jacales.— Lat. 20° 26'; long. 98° 27'. Village in the arm of Veracruz that ex- tends westward, north of Tulancingo, Hidalgo [9].
Jalacingo. — Lat. 19° 49'; long. 97° 19'. Village on road from Perote to Teziutlan, Puebla [17].
Jalapa. — Lat. 19° 32'; long. 96° 55'. City in central Veracruz, and capital of the state. [24].
Jaltipan. — Lat. 17° 58'; long. 94° 43'. A town on railroad from Jesus Carranza to Coatzacoalcos [61].
Jesus Carranza. — Lat. 17° 26'; long. 95° 01'. Also known as Santa Lucrecia. Town and railroad division point in central part of Isthmus of Tehuantepec in southern Veracruz [64].
Jico.— Lat. 19° 25'; long. 97° 00'. Village south of Jalapa [27].
Jimba. — Lat. 17° 56'; long. 95° 23'. Village on railroad from Cordoba to Jesus
Carranza [62]. La Capilla.— Lat. 18° 58'; long. 96° 19'. Village on road from Cordoba to
Veracruz [38]. La Cieba. — Lat. 18° 52'; long. 96° 51'. Area in cane field immediately west of
Potrero Viejo. La Joya.— Lat. 19° 36'; long. 97° 03'. Village on road from Perote to Jalapa;
not on maps. Approximately 3 miles south and 2 miles east of Las Vigas. La Pesca.— Lat. 18° 53'; long. 96° 49'. Village one mile northeast of Potrero
Viejo. Las Vigas. — Lat. 19° 39'; long. 97° 05'. Town on road from Perote to Jalapa
[19]. Laguna Cerro Pez. — Lat. 22° 10'; long. 98° 22'. Lake on boundary between
Veracruz and San Luis Potosi [1]. Limon. — Lat. 19° 30'; long. 97° 21'. Also known as San Antonio Limon.
Village on Veracruz boundary west of Perote [23]. Matacabresto. — Forbes locality; not on maps. Said to be below (east of)
Cuitlahuac. Mecayucan. — Lat. 18° 53'; long 96° 15'. Village on road from Cordoba to
Veracruz [43].
LowERY AND Dalquest: Birds From Veracruz, Mexico 541
Metlac. — Not on maps; a power house and canyon approximately three kilometers in a northerly direction from Fortin, near road from Cordoba to Orizaba.
Minatitlan.— Lat. 17° 59'; long. 94° 32'. Town in southern Veracruz, on rail- road from Jesus Carranza to Coatzacoalcos [60].
Mirador.— Lat. 19° 17' ; long. 96° 54'. Hacienda on road from Fortin to high- way between Jalapa and Veracruz [32].
Motzorongo.— Lat. 18° 39'; long. 96° 44'. Village on railroad from Cordoba to Jesus Carranza [54].
Nautla.— Lat. 20° 13'; long. 96° 46'. Village on Gulf of Mexico on road leading easterly from Teziutlan, Puebla [14].
Ojo de Agua.— Lat. 18° 56'; long. 96° 54'. Village, cave, and spring 6 miles northwest of Potrero Viejo.
Omealco.— Lat. 18° 45'; long. 96° 47'. Town on railroad from Cordoba to Jesus Carranza [48].
Orizaba.— Lat. 18° 51'; long. 97° 06'. City in central Veracruz [42].
Otatitlan.— Lat. 18° 11'; long. 96° 02'.^ Village beside Rio Papaloapan in southern Veracruz. Also called Otatlan [58].
Ozuluama.— Lat. 21° 40'; long. 97° 51'. Town in northern Veracruz, forty-eight miles south of Tampico, Tamaulipas [4].
Palma Sola. — Same as Arroyo de la Piedra.
Palo Gacho.— Lat. 18° 43'; long. 96° 20'. Not on maps. Situated above (west of) Piedras Negras.
Papantla.— Lat. 20° 27'; long. 97° 19". City in northern Veracruz [11].
Paraje Nuevo.— Lat. 18° 52'; long. 96° 52'. Village on railroad from Cordoba to Veracruz, 2 miles north of Potrero Viejo.
Paso del Macho.— Lat. 18° 59'; long. 96° 45'. Village on railroad from Cordoba to Veracruz, 9 miles approximately northeast of Potrero Viejo.
Paso de San Juan.— 19° 12'; long. 96° 12'. Village on road from Fortin to highway between Jalapa and Veracruz [34].
Paso del Toro.— Lat. 19° 03'; long. 96° 09'. Village on road from Cordoba to Veracruz, approximately 4 miles south and 2 miles west of Boca del Rio.
Pasa Nueva. — This locality was not found by us on any map; Allen (1904: 29) writes concerning a collection of mammals made in 1901 by E. A. Colburn that "Pasa Nueva" is "situated a short distance from Tlacotalpan, about sixty miles south of the city of Vera Cruz, in the low tropical coastal belt." Wetmore (1943: 216-217) refers to collections of birds made in part by the same collector in the same year as from "Paso Nuevo" and shows this place as being located on the Rio San Juan at latitude 17° 59' and longitude 95° U'. This is fifty-three air-line miles from Tlacotalpan and eighty-three miles south and east of Veracruz. We wonder if Allen may not have been more nearly correct than Wetmore, because there is a "Paso Nuevo" listed by Nava (Direccion General Correos y Telegrafos de los Estados Unidos Mexicanos, 1892: 203) as a rancho near Cosamaloapan. Presumably this Paso Nuevo is on or near the Rio Papaloapan instead of on the Rio San Juan. Cosamaloapan is only nineteen and a half miles in an air-line from Tlacotalpan and fifty-six and a half miles south-southeast of Veracruz.
Perote. — Lat. 19° 34'; long. 97° 14'. Town on the road extending from Jalapa
westward to Puebla boundary [21]. Piedras Negras. — Lat. 18° 46'; long. 96° 11'. Village on railroad from Veracruz
to Jesus Carranza [50].
Plan del Rio.— Lat. 19° 23'; long. 96° 36'. Village on road from Jalapa to Veracruz [30].
Potrero. — Lat. 18° 53'; long. 96° 47'. Also known as Potrero Nuevo. Village on railroad from Cordoba to Veracruz, approximately 1 mile north and 3 miles east of Potrero Viejo.
542 University of Kansas Publs., Mus. Nat. Hist.
Potrero Llano.— Lat. 21° W; long. 97° 43'. Village on road from Papantla to Tampico, Tamaulipas [5].
Potrero Viejo.— Lat. 18° 52'; long. 96° 50'. Village on railroad from Cordoba to Veracruz [41].
Presidio.— Lat. 18° 39^; long. 96° 46'. Village on railroad from Cordoba to Jesus Carranza [53].
Puenta Nacional.— Lat. 19° 19'; long. 96° 29'. Village on road from Jalapa to Veracruz [31].
Rincon Area. — Peninsula-like area west of Jimba, in southern Veracruz.
Rio Atoyac— From lat. 18° 58'; long. 96° 54', SE to junction with Rio Jamapa at lat. 18° 50'; long. 96° 40'. Some maps give the name Rio Atoyac to the lower part of the river, to which the name Rio Jamapa is applied in this account.
Rio Banderilla.— From lat. 19° 35'; long. 96° 55', draining eastward into the Rio Actopam.
Rio Blanco.— From lat. 18° 43'; long. 97° 18', east into Laguna Tlalixcoyan (a part of Laguna de Alvarado), at lat. 18° 45'; long. 95° 50'.
Rio Chalchijapa. River flowing across Oaxacan boundary at latitude 94° 46' (where it is called the Rio Alegro) and continuing northerly to Boca Chal- chijapa (lat. 17° 26'; long. 94° 50'), at confluence with Rio Coatzacoalcos.
Rio Coatzacoalcos. River flowing across Oaxacan boundary at lat. 17° 20' ; long. 94° 59^, and northeastward into the Gulf of Mexico at the city of Coatza- coalcos (lat. 18° 09'; long. 94° 25').
Rio Jamapa. — See Rio Atoyac.
Rio Metlac. — Tributary of Rio Blanco with junction at lat. 18° 50'; long.
96° 58'. Rio Solosuchi. — Also written Rio Solosuchil. A river flowing from boundary of
state at lat. 17° 14'; long. 94° 28', northwestward into Rio Chalchijapa at
lat. 17° 22'; long. 94° 47'.
Sala de Agua. — Lat. 18° 50' ; long. 96° 43'. A cave and village 2 miles south and 7 miles east of Potrero Viejo.
San Andres Tuxtla.— Lat. 18° 27'; long. 95° 13'. City at terminus of railroad spur running eastward to Tuxtla Mountains from the Cordoba-Jesus Car- ranza main line [56].
San Francisco de las Mesillas. Same as Yanga and San Lorenzo.
San Isidro.— Lat. 20° 56'; long. 97° 33'. Village west of Tuxpam, in northern Veracruz [7].
San Juan de la Punta.— Lat. 18° 45'; long. 96° 43'. Town on road from Cor- doba to Veracruz [46].
San Marcos.— Lat. 20° 12'; long. 96° 57'. Village on road from Teziutlan, Puebla, eastward to Gulf of Mexico [13].
Tampico Alto.— Lat. 22° 07'; long. 97° 48'. Village south of Tampico, Ta- maulipas [2].
Tehuatldn.— Lat. 20'° 43'; long. 97° 32'. Tihuatlan on maps. Village on road from Papantla to Tampico, Tamauhpas [8].
Teocelo.— Lat. 19° 23'; long. 96° 58'. Village south of Jalapa [29].
Tlacotepcc— Lat. 19° 11'; long. 96° 50'. Village on road from Fortin to high- way from Jalapa to Veracruz [33].
Tlapacoyan.— Lat. 19° 58'; long. 97° 13'. Village on road from Teziutlan, Puebla, eastward to the Gulf of Mexico [16].
Tuxpam. — Lat. 20° 57'; long. 97° 24'. City in northwestern Veracruz, beside Gulf of Mexico [6].
Valsequilla.— Lat. 19° 37'; long. 97° 11'. Village on road from Perote to Jalapa; not on maps [22].
Veracruz.— Lat. 19° 12'; long. 96° 08'. Large city on coast of the Gulf of Mexico [35].
LowERY AND Dalquesti Birds From Veracruz, Mexico 543
Volcan San Martin Tuxtla.— Lat. 18° 33'; long. 95° 13'. Volcanic peak in
Tuxtia Mountains [55]. Yanga.— Lat. 18° 50'; long. 96° 47'. Also known as San Lorenzo. A village on
road from Cordoba to Veracruz [44]. Zacualpilla.— Lat. 20° 25'; long. 98° 22'. Village in the arm of Veracruz that
extends westward, north of Tulancingo, Hidalgo [10].
Family TINAMIDAE
Tinamus major robustus Parker
Great Tinamou
Tinamus robustus Parker, Trans. Zool. Soc. London, 5, 1866: 205-232, pl. 34, fig. 8, pi. 39, pi. 40, figs. 1-7, pi. 41, figs. 1-3 (San Geronimo, Baja Verapaz, Guatemala.) C/. Brodkorb, Misc. Publ. Mus. Zool., Univ. Michi- gan, 55, January 30, 1943, pp. 20-21.
One 9. 20 km. E Jesus Carranza, 300 ft., April 14, 1948; 1 9 , 25 km. SE Jesus Carranza, March 30, 1949; 1 5 , 35 km. SW Jesus Carranza, February 18, 1948.
Perdiz grande, Perdiz real. This large tinamou was not found north of the Rio Coatzacoalcos, on the Isthmus of Tehuantepec, but it was abundant in the dense jungles south of the Coatzacoalcos. The species prefers the deep shade of the high forest where the dense crowns of the trees overhead so completely block the light that there is little understorj^ vegetation. The ground in such habitat is covered with dry leaves and shadows, and the gray colors of the tinamous blend with the dull surroundings. So well protected by their color are they that, in searching for them, one must look for the slight bobbing movement of the birds' heads rather than for the birds themselves. This species flushes far more readily than the smaller species of tinamous and, taking off with a roar of wings similar to that of the Ruffed Grouse, usually flies about 200 feet. On the three occasions that birds were seen to alight, they turned, just before landing, to face the direction from which they had come.
A nest of this species was found on April 4, 1949. Three violet eggs, almost round in shape (59x49 mm.) were placed at the base of one of the tall, sinuous, narrow roots of a wild fig tree in the jungle. This site was 100 feet from the shore of the Rio Solosuchi, thirty-five kilometers southeast of Jesus Carranza, at 350 feet elevation.
The nest consisted of a scarcely perceptible hollow in the dry leaves, about ten inches from the root. At this particular place there was a thin grove of slender, thorny ground palms, about twelve feet in height, surrounding the fig tree. Two birds were flushed here.
544 University of Kansas Publs., Mus. Nat. Hist.
Their presence in such relatively dense cover led to the investigation that disclosed the nest. The eggs were in an advanced state of incubation.
The mournful cry of this bird is heard most often in the late afternoon, but is commonly heard at any hour of the day or night. The natives say that the bird lives entirely on fireflys, a belief pos- sibly originating from its habit of calling at night.
The pale green flesh of this tinamou is tender and has a delicate flavor and, in the uninhabited southern jungles, formed one of the principal table meats of our field party. The birds were usually fat, and were more tender than the Guans and Curassows.
Crypturellus boucardi boucardi (P. L. Sclater) Boucard Tinamou
Tinamus boucardi P. L. Sclater, Proc. Zool. Soc. London, 1859: 391. (Teotalcingo, Oaxaca, Mexico.)
Two 5, 35 km. SW Jesus Carranza, 350-400 ft., February 18, 1948, and April 9, 1949; IS, 20 km. SE Jesus Carranza, 250 ft., May 5, 1949.
Perdiz chica. This smaller species was found in the same localities and habitat as Tinamus major, and nowhere else in Veracruz. It appeared to be less common than the larger species. Its mournful whistles were usually heard in the early morning and late afternoon. Rarely did it call at night. Except when chased by dogs, this bird refused to flush. When approached by man, it usually "freezes," and allows an approach to within ten to fifteen feet before dashing off into the jungle. The birds taken were seen only when attention was called to them by the rustling of leaves when the birds ran off, almost from beneath the collector's feet.
Twenty-five kilometers southeast of Jesus Carranza, on March 30, 1949, a nest containing two rounded (50x40 mm.) cream-colored eggs was found. The nest was a shallow depression in dry leaves be- neath the dense, tangled branches of a large fallen tree, in a small clearing in the jungle. The eggs were in an advanced state of incu- bation.
The flesh of this species, like that of the other tinamous, is a fine table meat.
One of these birds was found sleeping near midnight, fourteen kilometers southwest of Coatzacoalcos. Its retreat consisted of three or four dead coyole palm fronds that lay in a tangled mass on the ground.
LowERY AND Dalquest: Birds From Veracruz, Mexico 545
Crypturellus cinnamomeus sallaei (Bonaparte) Cinnamoneous Tinamou
Nothocernus [sic] sallaei Bonaparte, Compt. Rend. Acad Sci. Paris, 42, May, 1856: 954. (Cordoba, Veracruz, Mexico.)
One $, Rio Atoyac, 8 km. NW Potrero, March 3, 1946; 1 9 , 2 km. N Motzorongo, 1500 ft., December 8, 1946.
Perdiz. This tinamou is common in central Veracruz, both on the coastal plain and in the humid upland jungles. It prefers a far denser growth of plants for cover than the other tinamous in Vera- cruz. It was found most commonly in dense thickets, overgrown corn milpas, dense jungle along water-courses on the coastal plain, and in poorly-kept coffee groves. Although locally abundant, and highly desired as game, this species is so difficult to take that even the native hunters do not attempt it. When a hunter does obtain a shot at a perdiz, it is usually at very close range, and the shot charge is directed at the bird's legs in order to preserve the body flesh.
Family COLYMBIDAE
Colymbus dominicus brachypterus Chapman Least Grebe
Colymbus dominicus brachypterus Chapman, Bull. Amer. Mus. Nat. Hist., 12, 1899 : 256. (Lomita Ranch, Texas.)
One ^, 5 km. N Jalapa, 4500 ft., October 19, 1946; 1^,7 km. NW Potrero, 1700 ft., January 10, 1947; 1?, 3 km. N Presidio, 1500 ft., December 2, 1946.
This is the commoner grebe in the central part of the state. It was not noted on the rivers in the southern part of Veracruz, such as the Rio Coatzacoalcos, but was common on ponds and such rivers as the Rio Atoyac and Rio Blanco, even in their upper reaches where the water is cold and clear.
Podilymbus podiceps podiceps (Linnaeus)
Pied-billed Grebe
Colymbus Podiceps Linnaeus, Syst. Nat., ed. 10, 1, 1758: 136. Based on the Pie-Bill Dapchick, Podiceps minor, rostro vario Catesby, Carolina, 1, p. 91. (In America septentrionali = South Carolina.)
One 2 , 15 km. W Piedras Negras, 300 ft., January 18, 1947.
The specimen listed seems to be the first taken in Veracruz, though Loetscher (MS) saw the species on two consecutive dates at Isla.
546 University of Kansas Publs., Mus. Nat. Hist.
Family ANHINGIDAE
Anhinga anhinga leucogaster (Vieillot)
Water-turkey
Plotus leucogaster Vieillot, Nouv. Diet. Hist. Nat., nouv. ed., 1, 1816: 545. (Florida.)
One $ , 22 km. SSW Jesus Carranza, 300 ft., March 24, 1948.
Pato buzo. The Water-turkey is common on the broad lower reaches of the larger rivers of Veracruz. They were often seen sitting on dead stubs along the river banks or on driftwood, or swimming in pairs or small groups. In extreme southern Veracruz they were found far up the rivers, where the streams were swdft, cold, and clear.
Family ARDEIDAE
Butorides virescens virescens (Linnaeus)
Green Heron
Ardea virescens Linnaeus, Syst. Nat., ed. 10, 1, 1758: 144 (America; restricted to coast of South Carolina by H. C. Oberholser, Proc. U. S. Nat. Mus., 42, 1912, p. 534.)
One 5 and 1 5 (LSUMZ), Potrero Viejo, 1,700 ft., January 25 and April 20, 1938; 1 5 (skel.), Potrero Viejo, 1,700 ft., October 21, 1947; 1 $ (skel.), Rio Blanco, 20 km. W Piedras Negras, 400 ft., October 6, 1946.
Garcita. Green Herons are fairly common in the lowlands of Veracruz.
Butorides virescens anthonyi (Mearns) Green Heron
Ardea virescens anthonyi Mearns, Auk, 12, July, 1895: 257. (Seven Wells, Salton River, on the Colorado Desert, Baja California.)
One 9 (LSUMZ), Piedras Negras, December 19, 1937.
This speciman appears to be a typical example of the race anthonyi, the first to be recorded in Veracruz. The wing measures 195 mm., the under parts are pale, and the neck is decidedly fulves- cent.
Florida caerulea (Linnaeus) Little Blue Heron
Ardea caerulea Linnaeus, Syst. Nat., ed. 10, 1, 1758: 143. Based mainly on the Blew Heron, Ardea cacrtdea Catesby, Carolina, 1, p. 76. (In America septentrionaH = South Carolina.)
Four $ , Rio Atoyac, 8 km. NW Potrero, March 2 and 15, 1946; 19,15 (skels.), Rio Blanco, 20 km. W Piedras Negras, October 3 and 6, 1946; 1 2 (LSUMZ), Matacabresto, Rio Blanco above Piedras Negras, December 18, 1937.
Garza blanca. This seems to be the commonest heron in Vera- cruz. It occurs along the larger rivers, even in their upper reaches
LowERY AND Dalquest: Birds From Veracruz, Mexico 547
where the water is cold, clear, and swift, as well as near ponds and lakes. They are often seen singly, but more often are found in small groups. Three kilometers west of Boca del Rio, in February, 1946, these herons congregated in the evening to roost in the tall trees bordering the Rio Jamapa. The birds began to arrive at the roost about one hour before sundown, and the last individuals did not arrive until almost complete darkness had set in. Blue birds and white birds were about equal in numbers.
Natives commonly confuse white immatures of this species with the Snowy Egret. The Little Blue Heron is occasionally shot for food by hunters who fail to find more acceptable game, especially when an opportunity is presented to kill several birds with one shot. For the most part, however, they are scorned as game.
Casmerodius albus egretta (Gmelin)
American Egret
Ardca Egretia Gmelin, Syst. Nat., 1, pt. 2, 1789: 629. (In insula S. Dominici insulus Falkland et America australi ad Louisianam usque = Cayenne.)
One $ (LSUMZ), Potrero Viejo, February 27, 1938; 1 $, Matacabresto, NW Piedras Negras, December 19, 1937.
Garza blanca, garza real. The American Egret was found to be fairly common in marshes and along the larger rivers near their mouths. Occasional birds visited the temporary ponds that formed following hea\y rains.
This egret is still shot occasionally for its plumes, though there is no market for these feathers. In several native houses plumes were tacked on the walls as ornaments. These never seemed to be from more than one bird; possibly they came from an individual shot primarily for food.
Nyctanassa violacea violacea (Linnaeus)
Yellow-crowned Night Heron
Ardea violaceea Linnaeus, Syst. Nat., ed. 10, 1, 1758: 143. Based on the Crested Bittern, Ardea stellaris cristata americana Catesby, Carolina, 1, p. 79. (In America septentrionali = South Carolina.)
One $,5 km. SW Boca del Rio, 25 ft., October 12, 1946; 1 5 , 2 km. N Motzorongo, 1500 ft., December 10, 1946.
Candil. Fairly common along the lower reaches of the rivers. As many as three or four birds were seen on some piles of drift, but
2—3247
548 University of Kansas Publs., Mus. Nat. Hist.
usually the species was solitary. The night heron is an important food species in Veracruz, usually hunted from dugout canoes in the evening.
Heterocnus mexicanus mexicanus (Swainson) Tiger Bittern
Tigrisoma mexicana Swainson, in Murray's End. Geog., 1834: 1383 [Amer. ed., 3, 1839, p. 315, fig. 1034]. (Real del Monte, Hidalgo, Mc.xico.)
One 9, Puenta Nacional, 500 ft., October 22, 1946; 1 $,l 2 , Rio Blanco, 20 km. WNW Piedras Negras, May 28, 1946.
Cerrador. This species was common along the rivers of extreme southern Veracruz and less common along streams on the coastal plain of central Veracruz. On the ground these birds are difiBcult to see in the marsh grasses and dense streamside vegetation where they customarily stay. Along the rivers in southern Veracruz, however, they were found perched in trees along the river banks at midday. At the approach of our canoe, they would become alarmed and fly to another perch a few hundred yards farther along the river, to repeat this process when we again drew near. The birds were always solitary in their roosts, and we found about one bird to each mile of river bank. Occasionally several would be herded along ahead of the canoe. This was a source of great amusement to the Indian guides.
Family COCHLEARIDAE
Cochlearius cochlearius zeledoni (Ridgway)
Boat-billed Heron
Cancroma zeledoni Ridgway, Proc. U. S. Nat. Mus., 8, 1885: 93. (Ma- zatlan, Mexico.)
One unsexed, 20 km. S Jesus Carranza, January 6, 1948.
Candil. Common along the sluggish rivers of the southern part of the state. These birds were usually found in dense vegetation along the shores of the rivers, about fifteen feet from the ground. From one to five birds were usually seen. They could be found because the branches below the roost would be white with excreta. The species roosts in the daytime but is seen commonly along the shallows of the rivers in the evening. It is hunted regularly for food, and has the same local name as the night heron.
LowERY AND Dalquest: Birds From Veracruz, Mexico 549
Family CICONIIDAE
Mycteria americana Linnaeus
Wood Ibis
Mycteria americana Linnaeus, Syst. Nat., ed. 10, 1, 1758: 140. (In America calidiore = Brazil.)
Galemhou. A flock of approximately 300 individuals was seen on several occasions in the large march west of Boca del Rio. Small flocks were noted several times along the Rio Atoyac. All were perched in the tops of tall trees. This bird is much desired by natives as food, but is shy and war3^ On several occasions hunters were seen trying to stalk them but never with success.
Family THRESKIORNITHIDAE
Guara alba (Linnaeus)
White Ibis
Scolopax alba Linnaeus, Syst. Nat., ed. 10, 1, 1758: 145. (In America = South Carolina.)
One $ (skel.), 60 km. ESE Jesus Carranza, 450 ft., March 29, 1948. Pato coco. This species was found only once in the course of this field work.
Ajaia ajaja (Linnaeus) Roseate Spoonbill
Platalea Ajaja Linnaeus, Syst. Nat., ed. 10, 1, 1758: 140. (In America australi = Jamaica, from first citation.)
The Spoonbill was found only once, February 8, 1947, fourteen kilometers southwest of Coatzacoalcos.
Family PHOENICOPTERIDAE
Phoenicopterus ruber Linnaeus
Flamingo
Phoenicopterus ruber Linnaeus, Syst. Nat., ed. 10, 1, 1758: 139. (Africa, America, rarius in Europe = West Indies.)
One flock of approximately fifty birds was seen along the shore of a small lake between Jesus Carranza and Jimba on March 22, 1947.
550 University of Kansas Publs., Mus. Nat. Hist.
Family ANATIDAE
Anas discors Linnaeus Blue-winged Teal
Anas discors Linnaeus, Syst. Nat., ed. 12, 1, 1766: 205. (South Carolina.)
One $ , 25 km. ESE Jesus Carranza, 200 ft., March 27, 1949.
Canati. Migrant ducks and geese bear local names in southern Veracruz. The only species called pato, or duck, is the resident Muscovy,
Family CATHARTIDAE
Sarcoramphus papa (Linnaeus) King Vulture
Vultur Papa Linnaeus, Syst. Nat., ed. 10, 1, 1758: 86. (India occiden- talis = Surinam.)
One S, 63 km. ESE Jesus Carranza, 500 ft., April 12, 1948; 1 unsexed (skel.), 60 km. ESE Jesiis Carranza, 450 ft., March 29, 1948.
Rey nopo. The King Vulture seems not to occur north of the latitude of the Tuxtla Mountains, save perhaps as an accidental wanderer. Along the Rio Coatzacoalcos it is fairly common. One was seen near the town of Coatzacoalcos, feeding on the partially submerged carcass of a cow. Another was feeding on a dead gar on the riverbank, five kilometers northeast of Jesus Carranza. South of the Rio Coatzacoalcos, this is the commonest vulture, far outnumbering the Black Vulture in the deep jungles of the Isthmus of Tehuantepec. One to ten are in sight, soaring over the jungle, at almost all times of the day. A tapir, killed for a specimen, drew fifty or more of these birds to our camp. One King Vulture had its crop distended with the remains of a tree snake [Spilotes pidlatus mexicanus). These birds have a strong and unpleasant odor that does not seem to emanate solely from the carrion upon which they feed.
Coragyps atratus (Bechstein) Black Vulture
Vultur atratus Bechstem, Latham, Allg. Uebers, Vogel, 1, 1793: 655. St. Johns River, Florida, ex Bartram.)
Nopo, zopilote. The Black Vulture is abundant everywhere. In the cities and towns, these birds may be seen on the streets and in the markets much as domestic pigeons are seen in the United States. The vultures live principally on the carrion of large animals, and are rarely seen with the Turkey Vulture, but on one occasion
LowERY AND Dalquest: Birds From Veracruz, Mexico 551
two Turkey Vultures were seen to drive three Black Vultures from the carcass of an opossum. The Turkey Vultures soon left without eating much of the opossum, and the carcass was quickly cleaned by the Black Vultures. When carcasses of skinned animals were placed out in tall grass, Black Vultures began to arrive in a short time. The body of a monkey or coyote would be finished in an hour, and the sated birds would then retire to nearby trees to rest. Commonly their weight broke the branches on which they perched. The grass for five feet around the carcass was often flattened to the earth by the beating of the birds' wings. Twice this species was found feeding on the fruit of the coyole palm, eating the thin layer of pulp between the tough rind and the rock-hard seed.
These birds may kill other animals; twenty kilometers east of Jesus Carranza, Black Vultures were said to have killed and partly eaten a new-born calf.
Vultures that die are not eaten by other vultures or by mammals, thus leading the Indians to say that their flesh is poisonous. Dead vultures are eaten by insects, such as ants and fly maggots. In central Veracruz, it is said that the flesh of this vulture, especially the liver, is a cure for tuberculosis.
Cathartes aura (Linnaeus) Turkey Vulture
Viiltur A^ira Linneaus, Syst. Nat., ed. 10, 1, 1758: 86. (America cali- diore ; Veracruz, Mexico, substituted by Nelson, Proc. Biol. Soc. Washing- ton, 18, 1905, p. 124.)
Aura, zopilote real. The Turkey Vulture is common in the Upper Tropical Life-zone. It was often caught in traps baited with meat and set for carnivorous mammals. Some were taken in traps set beneath logs or in hollows at the bases of trees, where the baits were completely invisible from the air above.
This species was noted only once on the costal plain, where two were seen about the carcass of an iguana. The Turkey Vulture is never so domestic as the Black Vulture, and does not visit the cities and town in search of food as does the latter species.
Family ACCIPITRIIDAE
Elanus leucurus majusculus Banks and Penard White-tailed Kite
Elanus leucurus majusculus Bangs and Penard, Proc. New England Zool. CI., 7, 1920: 46. (San Rafael, California.)
One $, Mirador, 3,500 ft., December 8, 1947.
552 University of Kansas Publs., Mus. Nat. Hist.
Odontriorchis palliatus (Temminck)
Brazilian Kite
Falco palliatus "P. Max." Temminck, PI. Col., livr. 23, 1822: pi. 204. (Brazil and Guiana = Rio Peruhype, southern Bahia.)
One $ , 20 km. S Jesus Carranza, 300 ft., February 12, 1948.
Our single speciman has the wing 313 mm. in length (unflattened) and the under wing coverts are plain black, characters which up- hold Brodkorb's contention (1943:26-27) that 0. p. mexicanus Swann (Syn. Accipitres, ed. 2, 1922, p. 159; type locality, Tampico, Tamaulipas, Mexico) is a valid race. An adult male from Venezuela in the Louisiana State University Museum of Zoology has the wing only 312 mm., but the under wing coverts are noticeably barred.
Chrondrohierax uncinatus aquilonis Friedmann Hooked-billed Kite
Chrondrohierax uncinatus aquilonis Friedmann, Jour. Washington Acad. Sci. 24, 1934: 314. (Tamaulipas, Mexico.)
One $, Rio Blanco, 20 km. W Piedras Negras, October 2, 1946.
Buteo albonotatus albonotatus Kaup Zone-tailed Hawk
Buteo albonotatus Kaup, Isis, col. 329, 1847: 17. (Mexico.)
One 9,5 km. SW Boca del Rio, October 11, 1946.
So far as can be determined, this is the first record for the Zone-tailed Hawk in Veracruz.
Buteo platypterus platypterus (Vieillot) Broad-winged Hawk
Sparvius Platypterus Vieillot, Tabl. Encld. Meth., 3, 1823: 1273. (Pennsylvania = Schuylkill River, ex Wilson.)
One $ Rio Atoyac, 8 km. NW Potrero, March 3, 1946.
Buteo magnitrostris griseocauda (Ridgway) Gray-tailed Hawk
Rupornis magniroslris var. griseocauda Ridgway, Proc. Boston Soc. Nat. Hist., 16, 1873-1874 [= December, 1873]: p. 87 (in key), p. 88. (Mexico.)
One 2, Mirador, 3,500 ft., December 8, 1947; 1 $, Rio Atoyac, 8 km. NW Potrero, March 2, 1946; 2 $ and 2 ,5 (one skel.), Rio Blanco, 20 km. WNW Piedras Negras, May 16, 19, 29, and 31, 1946; 1 9 and 2 $ (skels.), Rio Blanco, 20 km. W Piedras Negras, September 30 and October 1 and 3, 1946; 1 9, 15 km., W Piedras Negras, 300 ft., January 13, 1947; 1 ? (skel.), 15 km. ESE San Juan de la Punta, 400 ft., September 27, 1946.
LowERY AND Dalquesti Birds From Veracruz, Mexico 553
Gavildn. This is the commonest hawk on the coastal plain of Veracruz. It is common also in the Upper Tropical Life-zone. Individuals are usually seen perched in isolated trees, from which they swoop down to catch the lizards upon which they feed. Usually they select a perch ten to thirty feet from the ground, and at in- tervals give their piercing, plaintive cry. The stomachs of several specimens were examined, and held only the remains of the common lizards, Ameiva sp. and Sceloporus variabilis.
Buteo nitidus plagiatus (Schlegel) Mexican Goshawk
Asturina plagiata Schlegel, Mus. Pays-Bas, 2, Asturinae, 1862: 1, note. (Veracruz, Mexico.)
Two $ (LSUMZ), Palma Sola, November 15 and December 18, 1937; 1 $ , Rio Atoyac, 8 km. NW Potrcro, February 25, 1946; 2 $ ,2 9,1 unsexed (4 skels.), Rio Blanco, 20 km. WNW Piedras Xegras, May 10, 14, 26, and 30, 1946; 2 9, Rio Blanco, 20 km. W Piedras Negras, October 1 and 3, 1946.
Gavildn. This hawk is abundant in central Veracruz. It is only slightly less common than the preceding species on the coastal plain. It was not seen in the southern part of the state.
The Mexican Goshawk, like B. magnirostris, feeds principally on lizards, and the stomachs of specimens examined contained many Ameiva, slender lizards about eight inches in length. This hawk commonly hunts from a low perch, like B. magirostris, but is seen soaring over clearings, apparently hunting on the wing.
Leucopternis albicollis ghiesbreghti (Du Bus) White Snake Hawk
Buteo ghiesbreghti Du Bus, Esq. Orni., livr. 1, 1845: pi. 1. (Hacienda Mirador, Veracruz.)
Two $ , 20 km. ENE Jesus Carranza, 200 ft., March 22 and April 28, 1949.
Gavildn bianco. This beautiful white hawk was found only in extreme southern Veracruz. It was not common, and was seen only in the deep, uninhabited jungles to the south of the Rio Coatza- coalcos. The species spends much of its time soaring in circles 200 or 300 feet in diameter and 100 feet or so above the tops of the jungle trees. The hawks seem to avoid passing over open places, such as rivers and sand bars. Only twice were individuals seen perched. Both were in open trees, approximately fifty feet from the ground. The stomach of one specimen held several lizards {Anolis and AiJieiva) and another held a small snake {Drymobius margaritiferus) . The species has a strong, unpleasant, skunk-like odor.
554 University of Kansas Publs., Mus. Nat. Hist.
Hypomorphnus urubitinga ridgwayi (Gurney) Ridgway Black Hawk
Urubitinga ridgwayi Gurney, List Diurn. Birds Prey, 1884 : 148. (Guatemala.)
One $ Rio Blanco, 20 km. WNW Piedras Negras, May 19, 1946; 1 $ (skel.), 3 km. SW Boca del Rio, 10 ft., December 18, 1947; 1 5 (skel.), 25 km. SE Jesus Carranza, 250 ft., March 29, 1949; 1 $ (skel.), SSE Jesus Car- ranza, 300 ft., May 8, 1949.
Aguila. The Ridgway Black Hawk seems to be rather uncommon in Veracruz ; it was seen only a few times on the coastal plain.
Concerning the specimen from three kilometers southwest of Boca del Rio field notes for December 18, 1947, read in part: "For the past two evenings, this hawk roosted in a large strangler fig tree, on an angular limb about thirty-five feet from the ground. Beneath the roost, on the ground, were many droppings composed of the remains of the exoskeletons of land crabs, especially a small, hard orange-colored variety. In one such pile of over 100 droppings only this particular land crab was detected. The stomach of the hawk also contained only these land crabs. Since this crab hides in burrows or under logs or rocks by day, the question arises as to how the hawk catches them."
Buteogallus anthracinus anthracinus (Lichtenstein) Mexican Black Hawk
Falco anthracinus Lichtenstein, Preis-Verz. Vogel Mex., 1830: 3 (Mexico.)
One 9 (skel.), Rio Blanco, 20 km. W Piedras Negras, October 2, 1946; 1 2 (skel.), 32 km. ESE Jesus Carranza, 350 ft., March 25, 1948.
Harpia harpyja (Linnaeus) Harpy Eagle
Vultur Harpyja Linnaeus, Syst. Nat., ed. 10, 1, 1758: 86. (Mexico, ex Hernandez.)
One unsexed, 38 km. SE Jesus Carranza, 450 ft., April 9, 1948.
Faisanero, aguila real. Although there are only two previous records from Veracruz for this species, both in the nineteenth cen- tury and both in the central part of the state (Loetscher MS) , the Harpy Eagle is actually not uncommon in extreme southern Vera- cruz. On one occasion a band of spider monkeys showed some alarm when one of these eagles passed over the tree in which they were feeding. Other than this, there was no evidence of their preying on monkeys, although both the spider monkey (Ateles 'pan) and the black howler monkey {Alouatta palliata) were common in the habi-
LowERY AND Dalquest: Birds From Veracruz, Mexico 555
tat of the Harpy Eagle. Those observed were feeding on large iguanas, and several were seen perched twenty or thirty feet from the ground with the body of a five- or six-foot iguana hanging over the limb beside them and securely held in one foot. Local Indians stated that the Harpy Eagle feeds at times on Guans and Curas- sows, and for that reason were called faisaneros.
Geranospiza nigra nigra (Du Bus) Blackish Crane Hawk
Ischnosceles niger Du Bus, Bull. Acad. Roy. Belg., 14, pt. 2, 1847: 102. (Mexico.)
One $ (skel.), 20 km. ENE Jesus Carranza, March 21, 1949.
Family FALCONIDAE
Herpetotheres cachinnans chapmani Bangs and Penard Laughing Falcon
Herpetotheres cachinnans chapmani Bangs and Penard, Bull. Mus. Comp. Zool., 62. 1918: 37. (Santa Luci, Rio Hondo, Quintana Roo.)
One 5 and 1 9 (skels.), Boca del Rio, 400 ft., October 11, 1946, and Decem- ber 8, 1948; 2 $ , Rio Blanco, 20 km. WNW Piedras Negras, 400 ft., May 19 and 26, 1946; 1 S (skel.), 19,1 unsexed, 20 km. W Piedras Negras, 400 ft., Sep- tember 24, October 6 and 11, 1946.
Pdjaro vaquero. This beautiful hawk is very common on the coastal plain and also in the dense jungle of the humid division of the Lower Tropical Life-zone of the southern part of the state. It was never seen nor heard in the Upper Tropical Life-zone. Its distinc- tive call is emitted in the morning and afternoon, frequently on moonlit nights, and less often at midday. Local people say that when the pdjaro vaquero calls from a dead tree, the next day will be clear; if it calls from a green tree, it will rain.
Three skins from Piedras Negras, listed above, have wings meas- uring 270, 271, and 273 mm., respectively. Brodkorb (1948a: 406- 410) regards populations of this species in Veracruz as intermediate between H. c. chapmani and H. c. excubitor. But Wetmore (1944: 37) does not recognize excubitor; instead he considers Mexican populations as highly variable in size without definite geographic correlation. Since our three specimens are nearly uniform and also well within the measurements assigned to the race chapmani, we are therefore using this name. Neither the opinions advanced by Brodkorb nor Wetmore appear conclusive, and the problem will have to be studied further with more material.
556 University of Kansas Publs., Mus. Nat. Hist.
Micrastur semitorquatus naso (Lesson) Collared Micrastur
Carnifex naso Lesson, Echo du Monde Savant, (6), 2, 1842: col. 1085. (Realejo, Nicaragua.)
One 9 , Rio Blanco, 20 km. WNW Piedras Negras, May 26, 1946; 1 $ im., 25 km. SE Jesus Carranza, 250 ft., March 29, 1949.
Juan de pe. Collared Micrasturs were found only in deep jungle, usually perched about twenty feet from the ground in leafy trees. They frequently allowed the hunter to approach within twenty feet or less before darting away. They were skillful in maneuvering in the dense, shaded jungle. Both of the specimens taken were shot in flight, while dodging through dense, low vegetation.
Native hunters state that they often shoot this hawk, which they mistake for a perched Chachalaca. The micrastur's dark colors and long tail could easily lead to such an error. The local Indians con- sider it to be a kind of owl because of its large eyes and shade-haunt- ing habits.
Daptrius americanus guatemalensis (Swann) Red-throated Caracara
Ibycter americanus guatemalensis Swann, Syn. Accip., ed. 2, 1921: 14. (Guatemala.)
One S , 15 km. SW Jimba, 750 ft., March 5, 1947.
This appears to be the first record for this species from Mexico. The specimen taken was one of several seen in high, dense jungle in the Rincon area of southern and western Veracruz. The birds were quite active, flying from tree to tree and soaring for short in- tervals just over the tree tops. They gave voice to a piercing scream that was most irritating to human ears. The specimen taken had many lizards {Anolis and Ameiva) in its stomach. The natives in the party stated that the bluish dust on the bird's feathers was a deadly poison, and refused to carry the specimen on the same horse with a Guan (Penelope) intended for food. The Red-throated Caracara was not seen in extreme southern Veracruz, but natives recognized the bird's description, and especially the imitation of its call.
Polyborus cheriway audubonii Cassin Audubon Caracara
Polybonis Audubonii Cassin, Proc. Acad. Nat. Sci. Philadelphia, 17, 1865: 2. (Florida.)
One 2, Rio Blanco, 20 km. WNW Piedras Negras, May 24, 1946; 1 5 , 20 km. S Jesus Carranza, 300 ft., February 14, 1948.
The Audubon Caracara is widespread on the coastal plain, but
LowERY AND Dalquest: Birds From Veracruz, IVIexico 557
not very common. They usually occur in pairs and often feed with the vultures on carcasses of dead animals. A few were seen in the extensive sugar cane fields near Potrero, at the lower edge of the Upper Tropical Life-zone.
Falco peregrinus anatum Bonaparte Duck Hawk
Falco Anatum Bonaparte, Geogr. and Corap. List, 1838: 4. (Great Egg Harbor, New Jersey, ex Wilson, Amer. Orni., 9, 1814, p. 120, pi. 76.)
One [$] (LSUMZ), Palma Sola, below Cuitlahuac, November 15, 1937.
This specimen, taken by ]\Ir. Dyfrig Forbes at Palma Sola, seems to be the first record of the Duck Hawk in Veracruz.
Falco femoralis septentrionalis Todd Aplomado Falcon
Falco femoralis septentrionalis Todd, Proc. Biol. Soc. Washington, 29, June 6, 1916: 98. (Fort Huachuca, Arizona.)
One ,J , 3 km. E San Andres Tuxtla, 1000 ft., January 18, 1948.
This species seems to be rare. A few were seen and only one was shot.
Falco sparverias sparverius Linnaeus Sparrow Hawk
Falco sparverius Linnaeus, Syst. Nat., ed. 10, 1, 1758: 90. Based on the Little Hawk, Accipiter minor Catesby, Carolina, 1, p. 5. (In America = South Carolina.)
One 9 , 5 km. N Jalapa, 4500 ft., October 19, 1946; 1 ,5 , Rio Atoyac, 8 km. NW Potrero, February 27, 1946; 19,7 km. NW Potrero, 1700 ft., January 7, 1947; 1 $,\ 9,1 [9], (all LSUMZ), Potrero Viejo, 1700 ft., March 3 and 10, 1938.
The Sparrow Hawk was common about open land wherever observations were made. In the state of Veracruz, this little hawk seems to be principally predaceous on birds. In the Upper Tropical Life-zone, Sparrow Hawks were observed chasing warblers and other small birds on several occasions. Two stomachs were ex- amined and each held only the remains of small birds.
Falco sparverius paulus (Howe and King) Sparrow Hawk
Ccrchneis sparverius pauhis Howe and King, Contri. N. Amer. Orni., 1, 1902: 28. (Miami, Florida.)
One $ , 5 km. N Jalapa, 4500 ft., October 19, 1946.
This male specimen with wing measuring 179 mm. and tail 114 mm., can hardly be regarded as anything other than paulus, a sub- species not previously recorded outside of the United States.
558 University of Kansas Publs., Mus. Nat. Hist.
Family CRACIDAE
Crax rubra rubra Linnaeus
Mexican Curassow
Crax ruber Linnaeus, Syst. Nat., ed. 10, 1, 1758: 157. (Western Ecuador; cj. Hellmayr and Conover, Publ. Field Mus. Nat. Hist., zool. ser., 13, pt. 1, no. 1, April 30, 1942, p. 130.)
Two 9, 32 km. SSW Jesus Carranza, 350 ft., March 26, 1948; 1^,19 (skel.), 55 km. SW Jesus Carranza, 400 ft., March 26, 1948; 1 5 , 60 km. SW Jesus Carranza, 450 ft., March 27, 1948.
Faisdn real. The Curassow is much hunted for food in Veracruz. North of the Tuxtla Mountains, in the northern three-fourths of the state, it has been almost exterminated. Even in southern Vera- cruz, wherever roads have been built into the previously inaccessible forests, for hauling out mahogany logs, this fine game bird is rapidly disappearing. It forms one of the principal game species wherever found. In the Rincon area, west of Jimba, great piles of feathers of Curassows and Guans were seen along the newly-built roads where previously few hunters had even been able to penetrate.
South of the Rio Coatzacoalcos, in the uninhabited jungles of extreme southern Veracruz, Curassows are abundant. On collecting trips up the Rio Solosuchi and Rio Chalchijapa, this species was observed daily. The birds were seen in pairs moving slowly along the open floor of the jungle, stopping often to look for possible enemies. When undisturbed, both males and females gave a low, hooting grunt at intervals of one minute or less. When frightened, they dashed off along the ground and only when closely pressed would they take wing. When flushed, they flew only to a nearby, low tree, and usually remained peering at their pursuer. The crops of specimens taken were never full. Stomachs contained princi- pally berries and small fruits, many of which were green, hard, and seemingly unripe.
No nests were found in Veracruz, but a downy chick was dis- covered on June 5, 1949. This chick, which later escaped, had a loud, piping call, which it uttered almost constantly. In spite of its loud call, it was extremely difficult to find, even by sharp-eyed Indians, so well did its warm, brown colors blend with that of the background of dead leaves.
Penelope purpurascens purpurascens Wagler Purplish Guan Penelope purpurascens Wagler, Isis, 23, 1830: col. 1110. (Mexico). One 9, 15 km. SW Jimba, 750 ft., March 5, 1947; 2 5 , 35 km. SW Jesus Carranza, 400 ft., February 17, 1948; 1^,19 (skels.), 63 km. ESE Jesus Carranza, 500 ft., March 30. 1948.
LowERY AND Dalquest: Birds From Veracruz, Mexico 559
Faisdn griton. The Purplish Guan has, like the Curassow, been hunted extensively throughout much of Veracruz. Where both birds are sought as game, the Guan seems slightly better able to survive than the Curassow. At least, in the Tuxtla Mountains and the Rin- c6n area, Guans are more common than Curassows, although in southern Veracruz, where they are little hunted, the two species seem to be about equally common.
The Guan is more arboreal than the Curassow. Although Guans were seen on the ground on a few occasions, they were far more often found in trees. They were not slow to fly, if in danger, but preferred to run off along the larger branches.
Ortalis vetula vetula (Wagler) Gray-headed Chachalaca
Penelope vetula Wagler, Isis, 23, 1830: col. 1112, (Mexico; Tamico, Tamaulipas, Mexico, designated as type locality by Miller and Griscom, Auk, 38, January, 1921, p. 46; corrected to "neighborhood of the City of Veracruz, Mexico," by Miller and Griscom, Auk, 38, July, 1921, p. 455.)
Two $, Rio Atoyac, 8 km. NW Potrero, February 24 and March 5, 1946; 1 S (LSUMZ), Potrero Viejo, January 25, 1938; 2 $ (one skel.), 2 $ , Rio Blanco, 20 km. WNW Piedras Negras, March 17 and May 11, 18, and 28, 1946; 1 5, 15 km. W Piedras Negras. 300 ft., January 16, 1946; 1 9, Jimba, 350 ft., March 2, 1947; I S , 25 km. ESE Jesus Carranza, 200 ft., March 27, 1949.
Chachalaca. This game bird is found throughout the tropical parts of Veracruz. It seems to be most abundant in the thicket-cov- ered lowlands of northern Veracruz, and in the low, open, thorny woods of the coastal plain. It is slightly less common in the tall forests of the Upper Tropical Life-zone; and in the deep jungles of the south, it was found only near streams.
In places where it is much hunted, the chachalaca becomes ex- tremely shy, slinking off at the least sign of human presence. Al- though the species is quite common near Potrero, the few specimens from this locality were obtained with great diflBculty.
In the thorny woods of the coastal plain, the resident peoples hunt this bird by driving. Two or three hunters are stationed in com- paratively open woods, while others circle out and away to a dis- tance of a kilometer, and then return slowly to their starting point, driving the birds ahead of them. The hunters stop at intervals, to chop on trees with machetes. The Chachalacas move ahead of the hunters by flying from tree to tree, but do not become frightened. The birds pause in almost every tree along the way, and when they pass the concealed hunters, they are shot. Several drives were ob- served, and one to three birds were killed on each drive.
560 University of Kansas Publs., Mus. Nat. Hist.
In southern Veracruz, Chachalacas are usually hunted from canoes, the birds being shot from trees and bushes along the bank. In the state of Veracruz as a whole, this bird is probably the most important single game species. Like the cottontail rabbit, it seems to hold its own in spite of steady, around-the-year, hunting. The birds are not sold to any great extent. The meat brings about one- half the price of a chicken, or three times the cost of a shotgun shell.
The Chachalaca usually calls from a perch considerable higher than the elevations otherwise visited. They call regularly in the early morning, shortly after dawn, throughout the year, and less regularly in the late afternoon or evening. Occasionally they are heard on bright, moonlit nights. The natives consider this a sign that bad weather is approaching.
Our material substantiates the conclusions of Wetmore (1943: 245-246) in that examples of this species from the Upper Tropical Life-zone of central Veracruz are intermediate between vetula and mccalli. In our three specimens from near Potrero, one has the tips of the lateral rectrices nearly white, whereas the other two have the tips brown as in examples from farther south in Veracruz.
Family PHASIANIDAE
Colinus virginianus pectoralis (Gould) Bob-white
Ortyx pectoralis Gould, Proc. Zool. Soc. London, 1842 [= 1843] : 182. (Mexico.)
One $ (LSUMZ), Potrero Viejo, 1700 ft, March 6, 1948.
Colinus virginianus godmani Nelson Bob-white
Colinus godmani Nelson, Auk, 14, 1897: 45. (Jaltipan, Veracruz, Mex- ico.)
Two $,29, Rio Blanco, 20 km. WNW Piedras Negras, May 11, 14, and 15, 1946; 2 ^ , 15 km. W Piedras Negras, 300 ft., January 15, 1947.
Cholina. The Bob-white is only fairly common on the coastal plain of Veracruz. Only rarely were its call-notes heard. Local people recognize this bird as a game species, but seemingly it is rarely hunted for food. The cost of ammunition is greater than the value of the meat of such a small bird. Probably Bob-whites are shot only when opportunity is offered to kill several with a single shell.
LowERY AND Dalquest: Birds From Veracruz, Mexico 561
Odontophorus guttatus (Gould) Spotted Wood Quail
Ortyx guttata Gould, Proc. Zool. Soc. London, 1837 [= 1838] : 79. ("Bay of Honduras.")
One $, 35 km. SE Jesiis Carranza, 400 ft., April 30, 1948; 1 $ (skel.), 38 km. SE Jesiis Carranza, 400 ft., April 30, 1948.
Totoloschoco. This jungle quail is locally common in extreme southern Veracruz. It was found only where there was open ground in the deep shade at the foot of limestone cliffs. In the early morning its call notes, from which the native name is derived, may be heard from numerous points along the cliffs. It was not heard at other times of the day.
This species was found only singly or in groups of four or less, and was always located by dogs. On every occasion the birds flew to low perches, ten to fifteen feet from the ground, and blinked their eyes stupidly. The natives captured our specimens by mak- ing a small vine into a noose and, with the aid of a slim pole, slip- ping the loop over the birds' heads. On some occasions, the quail ducked their heads out of the noose. If this occurred, and the birds seemed about to fly, the Indians barked like dogs, and the quail again settled down. The natives seemed more anxious to ob- tain the birds for pets than for food.
Family ARAMIDAE
Aramus guarauna dolosus Peters Limpkin
Ararmis pictus dolosus Peters. Occas. Papers Boston Soc. Nat. Hist., 5, 1925: 144. (Bolson, Costa Rica.)
One S (skel.), 20 km. ENE Jesus Carranza, 200 ft., May 16, 1949.
Pdjaro taniche. The Limkin is fairly common along the rivers in the jungles of southern Veracruz. Whenever these birds were seen the natives in the party usually laughed, and repeated the name given above. Presumably there is some amusing, but undis- closed, legend connected with this bird. Apparently the species is never eaten, but the Indians had no compunctions against shoot- ing a bird for a specimen.
The amount of white on the secondaries and the larger size are the critical characters distinguishing dolosus from pictus. Unfor- tunately, our single specimen, when examined as a carcass prior to being made into a skeleton, had all the primaries intact on the
562 University of Kansas Publs., Mus. Nat. Hist.
mani, but only two secondaries remained on one wing and none on the other. These, however, show much less white than is accredited to dolosus (in pictus the secondaries are virtually without white, save at their extreme base). The wing of our specimen measures 316 mm.; exposed culmen, 119; tarsus, 119. In size these measure- ments are somewhat intermediate between those of dolosus and pictus. Brodkord (1943:33) points out the same situation with respect to specimens from Tabasco and Campeche examined by him.
Family RALLIDAE
Rallus limicola limicola Vieillot Virginia Rail
Rallus limicola Vieillot, Nouv. Diet. Hist. Nat., 28, 1819: 558. (Penn- sylvania.)
One $ , Potrero Viejo, 1700 ft., November 26, 1948.
This seems to be the second locality record for this species in Vera- cruz. Chapman (1898:36) records four specimens taken by him near Jalapa.
Aramides cajanea mexicana Bangs Wood Rail
Aramides albiventris meoncanus Bangs, Amer. Nat., 41, 1907: 178 (in key) and 185. (Buena Vista, Veracruz, Mexico.)
One $,6 km. W Boca del Rio, 10 ft., February 23, 1946; 1^,29 (one skel.), 20 km. E Jesus Carranza, 300 ft., February 21, 1948.
Poposcala, poposcela. The Wood Rail is not common in central Veracruz. The specimen taken at Boca del Rio was seen to alight in a slim, dense-crowned tree, beside the Rio Jamapa at dusk. It perched in the top of the tree, fifteen feet from the ground, and could not be seen. Shouting and throwing sticks into the tree failed to make it fly. A tough vine hung down from the tree, but even shaking this did not cause the bird to take wing. The vine was then pulled down several feet, bending the tree, and when the vine was re- leased, the rail was catapulted into the air where it was shot.
In southern Veracruz this bird is abundant in marshy places and along rivers. In the early morning and late evening their loud calls, from which the native name is derived, drown out most other bird notes. The chorus is of short duration, usually about ten minutes. The alarm note is more rarely heard, and consists of a deep, menacing growl, similar to that of a large carnivore.
The Wood Rail is not much hunted for food in Veracruz, save in a few localities, as for example, near the city of Coatzacoalcos.
LowERY AND Dalquest: Birds From Veracruz, Mexico 563
Porzana Carolina (Linnaeus) Sora
Rallus caroliniLs Linnaeus, Syst. Nat., ed. 10, 1, 1758: 153. (Hudson Bay, Canada.)
One 5, 7 km. W Potrero Viejo, 1700 ft., October 26, 1946.
Sumichrast (1881:229) lists this species from Orizaba without citing any definite record. The specimen taken at Potrero appears to be the only other record for the species in Veracruz.
Family HELIORNITHIDAE
Heliornis fulica (Boddaert) Finfoot
Colymbus fulica Boddaert, Tabl. PI. Enl., 1783: 54. (Cayenne; based on Le Grebijoulque, de Cayenne Daubenton, PI. Enl., ix, pi. 893.)
One $, 35 km. SW Jesus Carranza, 400 ft., February 16, 1948; 1 $ (skel.), 22 km. ESE Jesus Carranza, 300 ft., March 23, 1948; 1 $ (skel.), 2 $ , 20-25 km. SE Jesus Carranza, March 28 and May 2, 1949.
Viudita. The Finfoot or Sun Grebe is common on the rivers of southern Veracruz, and slightly less so in central Veracruz, as, for example, on the Rio Atoyac. The natives say that it is called "Little Widow," because it is always alone. Indeed, the solitary habits of this species are worthy of mention. Never were more than one seen at a time. The Sun Grebes were surprisingly wary, as compared with other tropical birds. They rarely allowed a canoe to approach within fifty j'ards without showing evidence of alarm. They spent considerable time on land, and several were seen sitting on river banks or logs beside the rivers. These usually took to the water when alarmed, but if approached by a canoe or wounded, they swam to shore and were lost in the thickets fringing the rivers.
Family JACANIDAE
Jacana spinosa spinosa (Linnaeus) Mexican Jacana
Fulica sinnosa Linnaeus, Syst. Nat., ed. 10, 1, 1758: 152. (South Amer- ica, ex Edwards, Nat. Hist. Bds., p. 48, pi. 48 = Panama, suggested by Todd, Ann. Carnegie Mus., 10, 1916, p. 219. See also Brodkorb Misc. Publ. Mus. Zool. Univ. Michigan, no. 55, Jan. 30, 1943, p. 36.)
One ? , 3 km. N Presidio, 1500 ft., November 30, 1946; 1 5 , 20 km. S Jesus Carranza, 300 ft., February 10, 1948.
Jacanas were rarely seen in central Veracruz but were abundant on ponds and lakes in southern Veracruz. They were usually seen walking on floating vegetation, and rarely came near shore. 3—3247
564 University of Kansas Publs., Mus. Nat. Hist.
Family CHARADRIIDAE
Charadrius vociferus vociferus Linnaeus Killdeer
Charadrius I'ociferus Linnaeus Syst. Nat., ed. 10, 1758: 150. (North America = South Carolina, ex Catesby.)
Two $ (LSUMZ), Potrero Viejo, 1700 ft., November 20, 1938, and Decem- ber 20, 1937.
Family SCOLOPACIDAE
Actitis macularia (Linnaeus) Spotted Sandpiper
Tringa macularia Linnaeus, Syst. Nat., ed. 12, 1, 1766: 249. (Pennsyl- vania.)
One 5,4 km. WNW Fortin, 3200 ft., April 5, 1946; 1 $ , Rio Blanco, 20 km. WNW Piedras Negras, May 24, 1946.
Limnodromus scolopaceus (Say) Long-billed Dowitcher
Limosa scolopacea Say in Long, Exped. Rocky Mts., 1, 1823: 170. (Near Boyer Creek = Council Bluffs, Iowa.)
One $,l 9 , 20 km. S Jesus Carranza, 300 ft., March 21, 1948; 1 $,2 ? , all skels., 20 km. E Jesus Carranza, 200 ft., March 20, 1948.
The Dowitcher was present in flocks of hundreds on the banks of small ponds along the Rio Coatzacoalcos in April.
Capella gallinago delicata (Ord) Wilson Snipe
Scolopax delicata Ord, in reprint of Wilson, Amer. Orni., 9, 1825: CCXVIII. (Pennsylvania.)
One $ and 1 unsexed (LSUMZ), Potrero Viejo, 1700 ft., January 25 and March 10, 1938.
Erolia minutilla (Vieillot)
Least Sandpiper
Tringa -minutilla Vieillot, Nouv. Diet. Hist. Nat., nouv. ed., 34, 1819; 466. (Amerique jusq'au dela du Canada = Halifax, Nova Scotia.)
One 2 (LSUMZ), Potrero Viejo, 1700 ft.. May 25, 1938.
Family BURHINIDAE
Burhinus bistriatus bistriatus (Waglcr) Thick-knee
Charadrius bistriatus Wagler, Isis, 22, 1829: col. 648. (Mexico.)
One S , Rio Blanco, 20 km. W Piedras Negras, 400 ft., October 3, 1946.
A pair of these birds was seen on the open grasslands of the coastal plain. They were found in a low swale, and when first noticed were
LowERY AND Dalquest: Birds From Veracruz, Mexico 565
running swiftly along the ground. When one was shot, the other flushed with a whirr of wings. Two local natives who were present at the time stated that the species, for w^hich they had a name that was not recorded, is a regular but rare visitant in the area, and is not a food species.
Family LARIDAE
Larus delawarensis Ord
Ring-billed Gull
Larus Delawarensis Ord, in Guthrie, Geog., 2d Amer. ed., 2, 1815 : 319. (Delaware River, below Philadelphia.)
Strange to say there are no published records for the Ring-billed Gull in Veracruz. It occurs rather regularly in winter, however, on Laguna Cerro Fez, a small lake on the boundary line lying partly in the state of San Luis Potosi and partly in the state of Veracruz. Members of the Louisiana State University field party working in the state of San Luis Potosi have observed it there on numerous occasions, as for example, on February 25, 1948, flying back and forth across the lake.
Hydroprogne caspia (Pallas) Caspian Tern
Sterna caspia Pallas, Novi Comm. Acad. Petr., 14, 1770: p. 582, pi. 22. (Caspian Sea, South Russia.)
This species also has apparently not been recorded previously from Veracruz. On February 25, 1948, Robert J. Newman and Pren- tiss D. Lewis found two individuals, one of which they shot, on the Laguna Cerro Fez. These birds were flying up and down the lake and were seen on occasions to cross over landmarks known to lie in Veracruz.
Family COLUMBIDAE
Columba flavirostris flavirostris Wagler Red-billed Pigeon
Columba flavirostris Wagler, Isis, 1831: col. 519. (Mexico; restricted to state of Veracruz by van Rossem, Trans. San Diego Soc. Nat. Hist., 6, August 30, 1930, p. 198.)
One $, Rio Atoyac, 8 km. NW Potrero, February 28, 1948; 2 $, 1 5 (all skels.), Rio Blanco, 20 km. W Piedras Xegras, October 1 and 6, 1946.
Paloma mora. The Red-billed Pigeon is fairly common through- out the Tropical Region of Veracruz. It is an arboreal species, usually seen in flight or perched in the tops of tall trees, in flocks of five to twenty-five birds. It regularly visits certain sand bars along rivers to drink and obtain gravel for its crop. When certain favored
566 University of Kansas Publs,, Mus. Nat. Hist.
fruits, such as the nanchi, are ripe, these pigeons gather in large numbers. Near Piedras Negras, fully a thousand birds were seen in one afternoon, as single individuals and as flocks ranging up to one hundred individuals.
The paloma mora is shot for food whenever a hunter can be rea- sonably sure of killing several birds with a single shell. For the most part, however, single birds are not shot, for the value of the meat of one bird is less than the cost of the ammunition to kill it.
Columba speciosa Gmelin Scaled Pigeon
Columba speciosa Gmelin, Syst. Nat., 1, 1789: 783. (Cayenne, ex Daubenton, PI. Enl., pi. 213.)
One $ , 10 km. NW Minatitlan, 100 ft., February 4, 1947; 1 5 , 20 km. ENE Jesus Carranza, March 22, 1949.
Palonia real. This apparently rare pigeon may be more common than is generally believed. The two specimens were taken almost by accident. The one from near Minititlan was in a tall tree, fully seventy-five feet from the ground. A Chachalaca had been heard in the vicinity, and was being hunted. A small branch was seen to move in the leafy crown of the tree, and a shot, supposedly at a Chachalaca, brought down the pigeon that had been hitherto unseen and unsuspected. Another pigeon flew from the same tree. The specimen from near Jesus Carranza was in the lower part of a dense tree, perched about fifty feet from the ground. Attention was called to the tree by slight movements of twigs and leaves, but no bird was seen, even after considerable peering. When the motion again started, a shot was fired into the branch and one of these pigeons fell to the ground. Again, another flew from the same tree.
From this evidence, it would seem that speciosa is a rather seden- tary species, having a preference for dense trees and a reluctance to fly. Its color blends with the background so well that the bird is
easily overlooked.
Columba nigrirostris Sclater Short-billed Pigeon
Columba nigrirostris Sclater, Proc. Zool. Soc. London, 1859 [= 1860] : 390. (Oaxaca, Mexico.) Two $ (one skel.), 20 km. S Jesus Carranza, 300 ft., Februaiy 22 and 23, 1948; 1 S (skel.), 30 km. SSE Jesus Carranza, 300 ft.. May 8, 1949; 2 $ (skels.), 35 km. SE Jesus Carranza, 350 ft., April 4 and 8, 1949.
Bust-a-huei. This small, dark pigeon is abundant in extreme southern Veracruz. Save in the late morning and afternoon, it lives deep in the jungle, staying in the tops of tall trees. It is sedentary
LowERY AND Dalquest: Birds From Veracruz, Mexico 567
and rarely seen. The jungle may seem completely empty of pigeons until, suddenly, one bird will call a soft, rather mournful, "boost, boost-a-way." Abruptly the jungle seems to come alive with pigeons, as though one were calling from every tree. If the birds are close, a short, rather harsh "chirr" note is heard, following the call. In spite of the abundance of calling birds, it is almost impossi- ble to see them in the tall trees where they are perched, unless they flutter to another branch. Specimens shot were usually killed seventy-five or one hundred feet from the ground, so that the fall caused them to split open.
In the late morning and afternoon, the Short-billed Pigeons are seen along the shores of rivers, where they probably go to drink. At such times they are seen perched on low branches or in low trees beside the water. They choose places where the rivers are deep and still, where the low, fringing vegetation grows down to the water level and permits them to drink without alighting on the ground. None was ever seen on a sand or gravel bar.
Zenaidura macroura carolinensis (Linnaeus) Mourning Dove
Columba carolinensis Linnaeus, Syst. Nat., ed. 12, 1, 1766: 286 (America = Carolina, ex Catesby).
One $, Rio Blanco, 20 km. WNW Piedras Negras, March 17, 1947; 1 $ (LSUMZ), Matacabresto, above Piedras Negras, December 17, 1938.
Paloma. The Mourning Dove is an abundant winter resident on the coastal plain.
Zenaida asiatica (Linnaeus) White-winged Dove
Columba asiatica Linnaeus, Syst. Nat., ed. 10, 1, 1758: 163. Based mainly on The Brown Indian Dove Edwards, Nat. Hist. Birds, 2, p. 76. (In Indiis = Jamaica.)
Three $, 1 2 , all skcls., Rio Blanco, 20 km. W Piedras Negras, October 5 and 6, 1946.
Paloma con alas blancas. The White-winged Dove is an abundant winter resident on the coastal plain, rarely hunted by natives for food unless opportunity is offered to kill several with one shot. Sub- specific identification of the skeletons listed above was not practica- ble.
Scardafella squammata inca (Lesson) Inca Dove
Chamaepelia inca Lesson, Compl. Oeuvres Buffon, 20, 1847: 211. (Mexico [probably west coast].)
One $ (skel.), 4 km. W Paso de San Juan, 250 ft., December 15, 1947; 1 $ (skel.), 5 km. W Potrero, March 24, 1946; 1 9 , Rio Atoyac, 8 km. NW Potrero,
568 University of Kansas Publs., Mus. Nat. Hist.
February 21, 1946; 2 $ (skels.), 15 km. ESE San Juan de la Punta, September 28, 1946; 1 ,J , Rio Blanco, 20 km. WNW Piedras Negras, May 17, 1946.
Palomita. Fairly common in the extensive sugar cane fields about Potrero, and far more common on the coastal plain, these little doves are one of the few birds of the arid coastal plain that remain active in the intense heat of midday.
Columbigallina passerina pallescens (Baird) Ground Dove
Chamaepelia passerina? var. pallescens Baird, Proc. Acad. Nat., Sci., Philadelphia, [11], sig. 21-23, Oct.-Nov., 1859 [=Jan. 12, I860]: 305. (Cape San Lucas, Baja California.)
One $ (skel.), 15 km. ESE San Juan de la Punta, September 28, 1946; 2 S (skels.), 3 km. W Plan del Rio, 1000 ft., October 20, 1946; 2 9,2 ,$ , Rio Blanco, 20 km. WISTW Piedras Negras, March 17, 1946; 1 $ (skel.), 35 km. SE Jesus Carranza, 400 ft., February 17, 1948.
Palomita. This tiny dove is but slightly less common on the coastal plain than the Inca Dove.
Columbigallina talpacoti rufipennis (Bonaparte) Ruddy Ground Dove
Chamaepelia rufipennis "Gray" Bonaparte, Compt. Rend. Acad. Sci. Paris, 40, 1855: 22. (Martagena, Columbia.)
One $, Rio Atoyac, 8 km. NW Potrero, Februaiy 22, 1946; 1 5 , 5 km. W Potrero, 1700 ft., March 23, 1946; 1 5 , 20 km. S Jesus Carranza, 300 ft., March 20, 1948.
Palomita. The Ruddy Ground Dove is not uncommon in shady places and clearings near water in the Upper Tropical Life-zone.
Columbigallina minuta interrupta (Griscom) Plain-breasted Ground Dove
Chaemepelia minuta interrupta Griscom, Amer. Mus. Novit., no. 379, 1929; 4. (Secanquim, Guatemala.)
One 9 (LSUMZ), Potrero Viejo, July 1, 1938; 1 $, Rio Blanco, 20 km. WNW Piedras Negras, May 18, 1946.
Claravis pretiosa (Ferrari-Perez) Blue Ground Dove
Peristera pretiosa Ferrari-Perez, Proc. U. S. Nat. Mus., 9, 1886: 175. New name for Columba cinerea Temminck, 1811. (Brazil.)
Two $ ,2 9 , 20 km. E Jesus Carranza, 300 ft., February 6, 1948.
Tortolito. This dove was noted about houses and clearings along the Rio Coatzacoalcos, and on sandbars and islands in the Rio Solosuchi and Rio Chalchijapa.
LowERY AND Dalquest: Birds From Veracruz, Mexico 569
Leptotila verreauxi fulviventris (Lawrence) Verreaux Dove
Leptoptila fulviventris Lawrence, Ann. New York Acad. Sci., 2, 1882:
287. (Yucatan.)
Four $, I 9 , Rio Atoyac, 8 km. NW Potrero, February 20 and 25, March 1, 3, and 10, 1946; 1 9 (skel.), Potrero Viejo, 1700 ft., December 16, 1948; 1 S (LSUMZ), Paraje Nuevo, 1700 ft., February 2, 1938; 2 $ (one skel.), Rio Blanco, 20 km. WNW Piedras Negras, May 11 and 18, 1946; 1 9 , 15 km. W Piedras Negras, 300 ft., January 15, 1947; 1 $ (skel.), 20 km. ENE Jesus Carranza, 350 ft., April 5, 1949; 2 9 (skels.), 20 km. E Jesus Carranza, 300 ft., February 10 and March 20, 1948.
Paloma. In the Upper Tropical Life-zone we found this common species of dove shy and difficult to obtain. They remained in dense cover on the hillsides, calling continually from concealed perches near the ground. As the hunters approached they would flush with much fluttering and move to another concealed perch. On the coastal plain they were confined to the brushy areas and dense jungle fringing arroyos and streams. Here the species was less shy and could easily be obtained by walking along the sandy beds of the arroyos and shooting the doves that flushed from the banks.
The specimens from central Veracruz are mostly intermediate between fulviventris and angelica, with some examples being indis- tinguishable from one or the other subspecies. The series as a whole, however, is so nearly intermediate that its assignment to fulviventris is arbitrary, AVhether the population at Jesus Carranza is more nearly typical of this subspecies than of angelica cannot be ascertained from the material available to us,
Leptotila plumbeiceps plumbeiceps (Sclater and Salvin)
Plumbeous-headed Dove
Leptoptila plumbeiceps Sclater and Salvin, Proc. Zool. Soc. London, 1868: 59. ([Choctum], Vera Paz, Guatemala.)
One 9 (LSUMZ), Potrero Viejo, August 22, 1938; 1 S (LSUMZ), Paraje Nuevo, 1700 ft., February 2, 1938; 1 5 , Rio Atoyac, 8 km. NW Potrero. March 12, 1946.
Paloma. This dove seems to be rare in Veracruz. Its habits seem much like those of L. verreauxi, but L. plumbeiceps was found in denser vegetation, such as deep thickets, in the Upper Tropical Life- zone near Potrero.
Family PSITTACIDAE
Ara macao (Linnaeus)
Red and Blue Macaw
Psittacus Macao Linnaeus, Syst. Nat., ed. 10, 1, 1758: 96. (South America.)
One 3,1 9 , 35 km. ENE Jesus Carranza, 350 ft., March 18, 1947.
570 University of Kansas Publs., Mus. Nat. Hist.
Guacamayo. This red, yellow, and blue macaw w^as found only in extreme southern Veracruz. It seems to be irregular in its occur- rence. In March and April, 1948, one to three pairs were seen al- most daily in trips that one of us (Dalquest) made up the Rio Chalchijapa and Rio Solosuchi. In the same area in March and April of 1949 only one pair was heard and none was seen.
A nest of this species was found in deep jungle south of the Rio Coatzacoalcos on March 18, 1947. It was in a cavity in the stub of a dead tree. The opening of the cavity, as seen from below, was approximately two feet in diameter and thirty-five feet from the ground. Eggs or young birds were present, for the parents returned to the nest after being frightened away, even when the hunters were at the foot of the nest stub.
In its native habitat, the deep jungles, this bird is indescribably attractive. As viewed against a background of green jungle, bro- meliads, and vines, its bright colors seem softened and natural. In captivity, the colors of the birds seem so gaudy as to be in bad taste.
The Macaw in Veracruz is shot for food. The flesh is tender and well flavored. The birds are ordinarily so shy, however, that prob- ably very few are ever obtained by native hunters.
Aratinga holochlora holochlora (Sclater) Green Parakeet
Comirus holochlorus Sclater, Ann. and Mag. Nat. Hist., (3), 4, 1859: 224. (Jalapa, Veracruz, Mexico.)
One 2,4 km. WNW Fortin, 3200 ft., April 6, 1946; 1 5 , Rio Atoyac, 8 km. NW Potrero, February 18, 1946; 1 $ (skel.), 1 5 , Rio Blanco, 20 km. W Piedras Negras, 400 ft., October 6, 1946.
Pariko. This is the only parakeet that was found in the Upper Tropical Life-zone.
Aratinga astec astec (Souance) Aztec Parakeet Comirus astec Souance, Rev. et Mag. Zool., (2), 9, 1857: 97. (Mexico.)
One 5 , 4 km. E Papantla, 400 ft., November 9, 1947; 2 $ (one skel.), 1 $ , 15 km. ESE San Juan de la Punta, 400 ft., September 27, 1946; 1 <^ , 2 $ (one skel.), Rio Blanco, 20 km. WNW Piedras Negras, May 26 and 27, 1946; 2 $, 1 9, all skels., Rio Blanco, 20 km. W Piedras Negras, October 4 and 6, 1946; 1 <5, 15 km. W Piedras Negras, 300 ft., January 14, 1947; 1 $ (LSUMZ), El Faro, above Piedras Negras, November 18, 1938; 1 5 and 1 9 (LSUMZ), Palo Gacho, above Piedras Negras, 350 ft., December 19, 1937; 1 9 , 20 km. E Jesus Carranza, 300 ft., February 21, 1948.
Pariko. This species was not noted in the Upper Tropical Life- zone, but is common in the arid division of the Lower Tropical Life-
LowERY AND Dalquest: Birds From Veracruz, Mexico 571
zone. When the nanchi fruit is ripe, both this parakeet and Aratinga holochlora gather in great flocks, although the Aztec Parakeet is by far the more numerous. This season is known as the "time of the nanchi," or the "time of the parakeets." At other seasons of the year they were found feeding on the pods of the tamarind tree, which were eaten when very green. Young birds of this species are often offered for sale in the markets.
Pionus senilis senilis (Spix) White-crowned Parrot
Psittacus senilis Spix, Aves Brazil, 1, 1824: p. 42, pi. 31, fig. 1. (No locality; Veracruz, Mexico, designated by Griscom, Amer. Mus. Novit., no. 379, 1929, p. 6.)
One 9,5 km. S Tehuatlan, 700 ft., November 12, 1947; 1 5 , 35 km. SW Jesus Carranza, 400 ft., February 16, 1948.
Pariko real. The White-crowned Parrot was found to be rather uncommon in Veracruz. In the southern part of the state, a few pairs were noted in low trees along the banks of the Rio Coatza- coalcos, and one flock of about ten birds was seen in a zapote mamay tree. In northern Veracruz, as at Tehuatlan, the species is more common, and a number of flocks of six to ten birds was seen. The species is less noisy than most other parrots. There seems to be no commercial traffic in the young of this species.
Amazona viridigenalis (Cassin)
Red-crowned Parrot
Chrysotis viridigenalis Cas.sin, Proc. Acad. Nat. Sci. Philadelphia, 6, 1853: 371. (South America = northeastern Mexico.)
One ^, 9 km. NW Nautla, 10 ft., November 6, 1947; 1 ?, Potrero Llano, 350 ft., February 15, 1949.
Catorro. The Red-crowned parrot is the least abundant of the three common, large parrots of the genus Amazona found in central and southern Veracruz {viridigenalis, autumnalis, and ochrocep- hala). Unlike the other species, viridigenalis gathers in large flocks, of from twenty to one hundred birds in which pairing is not always evident. The large flocks are active throughout the day, usually resting or feeding in the tops of two or three adjoining trees, for varying lengths of time, before taking flight and wheeling about in a compact mass, to settle shortly on other trees, meanwhile scream- ing.
The Red-crowned Parrot was most often seen in northern Vera- cruz, where the jungle is low and dense. It was never seen in the Upper Tropical Life-zone, and was noted only a few times on the coastal plain of central Veracruz.
572 University of Kansas Publs., Mus. Nat. Hist.
Amazona autumnalis autumnalis (Linnaeus) Golden-cheeked Parrot
[Psittacus] autumnalis Linnaeus, Syst. Nat., ed. 10, 1, 1758: 102. (West Indies := southern Mexico.)
One $,19, Rio Atoyac, 8 km. NW Potrero, February 28 and March 14, 1946; 1 $ (LSUMZ), Matacabresto, above Piedras Negras, December 17, 1937.
Loro. This is the common parrot in the Upper Tropical Life-zone. Golden-cheeked Parrots were active in the morning and evening and to a lesser extent on dull or rainy days. They were usually found in pairs and were observed feeding in the tops of the taller trees. They were shy and difficult to obtain. One common food item is the fruit locally called "bull's testicles." Their call, when in flight, is a characteristic "clankety-clank-clank," distinctly more metallic in quality than the call of the Yellow-headed Parrot.
Amazona ochrocephala oratrix Ridgway Yellow-headed Parrot
Amazona oratrix Ridgway, Man. No. Amer. Birds, 1887: 587. (Petapa, Oaxaca, Mexico.)
One 9 , Rio Blanco, 20 km. WNW Piedras Negras, May 24, 1946; 15,15 (skel.), 20 km. E Jesus Carranza, 300 ft., February 14, 1948.
Catorro, Loro. This species is common in the arid division of the Lower Tropical Life-zone. It is usually seen in pairs and is com- monly found feeding in trees thirty-five feet or less in height. The mango is a favored food. Damage is done to bananas when the fruit is green.
On March 4, 1947, on the coastal plain about ten kilometers south- west of Jimba, literally hundreds of Yellow-headed Parrots were flying from their feeding grounds in the jungles of the humid division of the Lower Tropical Life-zone, to their roosting grounds on the coastal plain. For an hour before dusk, from a few to a hundred parrots were in sight at all times, in pairs and small flocks, all flying to the eastward.
Amazona farinosa guatemalae (Sclater) Blue-crowned Parrot
Chrysotis guatemalae "Hartlaub" Sclater, Ibis, 1860: 44. (Guatemala and Honduras.)
One 5 , 20 km. S Jesiis Carranza, 300 ft., February 10, 1948.
This species was found but once. In the jungle along the Rio Coatzacoalcos, an Indian guide showed the collector two parrots in a tall tree in deep forest. The birds were motionless, and re- mained so for more than ten minutes before the collector could
LowERY AND Dalquest: Birds From Veracruz, Mexico 573
discern them in the green vegetation. One was shot. The Indians had not seen this species before. Also, the species was not recorded by Wetmore or Brodkorb in the course of their work in the state.
Family CUCULIDAE
Coccyzus americanus americanus (Linnaeus) Yellow-billed Cuckoo
Cuculus americanus Linnaeus, Syst. Nat., ed. 10, 1, 1758: 111. (In Carolina = South Carolina, ex Catesby.)
One 9 , Rio Blanco, 20 km. WNW Piedras Negras, May 13, 1946.
The wing of this specimen measures 141 mm., and makes it refer- able, therefore, to the nominate subspecies.
Piaya cayana thermophila P. L. Sclater Central American Squirrel Cuckoo
Piaya thermophila P. L. Sclater, Proc. Zool. Soc. London, 1859: 368 (Jalapa, Veracruz, Mexico.)
One $ (LSUMZ), 3.5 mi. S Jalapa, May 28, 1949; 1 9 (skel.), Puenta Nacional, 500 ft., October 21, 1946; 1 5 and 1 9 (LSUMZ), Potrero Viejo, January 3, 1938; 1 5 and 1 9 (LSUMZ), Paraje Nuevo, August 4, 1938, and November 2, 1937; 1 9 (skel.) Rio Atoyac, 8 km. NW Potrero, March 3, 1946; 5 5,3 9 , Rio Atoyac, 8 km. NW Potrero, February 26— March 10, 1946; 1 $ , 2 9 (skels.), Rio Blanco, 20 km. WNW Piedras Negras, May 12-31, 1946; 1 9 (skel.), Rio Blanco, 20 km. W Piedras Negras, October 1, 1946; 1 9 , 10 km. NW Potrero, February 16, 1946; 19,2 km. N Motzorongo, 1500 ft., December 10, 1946.
Canela. This is a common species in the Upper Tropical Life- zone. It is far less common in the arid division of the Lower Tropi- cal Life-zone.
Crotophaga sulcirostris sulcirostris Swainson Groove-billed Ani
Crotophaga sulcirostris Swainson, Philos. Mag. (n. s.), 1827: 440. (Mexico.)
One $,2 9, Rio Atoyac, 8 km. NW Potrero, March 11 and 21, 1946; 1 $, 10 km. NW Potrero, February 16, 1946; 1 $ (skel.), 15 km. ESE San Juan de la Punta, 400 ft., September 27, 1946; 1 5 and 1 9 (LSUMZ), Potrero Viejo, 1700 ft., August 20 and December 28, 1937; 4 ^ , 3 9 , all skels., Rio Blanco, 20 km. WNW Piedras Negras, May 25, 27, and 28, 1946.
Pijul, garrapatero. The Groove-billed Ani is abundant through- out the Tropical Region of Veracruz. It was absent only in the deep jungles of the extreme southern part of the state. The species prefers an open habitat, with dense, low brush for protection. The native name, pijul, is derived from its call, "pee-hool." The name
574 University of Kansas Publs., Mus. Nat. Hist.
garrapatero is derived from its habit of associating closely with cattle, and riding about on the backs of cows, presumably in search of engorged ticks. At night, the anis retire to dense bushes or low trees to roost. On a number of occasions, they were found roosting in low orange trees.
Tapera naevia excellens (Sclater) Striped Cuckoo
Diplopterus excellens Sclater, Proc. Zool. See. London, 1857 [= 1858] : 229. (Southern Mexico.)
Two $, 15 km. ESE San Juan de la Punta, 400 ft., September 28, 1946.
This cuckoo is probably fairly common, but it is so secretive that it is rarely seen. The few individuals observed were found in dense brush on the coastal plain.
Family TYTONIDAE
Tyto alba pratincola (Bonaparte) American Bam Owl
Strix Pratincola Bonaparte, Geog. and Comp. List, 1938: 7. (Northern parts [of America] = Pennsylvania.)
One $ (LSUMZ), Potrero Viejo, 1700 ft., April 5, 1938.
This specimen is virtually indistinguishable from light-phased examples from the United States, thereby excluding the possibility of its being referable to Tyto alba guatemalae.
Family STRIGIDAE
Pulsatrix perspicillata saturata Ridgway Spectacled Owl
Puhatrix perspicillata saturata Ridgway, Bull. U. S. Nat. Mus., 50, pt. 6, 1914: 758. (Santo Domingo, Oaxaca, Mexico.)
One $, 2 km. N Motzorongo, 1500 ft., December 6, 1946.
Only two individuals of this large owl were seen in Veracruz in the course of this study. One was shot at night by the light of the moon, when it perched forty feet from the ground in a tree. The other was seen at midday, in a dim, shadowed, marshy place in deep jungle southwest of Jimba. It was sitting about thirty-five feet from the ground, on a moss-covered limb, about six inches in diameter and slanted at a forty-five degree angle. The owl held its body vertical, seemingly a difficult stance for it to maintain. The position of the owl so close to the limb, however, made it difficult to see.
LowERY AND Dalquest: Birds From Veracruz, Mexico 575
Glaucidium brasilianum ridgwayi Sharpe Ferruginous Pygmy Owl
Glaucidium ridgwayi Sharpe, Ibis, ser. 3, 5, Januarj^ 1875: 55. (Cen- tral America = Mexico.)
One $, 8 km. NW Potrero, 1500 ft., February 13, 1946; 2 S, Z 9 , Rio Atoyac, 8 km. NW Potrero, February 21— March 15, 1946; 1 9 (LSUMZ), Paraje Nuevo, December 15, 1937; 2 5 , Rio Blanco, 20 km. WNW Piedras Negras, May 11 and 12, 1946; 1 S , 15 km. W Piedras Negras, 300 ft., January
15, 1947; 1 $ (skel.) Rio Blanco, 20 km. W Piedras Negras, October 6, 1946, 1 5 , 20 km. ENE Jesus Carranza, 200 ft., March 26, 1949.
Tincalero. These tiny owls are common throughout the Tropical Region of Veracruz. Their monotonous whistle is heard regularly in the early morning, just at dawn, and in the evening, and may be noted occasionally at almost any hour of the day. Their call was never heard at night. In fact, the first call of the Wood Owl usually silenced all Pygmy Owls in the vicinity, and they were not heard again until the following morning. The species must be present in Veracruz in great numbers. At almost any thicket or the edge of almost any jungle, at any time of day, an imitation of their almost interminable short, whistled notes would cause one or more birds to answer. The stomachs of most of the specimens listed above were examined, and found to contain remains of scorpions and insects, principally Orthoptera.
Speotyto cunicularia hypugaea (Bonaparte) Burrowing Owl
Sfrix hypugaea Bonaparte, Amer. Orni., 1, 1825: 72, note. (Western United States =: Plains of the River Platte.)
One 9 (LSUMZ), Paraje Nuevo, 1800 ft.. May 17, 1938; 1 9 , 15 km. W Piedras Negras, 300 ft., January 18, 1947.
The specimen taken by Dalquest, on January 18, 1947, was the only one seen by him.
Ciccaba virgata centralis Griscom Wood Owl
Ciccaba virgata centralis Griscom, Bull. Mus. Comp. Zool., 69, 1929: 159. (Chivela, Oaxaca, Mexico.)
One 9 (skel.), 4 km. W Tlapacoj-an, 1700 ft., November 22, 1947; 4 $,2 9 , Rio Atoyac, 8 km. NW Potrero, February 24-March 15, 1946; 2 9 , Rio Blanco, 20 km. WNW Piedras Negras, May 18, 1946; I $ ,1 km. NW Potrero, 1700 ft., January 9, 1947; 2 $ (LSUMZ), Potrero Viejo, 1700 ft., December
16, 1937, and January 28, 1938; 1 $ (LSUMZ), Paraje Nuevo, February 4, 1938; 1 9, 3 km. E San Andres Tuztla, 1000 ft., Januarj- 15, 1948.
576
University of Kansas Publs., Mus. Nat. Hist.
Tecolote. A common resident of both the Upper Tropical Life- zone and the arid division of the Lower Tropical Life-zone, this owl rests by day in dense jungle, usually in the branches of trees about ten feet from the ground. A few were found in hollow trees. They begin to call shortly after dark. Most of the specimens taken were shot at night with the aid of a headlight. Two specimens from Piedras Negras are lighter and have more white spotting above than the light-phased examples from Potrero.
Rhinoptynx clamator forbesi new subspecies Striped Owl
Type. — Adult male; no. 233824, Museum of Comparative Zoology; Presidio, Veracruz, Mexico; May 3, 1925; collected by W. W. Brown.
Characters. — Similar to Rhinoptynx clamator clamator (Vieillot) of northern South America, but black streaks of upper parts and under parts, especially the latter, decidedly reduced in breadth. Resembles R. c. midas (Schlegel)
Table 1. — Measurements in Millimeters of the Races of Rhinoptynx clamator
R. c forbesi
2 males from Presidio and
Rio Blanco, Veracruz
1 female from Rio Blanco, Veracruz .
3 females from Chiapas
1 male from Costa Rica
3 females from Panamd and
Canal Zone
Avg. of all females from Mexico and Central America
R. c. clamator
2 males from Colombia and Venezuela
2 females from Venezuela
R. c. midas
1 male from Argentina
3 females from Argentina and southern Brazil
R. c. oberi (unique type)
unsexed, Tobago
Wing
222-230
(229)
230
242-257
(247.7)
231
233-234
(233.5)
239
230-241
(235.5)
245-256 (250.5)
267
269-282 (272.7)
276
Tail
126-128
(127)
140
131-141
(135.7)
127
127-142 (133.6)
135.4
129
142-150 (146)
139
145-159
(152)
151
Culmen from cere
20.2-20.3
(20.2)
20.3
19.5-21.5
(20.6)
20.7
20.7-23.1
(21.7)
21.3
20.0-21.3
(20.7)
21.8-22.7 (22.3)
22.2
20.6-23.2
(22.2)
LowERY AND Dalquest: Birds From Veracruz, Mexico 577
in general coloration, but size much smaller. Differs likewise from the unique type of Rhinoptynx clamator oberi E. H. Kelso by its much smaller size.
Measurements.— Type: wing, 230 mm.; tail, 128; culmen from cere, 20.2. For comparative measurements see Table I.
Distribution. — Southern Mexico (Veracruz and Chiapas) southward through Central America to Panama.
Remarks. — The name [Asio] niexicana Gmelin, 1788 (based on the Mexican Eared Owl Latham, Synop. Birds, 1, pt. 1, 123) is in- determinate and cannot justifiably be applied to any of the eared owls of Mexico. If it represents a Rhinoptynx, it is an earlier name than clamator; if it represents a Bubo, it is an earlier name than mayensis; if it represents an Asio, it is an earlier name than either wilsonianus or stygius. Since there is no way of ascertaining to which owl Gmelin's name applies, we believe that it should con- tinue to be regarded as unidentifiable.
This new geographical race of the Striped Owl is named for Mr. Dyfrig McH. Forbes, of Potrero Viejo, Veracruz, who, along with his family, has contributed greatly to scientific knowledge by mak- ing extensive collections of natural history objects and by assisting personnel connected with various museum expeditions. In the latter regard, the Hacienda Potrero has served as headquarters or temporary abode for numerous field parties of diverse zoological interest. There are indeed few zoologists who have worked in Vera- cruz in the last two decades who have not enjoyed the hospitality and helpful assistance of Mr. and Mrs. Forbes.
Specimens exam.incd. — Total number, 13, as follows:
Mexico: Veracruz: 1 $ (MCZ 233824), Presidio; 1 5 and 1 $ (UKNHM 24519 and 24520). 20 km. WNW Piedras Negras. Chiapas: 2 9 (U. Mich. 94983 and 107656), Finca Esperanza; 1 $ (U. Mich. 101967), El Corozal.
Costa Rica: San Jose: 1 S (MCZ il7501), San Jose. Guanacoste : 1 juv. 9 (MCZ 148164), Puntarenas. 1 unsexed, locality unspecified (USNM 90395).
Panama: Santos: 1 9 (USNM 400261), Parita. Panama: 1 juv. $ and 1 9 (USNM 409353 and 409354), Cerro Azul. Canal Zone: 1 9 (U. Mich. 56266), Ancon.
Comparative material. — Rhinoptynx cxamator clamator:
A'enezuela: Anzodfegui: 1 ^ (USNM 406473), Cantaura. Movagas: 2 J (USNM 406474 and 406475), Caicara.
Colombia: Santander del Xorte: 1 $ (USNM 372657), Petrolea.
RHINOPri'NX CLAMATOR MIDAS :
Argrentina: Tucumdn: 1 ^ (USNM 400051), Tucuman; 1 $ (USNM 400050), San Pedro de Colalao. Buenos Aires: 1 5 (USNM 55877), Buenos Aires. Brazil: Bahia: 1 $ (USNM 25841), locality unspecified.
Rhinoptynx clamator oberi (the type) : Brazil: Tobago Island (USNM 75112).
578 University of Kansas Publs., Mus. Nat. Hist.
Family NYCTIIBIDAE
Nyctibeus griseus mexicanus Nelson Potoo
Nyctibeus jamaicensis mexicanus Nelson, Auk, 17, 1900: 260. (Met- laltoyuca, Puebla, Mexico.)
One $ , 14 km. SW Coatzacoalcos, 100 ft., February 3, 1947; 1 $ (skel.), 20 km. E Jesus Carranza, 300 ft., Februaiy 5, 1948: 1 9 (skel.), 20 km. ENE Jesus Carranza, 200 ft., March 21, 1949.
Lechusa. Although this species has not been recorded from Vera- cruz in recent years, it is fairly common in the arid division of the Lower Tropical Life-zone, at least as far north as Piedras Negras. In the humid division of the Lower Tropical Life-zone, of extreme southern Veracruz, the Potoo is more abundant. Several individuals are usually seen each hour in a night's hunt. The bird's large, brilliant red eyes have a tremendous reflecting power, and can be seen for more than 200 yards, in the beam of a hunting light. Un- like most goatsuckers, the Potoo usually looks at the source of light with both of its eyes. It is not shy, and by keeping the light in the bird's eyes, one can usually be approached to within fifty feet. The eyes of the bird, when it flies, can be easily kept in the light. When thus blinded in flight, it becomes entangled in branches and strikes other objects. None was seen to strike hard enough to be knocked to the ground, but several almost did so.
The Potoo was never seen on the ground; birds were usually perched twenty-five to fifty feet high, in trees that were isolated or that were on the edges of clearings. In the beam of a flashlight, at close range, the body of this species appears gray, and stands out distinctly. When perched, the body is held across the limb and at almost a sixty-degree angle from the horizontal, its position re- sembling that of a hawk. There is little in the posture of the body to suggest a goatsucker.
In some areas the lechusa is considered a bird of ill omen. Accord- ing to one superstition, the gun of a person shooting one of these birds will burst, unless two sticks are held under the gun, in the form of a cross, when the gun is fired. Even this, it is agreed by most natives, is dangerous, and bad luck in other forms may visit the shooter.
LowERY AND Dalquest! Birds From Veracruz, Mexico 579
Family CAPRIMULGIDAE
Chordeiles acutipennis texensis Lawrence
Texas Nighthawk
Chordeiles texensis Lawrence, Ann. Lye. Nat. Hist. New York, 6, 1856: 167. (Texas = Rio Grande Valley.)
One ^ , 4 km. WNW Fortin, 3200 ft., March 28, 1946.
Nighthawks were seldom seen in Veracruz. This specimen has a wing and tail measurement of 184 mm. and 115 mm., respectively.
Nyctidromus albicollis merrilli Sennett Pauraque
Nyctidromus albicollis merrilli Sennett, Auk, 5, 1888: 44. (Nueces River, Nueces County, Texas.)
One 5 , 3 km. N Presidio, 1500 ft., December 2, 1946.
This specimen has the wing 171 mm. and the tail 174 mm. It may be regarded, therefore, as a migrant from farther north where the race merrilli occurs.
Nyctidromus albicollis yucatanensis Nelson
Pauraque
Nyctidromus albicollis yucatanensis Nelson, Proc. Biol. Soc. Washing- ton, 14, Sept. 25, 1901: 171. (Tunkas, Yucatan.)
One S (skel.), 5 km. N Jalapa, 4500 ft., October 19, 1946; 1 $ (skel.), 5 km. S Tehuatlan, 700 ft., November 14, 1947; 2 S, I 2 , all skels., Teocelo, 4000 ft., January 3 and 29, 1949; 2 5,1 unsexed, (all LSUMZ), Potrero Viejo, August 20, 1937, and February 25, 1938; 4 3 (two skels.), 1 9, Rio Blanco, 20 km. WNW Piedras Negras, March 17-May 11, 1946; 1 <5 , 15 km. W Piedras Negras, 300 ft., January 13, 1947; 1^,1$ (skel.), Rio Blanco, 20 km. W Piedras Negras, 400 ft., October 1, 1946; 3 $ (skels.), 25-30 km. SE Jesus Carranza, 350 ft., April 3 and 5, and May 31, 1949.
Tapacmnino. The call of the Pauraque is one of the most charac- teristic sounds of the night in the lowlands of Veracruz. The species is somewhat less common in the Upper Tropical Life-zone than in the Lower Tropical Life-zone. In the daytime Pauraques hide on the open ground beneath dense bushes, especially thorny species. Shortly after dusk they emerge and fly to favored hunting places. In most cases the birds rest upon the ground in the open; such situations as the dusting place of a Chachalaca, a road, a trail, a ploughed field, or a closely grazed field are favored stations. Oc- casionally, and strangely enough, only in certain limited localities, such as near Piedras Negras, the birds seem to prefer to rest on logs or low, horizontal branches. From their stations, they fly up, with a hopping motion and a few wing strokes, to intercept passing 4—3247
580 University of Kansas Publs., Mus. Nat. Hist.
insects. Their mournful call is given intermittently. It consists of a somewhat catlike note, that might be translated as "where-are- you." The natives of southern Veracruz translate the call as "pavo- real," in reference to a legend in which the peacock refused to lend its plumage to the tapacamino. There is also a children's song concerning this bird.
The varied degrees of shyness of the Pauraque seems to depend on the brilliance of the moon. On moonlit nights, it rarely allows one to approach to within fifty feet. On dark nights, if the hunting light is kept on the brilliant glow of the bird's eye (a single eye is usually seen in the light) it can usually be approached so closely as to be captured in the hand. Most of the specimens listed above were taken in this manner.
Several nests of the Pauraque were found in southern Veracruz in June. Each consisted of a scarcely perceptible depression in the leaf-littered sand beside a river. In every case, the nest was somewhat sheltered above by open, low trees, and on a slight slope. Two nests that were carefully examined each held two eggs.
Phalaenoptilus nuttallii centralis Moore Poor-will
Phalaenoptilus nuttallii centralis Moore, Proc. Biol. Soc. Washington, 60, 1947: 146. (Puerto de Guadalupe, 5 miles west of Ibarra, Guanajuato, Mexico.)
One 5 , 2 km. W Limon, 7500 ft., September 26, 1948.
Although this specimen has not been compared with the unique type of Moore's new race, centralis, it appears to agree with the description in most particulars. Moreover, it differs from material in the United States National Museum in the same way that Moore distinguishes centralis from the various named races. This appears to be the first record for this species in Veracruz. The specimen was shot at night on sandy soil in a corn field.
Caprimulgus serico-caudatus salvini Hartert Salvin Whip-poor-will
Caprimulgus salvini Hartert, Ibis, 1892: 287. New name for Capri- mulgus macromystax of American authors but not of Wagler and other European authors. (Mirador, Veracruz, Mexico, accepted as type locality, c/. Peters, Birds World, 1940, p 198.)
One S (LSUMZ), Potrero Viejo, 1700 ft., December 12, 1938.
The specimen supposedly taken by Sartorius probably at Mirador (Baird, et al., 1874:409) seems to be the only other record for this species in Veracruz.
LowERY AND Dalquest: Birds From Veracruz, Mexico 581
Caprimulgus vociferus Tociferus Wilson American Whip-poor-will
Caprimulgus vociferus Wilson, Amer. Orni., 5, 1812: 71, pi. 41, figs. 1-3. (Pennsylvania.)
One 5 , 2 km. N Motzorongo, 1500 ft., December 7, 1946.
Caprimulgus vociferus setosus van Rossem American Whip-poor-will
Caprimulgus vociferus setosxis van Rossem, Bull. Mus. Comp. Zool., 77, 1934: 408. (Galindo, Tamaulipas, Mexico.)
One 5, 6 km. SSE Altotonga, 9000 ft.. November 10, 1946; 1 $ (skel.), 7 km. NW Potrero, 1700 ft., January 9, 1947.
The specimen from Altotonga was shot from the limb of a pine tree, where it was calling approximately an hour after dusk. Both the presence of the bird on a limb thirty-five feet from the ground and the fact that it was calling in November are worthy of note.
Family APODIDAE
Chaetura pelagica (Linnaeus) Chimney Swift
Hirundo Pelagica Linnaeus, Syst. Nat., ed. 10, 1, 1758: 192. (America = South Carolina.)
One $ , 20 km. ENE Jesiis Carranza, 200 ft., April 10, 1949.
Chaetura vauxi richmondi Ridgway Vaux Swift
Chaetura richmondi Ridgway, Proc. Biol. Soc. Washington, 23, 1910: 53. (Guayabo, Costa Rica.)
One $ , 35 km. SE Jesiis Carranza, 350 ft., April 7, 1949.
Golondrina. On the Rio Solosuchi and Rio Chilchijapa in April, 1949, flocks of fifty to one hundred swifts were seen flying over the surface of the water. Both the specimen of vauxi and the specimen of pelagica were collected from these flocks.
Chaetura rutila brunnitorques Lafresnaye Chestnut-collared Swift
Chaetura brunnitorques Lafresnaye, Rev. Zool., 1844: 81. (Colombia.)
One $ ,1 5,5 km. N Jalapa, 4500 ft. October 18, 1946.
The only two specimens taken were captured at night in a small cave behind a waterfall in the Upper Tropical Life-zone.
582 University of Kansas Publs., Mus. Nat. Hist.
Family TROCHILIDAE
Phaethornis superciliosus veraecrucis Ridgway White-browed Hermit
Phaethornis longirostris veraecrucis Ridgway, Proc. Biol. Soc. Washiug- ton, 23, 1910: 54. (Buena Vista, Veracruz, Mexico.)
One 2 , 35 km. SW Jesus Carranza, 400 ft., February 17, 1948; 1 9 , 30 km. SSE Jesus Carranza, 300 ft., May 10, 1949.
These dull-colored hummers seem somewhat restricted in their feeding habits. They are primarily birds of the deep jungle, and do not gather with other species of hummingbirds at the flowering trees along the river banks. Usually they are solitary, moving about in the deep shade, and pausing at bromiliads, vines, and wild banana plants. One gathering of hummingbirds of this species was noted in the jungle near Jesus Carranza, where, in a limited area, there were many brilliant red flowers of a honeysuckle type. This was the only species of hummingbird noted feeding on the flower. Probably 100 of these birds were seen in one afternoon, and often there were ten or more in sight at once. They seemed to be less quarrelsome than most other species of hummingbirds. All trochilids in Veracruz are called chupa rosa or chupa miel by the natives.
Phaethornis longuemareus adolphi Gould Boucard Hermit
Phaethornis adolphi Gould, Monogr. Trochil., pt. 14, 1857: pi. [15] and [text =1, pi. 35 of volume]. (Cordoba, Veracruz, Mexico.)
One 5, Rio Atoyac, 8 km. NW Potrero, March 5, 1946; 1 9 (LSUMZ), Potrero, 1700 ft., January 4, 1938; 1 $ (skel.), Potrero Viejo, 1700 ft., Decem- ber 16, 1948; 1 $ (skel.), 13 km. ENE Potrero, 1700 ft., April 24, 1949; 2 9 (one skel.), 20 km. SE Jesus Carranza, 250 ft.. May 2, 1949.
The body of this tiny hummingbird is so plump that it appears nearly spherical. It is an inconspicuous species, usually found at the edges of patches of jungle, especially along small water-courses. One of its favored haunts is the small, still-surfaced pools of tiny streams in the forest. It spends considerable time skimming the surface of the pools and plunging into the water. The species is noteworthy in that it remains close to the ground. Probably most of its activities are carried on at a distance of three to five feet from the ground, and rarely does it go higher. Although not social, and rarely seen with other species of hummers, individuals of this species rarely fight as do most other kinds of hummers when they meet others of their kind. They feed in flowers of small size, in bromiliads, and in crevices in twigs and vines.
LowERY AND Dalquest: Birds From Veracruz, Mexico 583
Campylopterus curvipennis curvipennis (Lichtenstein) Curved-winged Saber-wing
Trochilus curvipennis Lichtenstein, Preis-Verz. Vogel Mexico, 1830: 1. (Mexico.)
One $ (skel), 5 km. ENE El Jobo, 600 ft., October 19, 1947; 3 S (one skel.), 19,4 km. W Tlapacoyan, 1700 ft., October 13 and 14, and November 22, 1947; 15,4 km. WNW Fortin, 3200 ft., March 27, 1946.
This large hummer is not common. It was usually solitary and found in dense jungle. Several specimens were shot that could not be found in the dense vegetation. On several occasions, individuals ceased the humming flight and flew in the normal, birdlike fashion. This was usually when they were about to perch, and the non- humming flight was only for a distance of six to eight feet.
Campylopterus excellens (Wetmore) Long-tailed Saber-wing
Pampa panipa exccUcnff Wetmore, Proc. Biol. Soc. Washington, 54, 1941 : 207. (Volcan San Martin, Sierra de Tu.xtla, Veracruz, Mexico.)
Four S (one skel.), 1 $ , 30 km. SSE Jesus Carranza, 300 ft.. May 8-14, 1949.
When Wetmore described excellens he believed that the form was confined to the Sierra de Tuxtla and that it was subspecifically re- lated to Campylopterus curvipennis despite its highly trenchant yet
Table 2. — Compar-\tive Me-^surements in Millimeters of Campylopterus c. curvipennis and campylopterus excellens
|
.2 |
'3 |
h |
||||
|
1 . |
. e3 |
a |
2co |
O CO |
||
|
.00 M g K go |
CO C |
CO 03 |
CO s o |
3 '^ S =3 |
a |
|
|
<a t.":! |
<*H w |
•■ C cc |
c a |
f—t |
||
|
irvip lesf i an ruz. |
ales I an ruz. |
liens es f r Jesii ruz. |
c3 . O § |
|||
|
^ CS cc o |
7fem Potos Verac |
«.J — ! - U |
||||
|
C. C. C 16 m Poto Vera |
C. exc 3 ma SSE Vera |
4 ma Mts. type |
Avg. excel |
|||
|
Wing |
62.9-69.1 (66.5) |
58. 4-62. 8 (60.6) |
67.1-69.8 (68.1) |
66.6 |
67.5-73.2 (70.3) |
69.4 |
|
Tail |
48.7-53.3 (50.6) |
42.6-46.2 (43.8) |
59.5-62.6 (60.7) |
49.6 |
60.3-63.5 (61.4) |
61.1 |
|
Exposed cidmen. . . |
24.1-27.5 |
23.2-26.3 |
28.8-29.4 |
26.9 |
26.9-29.2 |
27.9 |
|
(25.4) |
(24.3) |
(28.6) |
(27.7) |
|||
|
Width of bill at |
||||||
|
posterior edge of nostril |
3 5-5 9 |
4 0-4 6 |
5 0-6 0 |
4 0 |
||
|
(4.8) |
(4.3) |
(5.4) |
584
University of Kansas Publs., Mus. Nat. Hist.
subtle characteristics. Superficially, excellens is virtually a replica of curvipennis; the skins of excellens, as Wetmore points out, appear to be nothing more than overstuffed examples of curvipennis. In color the two are almost identical. On close inspection, however, one discovers that the tail feathers of excellens are decidedly longer and broader and that the head appears to be larger, even though the bill itself is only slightly longer and only slightly broader basally (see Table 2).
In the material recently acquired by Dalquest there are four skins and one skeleton of Campylopterus from a locality thirty kilometers SSE of Jesus Carranza. Somewhat surprising is the fact that these specimens are referable to excellens, even though they were taken approximately 140 kilometers from the Volcan San Martin in the Sierra de Tuxtla, at an elevation of only 300 feet. Since all were collected between May 8 and 14, there can be little doubt that they represent breeding birds at the Jesus Carranza locality and not altitudinal migrants from the higher elevations in the Sierra de Tuxtla.
Figs. 2-7. Skulls of Campylopterus curvipennis curvipennis and Campylop- terus excellens, to show difference in size and proportion. X 1.
Figs. 2, 4, and 6. Campylopterus excellens (Wetmore), $, UKNHM 28954, 30 km. SSE Jesus Carranza, Veracruz, May 14, 1949.
Figs. 3, 5, and 7. Campylopterus c. curvipennis (Lichtenstein), $, LSUMZ 13794, Xilitla, San Luis Potosi, Mexico, February 10, 1947.
LowERY AND Dalquesti Birds From Veracruz, Mexico 585 Table 3. — Skeletal Measurements in Millimeters of Campylopterus
C. CXJRVIPENNIS AND CaMPYLOPTERUS EXCELLBNS
|
•2 C3 |
a . |
— N |
2 |
||
|
s:3 |
-« N |
•^ 3 |
^ |
||
|
o" _ |
o |
||||
|
ipylopterus c. SUMZ 13794 in Luis Potos |
SUMZ 14206, ilitla, San Lu |
KNHM 2492 NE El Jobo, |
KNHM 2493 ^ Tlapocoyan |
ipylopterus ex KNHM 2895 5E Jesus Car eracruz. |
|
|
IhJm |
^y. |
PW |
Pl^ |
|P«2> |
|
|
o |
o |
||||
|
Greatest Length |
|||||
|
of Skull |
44.3 |
44.4 |
45.5 |
44.9 |
48.3 |
|
Greatest Width |
|||||
|
of Skull |
11.7 |
11.6 |
11.8 |
11.8 |
12.7 |
|
Interorbital |
|||||
|
Breadth |
6.1 |
6.1 |
6.6 |
6.5 |
7.3 |
|
Maxillary |
|||||
|
Breadth |
5.0 |
4.8 |
5.0 |
5.0 |
5.1 |
|
Quadrato-jugal |
|||||
|
Breadth |
9.7 |
9.6 |
9.6 |
9.6 |
10.5 |
|
Interlacrvmal |
|||||
|
Breadth |
6.1 5.9 |
6.1 6.1 |
6.6 6.0 |
6.5 5.9 |
7 3 |
|
Length of Palatines |
6.8 |
||||
|
Coracoid |
8.7 6.6 6.4 8.1 |
8,5 6.5 6.3 8.1 |
8.6 6.6 6.4 8.1 |
8.6 6.4 6.3 8.1 |
9.5 |
|
Humerus |
7.4 |
||||
|
Ulna |
7.3 |
||||
|
3rd Metacarpal |
8.7 |
||||
|
Femur |
9.8 14.0 |
9.8 14.0 |
9.6 14.1 |
11.2 |
|
|
Tibio-tarsus |
14.1 |
15.4 |
|||
|
Tarso-metatarsus . . . |
6.2 |
6.2 |
6.2 |
5.9 |
7.0 |
|
Length of mandi- |
|||||
|
bular ramus |
38.4 |
37.9 |
39.3 |
38.4 |
42.3 |
586 University of Kansas Publs., Mus. Nat. Hist.
At present there is no material on hand showing that excellens occurs sympatrically with curvipennis, but it almost certainly does so since the latter is known from localities in the humid division of the Lower Tropical Life-zone from Veracruz to Guatemala. Jesus Carranza, where our recent examples of excellens were taken, lies in this same zone.
The really interesting point brought to light by these specimens, however, is found in our study of the one male skeleton. The al- legedly greater head-size of excellens, which Wetmore discerned en- tirely by his comparison of skins, is a fact that can be expressed by actual measurements (Figures 2-7 and Table 3). The skull of excellens is actually markedly larger than that of curvipennis. Also, many of the skeletal elements in the two forms differ significantly in size. The larger over all size of the cranium, and the body skeleton of excellens, however, is not the result of an increase in size of all the component parts, for certain dimensions do not vary appreciably from those of curvipennis (see Table 3). Even though the tail length of excellens shows an absolute difference, the variation in wing length between the two forms is only an average difference.
The question arises as to whether excellens should be regarded as specifically distinct from curvipennis. In the first place, the fact that curvipennis is already polytypic is highly pertinent. The spe- cies ranges southward from San Luis Potosi through Veracruz and southern Mexico to Yucatan and Guatemala. Birds from the two last-named places are smaller than those from San Luis Potosi and central Veracruz and are currently recognized under the name C. c. pampa (Lesson). Intergradation occurs between the two in Cam- peche (Ridgway, 1911:356). Superimposed, in southern Veracruz, on this geographically variable and continuous population of curvi- pennis, is the large form, excellens, which seems to be reproductively isolated. These facts, coupled with the cranial and other differences in size noted above, point to the conclusion that excellens is a sibling species. We propose that it be so regarded until facts are adduced to the contrary.
Campylpoterus hemileucurus (Lichtenstcin) Delattre Saber-wing
Trochilus hemileucurus Lichtenstein, Preis-Verz. Vogel Mexico, 1830 : 1. (Mexico.)
Two $, 20 km. SE Jesus Carranza, May 1, 1939; 2 $ (one skel.), 1 $ , 30 km. SSE Jesus Carranza, May 12, 1949.
This species seems to be rather rare. In April, May, and June, in extreme southern Veracruz along the Rio Chalchijapa and Rio Solo-
LowERY AND Dalquest: Birds From Veracruz, Mexico 587
suchi, certain species of trees that are confined to sandy river banks come into bloom. The flowers draw swarms of hummers of several species, and usually a dozen or more are present at each tree. Per- haps one in 500 hummers seen at such trees is of this large, purple species. In the field, C. hemileucurus appears to be black with the white markings of the tail abrupt and distinct. Its large size sepa- rates it from other species with which it is usually seen.
Florisuga mellivora mellivora (Linnaeus) Jacobin Hummingbird
Trochilus mellivorits Linnaeus, Syst. Nat., ed. 10, 1, 1758: 121. (India, error = Surinam, ex Edwards.)
Four 9 (one skel.), 20 km. SE Jesus Carranza, 250 ft., May 1 and 2, 1949; 1 5 , 30 km. SSE Jesus Carranza, 300 ft.. May 12, 1949.
Florisuga mellivora was noted only in extreme southern Veracruz, where it was found in the same flowering trees with Campylopterus hemileucurus and other species of hummers. In the field, this form closely resembles Campylopterus hemileucurus, appearing black with abrupt white markings on the tail. It is, however, much smaller than the latter species. Florisuga was more common than Camply- opterus hemileucurus and was noted in an approximate ratio of four to one. It was, nevertheless, one of the less common species at the flowering trees, being outnumbered many times by the species of Amazilia.
Loetscher (MS) doubts the validity of the one previous record for this species in Veracruz, at Orizaba (Sumichrast, 1881:230), and therefore includes it in his list of species of hypothetical occurrence.
Anthracothorax prevostii prevostii (Lesson) Prevost Mango
Trochilus prevostii Lesson, Hist. Nat. Colibris, Suppl, [1832]: p. 87, pi. 24. (South America.)
Two S, Potrero Viejo, February 12 and March 23, 1946; 1 $, 1 9 , Rio Blanco, 20 km. WNW Piedras Negras, May 10 and 28, 1946; 5 $ , 1 $ , 20 km. ENE Jesus Carranza, 200 ft., March 26, 1949.
This large and beautiful hummingbird seems to be rather local in its distribution. It was not uncommon at Potrero Viejo, where two were shot from the same twig a few weeks apart, and others were seen. This was the only locality in the Upper Tropical Life-zone where the species was noted. Strangely enough, only males were seen at Potrero Viejo. The two specimens taken near Piedras Negras were the only ones seen in that locality. On the banks of the Rio Coatzacoalcos, this was one of the more numerous hummers.
588 University op Kansas Publs., Mus. Nat. Hist.
It was common about certain species of flowering trees, at thirty to forty feet from the ground, where it was associated with species of Amazilia. It was not present with other hummers in the deep jungles twenty-five kilometers to the southeast, where such species as Florisuga rnellivora and Campy lop terus hemileucurus, not seen on the Rio Coatzacoalcos, were found.
Chlorostilbon canivetii canivetii (Lesson) Canivet Emerald
Ornismya canivetii Lesson, Hist. Nat. Colibris, Suppl., [1832]: pp. 174 and 177, pis. 37 and 38. (Brazil, error = Jalapa, Veracruz, Mexico, ex Rev. Zool., 1839, p. 15.)
One 9, 9 km. E Papantla, 300 ft., November 16, 1947; 1 $ (LSUMZ), Potrero, 1700 ft., January 6, 1938; 1 ,J , Rio Blanco, 20 km. W Piedras Negras, 400 ft., October 1, 1946.
This tiny, brilliant green hummer is rare and inconspicuous. One individual was feeding in a bull's-horn acacia tree after a rainfall, seemingly on small insects floating out from the leaf bases of small bromeliads by the rain.
Amazilia Candida genini (Meise) White-bellied Emerald
Agyrtrina versicolor genini Meise, Bull Mus. Roy. Hist. Nat. Belg., 14, 1938: 2. (El Asterillo, "Veracruz, Mexico.)
One $ , 2 9 , Rio Atoyac, 8 km. NW Potrero, February 19 and 25, and March 21, 1946; 1 5 , 4 km. WNW Fortin, 3200 ft., April 5, 1946; 5 $ (one skel.), 20 km. ENE Jesus Carranza, March 23-26, and May 16, 1949; 5 $ (three skels.), 30 km. SSE Jesus Carranza, May 9, 12, and 14, 1949; 1 9 , 60 km. SW Jesus Carranza, 450 ft., March 27, 1948.
This small species is probably the most common hummingbird in the state of Veracruz. In the extreme southern part of the state, when the flowering trees along the river banks were in bloom, literally thousands were seen daily.
Amazilia cyanocephala cyanocephala (Lesson) Red-billed Azure-crown
Ornismya cyanoceph<ilus Lesson, Hist. Nat. Ois.-Mouches, [1829] : xlv. (Brazil = Veracruz, Mexico.)
One 9, 4 km. W Tlapacoyan, 1700 ft., October 15, 1947; 1 9 (LSUMZ), Jalapa, May 27, 1949; 1 $, Teocelo, 4000 ft., January 30. 1949; 1^,4 km. WNW Fortin, 3200 ft., April 5, 1946; 1 5 , 20 km. SE Jesus Carranza, 250 ft., May 2, 1949.
This species seems not to be common, although it might easily be overlooked and mistaken for the preceding form. It was often found
LowERY AND Dalquesti Birds From Veracruz, Mexico 589
with Campylopterus curvipennis, which it resembles except in its much smaller size.
Amazilia beryllina beryllina (Lichtenstein) Berylline Hummingbird
Trochilus beryllinus Lichtenstein, Preis-Verz. Vogel Mex., 1830: 1. (Mexico.)
One 5,4 km. N Jalapa, 4500 ft., October 13, 1946; 1 S,2 9 , (all LSUMZ), Jalapa, May 27 and 29, 1949; 1 S , 1 9 , Atoyac, 8 km. NW Potrero, March 5 and 20, 1946; 1 $, Ojo de Agua, 8 km. NW Paraje Nuevo, February 12, 1946; 1 ,5 , 4 km. WNW Fortin, 3200 ft., March 25, 1946.
Amazilia yucatanensis cerviniventris (Gould) Buff-bellied Hummingbird
Amazilius cerviniventris Gould, Proc. Zool. Soc. London, pt. 24, 1856: 150. (Cordoba, Veracruz, Mexico.)
One $ (LSUMZ), Potrero, 1700 ft., January 4. 1938; 1 $ , Potrero Viejo, 1700 ft., October 23, 1946; 2 $, Rio Blanco, 20 km. WNW Piedras Negras, May 12 and 13, 1947; 1 5 , 20 km. W Piedras Negras, October 1, 1946.
This and the two following species sometimes occur in the same areas and are so similar that they cannot always be separated with certainty from one another in the field.
, Amazilia tzacati tzacatl (De la Llave)
Rieffer Hummingbird
Trochilus Tzacatl De la Llave, Registro Trimestre, 2, no. 5, 1833: 48.
(Mexico.)
One $ (LSUMZ), Paraje Nuevo, February 12, 1938; 1 5 and 2 $ (one skel.), 20 km. ENE Jesiis Carranza, March 24-26, and April 12, 1949; 1 5 , 20 km. SE Jesus Carranza, 250 ft., May 2, 1949; 2 $ (one skel.), 3 9 (two skels.), 30 km. SSE Jesus Carranza, 300 ft., March 28 and May 9 and 12, 1949; 1 $ (skel.), 35 km. SE Jesus Carranza, 400 ft., February 19, 1948.
Eupherusa eximia nelsoni Ridgway Striped-tailed Hummingbird
Eupherusa eximia jielsoni Ridgway, Proc. Biol. Soc. Washington, 23, 1910: 54. (Motzorongo, Veracruz.)
One $ (skel.), Potrero Viejo, 1700 ft., March 13, 1949; 1 5 , 35 km. SE Jesus Carranza, 350 ft., April 5, 1949.
Heliomaster longirostris pallidiceps Gould Pale-crowned Star-throat
Heliomaster pallidiceps Gould, Intro. Trochil., 1861 : 139 (Mexico and Guatemala = Jalapa, Veracruz, Mexico.)
One 5, 7 km. W Potrero, 1700 ft., December 26, 1946.
Our only specimen was taken by a Mexican girl, and was brought to us bv Miss Loni Forbes.
590 University of Kansas Publs., Mus. Nat. Hist.
Calothorax lucifer (Swainson) Lucifer Hummingbird
Cynanthus Lucifer Swainson, Philos. Mag. (n. s.), 1, 1827:442 (Temas- scaltepec.)
One unsexed [= $] (skel.), Potrero Viejo, 1700 ft., December 16, 1948.
The specimen listed was found dead on the window sill in a house, where it had seemingly become exliausted and died in attempting to escape from, a room that it had entered by accident.
Archilochus colubris (Linnaeus) Ruby-throated Hummingbird
Trochilus Colubris Linnaeus, Syst. Nat., ed. 10, 1, 1758: 120. Based on The Hummingbird, Mellivora avis carolinensis Catesby, CaroHna, 1, p. 65. (In America imprimis septentrionaU = South Carolina.)
One S , 5 km. ENE El Jobo, 600 ft., October 18, 1947; 5 $ (two skels.), 3 9 ,
5 km. N Jalapa, 4500 ft., October 13-16, 1946; 1 5 , Rio Atoyac, 8 km. NW
Potrero, February 26, 1946; 1 5 , Rio Blanco, 20 km. W Piedras Negras. October 1, 1946.
This familiar North American migrant is abundant in the Upper Tropical Life-zone. It appears to be less common in the Lower Tropical Life-zone.
Atthis heloisa heloisa (Lesson and Delattre) Heloise Hummingbird
Orismya Helosia Lesson and De Lattre, Rev. Zool., 2, 1839; 15. (Jalapa and Quatepu [ = Coatepec], Veracruz, Mexico.)
Two 5 , 5 km. N Jalapa, 4500 ft., October 14, 1946.
Trogon massena massena Gould Massena Trogon
Trogon massena Gould, Monogr. Trogonidae, pt. 3, 1838: pi. [4] and text [ = pi. 16 of volume]. (Mexico.)
Three $ (one skel.), 2 $ (one skcl.), 20 km. ENE Jesus Carranza, 300 ft., March 20 and 23, 1949; 1 $ (skel.), 35 km. SE Jesus Carranza, 400 ft., February 19, 1948.
This large trogon was found only in extreme southern Veracruz. South of the Rio Coatzacoalcos, it was common in the deep, unin- habited jungles. Birds of this species are rather inactive, sitting almost motionless on vines or branches for long periods, where in spite of their brilliant colors, they are inconspicuous. Usually they are found singly or in pairs, but two flocks of six to ten birds were seen. Possibly there was a fruit-laden tree in the vicinity that brought separate birds together from some distance. In Veracruz, trogons arc called mira-el-sol by the natives.
LowERY AND Dalquest: Birds From Veracruz, Mexico 591
Trogon citreolus melanocephalus Gould
Citreoline Trogon
Trogon melancephala Gould, Monogr. Trogonidae, pt. 2, 1S35: pi. [6] [and text = pi. 12 of volume.] (State of Tamaulipas, Mexico.)
Two $ (one skel.), 2 9 (one skel.), Rio Blanco, 20 km. WNW Piedras Negras, May 11 and 18, and September 30, 1946; 1 5 , 15 km. W Piedras Negras, 300 ft., January 15, 1947; 1 9, Jimba, 350 ft., March 2, 1947; 1 S (skel), 20 km. ENE Jesus Carranza, 200 ft., March 23, 1949; 1 5 , 20 km. S Jesus Carranza, 300 ft., February 12, 1948.
This comparatively dull-colored trogon is common in the arid di- vision of the Lower Tropical Life-zone. In the low jungle and thorny brushlands of the coastal plain, it lives closer to the ground than the trogons occupying the higher forests. It was never seen in the Upper Tropical Life-zone, where the other yellow-breasted species, Trogon violaceus, is common. Only a few Citreoline Tro- gons were seen in the jungles of extreme southern Veracruz. Like other trogons, citreolus is a rather sedentary species, and spends most of the day perched on some twig or vine.
Trogon collaris puella Gould Jalapa Trogon
Trogon -puella Gould, Proc. Zool. Soc. London, pt. 13, 1845: 18. (Es- cuintla. South America [probably Escuintla, Guatemala].)
One 9, Canada Blanca, 7 km. NW Paraje Nuevo, February 12, 1946; 1 9, 4 km. W Tlapocoyan, 1700 ft., October 15, 1947; 2 ^ , Rio Atoyac, 8 km. NW Potrero, March 3 and 10, 1946; 1 9,1 5,7 km. NW Potrero, 1700 ft., Decem- ber 28, 1946, and January 10, 1947; 1 $ ,Z 9,3 km. N Presidio, 1500 ft., De- cember 3-12, 1946; 2 9 (one skel.), 35 km. SE Jesiis Carranza, 400 ft., Feb- ruaiy 16, 1948.
The Jalapa Trogon is most common in the Upper Tropical Life- zone, but is far less numerous there than the Gartered Trogon. It was seen only a few times in extreme southern Veracruz, and never on the coastal plain.
Near Presidio, trogons of this species and the following w^ere watched while they are feeding in a low tree in the jungle. The tree bore an abundance of small, berrylike fruit, upon which other birds, especially thrushes and tanagers, were feeding. The Jalapa Trogons confined their attentions to the berries near the top of the tree, about ten feet from the ground. They did not perch in the fruit tree it- self, but instead in nearby trees, at a slightly higher elevation. The fruit tree was approached with a silent glide and an upward swoop, so that the bird leaving its perch, dropped well below the level of the fruit and came up to it from below. They hovered a moment w4th a
592 University of Kansas Publs., Mus. Nat. Hist.
loud whirring and fluttering of wings in front of the berries as they snipped them from the cluster. The fruit seemed to be swallowed while the trogon was on the wing.
Near Potrero, a trogon of this species was seen to make two fly- catcherlike flights outward away from its perch on a horizontal vine approximately fifty feet from the ground. After seeming to catch an insect, the bird returned to the vine. A Mexican assistant who retrieved a wounded trogon of this kind received a severe bite from the bird's powerful bill.
Trogon violaceus braccatus (Cabanis and Heine) Gartered Trogon
Aganus braccatus Cabanis and Heine, Mus. Hein., Th. 4, Heft 1, 1862- 63 [==1863]: 184. (Mexico; cf. Peters, Birds World, 5, 1945, p. 158 (note), and Carriker and de Schauensee, Proc. Acad. Nat. Sci. Phila., 87, 1935, p. 424.)
One 9, 7 km. NW Potrero, 1700 ft., December 29, 1946; 6^,39 (one skel.), Rio Atoyac, 8 km. NW Potrero, February 17-21, and March 10-20, 1946; 2 $ (LSUMZ), Potrero Viejo, January 23, 1938, and August 22, 1937.
The trim, little Gartered Trogon is abundant in the Upper Tropical Life-zone of Veracruz. Its small size and sedentary habits make it easy to overlook in the tall forest. Even when the birds are calling steadily, it is difficult to make out their plump forms in the leafy vegetation near the tops of the tall trees. Often, however, they feed in low trees, with thrushes and tanagers, on ripe berries and other fruit.
Family ALCEDINIDAE
Megaceryle torquata torquata (Linnaeus)
Ringed Kingfisher
Alcedo torqiuxta Linnaeus, Syst. Nat., 1, ed. 12, 1766: 180. (Mexico.)
One S, Puente Nacional, 500 ft., October 22, 1946; 1 $ (LSUMZ), Rio Blanco, Matacabresto, 20 mi. NE Piedras Negras, December 19, 1937; 1 $, Rio Blanco, 20 km. WNW Piedras Negras, May 15, 1946; 1 5 , 20 km. ENE Jesus Carranza, 20O ft., April 28, 1949; 1 9 (skel.), 35 km. SE Jesus Carranza, 400 ft., February 19, 1948.
Martin pescador. The Ringed Kingfisher seems to be confined to the larger rivers. It is not uncommon on such rivers as the Antiguo, Jamapa, and Blanco in central Veracruz, and is abundant on the rivers Coatzacoalcos, Solosuchi, and Chalchijapa in southern Vera- cruz. When one travels by canoe on these rivers, the large king- fishers are almost constantly in sight, flying along the rivers or from perch to perch along the shores. Their rattling call is usually made when the bird is flying. When it plunges into the water after a
LowERY AND Dalquest: Birds From Veracruz, Mexico 593
fish, the splash caused by the kingfisher's large body can be heard for some distance. Several nests were seen in vertical banks of sand or claylike soil beside the rivers. Birds were entering and leaving the holes in May.
Chloroceryle amazona mexicana Brodkorb Amazon Kingfisher
Chloroceryle amazona mexicana Brodkorb, Auk, 57, 1940: 543. (Barra de Cahuacan, Chiapas, Mexico.)
One $ , Puente Nacional, 500 ft., October 21, 1946; 1 $ , 5 km. SE Boca del Rio, October 10, 1946.
This is the least common of the four species of kingfisher found in the course of this study. It occurs, like Megaceryle torquata, most commonly at the lower elevations where the rivers are broad and sluggish.
Chloroceryle americana septentrionalis (Sharpe) Green Kingfisher
Ceryle septentrionnlis Sharpe, Cat. Birds Brit. Mus., 17, 1892 : 134. (Teapa, Tabasco, Mexico.)
One $ (skel.), Puente Nacional, 500 ft., October 21, 1948; 1 5 , 5 km. SW Boca del Rio, October 11, 1946; 1 9 , Rio Atoyac, 8 km. NW Potrero, February 19, 1946; 1 5 and 1 $ (LSUMZ), Potrero Viejo, Rio Atoyac, October 25, 1938; 1 S (LSUMZ), Paraje Nuevo, January 25, 1938; 1 $, 1 $ , Rio Blanco, 20 km. WNW Piedras Negras, May 16 and 17, 1946; 1 5 , 15 km. W Piedras Negras, 300 ft., January 15, 1946; 1 9,1 S, (skels.), Rio Blanco, 20 km. W Piedras Negras, September 29 and October 1, 1946; 1 9 (skel.), 35 km. SE Jesus Carranza, 350 ft., April 6, 1949.
Martin del no. This is the commonest kingfisher found in Vera- cruz. It occurs throughout the Tropical Region, from the cold, crys- tal-clear streams at higher elevations to the broad, brown, sluggish rivers of the lowlands. It was seen also at ponds and lakes, and even at some temporary pools of rain water. It feeds principally on small fish, but may eat also insects, for occasionally it was seen to dive into the pools of rainwater where there could hardly have been any fish.
Chloroceryle aenea stictoptera (Ridgway) Least Kingfisher
Ceryle superciliosa stictoptera Ridgway, Proc. Biol. Soc. Washington, 2, 1884:95. (Sisal, Yucatan.)
One $, Rio Blanco, 20 km. WNW Piedras Negras, May 16, 1946; 1 5 , 30 km. ESE Jesus Carranza, 300 ft.. May 10, 1949; 1 9 , 35 km. SE Jesus Carranza, 350 ft., April 7, 1949; 2 9 (skels.), 30 km. SSE Jesus Carranza, 300 ft., May 12-13, 1949.
Martin del arroyo. This tiny species of kingfisher is rare in cen-
594 University of Kansas Publs., Mus. Nat. Hist.
tral Veracruz, and not common elsewhere. Only in the small, sluggish streams leading into tributaries of the Rio Coatzacoalcos near its mouth, was it found to be even fairly common. Along the Rio Chalchijapa and Rio Solosuchi, a few birds were seen perched over pools left by the river as it receded during the dry season. As the water in the pools diminished, tiny fish and fish fry concentrated in great numbers, and dead and dying fish formed a skum over the surface. These probably served as food for the tiny kingfishers. Never was the species seen over the open water of the rivers. This, the natives stated, was because they feared the large fish. Certainly the abundant gars and other carnivorous fish could swallow C. aenea in a single gulp, but the same could be said for C. americana, which customarily fishes in the open rivers.
Family MOMOTIDAE
Hylomanes momotula momotula Lichtenstein
Tody-motmot
Hylomanes Momotula Lichtenstein, Abh. Akad. Wiss. Berlin for 1838 [= 1839] : p. 449, pi. 4. (Valley Real, Mexico.)
One $,2 9 , 35 km. SE Jesiis Carranza, 300 ft., February 12, 1948, and April 7 and 9, 1949.
The Tody-motmot was found only in extreme southern Veracruz. It is probably fairly common, but is a species that may be easily overlooked. It is, like most motmots, a sedentary bird. In addition it lives in the deepest, most shaded part of the jungle, and its dis- proportionately large head gives it a profile so unlike a bird that the collector is likely to overlook the species. Two of the three specimens listed were discovered almost by accident; one in a thicket of thorny ground palms beneath tall, dense jungle, and another in a gloomy gulch in the forest. Both birds were seen as they sat motionless on their perches, about six feet from the ground and approximately the same distance from the collector's eye. The third specimen was taken at night in a brushgrown, sandy delta of a dry arroyo beside the Rio Solosuchi. The bird was roosting on a twig at about shoulder height, and looked pale in the beam of the hunting light. It was captured in the hand.
Electron carinatum (Du Bus)
Keel-billed Motmot
Prionites carinatus Du Bus, Bull. Acad. Roy. Sci. Belg., 14, 1847: 108. (Guatemala.)
One $ , 30 km. SSE Jesiis Carranza, 300 ft., May 12, 1949.
Gallina. While the party was traveling on the Rio Chalchijapa
LoTVERY AND Dalquest: Birds From Veracruz, Mexico 595
and Rio Solosuchi in 1948, a bird was heard on a number of occasions that the natives called "chicken." The call was a loud, far-reaching "cut-cut-cadack," strikingly like the cackle of a hen. It usually seemed to emanate from tall jungle trees on hillsides near the river. One native thought the bird was a kind of pigeon; an- other, a kind of pheasant. On several occasions, the canoe was beached and a hasty path slashed out in the direction of the sound. Each time, the calling would cease when we came near, and an hour of hard work would have been spent in vain. Throughout that season's work in the south, in spite of many efforts, the identity of the gallina remained a mystery. Early in 1949, the same kind of bird was once heard calling, but attempts to find it again ended in failure. Finally, on May 12, the field party was resting after a hurried scramble up a steep hillside after a gallina that ceased to call as soon as the top of the hill was reached. The hunters paused for about five minutes, to rest and to smoke cigarettes, and the bird was given up as lost; for past attempts to wait out the callers had always failed. Suddenly the gallina called from a tree overhead, perhaps forty feet from the ground. The sharp eyes of the na- tive guide detected the bird, and a shot brought it down. The mysterious gallina was thus found to be this strange motmot, with its enormous serrated bill.
Momotus momota lessonii Lesson Lesson Motmot
Momotus Lessonii Lesson, Rev. Zool., 1842; 174. (Realejo, Nica- ragua.)
One $ (LSUMZ), Paraje Nuevo, Ojo de Agua, November 2, 1937; 2 $, Canada Blanco, 7 km. NW Paraje Nuevo, February 12, 1946, and January 7, 1947; 2 S ,2 9 , Rio Atoyac, 8 km. NW Potrero, February 17— March 12, 1946; 2 unsexed (one skel.), Sala de Agua, 1500 ft., 8 km. S Potrero, February 10, 1946; 15,3 km. N Presidio, 1500 ft., December 4, 1946; 4 5 , Rio Blanco, 20 km. WNW Piedras Negras, March 17 and May 11, 1946; 1 5 , 15 km. W Piedras Negras, 300 ft., January 12, 1947; 2 $, Rio Blanco, 20 km. W Piedras Negras, October 1, 1946; 1 9 , 20 km. S Jesus Carranza, 300 ft., February' 13, 1948.
Pdjaro aziil; pdjaro coo; barrancana. The Lesson Motmot is abundant throughout the Tropical Region of Veracruz. It is found in cool, shaded places in the jungle. Favored haunts include bases of cliffs, rocks, arroyos, dimly-lighted gulches, and deep forest. It is a sedentary bird and is by no means shy. It calls regularly in the morning and, less regularly, throughout the day. Its note is reminiscent of the call of a pigeon or owl, a hollow "hoot-hoot, hoot-
5—3247
596 University of Kansas Publs., Mus. Nat. Hist.
hoot-hoot." The sound is difficult to locate. Calling birds are usually well concealed in densely-leaved trees.
There is a rather long and involved Indian legend concerning the motmot, of which several versions were heard. According to the legend, the feathers of the motmot were contributed by all the other birds of the jungle, and were put in place on the motmot by the owl. In the light of the dawn the ow4 became blinded and the last feathers used included by mistake two worn-out feathers from the macaw, which went into the tail, and a pinch of feathers from the blackbird, which were inserted on the breast. The two anomalies in the feathering of the motmot are thus accounted for!
We follow Wetmore (1943: 268) in not recognizing goldmani. Two of the three specimens from Piedras Negras have no black on the crown, and in the third specimen only a trace is evident. None of the specimens shows an approach toward coeruliceps.
Family GALBULIDAE
Galbula melanogenia Sclater Black-chinned Jacamar
Galbula melanogenia Sclater, Jardine's Contr. Orni., 1852: p. 61, pi. 90. (No locality given = Vera Paz, Guatemala; c/. Sclater, Mon. Jacamars and Puffbirds, 1879, p. 19.)
One 5, 35 km. SE Jesus Carranza, 400 ft., February 18, 1948; 1 $ (skel.), 25 km. ESE Jesiis Carranza, 400 ft., March 24, 1948.
The Jacamar was seen only in extreme southern Veracruz, near rivers and clearings in the jungle, where there was an abundance of low and unusually dense brush. In flight, the brilliant, iridescent green plumage of this bird is striking, but at rest the bird is almost impossible to see. Four individuals of this species were shot, but only two were recovered, so dense and thorny is the habitat where they customarily live. Probably less than ten others were seen.
Family RAMPHASTIDAE
Aulacorhynchus prasinus prasinus (Gould)
Emerald Toucanet
Pteroglossus prasinus Gould, Proc. Zool. Soc. London, 1834: 78. (Mexico.)
One 5, Rio Atoyac, 8 km. NW Potrero, March 3, 1946; 19,7 km. NW Potrero, 1700 ft., December 29, 1946; 1 $ and 1 $ (LSUMZ), Potrero Viejo, December 28, 1937; 1 $ (LSUMZ), El Faro, NW of Piedras Negras, May 4. 1938.
Pico verde. Toucanets are probably common in the forest of the Upper Tropical Life-zone. They occur in small groups, six to eight
LowERY AND Dalquesti Birds From Veracruz, Mexico 597
birds usually being seen together. The brilliant green color of this species matches closely the green of the tropical vegetation, and unless the birds are moving about, they are difficult to see. Those observed were usually only about thirty feet from the ground. The specimen shot by Forbes at El Faro was taken in the arid division of the Lower Tropical Life-zone, but this is the only record known to us from that area. \
Pteroglossus torquatus torquatus (Gmelin) Collared Aragari
Ramphastos torquatus Gmelin, Syst. Nat., 1, pt. 1, 178S: 354. (Central America.)
Four 5,3$ (one skel.), Rio Atoyac, 8 km. NW Potrero, February 20— March 12, 1946; 1 $ , 8 km. NW Potrero, 1500 ft., February 13, 1946; 2 5,19 (all LSUMZ), Potrero Viejo, March 7. 1938; 1 $ Rio Blanco, 20 km. WNW Piedras Negras, May 12, 1946; 15,3 km. N Presidio, 1500 ft., December 4, 1946; 1 9 (skel.), 20 km. E Jesus Carranza, February 14, 1948.
Pico real. The Collared Aragari is common in the Upper Tropical Life-zone; is fairly common in the humid division of the Lower Tropical Life-zone; and was seen but once in the arid division of the Lower Tropical Life-zone. The species usually occurs in small flocks of five to fifteen birds. The birds are usually conspicuous, and follow each other from tree to tree at short intervals. Most of their activities take place fifty feet or more above the ground. Rarely were birds of this species seen at less than thirty feet above the ground.
Ramphastos sulfuratus sulfuratus Lesson Keel-billed Toucan
Ramphastos sulfuratus Lesson, Traite Orni., livr. 3, 1830: 173. (Mexico.)
One unsexed (skel.), 3 km. W San Marcos, 200 ft., November 4, 1947; 3 $ (one skel.), 1 9 (skel.), 4 km. W Tlapacoyan, 1700 ft., October 16 and 17, 1947; 1 5, Rio Blanco, 15 km. W Piedras Negras, 300 ft., November 20, 1946; 1 $, Rio Blanco, 20 km. WNW Piedras Negras, 300 ft., November 25, 1946; 1 $ (LSUMZ), Piedras Negras, El Faro, 550 ft., December 18, 1937; 1 5 (skel.), 20 km. E Jesus Carranza. 300 ft., March 19, 1948; 1 9 (skel.), 25 km. SE Jesus Carranza, 250 ft., April 1, 1949.
Pico canoa. The big toucan is not now found near Potrero Viejo, but there are old records from that general area, and several flocks were seen in the Upper Tropical Life-zone near Tlapacoyan. These birds were fairly common along the Rio Blanco, and were abundant in southern Veracruz. Only near Tlapacoyan were they seen in flocks. Elsewhere only pairs and single birds were seen, although, it is true, most of the observations in southern Veracruz and on the
598 University of Kansas Publs,, Mus. Nat. Hist.
coastal plain were made in April, May, and early June, when, per- haps, the species is paired for the breeding season. These toucans are rather inactive, and spend considerable time perched promi- nently on high limbs in the treetops, occasionally swinging their big bills from side to side. Their call, a low "croak-croak-croak," is given throughout the day. Toucans were found roosting at night on several occasions. They were not high in the trees as might be expected, but instead were on isolated branches of jungle trees, only twenty to twenty-five feet from the ground. In southern Veracruz the Keel-billed Toucan is considered a game species, and is shot for food on occasions.
Family PICIDAE
Centurus aurifrons veraecrucis (Nelson) Golden-fronted Woodpecker
Melanerpes dubius veraecrucis Nelson, Auk, 27, 1900: 259. (Coatza- coalcos, Veracruz, Mexico.)
One 9 (skeI.),Teocelo, 4500 ft., January 2, 1949; 1 $ (skel.), 5 km. ENE El Jobo, 600 ft., October 19, 1947; 1 $ (skel.), Potrero Viejo, 1700 ft., October 23, 1947; 2 $ (LSUMZ), Paraje Nuevo, 1700 ft., April 4, 1938; 2$ (one skel.),
1 ?, Rio Atoyac, 8 km. NW Potrero, February 21— March 20, 1946; 1 2,
2 $ (one skel.), Rio Blanco, 20 km. WNW Piedras Negras, May 13 and 14, and September 29, 1946; 2 $ (one skel.), 3 km. E San Andres Tuxtla, 1000 ft., January 19, 1948; 1 5 (skel.), 20 km. ENE Jesus Carranza, 200 ft., March 26, 1949.
Chincheri. The Golden-fronted Woodpecker occurs throughout the Tropical Life-zones of Veracruz, and is usually the commonest woodpecker found in any locality. Although other races are known to occur in the state, only veraecrucis is represented among the skins available in the present collections; the skeletal material was not determined subspecifically.
Balanosphyra formicivora formicivora (Swainson) Ant-eating Woodpecker
Picus jormicivorus Swainson, Philos. Mag., n. s., 1, no. 6, June, 1827: 439. (Temascaltepec, Me.xico.)
Two $ (skels.), 15 km. ENE Tlacotepec, 1500 ft., December 10 and 12, 1947.
This wide-ranging species is fairly common in the low-lying oak forest between the Upper Tropical Life-zone and the arid division of the Lower Tropical Life-zone. The oak belt is now much reduced in area both because it is the most fertile part of the level land, and consequently is much utilized for agriculture, and because the oak
LowERY AND Dalquest: Birds From Veracruz, Mexico 599
makes a superior grade of charcoal that has resulted in processing most of the trees.
Piculus rubiginosus aeruginosus (Malherbe) Lichtenstein Woodpecker
Chrysopicus aeruginosus Malherbe, Mon. Picidae, 2, 1862: 171. (Mexico.)
One $ (LSUMZ), Potrero Viejo, August 20, 1937.
This specimen from Potrero Viejo shows no approach to Picidus rubiginosus yucatanensis, which is known to occur in southern Vera- cruz, at least as far north as Presidio and Montzorongo, and possibly as far as Orizaba (Ridgway, 1914: 132). If P. rubiginosus does occur at Orizaba, only twenty miles east of Potrero Viejo, the two forms may be sympatric and further investigation may show that rubiginosus and aeruginosus are not conspecific.
Celeus castaneus (Wagler)
Chestnut-colored Woodpecker
Picus castaneus Wagler, Isis, 1829: 515. (No locality given; restricted to Veracruz, Mexico, by Corv', Publ. Field Mus. Nat. Hist., zool. ser., 13, pt. 2, Dec. 31, 1919, p. 453!)
One 5, 3 km. ESE Jesus Carranza, 300 ft., May 10, 1949; 1 $ , 35 km. SE Jesus Carranza, 400 ft., April 30, 1948.
This woodpecker was found only in the deep jungles of extreme southern Veracruz. A few individuals were seen each day spent along the Rio Solosuchi and Rio Chalchijapa, in April, May, and June, in 1948 and 1949.
Dryocopus lineatus similis (Lesson) Lineated Woodpecker
Picus similis Lesson, Oeuvres completes de Buffon, 20, April, 1847: 204. (San Carlos, El Salvador.)
Two $ , Rio Atoyac, 8 km. NW Potrero, March 9 and 13. 1946; 3 S (skels.) 1 $, Rio Blanco, 20 km. WNW Piedras Xegras, May 11-26; 1946; 1 $, Rio Blanco, 20 km. W Piedras Negras, October 1, 1946.
Carpinfero. This large species is found throughout the Tropical Region of Veracruz and, next to the Golden-fronted Woodpecker, is the most common woodpecker found in the state. The wing meas- urements of our skins are 172, 172, 175, 181 mm.
Phloeoceastes guatemalensis regius (Reichenbach) Guatemalan Ivory-billed Woodpecker
Campephilus regius Reichenbach, Handb. Spec. Orni., Scansores, Pic- inae, 1854: p. 393, pi. 649, figs. 4331-4332. (Papantla, Veracruz.)
One $ (skel.), 20 km. ENE Jesus Carranza, 200 ft., March 21, 1949.
600 University of Kansas Publs., Mus. Nat, Hist.
Veniliornis fumigatus oleaginus (Lichtenstein) Oleaginous Woodpecker Picus oleaginus Lichenstein, Preis-Verz. Vogel. Mex., 1830: 1. (Mexico.) One $ , Rio Atoyac, 8 km. NW Potrero, March 3, 1946.
This interesting woodpecker was seen only once, on a dead tree trunk, about six feet from the ground in the dense forest of a hill- side in the Upper Tropical Life-zone.
Dendrocopos villosus jardinii (Malherbe) Hairy Woodpecker
Picus (Leuconotopicus) jardinii Malherbe, Rev. Zool. 1845: 374. (Veracruz, Mexico; cf. Cory, Field Mus. Nat. Hist., zool. ser., 13, pt. 2, Dec. 31, 1919, p. 488.)
One 5 , 3 km. E Las Vigas, 8000 ft., November 1, 1946.
The Hairy Woodpecker is not uncommon in the pine forest near Las Vigas. It was not seen elsewhere.
Dendrocopos scalaris ridgwayi (Oberholser) Ladder-backed Woodpecker
Dryobates scalaris ridgwayi Oberholser, Proc. U. S. Nat. Mus., 41, 1911 : p. 140, in key, p. 143. (Jaltipan, Veracruz, Mexico.)
One 9, 4 km. W Paso de San Juan, 250 ft., December 15, 1948; 1 5 , Rio Blanco, 20 km. WNW Piedras Negras, May 27, 1946; 1 $ (skel.), Rio Blanco, 20 km. W Piedras Negras, October 6, 1946; 1 $ (LSUMZ), Matacabresto, above Piedras Negras, 500 ft., November 4, 1937.
The Ladder-backed Woodpecker was noted on only a few occa- sions, in the jungle at low elevations.
Family DENDROCOLAPTIDAE
Xiphorhynchus flavigaster saltuarius Wetmore Ivory-billed Woodhewer
Xiphorhynchus flavigaster saltuMrius Wetmore, Auk, 59, 1942: 266. (Altamira, Tamaulipas, Mexico.)
One $ , 4 km. E Papantla, 400 ft., November 10, 1947.
This specimen is definitely paler on the under parts than speci- mens from farther south in Veracruz, and is therefore assignable to saltuarius instead of eburneirostris. In this regard the examples taken in the Potrero region are somewhat intermediate, but they are closer to the latter, to which they are here provisionally referred.
LowERY AND Dalquest: Birds From Veracruz, Mexico 601
Xiphorhynchus flavigaster eburneirostris (Des Murs) Ivory-billed Woodhewer
Dryocopus eburneirostris Des Murs, Iconog. Orni., livr. 9, 1847: pi. 52, with text. (Realejo, Nicaragua.)
One $ (skel.) 5 km. ENE El Jobo, 600 ft., October 19, 1947; 1 9, Rio Atoyac, 8 km. NW Potrero, February 19, 1946; 1 5 , 7 km. NW Potrero, 1700 ft., January 10, 1947; 15,2 km. N Motzorongo, 1500 ft., December 7, 1946, 1 $ (skel.), 20 km. E Jesus Carranza, 300 ft., February 10, 1948.
Lepidocolaptes souleyetii insignis (Nelson) Streaked-headed Woodhewer
Picolaptes compressus insignis Nelson, Auk, 14, 1897: 54. (Otatitlan, Veracruz, Mexico.)
One $, 2 km. N Motzorongo, 1500 ft., December 10, 1946; 1 9 , 20 km. S Jesus Carranza, 300 ft.. March 20, 1948; 1 $, I 9, skels., 20 km. ENE Jesus Carranza, 200 ft., March 20-21, 1949.
This species seems to be far less common than Xiphorhynchus flavigaster and was not detected north of the vicinity of Motzorongo.
Glyphorynchus spirurus pectoralis Sclater and Salvin
Wedgebill
Glyphorhynchus pectoralis Sclater and Salvin, Proc. Zool. Soc. London. 28, 1860: 299. (Vera Paz, Guatemala.)
One 9 , 35 km. SW Jesus Carranza, 400 ft., February 19, 1948.
The specimen taken was working up the side of a tree in the deep jungle, in much the manner of a Brown Creeper.
Sittasomus griseicapillus sylviodes Lafresnaye Mexican Sittasomus
Sitta-somits sylviodes Lafresnaye, Rev. et Mag. Zool., (2), 2, 1850: 590. (Mexique; restricted to Veracruz. Mexico, by Bangs and Peters, Bull. Mus. Comp. Zool.. 68, 1928, p. 392.)
One unsexed (LSUMZ). Potrero Viejo, August 22, 1938; 1 5 , Rio Atoyac, 8 km. NW Potrero, March 3, 1947.
This little dendrocolaptid was seen several times, always near the bases of tall trees in dense forest in the Upper Tropical Life-zone.
Family FURNARIDAE
Synallaxis erythrothorax furtiva Bangs and Peters Rufous-breasted Synallaxis
Synallaxis erthrothorax furtiva Bangs and Peters, Bull. Mus. Comp. Zool., 67, Jan., 1927: 476. (Presidio, Veracruz, Mexico.)
One S, Rio Atoyac, 8 km. NW Potrero, March 11, 1947; 1 9 (LSUMZ) and 15,1 unsexed, (skels.), Potrero Viejo, August 22, 1938, and April 24,
602 University of Kansas Publs., Mus. Nat. Hist.
1949; 1 ?, 20 km. ENE Jesus Carranza, 200 ft., April 10, 1949; 19,15 (skel.), 20 km. ENE Jesus Carranza, 300 ft., February 23 and March 20, 1948.
These little birds are fairly common in the Upper Tropical Life- zone and in the humid division of the Lower Tropical Life-zone. They are inconspicuous, living in thickets and brushy places. Their actions are much like those of wrens, which they resemble also in appearance and the display of curiosity. One specimen was taken on the ground, in a banana-baited mouse trap set beneath a bush. One nest was placed in a mass of dense vines beside a stump, about three feet from the ground. Another was about six feet from the ground in a dense bush, which was one of six or eight bushes grow- ing as an isolated thicket, twenty-five feet from a spring. A third nest was placed on the horizontal dead limb that jutted out about three feet over the water of a river. The shore was covered with dense brush that enveloped the base of the limb. This nest was about six feet from the surface of the water. All three of the nests were found in April and May. All were crude, unlined structures, composed of small twigs with numerous openings extending com- pletely through the nest. The one on the horizontal limb, perhaps unfinished, was in the form of a cup, about four inches in diameter by three inches in height. The other two nests were nearly spherical, and about five inches in diameter.
Our Veracruz specimens, as well as the large series in Washington, fail to demonstrate clearly the alleged characters of this race. Indeed, nearly all of the specimens examined by us are indistinguish- able from examples of erythrothorax taken in Guatemala.
Automolus ochrolaemus cervinigularis (Sclater) Buff-throated Automolus
Anabates cervinigularis P. L. Sclater, Proc. Zool. Soc. London, 24, 1856 [= 1857] : 288. (Cordoba, Veracruz, Mexico.)
One ?, 30 km. SSE Jesus Carranza, 300 ft.. May 11, 1949.
Sclerurus mexicanus mexicanus Sclater Mexican Sclerurus
Sclerurus mexicanus P. L. Sclater, Proc. Zool. Soc. London, 24, 1856 [= 1857] : 290. (Cordoba, Veracruz, Mexico.)
One 9 (LSUMZ), Potrero Viejo, August 22, 1938.
Sclerurus guatemalensis guatemalensis (Hartlaub) Guatemalan Sclerurus
Tinnctor guatemalensis Hartlaub, Rev. Zool., 7, 1844: 370. (Guate- mala.)
One 5, 30 km. SSE Jesus Carranza, 300 ft., May 12, 1949.
LowERY AND Dalquest: Birds From Veracruz, Mexico 603
This species has been recorded only from Jico, on such an un- certain basis that Loetscher (MS) hesitated to include it in the state list. The above specimen may, therefore, be regarded as the first indubitable record.
Family FORMICARIDAE
Thamnophilus doliatus intermedius Ridgway Mexican Antshrike
Thamnophilus doliatiis intermedius Ridgwav, Proc. U. S. Nat. Mus., 10, Aug., 1888: 581. (Trujillo, Honduras; c/. 'Peters, Bull. Mus. Comp. Zool., 69, 1929, p. 439.)
One $, 15 km. W Piedras Negras, 300 ft., January 12, 1946; 1 5 , 20 km. ENE Jesus Carranza, 200 ft., March 23, 1949.
This species seems to be uncommon in Veracruz. Perhaps a dozen individuals were seen, always in dense, thorny brush of less than five feet in height.
Microrhopias quixensis boucardi (Sclater) Boucard Antwren
Formicivora boucardi P. L. Sclater, Proc. Zool. Soc. London, 26, 1858: 300. (Acatepec, Oaxaca, Mexico.)
One S , 25 km. SE Jesus Carranza, 250 ft., April 1, 1949.
This antwren was noted only a few times, in extreme southern Veracruz. It resembles a small warbler or flycatcher in its actions.
Formicarius analis moniliger Sclater Mexican Antthrush
Formicarius moniliger P. L. Sclater, Proc. Zool. Soc. London, 24, 1856 [=Jan., 1857]: 294. (Cordoba, Veracruz, Mexico.)
One 5,1 [ 5 ], 35 and 60 km. SE Jesus Carranza, 350 and 450 ft., April 9, 1949, and March 20, 1948.
The Mexican Antthrush is a common species in the jungles of extreme southern Veracruz. It is a ground-inhabiting species, quick to run when alarmed, but taking flight only when surprised. When running, antthrushes resemble tiny tinamous. They are in- conspicuous birds, and can be seen only by sitting motionless in the forest for some time.
Grallaria guatimalensis guatimalensis Prevost and Des Murs Guatemalan Antpitta
Grallaria guatimalensis Prevost and Des Murs, Voyage autour du Monde sur . . . la Venus, Zool., Atlas, 1, 1846 [=1842]: pi. 4. (Guatemala.)
One 5 , 30 km. SSE Jesus Carranza, 300 ft., May 7, 1949.
604 University of Kansas Publs., Mus. Nat. Hist.
The only specimen taken was shot in a thicket beside a river, deep in the jungle. The observer was sitting motionless, with his back against a tree, when the bird came from behind with a swish of wings and perched on a twig six feet away. It resembled a large thrush that had lost its tail.
Family COTINGIDAE
Rhytipterna holerythra holerythra (Sclater and Salvin) Rufous Mourner
Lipaugus holerythnis Sclater and Salvin, Proc. Zool. Soc. London, 28, 1860: 300. (Choctum, Vera Paz, Guatemala.)
One 9, 35 km. SE Jesus Carranza, 400 ft., February 18, 1948; I S , 20 km. SE Jesus Carranza, 250 ft.. May 2, 1949.
Aside from a single Verreaux specimen in the United States Na- tional Museum labeled "Mexique," and Salvin and Godman's non- committal ascription of the species to Mexico (1891: 131) based probably on a skin in the British Museum also labeled simply "Mex- ico," there seem to have been no other records for the Republic until its recent collection in Oaxaca (Blake, 1949: 2) and in Vera- cruz in the course of the present study, when it was observed in num- bers in the jungles of extreme southern Veracruz. There, beside the Rio Chalchijapa and Rio Solosuchi, it was one of the more common birds. It was usually seen on or near the ground, and in the shade. It resembles somewhat an ant tanager in its actions. Birds in copu- lation were noted in early May, 1949.
Platypsaris aglaiae sumichrasti Nelson
Rose-throated Becard
Platypsaris aglaiae sumichrasti Nelson, Auk, 1897: 52. (Otatitlan, Veraciuz, Mexico.)
One 5, 5 km. N Jalapa, 4500 ft., October 18, 1946; 1 S (skel.), 1 $ , Rio Blanco, 20 km. WNW Piedras Negras, May 17 and 25, 1946.
Copeton. This bird is common in the arid division of the Lower Tropical Life-zone. It was seen only once in the Upper Tropical Life-zone.
The Jalapa specimen is immature and cannot be subspecifically identified with certainty. It is, however, somewhat darker through- out than immature examples of gravis and is therefore assigned to the southern race.
LowERY AND Dalquest: Birds From Veracruz, Mexico 605
Tityra semifasciata personata Jardine and Selby Mexican Tityra
Tityra -personata Jardine and Selby, lUust. Omi., 1, pt. 2, June, 1827: pi. 24. (Real del Monte, Hidalgo, Mexico.)
One $ (skel.), Potrero Llano, 350 ft., February 11, 1949; 2 $ (one skel.), 1 9, Rio Atoyac, 8 km. NW Potrero, February 18— March 3, 1946; 1 $, Ojo de Agua, 8 km. NW Paraje Nuevo, February 12, 1946; 3 5,22 (one skel.), Rio Blanco, 20 km. WNW Piedras Negras, May 16—25, 1946; 1 $ (LSUMZ), below Yanga toward Cuitlahuac, January 12, 1938; 1 $ (LSUMZ), San Fran- cisco de las Mesillas near Yanga, January 12, 1938.
Goyo. This tityra was one of the most common and prominent birds noted in the forests and jungles of Veracruz. It was found throughout the Tropical Region, wherever there were tall trees. Most of its activities were carried on at fifty feet or more above the ground, but occasionally, where there were low trees or high bushes laden with fruit, it descended to feed with the ant tanagers and thrushes. It is not a shy species, but is relatively inactive save when feeding.
Erator inquisitor fraserii (Kaup) Inquisitive Tityra
Psaris fraserii Kaup, Proc. Zool. Soc. London, 19, 1851 [= Oct., 1852] : p. 47, pis. 37 and 38. (Locality unknown = Veracruz, Mexico; cf. Hellmayr, Publ. Field Mus. Nat. Hist., zool. ser., 13, pt. 6, 1929, p. 223.)
One 9 (LSUMZ), Paraje Nuevo, 1700 ft., March 6, 1938.
Family PIPRIDAE
Pipra mentalis mentalis Sclater Yellow-thighed Manakin
Pipra mentalis Sclater, Proc. Zool. Soc. London, 24, 1856 [= January, 1857] : p. 299, pi. 121. (Cordoba, Veracruz, Mexico.)
Two $, 15 km. SW Jimba, 750 ft., March 5, 1947; 3 <5 , 35 km. SE Jesus Carranza, 350 ft., April 7, 1949.
A few small bands of these little birds were seen in extreme southern Veracruz, near Jesus Carranza. They seem to buzz, almost crackle, when they fly. The sound is much sharper than that of a hummingbird.
Manacus candei (Parzudaki) Cande Manakin
Pipra candei Parzudaki, Rev. Zool., 4, 1841 : 306. (Trujillo, Honduras.)
One 5 , 20 km. ENE Jesus Carranza, 200 ft., March 22, 1949.
606 University of Kansas Publs., Mus. Nat. Hist.
On the Rio Coatzacoalcos, the Indians stated that a very pretty little bird lived by a small spring in the jungle. They commonly watered their horses at this spring, and the bird always flew up and perched on a twig to chirp at them. When the spring was visited in search of the bird, the Indian guide swung off his horse, walked down the slight declivity to the spring, looked about for a moment, lifted his gun, fired, and brought down the handsome little mana- kin listed above. The purpose of an hour's ride was achieved in about two minutes. The species seems to be solitary and confined to a small home range. No others were seen, nor were others known to the natives.
Family TYRANNIDAE
Sayornis nigricans nigricans (Swainson) Black Phoebe
Tyrannula nigricans Swainson, Philos. Mag. (n. s.), 1, May, 1S27: 367. (Tableland of Mexico.)
One 9,4 km. WNW Fortin, 3200 ft., April 5, 1946; 1 $ (skel.), Rio Blanco, 20 km. WNW Piedras Negras, May 27, 1946.
In Veracruz, Black Phoebes are restricted to the vicinity of water. They perch on stones, logs, and twigs, usually over the surface of streams and ponds. Nests of the species were found in niches in limestone cliffs and clay banks beside the Rio Solosuchi and Rio Chalchijapa in April and May, 1948 and 1949.
In S. n. semiatra the under tail coverts are pure white, whereas in S. n. nigricans the under tail coverts are well marked with shaft streaks of black (never pure white). The latter condition obtains in our specimens that are listed above. Also, since in these speci- mens the under tail coverts are not solid black and since the white of the abdominal patch is not restricted, there is no basis for con- sidering them to be S. n. aquaticus, a race recently recorded from Chiapas by Blake (1949: 2).
Sayornis saya saya (Bonaparte)
Say Phoebe
Musdcapa saya Bonaparte, Amer. Orni., 1, 1825: p. 20, pi. 11, fig. 3. (Arkansas River, twenty miles from the Rocky Mountains.)
One 9, 2 km. W Limon, 7500 ft., September 26, 1948.
The Say Phoebe is common on the desert near Limon and Perote in October and November.
LowERY AND Dalquest: Birds From Veracruz, Mexico 607
Pyrocephalus rubinus blatteus Bangs Vermilion Flycatcher
Pyrocephalus rubinus blatteus Bangs, Proc. Biol. S'oc. Washington, 24, 1911: 189. (Sabune District, British Honduras.)
One $ (skel.), 5 km. W Tehuatlan, 700 ft., November 15, 1947; 19,5 km. ENE El Jobo, 600 ft., October 18, 1947; 1 $ (LSUMZ), Potrero Viejo, February 10, 1938; 2 ? , 1 5 , all LSUMZ, Paraje Nuevo, January 6 and February 5 and 10, 1938; 1 5, 15 km. ESE San Juan de la Punta, 400 ft., September 28, 1946; 3 S (one skel.), Rio Blanco, 20 km. WNW Piedras Negras, May 12 and 13, 1946; 1 $, Rio Blanco, 20 km. W Piedras Negras, 400 ft., October 3, 1946; 1 5 , 15 km. W Piedras Negras, 300 ft., January 12, 1947.
Lagunita. This species is uncommon in the Upper Tropical Life-zone. On the coastal plain, however, with its grasslands and scattered trees, it is abundant. On the basis of the material listed above, as well as on the basis of that which was examined in the United States National Museum, it is apparent that size (notably wing length) diminishes from north to south in the state of Vera- cruz. Wetmore (1943: 284) has already shown that material from San Andres Tuxtla is definitely referable to blatteus, as evidenced by wing measurements in his material ranging from 74 to 77 mm. in males, and 72.7 to 73.4 mm. in females. In the present material, the wings of males from Piedras Negras range from 73.4 to 76.3, and hence may be regarded also as blatteus. Farther north, how- ever, in the region of Potrero, males average considerably larger — 77.6 to 81.9 mm. A single female from El Jobo has the wing ex- ceptionally long (84.0 mm.) and is possibly a migrant from still farther north in Mexico where birds of this size occur and are known under the name of mexicaniis.
Muscivora forficata (Gmelin) Scissor-tailed Flycatcher
Muscicapa forficata Gmelin, Syst. Nat., 1, pt. 2, 1789: 931. (In nova Hispania = Mexico.)
One S ,1 9 (skel.), Rio Blanco, 20 km. W Piedras Negras, 400 ft., October 3, 1946; 1 S (LSUMZ), Piedras Negras, below El Faro, 450 ft., October 20, 1937; 2 $ (LSUMZ), Piedras Negras, below El Faro, 500 ft., October 20, 1937, and December 17, 1938.
Tijereta. In the first week of October these birds were abundant on the coastal plain near Piedras Negras. Flocks of ten to 100 were perched on weed stalks, bushes, and other low vegetation. A few individuals of Muscivora tyrannus were with them. The December 17 specimen listed above probably constitutes the first winter rec- ord for the species in the state.
608 University of Kansas Publs., Mus. Nat. Hist.
Muscivora tyrannus monachus (Hartlaub) Forked-tailed Flycatcher
Tyrannus (Milvulus) monachus Hartlaub, Rev. Zool., 7, 1844: 214. (Guatemala.)
One $, Rio Blanco, 20 km. WNW Piedras Negras, May 27, 1946; 1 S (skel.), Rio Blanco, 20 km. W Piedras Negras, October 3, 1946.
Tijereta. This species is an uncommon resident on the coastal plain. One pair that lived near Piedras Negras in May, 1946, fed almost entirely over water. A few were noted there with large flocks of Muscivora forficata in October, 1946.
Tyrannus melancholicus chloronotus Berlepsch Melancholy Kingbird
Tyrannus chloronotus Berlepsch, Ornis, 14, 1907: 474. (Temax, Yuca- tan.)
One 5,25 (one skel.), Rio Blanco, 20 km. WNW Piedras Negras, May 27- 30, 1946; 1 9 , 15 km. W Piedras Negras, January 15, 1947; 1 $ (LSUMZ), Piedras Negras, 450 ft.. May 2, 1938.
This species was found only in the arid division of the Lower Tropical Life-zone.
Legatus leucophaius variegatus (Sclater)
Striped Flycatcher
Elaenia variegata P. L. Sclater, Proc. Zool. Soc. London, 1856 [= 1857] : 297. (Cordoba, Veracruz, Mexico.)
One $ , Rio Blanco, 20 km. WNW Piedras Negras, May 13, 1946.
Myiodynastes luteiventris luteiventris Sclater Sulphur-bellied Flycatcher
Myiodynastes luteiventris P. L. Sclater, Proc. Zool. Soc. London, 1859: 42. (Orizaba, Veracruz, Mexico.)
Four S (three skels.), 3 ? (skels.), Rio Blanco, 20 km. WNW Piedras Negras, May 13-27, 1S46.
Copeton. This species is common on the coastal plain. It is aggressive, chasing even large birds from the vicinity of its perch.
Megarhynchus pitangua mexicanus (Lafresnaye) Boat-billed Flycatcher
Scaphorhynchus mexicanus Lafresnaj^e, Rev. et Mag. Zool., 3, 1851: 473. (Mexico.)
One 9 (skel.), 5 km. ENE El Jobo, 600 ft., October 18, 1947; 2 $ (one skel.), 1 9, Rio Atoyac, 8 km. NW Potrero, February 17— March 13, 1946; 1 $ (LSUMZ), Potrero, August 20, 1937; 1 S (LSUMZ), Piedras Negras, N El Faro, May 2, 1938.
Peche amarillo. This flycatcher is common in the Upper Tropical
LowERY AND Dalquest: Birds From Veracruz, Mexico 609
Life-zone. The Forbes specimen from Piedras Negras, on the coastal plain, shows the presence of the species in the arid division of the Lower Tropical Life-zone, although it seems to be rare there.
Myiozetetes similis texensis (Giraud) Social Flycatcher
Muscicapa texensis Giraud, Desc. Sixteen New Species Birds, 1841: pi. 1. (Texas = [probably] Veracruz, Mexico.)
One S (LSUMZ), Potrero Viejo, February 24, 1938; 2 $ (one skel.), Potrenj Viejo, 5 km. W Potrero, May 23, 1946, and September 30, 1947; 1 $ , Rio Blanco, 20 km. WNW Piedras Negras, May 25, 1946.
Pitangus sulfuratus texanus van Rossem Derby Flycatcher
Pitangus sulfuratus texanus van Rossem, Trans. San Diego Soc. Nat. Hist., 9, April 30, 1940: 82. (Brownsville, Texas.)
One 9 (skel.), 3 km. W Boco del Rio. 25 ft., September 28, 1947; 1 $ , Rio Atoyac, 8 km. NW Potrero, March 8, 1946; 1 $ (skel.), Potrero Viejo, 1700 ft., October 23, 1947; 1 $ (skel.), 15 km. ESE San Juan de la Punta, 400 ft., September 28, 1946; 2 $ , Rio Blanco, 20 km. WNW Piedras Negras, March 17 and May 25, 1946.
Peche amarillo. This species is probably the commonest and most prominent flycatcher in the arid division of the Lower Tropical Life-zone; it is slightly less common in the Upper Tropical Life-zone than Megarhynchus pitangua, which it closely resembles in life.
As stated by van Rossem (1940: 82), central Veracruz lies in the region of intergradation between texanus and guatimalensis. The skins listed above bear out this statement. One female from near Piedras Negras has as much white on the frontal area as most speci- mens of texanus examined in the present connection, whereas the Rio Atoyac female, actually taken farther north, is almost identical with certain specimens of guatimalensis that we have studied. Our three specimens from Rio Atoyac and Piedras Negras measure as follows: wing, 111, 119, 124 mm.; tail, 88, 89, 97 mm., respectively. This again indicates the variability of the species in central Vera- cruz.
Myiarchus tyrannulus cooperi Baird Mexican Crested Flycatcher
Myiarchus cooperi Baird, in Baird, Cassin. and Lawrence, Rept. Expl. R. R. Pac, 9, 1858: 180. (Mexico; cj. Deignan, Condor, 51, 1949, p. 270.)
One $ (skel.), 4 km. W Paso de San Juan, 250 ft., December 16, 1947; 1 $ , 15 km. ESE San Juan de la Punta, 400 ft., September 27, 1946.
The Paso de San Juan specimen listed above appears to constitute the first winter record for this species in Veracruz.
610 University of Kansas Publs., Mus. Nat. Hist.
Myiarchus tuberculifer lawrenceii (Giraud) Dusky-capped Flycatcher
Muscicapa lawn-enceii Giraud, Desc. Sixteen New Species Birds, 1841 : p. [9], pi. [2], fig. 1. (Texas = Nuevo Leon, Mexico.)
One 9, Rio Atoyac, 8 km. NW Potrero, March 5, 1946; 1 $, Rio Blanco, 20 km. WNW Piedras Negras, May 27, 1946.
Nuttallornis borealis (Swainson) Olive-sided Flycatcher
Tyrannus borealis Swainson, Fauna Bor.-Amer., 2, 1831 [^Feb., 1832] : p. 141, pi. 35. (Cumberland House [=Carelton House], Saskatchewan.)
One S (LSUMZ), Potrero Viejo, January 23, 1938.
The specimen taken by Forbes at Potrero Viejo on January 23 appears to be the first known instance of the occurrence of this spe- cies in Veracruz in winter.
Contopus richardsonii sordidulus Sclater Western Wood Pewee
Contopus sordidulus Sclater, Proc. Zool. Soc. London, 27, June, 1859: 43. (Southern Mexico and Guatemala; the marked type in British Museum is from Orizaba.)
One unsexed (skel.), 15 km. ESE San Juan de la Punta, 400 ft., September 28, 1946.
This specimen is a skeleton. Before being cleaned, however, the wing (with primaries intact) was carefully measured, as was also the tail. These measurements fell within the limits assigned by Ridg- way (1907: 525) to this resident race.
Contopus pertinax pertinax Cabanis and Heine Swainson Pewee
Contopus pertinax Cabanis and Heine, Mus. Hein., 2, 1859: 72. (Jaiapa, Veracruz, Mexico.)
One $,4 ? , all LSUMZ, Jaiapa, May 27 and 29, 1949.
Empidonax flaviventris (Baird and Baird)
Yellow-bellied Flycatcher
Tyrannula flaviventns W. M. and S. F. Baird, Proc. Acad. Nat. Sci. Philadelphia, 1, July— Aug. [Sept. 18], 1843: 283. (Carlisle, Pennsyl- vania.)
One 5 ,4 km. W Tlapacoyan, 1700 ft., October 15, 1947; 1 $ (LSUMZ), 3.5 mi. S Jaiapa, May 28, 1949; 1 9, Rio Atoyac, 8 km. NW Potrero, March 21, 1946; 1 $, Potrero Viejo, 1700 ft., October 24, 1948; 1 $ (LSUMZ), 1 mi. W Fortin, 3200 ft.. May 25, 1949; 2 unsexed (skels.), 15 km. ESE San Juan de la Punta, 400 ft., September 28, 1946; 1 S , Rio Blanco, 20 km. WNW Piedras Negras, May 18, 1946.
LowERY AND Dalquest: Birds From Veracruz, Mexico 611
Empidonax traillii brewsteri Oberholser Alder Flycatcher
Empidonax traillii brewsteri Oberholser, Ohio Joum. Sci., 18, 1918: 93. (Cloverdale, Nye County, Nevada.)
One $ (LSUMZ), Jalapa, May 27, 1949.
The specimen taken at El Conejo on May 15, 1940, and identified by Wetmore (1943: 290) as E. t. traillii appears to be the only previous example of the Alder Flycatcher from Veracruz. The individual listed above, shot by Robert J. Newman at Jalapa on the late date of May 29, is apparently, therefore, only the second of its species to be collected in the state, and the first of the race brewsteri, to which it is definitely assignable. Previous to either of these records, however, Loetscher (MS) had included the Alder Flycatcher in the Veracruz list on the basis of a bird seen and heard by him at Coatepec on May 5, 1939.
Empidonax minimus (Baird and Baird) Least Flycatcher
Tyrannula minima W. M. and S. F. Baird, Proc. Acad. Nat. Sci. Phila- delphia, 1, July-Aug. [Sept. 18], 1843: 284. (Cumberland County, Pa. = Carlisle, Pennsylvania.)
Two $ , Rio Atoyac, 8 km. NW Potrero, March 8 and 11, 1946; 1 9 , 15 km. ESE San Juan de la Punta, 400 ft., September 27, 1^6; 1 $, 3 km. E San Andres Tuxtla, 1000 ft., January 19, 1948.
Empidonax albigularis axillaris Ridgway White-throated Flycatcher
Empidonax axillaris Ridgway, in Baird, Brewer, and Ridgway, Hist. N. Amer. Birds, 2, 1874: 363. (Orizaba, Veracruz.)
One 9 (LSUMZ), Jalapa, May 27, 1949.
Empidonax fulvifrons rubicundus Cabanis and Heine Fulvous Flycatcher
Empidonax rubicundus Cabanis and Heine, Mus. Hein., 2, 1859: 70, footnote. (Mexico.)
One 9 (LSUMZ), Las Vigas, 7800 ft.. May 30, 1949.
Since, as is well known, Giraud's type of Muscicapa fulvifrons (1841) in all probability did not come from Texas as stated, but instead was taken somewhere in Mexico, it would be desirable to restrict the type locality by referring the type specimen to some definite population in that Republic. One reason why this has not been done already is the fact that the type of fulvifrons long has been considered unique in respect to length of wing and tail. The wing measures 66.5 mm., the tail, 57 mm. (Ridgway, 1907, p. 588,
6—3247
612 University of Kansas Publs., Mus. Nat. Hist.
gives the wing as 68 mm.), whereas other examples of this species from Mexico and the United States are much smaller. Since all of the birds described by Giraud are of the sort that could have been taken on one of the two roads existing in his time that lead from Veracruz to Mexico City, it would seem desirable on first con- sideration to restrict the type locality of each of Giraud's new birds to some place located on either of these routes. This has been done in some cases. However, there is actually no sound basis for assuming that all sixteen of the "new" birds were of common origin. Indeed, Giraud possessed a number of collections that obviously came from diverse sources. Moreover, to assume that this type of Muscicapa fulvifrons came from Veracruz imposes serious difficulties, since examples of the species from that general area are assignable to E. /. rubicundus. Recently we have examined a specimen of fulvifrons belonging to the Museum of Comparative Zoology, which was taken by W. W. Brown at Miquiahuana, Ta- maulipas. The measurements of this specimen meet fairly satis- factorily the specifications of the type in that the wing is 64.1 mm. in length and the worn tail is 51.4 mm. If populations from north- eastern Mexico are given the name fulvifrons, the argument might, nevertheless, be advanced that Giraud's type was a migrant, for instance, from Tamaulipas, that in winter had moved down to that part of the state of Veracruz that was transected by one of the two highways which then extended from the east coast to Mexico City; in other words the type of fulvifrons still might have come from Veracruz. Since the subspecies rubicundus also has an indefinite type locality (that is to say, "Mexico"), and since, as noted above, breeding populations from central Veracruz are referable to this subspecies, it would be logical, eventually, to restrict the type locality of rubicundus to some locality outside of that state so as to avoid placing the type locality of both fulvifrons and rubicundus in the same area. The best course, however, would seem to be to restrict fidvifrons to Miquiahuana, Tamaulipas, the only locality from which there is a specimen that agrees closely with the type.
If the name fulvifrons is applied to the populations of the species occupying northeastern Mexico, a re-characterization of this popu- lation is necessary since Ridgway's description {loc. cit.) was based on Giraud's greatly faded type, which was at one time a mounted specimen. We now have for study a male and female speci- men from Miquiahuana, Tamaulipas (M. C. Z.), and one male and three females, in unworn plumage, from Pendencia, San Luis Po- tosi (LSUMZ). Both of these two population samples average
LowERY AND Dalquest: Birds From Veracruz, Mexico 613
larger (four females: wing, 59.1-60.7, avg. 60.1 mm.; tail, 48.9-53.3, avg. 50.5 mm.) than populations of ruhicundus (four females from Morelia, Las Vigas, and Rio Frio: wing, 56.6-59.2, avg. 57.9 mm.; tail, 47.0-50.2, avg. 48.3 mm.). The San Luis Potosi material comes from a locality only sixty-five miles from Miquiahuana and from the same range of mountains, agrees closely in color with the Miquiahuana material, even though the latter is somewhat worn, and the two lots of material seem to represent the same geographic race.
Notes made in Washington, D. C, by Robert J. Newman and Lowery in the course of identifying their material from the Mexican state of San Luis Potosi, are as follows: "The San Luis Potosi series (fulvifrons) is darker above than either pygmaeus or rubi- cundiis as each is represented in the United States National Museum collections. The greenish brown of the back is between Olive and Buffy Olive, the color less uniform, being more or less broken by spaces of grayish, and contrasting with the Medal Bronze and Buffy Citrine in pygmaeus. The top of the head is more dusky (less brownish) than in either pygmaeus or ruhicundus. Beneath they are nearest to rubiciindus, with the color of the breast ochra- ceous tawny, as in ruhicundus, but the flanks and abdomen are nearly as pale as in pygmaeus."
Mitrephanes phaeocercus hidalgensis Sutton and Burleigh Tufted Flycatcher
Mitrephanes phaeocercus hidalgensis Sutton and Burleigh, Wilson Bull. 52, 1940: 30. (Along main highway, 6 mi. S Jacala, at elev. 6000 ft., at La Placita, Hidalgo.)
One 5,4 km. W Tlapacoyan, 1700 ft., October 16, 1947; 1 $ (I^UMZ), 3.5 mi. S Jalapa, May 28, 1949; 1 5, Ojo de Agua, 8 km. NW Paraje Nuevo, February 12, 1946.
The Tufted Flycatcher is not uncommon in the LTpper Tropical Life-zone. The specimens from Tlapacoyan and Ojo de Agua might be M. p. hidalgensis, since they are decidedly green-backed; the backs are even greener than in near-topotypical specimens of hidal- gensis from San Luis Potosi examined in the present connection. The type of M. p. phaeocercus came from Cordoba, a place only nine kilometers northeast of Ojo de Agua. The specimen from Ojo de Agua, therefore, might be a near-topotype of phaeocercus. In San Luis Potosi, however, the species seems to be migratory, since members of field parties from Louisiana State University have not found it there in winter. Consequently, the specimen taken at Ojo de Agua, Veracruz, might be a migrant from farther north. A
614 University of Kansas Publs,, Mus. Nat. Hist.
small series of breeding birds from San Luis Potosi in the Louisiana State University Museum of Zoology demonstrates a considerable degree of progressive fading in which worn June-specimens are decidedly more brownish and hence not unlike certain examples of phaeocercus taken at a comparable season in Veracruz and Guate- mala. If the examples referred to above from Tlapocoyan and Ojo de Agua are not migrants, then hidalgensis possibly is nothing more than the fresh unfaded plumage of phaeocercus. This possibility is further strengthened by the fact that specimens of this species that have been in museums for a long time obviously have under- gone postmortem fading.
Myiobius sulphureipygius sulphureipygius (Sclater) Sulphur-rumped Flycatcher
Tyrannula sulphur ei-pygia P. L. Sclater, Proc. Zool. Soc. London, 1857: 296. (Cordoba, Veracruz.)
One $, 20 km. ENE Jesiis Carranza, 200 ft., March 20, 1949; 1 $ (skel.), 25 km. SE Jesus Carranza, 250 ft., April 2, 1949.
A common species in the deep jungles of southern Veracruz, this flycatcher is seen perched on twigs of bushes and vines, usually about three feet from the ground.
Platyrinchus cancrominus Sclater and Salvin
Mexican Flat-bill
Platyrhynchus cancromin'us Sclater and Salvin, Proc. Zool. Soc. Lon- don, 1860: 299. (Choctum, Vera Paz, Guatemala.)
One ?, 30 km. SSE Jesus Carranza, 300 ft.. May 11, 1949.
This is an uncommon species which is usually found in the same localities and habitats as Myiobius sulphureipygius. Like that species, it was seen near the ground, and was rather sedentary.
Rhynchocyclus brevirostris brevirostris (Cabanis) Short-billed Flat-bill
Cyclorhynchu^ brevirostris Cabanis, Arch. Naturg., 13, 1848: 249. , (Jalapa, Veracruz, Mexico.)
One $ (LSUMZ), Potrero Viejo, August 22, 1938.
This appears to be the northernmost record for this apparently
rare flycatcher in recent years.
Elaenia flavogaster subpagana (Sclater and Salvin)
Yellow-bellied Elaenia
Elainia subpagana Sclater and Salvin, Ibis, 2, 1860: 36. (Duenas, Guatemala.)
One $ (LSUMZ), Boca del Rio, May 26, 1949.
The male listed above weighed 26 grams (no fat) .
LowERY AND Dalquest: Birds From Veracruz, Mexico 615
Elaenia viridicata placens (Sclater) Placid Elaenia
Elainia placens Sclater, Proc. Zool. Soc. London, 27, 1859: 46. (Cordoba, Veracruz, Mexico.)
One 2 (LSUMZ), Jalapa, just E of city, 4500 ft., May 27, 1949.
Although there are old records of the species from Jalapa, in re- cent years it was thought that the species was confined to much lower altitudes. This specimen was taken by Newman immediately east of the city and at approximately the same elevation as Jalapa. The weight was 13.6 grams.
Camptostoma imberbe imberbe Sclater Beardless Fh'catcher
Cam-ptostoma imberbe Sclater, Proc. Zool. Soc. London, 25, no. 339, 1857 [Nov. 16] : 203. (In vicinitate urbis S. Andres Tuxtla, in rep. Mexicana [Veracruz].)
One 9 , Rio Blanco, 20 km. WNW Piedras Negras, May 17, 1946.
This specimen appears to be the third record of occurrence for the species in Veracruz.
Family ALAUDIDAE
Eremophila alpestris chrysolaema (Wagler) Horned Lark
Alauda chrysolaema Wagler, Isis, 1831: 530. (Mexico.)
One $ (LSUMZ), 1 mi. E Perote, May 30, 1949.
This specimen, taken from a small flock feeding beside the high- way, weighed 25.7 grams (no fat) and the testes were greatly en- larged.
Family HIRUNDINIDAE
Stelgidopteryx ruficollis fulvipennis (Sclater) Rough-winged Swallow
Cotyle julvipennis Sclater. Proc. Zool. Soc. London, 27, 1859 [= 1860] : 364. (Vicinity of Jalapa, Veracruz, Mexico.)
One $, 3 km. SE Orizaba, 5500 ft.. December 20, 1946; 1 $, Rio Atoyac, 8 km. NW Potrero, February 17, 1946; 1 $,2 ?, all LSUMZ, Jalapa, May 28, 1949.
We have assigned our specimens of the Rough-winged Swallow to fulvipennis after considerable hesitation, for we seriously ques- tion if the race can be maintained. The topotypical material from Jalapa can be matched by specimens in a large series of serripennis from the eastern United States. The wing of the male measures only
616 University of Kansas Publs., Mys. Nat. Hist.
110 mm., which corresponds almost exactly with the average for populations in the United States (Ridgway, 1904: 59; Brodkorb, 1942: 215). The wing measures in the two females, 114 and 115 mm., respectively. This is more than the average for females of serripennis, but is within the range of variation exhibited by the species. The throat of fulvipennis is allegedly strongly tinged with cinnamon. However, even though the plumage of our three speci- mens from Jalapa is not excessively worn, one of them has no cinnamon on the throat, and two show only the barest trace of this color; many examples of serripennis have as much. None of the three topotypes is darker than most specimens of serripennis. The terminal spotting on the under tail-coverts is also lacking. The naming by Brodkorb (op. cit., 217) of an additional race of this species, >S. r. stuarti, from Central America with a range extending into Veracruz as far north as Motzorongo, further complicates the problem. Accordingly, we have tentatively referred our material to fulvipennis until the geographical variation in Mexican popula- tions of the species can be reviewed in its entirety.
Hirundo rustica erythrogaster Boddaert Bam Swallow
Hirundo erythrogaster Boddaert, Tabl. PI. Enl., 1783: 45. Based on Hirondelle a ventre de Cayenne Daubenton, PI. Enl., pi. 724, fig. 1. (Cayenne.)
One S , Potrero Viejo, 7 km. W Potrero, September 24, 1946; 2 9 (LSUMZ), Cuitlahiiac, 1300 ft., April 30, 1938.
The above-listed specimens appear to be the first examples taken in the state, although Loetscher (MS) observed the species as an abundant migrant in both spring and fall.
Iridoprocne albilinea albilinea (Lawrence) Mangrove Swallow
Petrochelidon albilinea Lawrence, Ann. Lye. Nat. Hist. New York, 8, 1863: 2. (Panama.)
One $ , 20 km. ENE Jesiis Carranza, 200 ft., April 10, 1949.
This swallow is abundant in extreme southern Veracruz, where it is confined to the vicinity of water. Nests are placed in cavities in logs that project above the water. The nests are composed of dry grass, in a rough mass within the cavity, and have a deep lining of feathers. Birds were nesting in late May and early April, 1949, when a flash flood on the Rio Chalchijapa washed away most of the nests. As soon as the water receded and the cavities became dry, nests were rebuilt.
LowERY AND Dalquest: Birds From Veracruz, Mexico 617
Family CORVIDAE
Psilorhinus morio morio (Wagler) Northern Brown Jay
Pica morio Wagler, Isis, 1829: 751. (Jalapa, Veracruz, Mexico; cj. van Rossem, Bull. Mus. Comp. Zool., 77, 1934, p. 414-416.)
One $ (LSUMZ), 3 mi. S Jalapa, May 29, 1949, 1 5 (skel.), 10 km. NW Potrero, February 16, 1946; 2 ^,1$, Rio Atoyac, 8 km. NW Potrero, March 8 and 12, 1946; 1 9 (LSUMZ), Potrero Viejo, August 20, 1937; 2 $ (LSUMZ), Palma Sola, SE of Cuitlahuac, October 30 and December 11, 1937; 2 $ (skels.), Rio Blanco, 20 km. W Piedras Negras, 400 ft., October 5, 1946.
Psilorhinus mexicanus mexicanus Riippell
Southern Brown Jay
Psilorhinus mexicanus Riippell, Museum Senckenbergianum, 2, pt. 2, 1837: p. 189, pi. 11, fig. 2. (Tamaulipas, Mexico.)
One $ (LSUMZ), Arroyo de Piedra, 1.7 mi NE La Capilla, 175 ft.. May 25, 1949; 2 $ (one skel.), 1 $, Rio Blanco, 20 km. WNW Piedras Negras, May 12-30, 1946; 1 $ , 3 km. W Boca del Rio, 10 ft., December 17, 1947; 1 $ (skel.), 1 km. E Mecayucan, 200 ft., December 19, 1947; 1 $ (skel.), 5 km. ENE El Jobo, 600 ft., October 19, 1947; 1 $ (skel.), 20 km. ESE Jesus Carranza, 400 ft., March 23, 1948; 3 S (LSUMZ), Palma Sola, November 15-30, 1937.
Pepe. The two kinds of Brown Jays are together one of the most prominent features of the avifauna of Veracruz. They are absent from the Tropical Region only in the deep jungles of extreme south- ern Veracruz. Their screams and "popping" notes are among the first bird calls heard in the morning and the last heard in the eve- ning. Brown Jays are quick to take alarm, and any unusual oc- currence starts them screaming. They are most annoying to hunters, whom they will follow through the forest, and whose pres- ence they announce to all the game in the jungle. Often they are shot by angry hunters, but seemingly are never used as food. In the vicinity of a fruit tree where these birds are feeding, they are relatively quiet, and seem anxious not to draw the attention of other birds to their feeding place. In the vicinity of their nests, which are placed in densely-leaved trees, such as sour orange trees, they are completely silent. At feeding-trees, they do not quarrel with small species of birds, such as thrushes, tanagers, and trogons, but they do fight with parakeets. Sometimes a jay will select one particular parakeet, from a feeding flock, and attack it, forcing the parakeet to move along the branch on which it is perched, and eventually caus- ing it to fly. The jay will pursue the parakeet for a short distance, and then return to the tree to annoy another feeding parakeet. The first parakeet will immediately return to the tree and resume its
618 University of Kansas Publs., Mus. Nat. Hist.
interrupted feeding. In the "time of the nanchi fruit" on the coastal plain, this procedure is repeated throughout the day. The para- keets seem to accept the jays as a mere annoyance that only inter- rupts momentarily their serious business of feeding.
It is our opinion that Psilhorinus morio and P. mexicanus should be considered as full species. In the Upper Tropical Life-zone, only morio was found. On the arid coastal plain, both forms were ob- served, often in the same flock. In the breeding season, however, when the birds were mated, the pairs that were observed closely, consisted always of individuals of the same type only.
There is no detectable difference in the call notes of the two spe- cies, but each is distinctively colored. Possibly there is some hy- bridization between the two forms on the coastal plain of Veracruz, where the two species come together. Since, in the Upper Tropical Life-zone only morio was found, and since, along the Rio Coatza- coalcos, in southern Veracruz, only mexicanus was found, and since, in the intermediate area, the coastal plain, the two forms occur to- gether but retain their specific distinctions, the two jays seem to be full species, instead of only subspecies of a single species, as some authors have suggested. That they are very closely related is never- theless certain.
The female example of P. morio taken near Jalapa on May 29, 1949, weighed 277.5 grams (no fat), whereas the female of this spe- cies, taken at Arroyo de Piedra, on May 25, 1949, weighed only 224 grams (no fat).
Xanthoura yncas vivida Ridgway Green Jay
Xanthoura hixuosa vivida Ridgway, Auk, 17, 1900: 28. (Pluma, Oaxaca, Mexico.)
One S,l 9 , 4 km. WNW Fortin, 3200 ft., March 26, and April 2, 1946; 2 $ (LSUMZ), Potrero Viejo, La Cieba, 1400 ft., March 25, 1938.
Esmeralda. The Green Jay is rather uncommon in most of Vera- cruz. It was noted regularly in Metlac Canyon, near Fortin, and casually in a few other localities in the Upper Tropical Life-zone.
Family CINCLIDAE
Cinclus mexicanus mexicanus Swainson Mexican Dipper
Cinclus mexicanus Swainson, Philos. Mag., (n. s.), 1, 1827: 368. (Temascaltepec, Mexico.)
One 5 , 4 km. WNW Fortin, 3200 ft., March 30, 1946.
The Dipper was noted only along the clear, cold streams of the
LowERY AND Dalquest: Bmos From Veracruz, Mexico 619
Upper Tropical Life-zone, and then not commonly. It was seen along the Rio Banderilla, near Jalapa, and the Rio Metlac, near Fortin.
Family TROGLODYTIDAE
Campylorhynchus zonatus zonatus (Lesson) Banded Cactus Wren
Picolaptes zonatus Lesson, Cent. Zool., 1832: p. 210, pi. 70. (Californie = Orizaba, Veracruz, Mexico, fide Brodkorb, Condor, 49, p. 242.)
One $,\ 9 , 4 km. WNW Fortin, 3200 ft., March 26 and April 5, 1946; 2 $ , 2 9, Rio Atoyac, 8 km. NW Potrero, Februar>' 23, and March 15-21, 1946; 1 5, 3 km. W Boca del Rio, 10 ft., December 17, 1947; 1 5 and 1 9 (LSUMZ), Potrero Viejo, 1700 ft., March 5 and April 27, 1938; 1 9 (LSUMZ), 8.8 mi. SW Paso del Toro, 350 ft.. May 25, 1949; 1 $ (LSUMZ), 3.9 mi. S Coatepec, May 28, 1949.
This large wren is particularly abundant in the dense thickets and brush of the Upper Tropical Life-zone. It lives in such dense cover that it would be an inconspicuous species were it not so in- quisitive and noisy. A male and a female weighed, respectively, 35.8 and 36.5 grams (no fat).
Campylorhynchus rufinucha rufinucha (Les.son) Rufous-naped Cactus Wren
Picolaptes rufinucha Lesson, Ann. Sci. Nat., (2), Zool., 9, 1838: 168. (Veracruz, Mexico.)
One $ (LSUMZ), Boca del Rio, May 26, 1949, 1 9 (LSUMZ), 11 mi. SW Paso del Toro, May 25, 1949; 1 $ (LSUMZ), 24 mi. E Cordoba, May 25, 1949; 15,19 (skel.), Rio Blanco, 20 km. WNW Piedras Negras, May 14 and 28, 1946; 1 unsexed (skel.), 15 km. ESE San Juan de la Punta, 400 ft., September 28, 1946.
This cactus wren is fairly common in the arid division of the Lower Tropical Life-zone. The three Louisiana State University Museum of Zoology specimens listed above weighed, respectively, 30.8, 28.2, and 30.7 grams.
Thryomanes bewickii bairdi (Salvin and Godman) Bewick Wren
Thryothorus bairdi Salvin and Godman, Biol. Centr. — Amer., Aves, 1, 1880: 95. (Oaxaca, Mexico.)
One $ , 10 km. SW Jacales, 6500 ft., November 6, 1948.
This specimen is darker than examples of bairdi that we have examined, but this difference may be the result of either dichroma- tism, or postmortem change in the comparative material. In size it is closer to bairdi than to murinus. The measurements are as fol- lows: wing, 54.8; tail, 53.2; exposed culmen, 15.3 mm.
620 University of Kansas Publs., Mus. Nat. Hist.
Troglodytes aedon parkmanii Audubon House Wren
Troglodytes Parkmanii Audubon, Onii. Biog., 5, 1839: 310. (Columbia River [= near Fort Vancouver, Washington].)
One $ , Las Vigas, 8500 ft., October 9, 1948; 19,1 km. E Jalacingo, 6500 ft., November 13, 1948; 1 $ (skel.), 1 km. E Mecayucan, 200 ft., December 20, 1947; 1 5, 7 km. NW Potrero, 1700 ft., April 2, 1946.
Also examined were seven skeletons (5 ^ and 2 5 ) of the House Wren collected on dates between October 9, 1938, and February 12, 1949, at Tampico Alto, Potrero Llano, Teocelo, and Las Vigas. These specimens are not identifiable with certainty to the sub- species.
Troglodytes brunneicollis brunneicollis Sclater Brown-throated Wren
Troglodytes brunneicollis Sclater, Proc. Zool. Soc. London, 26, 1858 : 297. (La Parada, six leagues from Oaxaca, Mexico.)
Two $ , Las Vigas, 8000 and 8500 ft., October 15 and 17, 1948.
Henicorhina leucosticta prostheleuca (Sclater)
White-breasted Wood Wren
Scytalopus prostheleucus P. L. Sclater, Proc. Zool. Soc. London, 1856 [= 1857] : 290. (Cordoba, Veracruz, Mexico.)
One 5, 4 km. WNW Fortin, 3200 ft., April 2, 1946; 1 $ , 35 km. SE Jesus Carranza, 350 ft., April 9, 1949.
This wren is an inconspicuous resident on the ground in deep, shaded forest or jungle.
Henicorhina leucophrys mexicana Nelson
Gray-breasted Wood Wren
Henicorhina mexicana Nelson, Auk, 14, 1897: 73. (Jico, Veracruz, Mexico.)
One S (skel.), Teocelo, 4500 ft., January 30, 1949; 1 $ (LSUMZ), 3.5 mi. S
Jalapa, May 28, 1949.
The specimen from Teocelo was taken in a mouse trap.
Salpinctes obsoletus (Say) ' Common Rock Wren
Troglodytes obsoleta Say, in Long, Exped. Rocky Mts., 2, 1823: 4. (Douglas County, Colorado.)
One 2 (skel.), 6 km. WSW Zacualpilla, 6500 ft., November 9, 1948.
Our specimen is not identifiable to subspecies.
Catherpes mexicanus mexicanus (Swainson)
Canyon Wren
Thryothorus mexicanus Swainson, Zool. 111., (2), 1, 1829: pi. 11. (Real del Monte, Hidalgo, Mexico.)
One $ (skel.), 3 km. W Acultzingo, 7000 ft., October 6, 1947.
LowERY AND Dalquest: Birds From Veracruz, Mexico 621
The specimen taken was shot on a cliff where there is a distinct break between the faunas of the Sonoran and Tropical Life-zones.
Family MIMIDAE
Melanotis caerulescens caerulescens (Swainson) Blue Mockingbird
Orpheus caerulescens Swainson, Philos. Mag., n. s., 1, May, 1827: 369. (Mexico.)
One 5, 5 km. N Jalapa, 4500 ft., October 19, 1946; 1 $ (skel.), 3 km. W Acultzingo, 7000 ft.. October 8, 1947.
The Blue Mockingbird is a fairly common resident of the higher parts of the Upper Tropical Life-zone.
Dumetella carolinensis (Linnaeus) Catbird
Muscicapa carolinensis Linnaeus, Syst. Nat., ed. 12, 1, 1766: 328. (In Carolina = Virginia.)
One 5,1$, skels., Teocelo, 4000 ft., January 8 and 9, 1949; 2 $ (one skel.), Rio Atoyac, 8 km. NW Potrero, February 18 and March 12, 1946; 2 $ (one skel.), 3 km. W Gutierrez Zamora, 300 ft., November 7, 1947.
The Catbird is a common winter visitor in Veracruz, where it frequents more open situations than it does in its summer range in the United States.
Mimus polyglottos leucopterus (Vigors) Common Mockingbird
Orpheus leucopterus Vigors, in Zool. Beechey's Voy., 1839: 17. (West coast of America = Monterey, California.)
One $ , 15 km. W Piedras Negras, 300 ft., January 12, 1947.
This species is rare in Veracruz. The specimen listed was the only individual noted during the present field work in that state.
Family TURDIDAE
Turdus migratorius phillipsi Bangs American Robin
Turdus migratorius phillipsi Bangs, Proc. Biol. Soc. Washington, 37, 1915: 125. (Las Vigas, Veracruz, Mexico.)
One S (LSUMZ), Valsequillo, 15 km. W Las Vigas, May 30, 1949; 1 9 (LSUMZ), La Joya, 6 km. SE Las Vigas, 7000 ft., May 30, 1949.
The male listed above weighed 66.6 grams, the female, 78.2. The latter had a conspicuous brood patch and had a small amount of body fat, whereas the male showed no fat.
622 University of Kansas Publs., Mus. Nat. Hist.
Turdus assitnilis assimilis Cabanis Jalapa Robin
Turdus assimilis Cabanis, Mus. Hein., 1, 1850 [= October, 1851]: 4. (Jalapa, Mexico.)
One S, Rio Atoyac, 8 km. NW Potrero, February 18, 1946; 1 $ (skel.), 1 $ , 4 km. W Tlapocoyan, 1700 ft., October 14, 1947.
Primavera. The Jalapa Robin is a shy, retiring species, which was usually seen in dense forest on hill-sides. It was found only in the Upper Tropical Life-zone. The specimen from Potrero is considerably darker above than examples of assimilis from San Luis Potosi. In this respect it shows a tendency toward T. a. leu- cauchen Sclater, which Wetmore discovered to be the race occur- ring in the Tuxtla Mountains of southern Veracruz. Unfortunately, we have seen no fresh, topotypical examples of assimilis from Jalapa with which to compare our extensive series from San Luis Potosi; consequently this latter material may not represent typical assimilis.
Turdus grayi grayi Bonaparte Gray Robin
Turdus grayi Bonaparte, Proc. Zool. See. London, 1837 [June 14, 1838] : 118. (Alta Vera Paz, Guatemala; cj. Griscom, Amer. Mus. Novit., no. 438, 1930, p. 6.)
One $ (LSUMZ), Cordoba, January 6, 1938; 1 9 (skel.), 10 km. NW Potrero, February 16, 1946; 4^,1$ (skel.), Rio Atoyac, 8 km. NW Potrero, February 17-March 20, 1946; 1 $ (LSUMZ), 1 mi. W Fortin. 3000 ft., May 25, 1949; 1 9, 4 km. WNW Fortin, 3200 ft., March 28, 1946; 1 9 , 10 km. NW Potrero, February 16, 1946; 2 ^, 5 km. N Jalapa, 4500 ft., October 17 and 18, 1946; 1 5, 4 km. W Tlapacoyan, 1700 ft., October 16, 1947.
Primavera. This thrush is abundant in the forests of the Upper Tropical Life-zone. It makes much noise when it flushes. Near Potrero, in the quarter hour before dark, the bushes in which a roost was situated resounded with a dull roar, caused by these thrushes. Seemingly each new arrival startled the entire flock, causing each bird to fly with much fluttering, to another twig or branch. The birds roosted approximately three feet from the ground. About thirty birds were present, as far as could be determined.
Hylocichla mustelina (Gmelin)
Wood Thrush
Turdus mustelinus Gmelin, Syst. Nat., 1, pt. 2, 1789: 817. (New York.)
One $, 25 km. SE Jesus Carranza, 250 ft., April 3, 1949; 1 5 , 35 km. SE Jesiis Carranza, 200 ft., April 2, 1949.
LowERY AND Dalquest: Birds From Veracruz, Mexico 62'3
On the nights of April 2 and 3, 1949, several of these birds were found roosting on branches of jungle trees, ten to twenty feet from the ground. These two nights were the only occasions on which this species was noted in Veracruz.
Hylocichla ustulata ustulata (Nuttall) Russet-backed Thrush
Turdtis cestulatus [= ustulnttis'l Nuttall, Man. Orni. U. S. and Canada, 1, ed. 2, 1840 : 400 and 830 and errata, p. vi. (Forests of Oregon = Fort Vancouver, Washington.)
One 9, Rio Atoyac, 8 km. NW Potrero, February 25, 1946; 1 $ (LSUMZ), Potrero Viejo, May 15, 1938.
Although Veracruz has been included in the winter range of the Russet-backed Thrush, Loetscher (MS) failed to find any record of the subspecies from the state. The two examples mentioned above may constitute the first definite record of this bird in Veracruz.
Catharus aurantiirostris melpomene (Cabanis) Orange-billed Nightingale Thrush
Turdus melpomene Cabanis, Mus. Hein., 1, Oct., 1851: 5. (Jalapa, Veracruz, Mexico.)
One 5,5 km. N Jalapa, 4500 ft., October 19, 1946; 2 $ (LSUMZ), Jalapa, May 27, 1949; 1 $ (LSUMZ), 3 mi. S Jalapa, May 28, 1949; 2 $ (skels.), 3 9 (two skels.), Teocelo, 4500 ft., January 2-10, 1949.
The Orange-billed Nightingale Thrush was found only in the higher part of the Upper Tropical Life-zone. It is an inconspicuous species, found in dense brush and thickets. The specimens from Teocelo were taken in mouse traps set in thickets beside tiny,
cold streams.
Sialia sialis gfuatemalae Ridgway Common Bluebird
Siolia sialis guatemalae Ridgway, Proc. U. S. Nat. Mus., 5, 1882: 13. (Guatemala.)
One $ (LSUMZ), Piedras Negras, El Faro, December 17, 1938.
The Bluebird is fairly common in high country. It was found in small flocks at the upper edge of the Upper Tropical Life-zone, and in the pine forests on the mountains.
Sialia sialis episcopus Oberholser Common Bluebird
Sialia sialis episcopus Oberholser, Proc. Biol. Soc. Washington, 30, 1917: 27. (Santa Engracia, Tamaulipas, Mexico.)
One $,5 km. N Jalapa, 4500 ft., October 18, 1946.
624 University of Kansas Publs., Mus. Nat. Hist.
Although the breeding population at Jalapa is probably guate- malae, the only specimen obtained there in the course of this study proves to be an example of episcopus.
Family SYLVIIDAE
Polioptila caerulea caerulea (Linnaeus) Blue-gray Gnatcatcher
Motacilla caerulea Linnaeus, Syst. Nat., ed. 12, 1, 1766: 337. (Phila- delphia, Pennsylvania.)
One $, Rio Atoyac, 8 km. NW Potrero, March 5, 1946; 1 $ (skel.), Rio Blanco, 20 km. W Piedras Negras, October 1, 1946; 1 $ (skel.), 1 9 , 5 km. ENE El Jobo, 600 ft., October 18 and 19, 1947; 2 5 , 5 km. S Tehuatlan, 700 ft., November 12 and 14, 1947.
Polioptila caerulea deppei van Rossem Blue-gray Gnatcatcher
Polioptila deppei van Rossem, Bull. Mus. Comp. Zool., 77, December, 1934: 402. (Rio Lagartos, Yucatan, Mexico.)
One $ , Potrero Viejo, 5 km. W Potrero, Febraury 12, 1946.
Family PTILOGONATIDAE
Ptilogonys cinereus cinereus Swainson Mexican Ptilogonys
Ptilogonys cinereus Swainson, Cat. Exhib. called Modern Mexico, App., 1824: 4. (Mexico.)
One S, 5 km. N Jalapa, 4500 ft., October 18, 1946; 2 nestlings in alcohol (LSUMZ), 7.8 mi. NW Acultzingo, 7700 ft., May 24, 1949.
This species was found at the very edge of the Upper Tropical Life-zone and at the lower level of the conifers. The finding of a nest of this species near Acultzingo on May 24, 1949, by a field party from the Louisiana State University Museum of Zoology, was a fortunate ornithological discovery. The description of the nest and its location will be published elsewhere.
Family LANIIDAE
Lanius ludovicianus mexicanus Brehm
Loggerhead Shrike
Lanius mexicanus Brehm, Jour. f. Orni., 2, 1854: 145, 148. (Mexico r= probably highlands of Veracruz whence came most of Deppe's birds.)
One $ (skel.), Rio Blanco, 20 km. W Piedras Negras, 400 ft., October 3, 1946.
Shrikes were noted on few occasions, and then only on the arid coastal plain.
LowERY AND Dalquest: Birds From Veracruz, Mexico 625
Family VIREONIDAE
Vireo griseus noveboracensis (Gmelin) White-eyed Vireo
Muscicapa noveboracensis Gmelin, Syst. Nat., 1, pt. 2, 1789: 947. (New York.)
One $, 5 km. S Tehuatlan, 700 ft., November 15, 1947; 1 9 , 9 km. E Papantia, 300 ft., November 16, 1947; 1 $ (skel.), Teocelo, 4000 ft., January 6, 1949.
Vireo solitarius solitarius (Wilson)
Blue-headed Vireo
Mriscicapa solitaria Wilson, Amer. Orni., 2, 1810: p. 143, pi. 17, fig. 6. (Philadelphia, Pennsylvania.)
Two 5, Rio Atoyac, 8 km. NW Potrero, March 13 and 21, 1946.
Vireo olivaceus flavoviridis (Cassin)
Red-eyed Vireo
Vireosylvia flavomridis Cassin, Proc. Acad. Nat. Sci. Philadelphia, 5, 1851 : 152. (San Juan de Nicaragua, Nicaragua.)
One S (skel.), 5 km. ENE El Jobo, 600 ft., October 18, 1947.
This is an exceptionally late date for this vireo, which is not known to be a winter resident in Veracruz.
Vireo gilvus amauronotus Salvin and Godman
Warbling Vireo
Vireo amauronotus Salvin and Godman, Biol. Centr. — Amer., Aves, 1, 1881 : 193. (Orizaba, Veracruz, Mexico.)
One S and 1 $ (LSUMZ), 3.5 mi. S Jalapa, May 28 and 29, 1949.
The two specimens listed above weighed, respectively, 12.2 and 13.5 grams (no fat).
Family COEREBIDAE
Cyanerpes cyaneus carneipes (Sclater) Blue Honey-creeper
Coereba carneipes P. L. Sclater, Proc. Zool. Soc. London, 27, 1859 [=: February, 1860] : 376. (Playa Vicente, Oaxaca, Mexico.)
One 5 , 30 km. SE Jesus Carranza, 500 ft., April 10, 1948.
The Blue Honey-creeper seems to be surprisingly rare in Veracruz. In the extreme southern part of the state, one of a pair, perched on a twig overhanging a stream, was shot. Another was seen in the same general area in May, 1949.
626 University of Kansas Publs., Mus. Nat. Hist.
Family PARULIDAE
Mniotilta varia (Linnaeus) Black and White Warbler
Motacilla varia Linnaeus, Syst. Nat., ed. 12, 1, 1766: 333. (Jamaica, Dominica = Santo Domingo.)
One 5 (skel.), Potrero Llano, 350 ft., February 10, 1949: 1 ? (LSUMZ), Potrero Viejo, August 22, 1937; 1 $ (LSUMZ), Rio Atoyac, March 12, 1938; 1 5 , 5 km. ENE El Jobo, 600 ft., October 18, 1947; 1 $ , 9 km. E Papantla, 300 ft., November 18, 1947; 1 $ (skel.), 20 km. ENE Jesiis Carranza, 200 ft., March 29, 1949.
Feucedramus olivaceus olivaceus (Giraud)
Olive Warbler
Sylma olivacea Giraud, Desc. Sixteen New Species Birds, 1841 : [p. 16, pi. 7, fig. 2]. ("Texas," error; Las Vigas, Veracruz, Mexico, suggested as type locality by Miller and Griscom.)
One 9 (LSUMZ), Valsequillo, 15 km. W Las Vigas, May 30, 1949; 1 9 (LSUMZ), La Joya, 6 km. SE Las Vigas, 7000 ft., May 30, 1949.
Dendroica petechia aestiva (Gmelin) Yellow Warbler
Motacilla aestiva Gmelin, Syst. Nat., 1, pt. 2, 1789: 996. (In Gujana, aestate in Canada = Canada.)
One S (LSUMZ), Boca del Rio, May 26, 1949.
This belated migrant weighed 10.0 grams and was notably fat.
Dendroica petechia amnicola Batchelder Yellow Warbler
Dendroica aestiva amnicola Batchelder, Proc. New England Zool. Club., 6, 1918: 82. (Curslet, Newfoundland.)
One S (LSUMZ), Paso del Macho, 1200 ft., April 5, 1938; 1 $ (LSUMZ), Potrero Viejo, 1650 ft.. May 3, 1938.
Dendroica magnolia (Wilson) Magnolia Warbler
Sylvia magnolia Wilson, Amer. Orni., 3, 1811: p. 65, pi. 23, fig. 2. (Fort Adams, Mississippi.)
Two S (skels.), Teocelo, 4000 ft., January 9 and February 1, 1949; 15,5 km. ENE El Jobo, 600 ft., October 18, 1947; 1 ^ , 9 km. E Papantla, 300 ft., No- vember 16, 1947; 1 S (skel.), 5 km. ENE El Jobo, 600 ft., October 19, 1947; 1 S (skel.), Boca del Rio, 10 ft., December 8, 1948: 1 $ (skel.), Potrero Viejo, 1700 ft., October 23, 1947; 1 $ (LSUMZ), Paso del Macho, April 5, 1938.
Dendroica virens virens (Gmelin) Black-throated Green Warbler
Motacilla virens Gmelin, Syst. Nat., 1, pt. 2, 1789: 985. (In Pennsyl- vania = Philadelphia, Pennsylvania.)
LowERY AND Dalquest: Birds From Veracruz, Mexico 627
One $ (skel.), Teocelo, 4000 ft., January 30, 1949; 1 $,\ 2 , Rio Atoyac, 8 km. NW Potrero, March 4 and March 21, 1946.
Dendroica occidentalis (Townsend)
Hermit Warbler
Sylvia occidentalis J. K. Townsend, Jour. Acad. Nat. Sci. Philadelphia, 7, pt. ii [November 21, 1837]: 190. (Forests of the Columbia River = Fort Vancouver, Washington.)
One $ (skel.), 3 km. E Las Vigas, 8000 ft., November 4, 1946; 1 $ (LSUMZ), Las Vigas, April 2, 1939.
Dendroica dominica albilora Ridgway Yellow-throated Warbler
Dendroica domiynca var. albilora (Baird MS) Ridgway, Amer. Natur., 7, October, 1873: 606. (Belize, British Honduras.)
One $, Potrero Vicjo, 1700 ft., October 21, 1947.
Dendroica pensylvanica (Linnaeus)
Chestnut-sided Warbler
Motacilla pensylvanica Linnaeus, Syst. Nat., ed. 12, 1, 1706: 333. (Pennsylvania.)
One S , 20 km. SE Jesus Carranza, 250 ft.. May 4, 1949.
Seiurus aurocapillus aurocapillus (Linnaeus) Oven-bird
Motacilla aurocapilla Linnaeus, Syst. Nat., ed. 12, 1, 1766: 334. (Pennsylvania = at sea approximately 30 miles off Haiti.)
One 9,5 km. N Jalapa, 4500 ft., October 17, 1946; 1 $ (skel.), 25 km. SE Jesus Carranza, 250 ft., March 30, 1949.
Seiurus motacilla (Vieillot) Louisiana Water-thrush
Turdus mofncilln Vieillot, Oiseaux Amer., September, 1807 [ = 1S08] : p. 9, pi. 65. (Kentucky.)
One S, Rio Atoyac, 8 km. NW Potrero, February 24, 1946.
Oporornis formosus (Wilson)
Kentucky Warbler
Sylvia formosa Wilson, Amer. Orni., 3, 1811: p. So, pi. 25; fig. 3. (Kentucky.)
One $ (skel.), 25 km. SE Jesus Carranza, 250 ft., March 30, 1949.
Oporornis tolmiei austinsmithi Phillips
Macgillivray Warbler
Oporornis tolmiei ausdnsynithi Phillips, Auk, 64, 1947: 298. (Emi- grant Gulch, 6500 ft., 3 mi. SE Chico, Montana.)
One $ (LSUMZ), Jalapa, May 29, 1949.
The subspecific allocation of our single specimen to one of the races described by Phillips is provisional, since the differences 7—3247
628 University of Kansas Publs., Mus. Nat. Hist.
exhibited by the various races are both minute and subject to con- siderable individual variation. In our specimen the wing measures 60 mm. and the tail, 53.5 mm. It weighed 11.7 grams and pos- sessed only a small amount of body fat. This seems to be the latest spring record for the species in the state; Loetscher (MS) did not find it present after May 10.
Geothlypis trichas trichas (Linnaeus) Common Yellow-throat
TurdxLs trichas Linnaeus, Syst. Nat., ed. 12, 1, 1766: 293. (In America septentrionali = Maryland.)
One $, i km. WNW Fortin, 3200 ft., March 23, 1946.
Geothlypis trichas typhicola Burleigh Common Yellow-throat
Geothlypis trichas typhicola Burleigh, Proc. Biol. Soc. Washington, 47, 1945: 21. (Athens, Georgia.)
One $ (LSUMZ), Boca del Rio, May 26, 1949.
The specimen from Boca del Rio is referred with some reluctance to the race typhicola, even though it agrees closely with a large series of examples of this race studied in the present connection. The specimen was exceedingly fat, as might be expected of a migrant, and it weighed 11.4 grams. The puzzling fact is that on May 26 yellow-throats within the breeding range of typhicola, in the southeastern United States, are well advanced in their nesting ac- tivities; yet here is evidence of an example of that race loitering far to the south when apparently it should have been on its breed- ing ground. Since the sex organs, according to the information on the label, showed only slight development, we can rationalize its belated occurrence in the tropics on the basis of a delayed migra- tory urge, a phenomenon that may be far more common in passer- ines than the present available facts would indicate.
Geothlypis trichas occidentalis Brewster Common Yellow-throat
Geothlypis trichas occidentalis Brewster, Bull. Nuttall Orni. Club, 8, 1883: 159."^ (Truckee River, Nevada.)
One $, 20 km. ENE Jesus Carranza, 200 ft., March 23, 1949; 1 ? , 20 km. E Jesus Carranza, May 3, 1938.
Geothlypis nelsoni nelsoni Richmond Nelson Yellow-throat
Geothlypis nelsoni Richmond, Auk, 17, 1900: 197. (Cofre de Perote, Veracruz, Mexico.)
One $ , Las Vigas, 8500 ft., October 9, 1948.
LowERY AND Dalquest: Birds From Veracruz, Mexico 629
Icteria virens auricollis (Lichtenstein)
Yellow-breasted Chat
Tanagra auricollis Lichtenstein, Preis-Verz. Vogel Mexico, September 1, 1830: 2. (Mexico = City of Mexico.)
One $,19 (skels.), 4 km. WNW Fortin, 3200 ft., March 31, 1946; 1 $, 10 km. NW Potrero, February 16, 1946; 1 $ (skel.), Rio Blanco, 20 km. W Piedras Negras, 400 ft., October 3, 1946; 1 $ (skel.), 5 km. S Tehuatlan, 700 ft., November 14, 1947; 1 $ (skel.), Potrero Viejo, 1700 ft., December 3, 1947. Also examined but not identified to subspecies: 1 $ (skel.), Tampico Alto, 50 ft., February 8, 1949; 1 S (skel.), Potrero Llano, 350 ft., February 12, 1949; 1 $, 1 9, skels., San Isidro, 100 ft., February 4, 1949; 1 $ (skel.), Puenta Nacional, 500 ft., December 12, 1948.
This species is a common winter resident in the Upper Tropical Life-zone. It is fully as retiring and shy in Veracruz as it is in the United States. Individuals were commonly taken in mouse traps set for small mammals. In brushy cover, they were some- times caught at an average of approximately one to each 100 traps set. All examples of this species from previous collections from Veracruz have been identified as /. v. virens. Ratio of length of wing to length of tail is the most reliable basis for distinguishing /. V. auricollis from /. v. virens. On some of our specimens the feathers were so broken that the critical ratio could not be ascer- tained. Consequently, some of our specimens may actually be I. V. virens.
Wilsonia citrina (Boddaert) Hooded Warbler
Muscicapa Citrina Boddaert, Table PI. Enl., 1783: 41. Based on the Gobe-mouche, de la Louisiane Daubenton, PI. Enl., pi. 666, fig. 2. (Louisiana.)
One $ , 2 km. N Motzorongo, 1500 ft., December 9, 1946.
Wilsonia pusilla pusilla (Wilson)
Wilson Warbler
Muscicapa pusilla Wilson, Amer. Orni., 3, 1811: 103. pi. 26, fig. 4. (Southern States, and lower parts of the states of New Jersey and Delaware = southern New Jersey.)
One 2 , 4 km. WNW Fortin, 3200 ft., March 26, 1946; 1 $ (skel.), Teocelo, 4000 ft., February 5, 1949; 1 $ (skel.), Potrero Viejo, 1700 ft., October 21, 1947; 1 $ (skel.), 4 km. W Tlapocoyan, 1700 ft., October 16, 1947; 1 $, Rio Atoyac, 8 km. NW Potrero, March 5, 1946; 1 $ Potrero Viejo, 1700 ft., September 30, 1947.
Wilsonia pusilla pileolata (Pallas) Wilson Warbler
Motacilla pileolata Pallas, Zoogr. Rosso-Asiatica, 1, 1811: 497. (In insula Kadiak = Kodiak Island, Alaska.)
One $ (skel.), 1 $ , 4 km. W Tlapacoyan, 1700 ft., October 16, 1947; 1 $
630 University of Kansas Publs., Mus. Nat. Hist.
(skel.), 5 km. ENE El Jobo, 600 ft., October 19, 1947; 1 5 , 9 km. E Papantla, 300 ft.', November 16, 1947; 1 ^ , 2 km. W Perote, 7500 ft., September 24, 1948; 1 S (skel.), Teocelo, 4000 ft., February 1, 1949.
Wilsonia canadensis (Linnaeus) Canada Warbler
Muscicapa canadensis Linnaeus, Syst. Nat., ed. 12, 1, 1766: 327. (Canada.)
One 9 (LSUMZ), 3.5 mi. S Jalapa, May 28, 1949; 1 9 , Rio Atoyac, 8 km. NW Potrero, March 8, 1946.
The May 28 specimen listed above represents an unusually late date for the occurrence of the species in Veracruz.
Setophaga ruticilla tricolora (Miiller) American Redstart Motacilla tncolora Muller, Natursyst., SuppL, 1776: 175 (Cayenne.) One 2 , Rio Atoyac, 8 km. NW Potrero, March 8, 1946. Alexander Wetmore kindly studied this specimen and identified it as an example of the northern race.
Myioborus miniatus miniatus (Swainson) Red-bellied Redstart
Setophaga miniata Swainson, Philos. Mag., (n. s.), 1, May, 1827: 368. (Woods of Valladolid [Michoacan], Tableland of Mexico.)
One $ (LSUMZ), La Joya, 6 km. SE Las Vigas, 7000 ft., May 30, 1949; 2 5 , 5 km. N Jalapa, 4500 ft., October 16 and 19, 1946; 1 $ (skel.), 3 km. SE Orizaba, 5500 ft., December 22, 1946.
The Red-bellied Redstart was found mainly in low, dense trees
at the upper edge of the Upper Tropical Life-zone. Sometimes birds
were separate or in pairs, but usually they were in flocks of from
six to ten.
Myioborus miniatus molochinus Wetmore
Red-bellied Redstart
Myioborus miniatus molochinus Wetmore, Proc. Biol. Soc. Washington, 55, August 13, 1942: 105. (Between 3,000 and 4,000 feet elevation on Volcan San Martin, Sierra de Tuxtla, Veracruz, Mexico.)
One 5 , 7 km. SE Volcan San Martin, Tuxtla Mts., 3000 ft., January 16, 1948.
Ergaticus ruber (Swainson)
Red Warbler
Setophaga rubra Swainson, Philos. Mag., (n. s.), 1, May, 1827: 368. (Valladolid, Michoacan, Mexico.)
Two $ (one skel.), 1 $ , 3 km. E Las Vigas, 8000 ft., November 1 and 2, 1946, and October 2, 1948; 1 $ (skel.), 6 km. SSE Altotonga, 9000 ft., November 11, 1946.
LowERY AND Dalquest: Birds From Veracruz, Mexico 631
This species is fairly common in the pine forests on the mountains. Its call note is somewhat reminiscent of the call of the Golden- crowned Kinglet. One of the specimens from Las Vigas was brought in by a domestic cat.
Basileuterus culicivorus culicivorus (Lichtenstein) Lichtenstein Warbler
Sylvia culicivora Lichtenstein, Preis-Verz. Vogel Mexico, 1830: 2. (Jalapa, Veracruz, Mexico.)
One 9,4 km. E Tlapacoyan, 1700 ft., October 16, 1947; 1 9 (LSUMZ), 3 mi. S Jalapa, May 29, 1949; 2 $ (skels.), Teocelo, 4000 ft., January 6 and 8, 1949; 1 $ (skel.), 3 km. E San Andres Tuxtla, 1000 ft., January 10, 1948.
The specimen taken near Jalapa on May 29 weighed 13 grams and held an egg about ready to be laid.
Basiletuterus rufifrons rufifrons (Swainson) Rufous-capped Warbler
Setophaga rufifrons Swainson, Anim. Mcnag., 1837: 294. (Mexico; restricted to Real Ariba, Veracruz, by Todd, Proc. U. S. Nat. Mus., 74, 1929, p. 92.)
Two $ (LSUMZ), Jalapa, May 27 and 29, 1949.
These two specimens weighed, respectively, 10.6 and 10.4 grams and possessed no body fat.
Family PLOCEIDAE
Passer domesticus (Linnaeus) English Sparrow
Fringilla domestica Linnaeus, S'yst. Nat., ed. 10, 1, 1758: 183. (In Europa = Sweden.)
Although there seems to be no previous published record of its occurrence in the state, the English Sparrow is abundant about human habitations in central Veracruz. A large flock lived about the stables at Potrero Viejo and built nests in the roofs of sheds nearby.
Family ICTERIDAE
Gymnostinops montezuma (Lesson) Montezuma Oropendola
Cacicus montezuma Lesson, Cent. Zool., livr. 2, October, 1830: p. 33, pi. 7. (Mexico.)
One $ (skel.), Teocelo, 4000 ft., Februaiy 1, 1949; 2 ^, 2$, Rio Atoyac, 8 km. NW Potrero, February 21-March 12, 1946; 1 $ , Rio Blanco, 20 km. WNW Piedras Negras, 400 ft., October 2, 1946; 2 $ (skels.), Rio Blanco, 20 km. W Piedras Negras, October 2, 1946; 2 9 (LSUMZ), Piedras Negras near El Faro, December 18, 1937, and December 18, 1938.
632 University of Kansas Publs., Mus. Nat. Hist.
Sakhua. The Oropendola is fairly common throughout the Tropi- cal Region, where it is usually seen in pairs, but sometimes in large flocks numbering up to fifty birds. In June, colonies of this species were nesting along the Rio Solosuchi. The long, pendulous nests were abundant, and usually were placed in two or three adjacent trees. From ten to twenty nests were seen in a colony. Isolated trees were utilized, especially those near rivers.
Amblycercus holosericeus holosericeus (Litchtenstein) Prevost Cacique
Sturnus holosericeus Lichtenstein, Pries-Verz. Vogel Mexico, 1830: 1. (Mexico = Alvarado, Veracruz, Mexico.)
One $ (LSUMZ), Potrero Viejo, 1700 ft., March 5, 1938; 1 ^ , Rio Atoyac, 8 km. NW Potrero, March 2, 1946; 1 $ (skel.), 20 km. E Jesus Carranza, 300 ft., February 12, 1948; 1 2 , 30 km. ESE Jesiis Carranza, 300 ft., May 10, 1949.
The Prevost Cacique is not a common bird in Veracruz. It was found in the Upper Tropical Life-zone and in the humid division of the Lower Tropical Life-zone. The species was usually observed near the ground, in dense thickets from which it flushed with a loud whirr of wings. Twenty kilometers east of Jesus Carranza, a flock of these birds was seen in the same locality almost daily, but else- where only single birds were noted.
Psomocolax oryzivorus impacifus Peters Mexican Rice Crackle
Psomocolax oryzivorus impacifus Peters, Proc. Biol. Soc. Washington, 42, 1929: 123. (Pasa Nueva, Veracruz, Mexico.)
One $ (LSUMZ), Potrero Viejo, 1700 ft., February 15, 1938.
Neither Dalquest nor Loetscher found this icterid which seems to be rare in Veracuz. Our only record is the specimen obtained by Forbes at Potrero Viejo.
Tangavius aeneus aeneus (Wagler) Red-eyed Cowbird
Psarocolius aeneus Wagler, Isis, 22, Heft 7, July, 1829: col. 758. (La- guna, Veracruz, Mexico.)
Eleven $ (seven skels.), 4 $, Potrero Viejo, 6 km. W Potrero, February 13 and 14, 1946; 3 5 , 5 km. W Potrero, February 12, 1946; 1 $ (LSUMZ), Potrero Viejo, October 14, 1938; 1 $ , Rio Blanco, 20 km. WNW Piedras Negras, May 13, 1946; 1 $ (skel.), 3 km. W Boca del Rio, 25 ft., September 28, 1947.
Tongonito. The Red-eyed Cowbird is abundant about clearings in the Upper Tropical Life-zone; it is less common on the coastal plain. Like Cassidix mexicanus, this bird does some damage to crops, especially corn and rice.
LowERY AND Dalquest: Berds From Veracruz, Mexico 633
Cassidix mexicanus mexicanus (Gmelin) Great-tailed Crackle
Corvus mexicanus Gmelin, Syst. Nat., 1, pt. 1, 1788: 375. (Mexico; restricted to Veracruz, Mexico by Lowery, Occas. Papers Mus. Zool. Louisiana State University, 1, May 4, 1938, p. 4.)
Fifteen ^,16$ (21 skels. in all), Teocelo, Potrero and environs, and 20 km. E Jesus Carranza, on numerous dates (LSUMZ and UKNHM).
Tordo. This species is abundant about clearings, villages, and water in the Upper Tropical Life-zone. It is less common in the arid division of the Lower Tropical Life-zone. This bird does some damage, especially to seedling corn. The nest is usually placed at the bases of palm fronds. Nesting is in full swing by late Feb- ruary.
Dives dives dives (Lichtenstein) Sumichrast Blackbird
Icterus dives Lichtenstein, Preis-Verz. Vogel Mexico, 1830: 1. (Mexico.)
One $ (skel.), Rio Blanco, 20 km. WNW Piedras Negras, May 28, 1946; 1 $ (skel.), Rio Atoyac, 8 km. NW Potrero, March 2, 1946.
Tongonito. This sweet-voiced icterid is common about water and open places in both the Upper Tropical Life-zone and the arid divisions of the Lower Tropical Life-zone. It is inconspicuous, but its clear call is one of the most typical notes of the bird chorus in the forests and jungles of Veracruz.
Icterus galbula (Linnaeus) Baltimore Oriole
Coracias Galbula Linnaeus, Syst. Nat., ed. 10, 1, 1758: 108. (America = Virginia.)
One 9 (skel.), Potrero Viejo, 1700 ft.. September 30, 1948; 2 5,1$ (all LSUMZ), 20 mi. SE of Cuitlahuac, 800 ft.. May 1, 1938; 1 5 , 3 km. E San Andres Tuxtla, 100 ft., Januar\' 12, 1948; 1 9 (skel.), 20 km. ENE Jesus Carranza, 200 ft., April 28, 1949.
The specimen taken on May 1 provides a date that seems to be unusually late for this species in Veracruz.
Icterus spurius (Linnaeus) Orchard Oriole
Oriolus spurius Linnaeus, Syst. Nat., ed. 12, 1, 1766: 162. (America septentrionali = South Carolina.)
One $ , 5 km. W Potrero, March 18, 1946; 1 9 (LSUMZ), Paso del Macho, April 5. 1938; 1 $ (LSUMZ), Omealca, May 15, 1938; I S , 20 km. ENE Jesus Carranza, 200 ft., April 13, 1949; 1 9 (skel.), 20 km. E Jesus Carranza, 300 ft., March 20, 1948; 1 $ (skel.), 25 km. SE Jesus Carranza, 250 ft., April 2, 1949.
634 University of Kansas Publs., Mus. Nat. Hist.
This species was not often seen. A flock of approximately fifty birds, however, was observed at Cosamaloapan on April 1, 1947.
Icterus mesomelas mesomelas (Wagler) Yellow-tailed Oriole
Psarocolius mesomelas Wagler, Isis, 1829; col. 755. (Mexico.)
One 9 (LSUMZ), Palma Sola, 1750 ft., May 1, 1938; 1 5 (LSUMZ), Paraje Nuevo, May 1, 1938; 1 $ (skel.), 20 km. E Jesus Carranza 300 ft., February 21, 1948; 1 S,l 9 , 30 km. SSE Jesus Carranza, 300 ft.. May 11, 1949.
Calandria. This oriole is common in thickets near water, usually in bushes that actually overhang the water. It is a noisy bird.
Icterus gularis tamaulipensis Ridgway Lichtenstein Oriole
Icterus gularis tamaulipensis Ridgway, Proc. Washington Acad. Sci., 3, 1901 : 152. (Alta Mira, Tamaulipas, Mexico.)
One 5,19, Rio Blanco, 20 km. WNW Piedras Negras, May 11 and 26, 1946; 1 $ (LSUMZ), Piedras Negras, March 12, 1938; 1 $ (LSUMZ), El Faro near Rio Blanco, March 12, 1938; 1 $ (LSUMZ), 15 mi. SE Palma Sola, 800 ft., May 1, 1938; 1 $ (LSUMZ), 20 mi. SE Cuitlahuac, 800 ft.. May 1, 1938; 1 $ (skel.), Potrero Llano, 350 ft., February 15, 1949; 1 $ (skel.), 1 km. W Mecayucan, 200 ft., December 19, 1947.
Calandria. The Lichtenstein Oriole is a common species, both in the Upper Tropical Life-zone and in the arid division of the Lower Tropical Life-zone.
Sturnella magna mexicana Sclater Eastern Meadowlark
Sturnella mexicana Sclater, Ibis, 3, 1861 : 179. (Jalapa, Veracruz, Mexico.)
One <5 , 2 9 (one skel.), Rio Blanco, 20 km. WNW Piedras Negras, March 17 and May 26 and 27, 1946; 1 S (skel.), 20 km. W Piedras Negras, October 1, 1946; 1 6 (LSUMZ), Yanga, November 14, 1938.
Tortilla con chile. Meadowlarks were fairly common on the extensive grasslands of the coastal plain. They were not found in the Upper Tropical Life-zone.
Specimens of this species from Veracruz are difficult to place sub- specifically. Wetmore refers his material from Tres Zapotes, with- out comment, to mexicanus instead of inexpectata, the subspecies to which we might have assigned our specimens because of their small size. Those listed above from Veracruz, and those recorded by Wetmore from Tres Zapotes, average small (wing of males, 100.3-104.0, average, 101.8 mm.; females, 91.2-97.2, average, 94.9). But, the si bspecies mexicanus, as a population, is probably no more
LowERY AND Dalquest: Birds From Veracruz, Mexico 635
than a stage in the cline between the large birds of the southern end of the Mexican plateau, named auropect oralis, and the small birds of Central America, named inexpectata. Therefore, no advantage is to be gained redefining the geographical limits of the three races in advance of a thorough revision of Mexican representatives of the species. Unfortunately, the name first proposed was mexicanus, with type locality at Jalapa, and this name must be preserved, even though it may be shown to apply to an intermediate popu- lation. The alleged difference between these two races in the color of the rectrices (Griscom, 1934:404) seems not to hold up in the materials examined by us.
Family THRAUPIDAE
Chlorophonia occipitalis occipitalis (Du Bus) Mexican Chlorophonia Euphonia occipitalis Du Bus, Esq. Orni., livr. 3. 1847 :pl. 14. (Mexico.) One $, 1 $, 3 km. E San Andres Tuxtla, 1900 ft., January 18, 1948.
This species seems to be rare in Veracruz, having been recorded previously only from Jalapa and Orizaba, more than a half cen- tury ago. Our two specimens of this brilliant-green tanager were shot with a pistol in the top of an "amate copulene" tree. The col- lector (Dalquest) had perched himself near the top of this tree to select desirable birds that were needed for specimens. The Mexican Chlorophonias were feeding with a large aggregation of euphonias, ant tanagers, and (especially) Abbot Tanagers. In the same tree, three wooly opossums (Caluromys), two tree-porcupines (Coendou), a climbing rat (Tylomys), and numerous squirrels and common opossums were obtained. The berry-like fruit in this tree was visited day and night by scores of birds and mammals and was a continuing source of valuable specimens.
Tanagra affinis Lesson Lesson Euphonia
Tanagra [Euphonia] affinis Lesson, Rev. Zool., 5, 1842: 175. (Realejo,
Nicaragua.)
One $ (skel.), 20 km. E Jesiis Carranza, 300 ft., February 23, 1948.
Tanagra lauta lauta Bangs and Penard Bonaparte Euphonia
Tanagra lauta lauta Bangs and Penard, Bull. Mus. Comp. ZooL, 63, 1919: 35. (Guatemala.)
Three $ (one skel.), 1 9 (skel.), 3 km. E San Andres Tuxtla, 1000 ft., Jan- uary 19, 1948; 1 5 , 20 km. ENE Jesus Carranza, 200 ft., March 26, 1949.
636 University of Kansas Publs., Mus. Nat. Hist,
These little tanagers, which are usually seen in flocks, were ob- served infrequently in the course of this field work. The species was found feeding on the fruit of the "amate copulene" tree.
Thraupis episcopus diaconus (Lesson) Bishop Tanager
Tanagra (Aglaia) diaconus Lesson, Rev. Zool., June, 1842: 175. (Real- ejo, Nicaragua.)
One $, Rio Blanco, 20 km. WNW Piedras Negras, May 14, 1946; 1 $ (skel.), 5 km. SW Boca del Rio, October 11, 1946; 1 5 and 1 9 (skels.), Potrero Viejo, 1700 ft., October 21-22, 1947; 1 5 and 1 $ (LSUMZ), Potrero Viejo, October 18, 1938; 1 $ (skel.), 20 km. ENE Jesus Carranza, 200 ft., March 26, 1949.
The Bishop Tanager was not uncommon, either in the Upper Tropical Life-zone or in the arid division of the Lower Tropical Life-zone. The bills of this species are sometimes so thickly covered with the dried juice and pulp of the mango fruit as to be ball- shaped.
Thraupis abbas (Lichtenstein) Abbot Tanager
Tanagra abbas Lichtenstein, Preis-Verz. Vogel Mexico, 1830: 2. (Mex- ico =rOaxaca, Mexico; cf. van Rossem, Bull. Mus. Comp. Zool., 77, 1934, p. 419.)
One $ (skel.), 4 km. W Tlapacoyan, October 15, 1947; 1 $ (skel.), Boca del Rio, 10 ft., December 17, 1947; 1 $,l $ , 5 km. N Potrero, 1500 ft., February 11, 1946; 3 $ (two LSUMZ), Potrero Viejo, January 12 and November 12, 1938, and October 23, 1947; 1 5 , Rio Blanco, 20 km. WNW Piedras Negras, May 17, 1946; 2 $ (skels.), 3 km. E San Andres Tuxtla, 1000 ft., January 19, 1948; 1 $ (skel.), 20 km. E Jesus Carranza, February 14, 1948.
Medio. This tanager is a common and prominent bird wherever there are tall trees.
Phlogothraupis sangulnolenta sanguinolenta (Lesson) Crimson-collared Tanager
Tanagra (Tachyphornis) sanguinolentus Lesson, Cent. Zool., March, 1831: p. 107, pi. 39. (Mexico.)
One $ (skel.), 20 km. ENE Jesus Carranza, 200 ft., April 28, 1949; 1 9 , 35 km. SE Jesus Carranza, 400 ft., February 15, 1948.
This brilliantly-colored tanager was found only in extreme southern Veracruz. It was not a common species, and was noted only along watercourses. In the early mornings, these birds fed in bushes and low trees, but later in the day they were seen only in higher trees. They were shy and difficult to approach.
LowERY AND Dalquest: Birds From Veracruz, Mexico 637
Piranga rubra rubra (Linnaeus)
Summer Tanager
Fringilla rubra Linnaeus, Syst. Nat., ed. 10, 1, 1758: 181. (Carolina and Virginia = South Carolina.)
One $, Rio Blanco, 20 km. WNW Piedras Negras, March 17, 1946; 1 $ (skel.), 25 km. SE Jesus Carranza, 250 ft., March 31, 1949; 1 5, 35 km. SE Jesus Carranza, 350 ft., April 7, 1949.
On the three or four occasions when these North American mi- grants were observed, they were in tamarind trees, on the coastal plain.
Piranga leucoptera leucoptera (Trudeau)
White-winged Tanager
Pyranga leucoptera Trudeau, Jour. Acad. Nat. Sci. Philadelphia, 8, 1839: 160. (Mexico.)
One $ (skel.), 4 km. W Tlapacoyan, 1700 ft., November 21, 1948; 1 $ , Rio Atoyac, 8 km. NW Potrero, March 3, 1946.
The White-winged Tanager was observed only in the Upper Tropical Life-zone, where it docs not appear to be common.
Habia rubica rubicoides (Lafresnaye) Red Ant Tanager Saltator rubicoides Lafresnaye, Rev. Zool., 7, 1844: 41. (Mexico.) Two 9 , 3 km. E San Andres Tuxtla, 1000 ft., January 12 and 19, 1948.
Habia gutturalis salvini (Berlepsch) Sclater Ant Tanager
Phoenicothraupis salvini Berlepsch, Ibis, 1883: 487. (Vera Paz, Guate- mala.)
Two $ (one skel.), 9 km. E Papantla, 300 ft., November 17, 1947; 2 $, 1 $ (two LSUMZ), Rio Atoyac, 8 km. NW Potrero, March 10 and 12, 1946; 1 ? (skel.), Potrero, 1700 ft., October 27, 1947; 2 $ (LSUMZ), Potrero Viejo, December 30, 1937, and February 15, 1938; 1 5 , 4 km. WNW Fortin, March 28, 1946; 1 $ (skel.), 1 9 , San Andres Tuxtla, January 18 and 19, 1948; 1 $ (skel.), 20 km. E Jesus Carranza, 300 ft., March 20, 1948.
This ant tanager is abundant in brushy areas and low forest throughout most of the Tropical Region. Neither the male speci- mens nor the skeletons listed above can be identified subspecifically with certainty. Better material from Papantla and Jesus Carranza probably will show that the birds there are assignable to H. g. littoralis, the lowland subspecies of this species in Veracruz.
638 University of Kansas Publs., Mus. Nat. Hist.
Cholorospingus ophthalmicus ophthalmicus (Du Bus) Brown-headed Chlorospingus
Arremon opthalmicus Du Bus, Bull. Acad. Roy. Sci. Lettres et Beaux- Arts Belgique, 14, 1847: 106. (Mexico; restricted by Lowery and Newman to Jalapa, Veracruz, Mexico, Occas. Papers Mus. Zool., Louisiana State Univ., 22, 1949, p. 8.)
Three S (one skel.) 1 $ , 4 km. W Tlapacoyan, 1700 ft., October 13— Novem- ber 21, 1947; 15,22 (all LSUMZ), 3—5.4 mi. S Jalapa, May 28, 1949; 1 $, 1 9 , 4 km. WNW Fortin, 3200 ft., April 5, 1946.
In the brush and thickets of the arroyos at the upper edge of the Upper Tropical Life-zone, the chlorospingus is fairly common. The male from near Jalapa weighed 18.4 grams (no fat) ; the two females from the same place weighed 18.9 and 17.2 grams (no fat).
Family FRINGILLIDAE
Saltator atriceps atriceps (Lesson) Black-headed Saltator
Tanagra (Saltator) atriceps Lesson, Cent. Zool., 1832: p. 208, pi. 69. (Mexico; restricted to Veracruz, Griscom, Auk, 54, April 8, 1937, p. 198.)
One $ , 10 km. NW Potrero, February 16, 1946; 1 $ , 7 km. NW Potrero, 1700 ft., January 9, 1947; 2 5,3? (one LSUMZ), Rio Atoyac, 8 km. NW Potrero, March 28, 1938, and March 1-15, 1946; 3 $,1 $, (all LSUMZ), Paraje Nuevo, 1700 ft., January 24 and May 20, 1938; 1 2 (LSUMZ), Paraje Nuevo, May 20, 1938; 2 $ (one skel.), 1 2 (skel.), Rio Blanco, 20 km. WNW Piedras Negras, May 16 and October 1, 1946.
This species is abundant and prominent in the Upper Tropical Life-zone, especially where there are many thickets and much low vegetation. It is less common in tall forest. Along the arroyos of the arid division of the Lower Tropical Life-zone of the coastal plain, it is uncommon.
Saltator coerulescens grandis (Lichtenstein) Grayish Saltator
Tanagra grandis Lichenstein, Preis-Verz. Vogel Mexico, 1830: 2. (Jalapa, Veracruz, Mexico.)
One S ,1 9 skels., Potrero Viejo, 1700 ft., October 21 and 22, 1947; 1 $,19, Rio Blanco, 20 km. WNW Piedras Negras, May 27, 1946.
The Grayish Saltator was noted twice near Potrero, along the edges of sugar cane fields. It was more common in brushy places on the coastal plain. This species is more shy than Saltator atric&ps, and less noisy. The two species of saltators are sometimes called "primavera" in Veracruz.
LowERY AND Dalquest: Birds From Veracruz, Mexico 639
Caryothraustes poliogaster poliogaster (Du Bus) Bishop Grosbeak
Pitylus -poliogaster Du Bus, Bull. Acad. Roy. Sci. Belgique, 14, 1847: 105. (Guatemala.)
One $,\ 9 , Rio Atoyac, 8 km. NW Potrero, March 10, 1946.
The two specimens listed were shot on the same day but in dif- ferent parts of the forest. They were in small flocks in the tops of tall trees. The species was not seen again.
Richmondena cardinalis coccinea (Ridgway) Cardinal
Cardinalis virginianus var. cocdneus Ridgway, Amer. Jour. Sci. (3), 5, no. 25, January, 1873: 39. (Mexico; probably Playa Vicente.)
One 9 (skel.), Mirador, 3500 ft., December 9, 1947; 4 $ (one skel.), 1 9 , Rio Blanco, 20 km. WNW Piedras Negras, May 11-31, 1946; 1 5 , 15 km. W Piedras Negras, January 12, 1947.
Cardinal. This wide-spread species is common on the coastal plain,
Pheucticus ludovicianus (Linnaeus) Rose-breasted Grosbeak Loxia ludoviciana Linnaeus, Syst. Nat., ed. 12, 1, 1766: 306. (Louisiana.) One $, 15 km. W Piedras Negras, 300 ft., January 14, 1947.
The Rose-breasted Grosbeak was seen but once in the course of our field work in Veracruz.
Passerina cyanea (Linnaeus) Indigo Bunting
Tanagra cyanea Linneaus, Syst. Nat., ed. 12, 1, 1766: 315. (Carolina = South Carolina.)
Two 9, Rio Atoyac, 8 km. NW Potrero, March 11, 1946; 1 9, 4 km. WNW Fortin, 3200 ft., April 1, 1946.
This migrant species is common in open fields in spring, usually in flocks of ten to fifty individuals. Sparrow Hawks were seen hunting this species on several occasions.
Passerina versicolor versicolor (Bonaparte) Varied Bunting
Spiza versicolor Bonaparte, Proc. Zool. Soc. London, 5, 1837 [= June 14, 1838] : 120. (Near Temascaltepec, Mexico.)
One 9 , Rio Blanco, 20 km. WNW Piedras Negras, May 30, 1946.
The specimen taken was the only one seen. It was shot from a bush on the coastal plain.
640 University of Kansas Publs,, Mus. Nat. Hist.
Passerina ciris ciris (Linnaeus)
Painted Bunting
Emberiza Ciris Linnaeus, Syst., Nat., ed. 10, 1, 1758: 179. (South Carolina.)
Two $, Rio Atoyac, 8 km. NW Potrero, February 19 and March 4, 1946; 2 $ (LSUMZ), El Faro, above Piedras Negras, May 2, 1938; 1 $ (skel.), 25 km. SE Jesus Carranza, 250 ft., April 2, 1949.
The Painted Bunting is a fairly common winter resident in thickets near water in the Upper Tropical Life-zone. The specimen taken on March 4, 1946, suggests P. c. pallidor, but the head is too dark.
Tiaris olivacea pusilla Swainson
Yellow-faced Grassquit
Tiaris pusillus Swainson, Philos. Mag., (n. s.), 1, no. 6, June, 1827: 438. (Temascaltepec and Real del Monte, Mexico.)
One 5 (skel.), 15 km. E Tlacotepec, 1500 ft., December 12, 1947.
Carpodacus mexicanus mexicanus (P. L. S. Miiller) House Finch
Fringilla mexicana P. L. S. Miiller, Natursyst., Suppl., 1766: 165. (Mexico.)
One $ (LSUMZ), Las Vigas, 7800 ft.. May 30, 1949; 1 5 , 2 km. E Perote, 7000 ft., October 12, 1947.
This species was noted on the spines of the maguey plants on the desert near Perote. The specimen taken at Las Vigas weighed 23.6 grams.
Sporophila torqueola morelleti (Bonaparte) Cinnamon-rumped Seed-eater
Spermophila morelleti Bonaparte, Consp. Gen. Aves, 1, 1850 [near end] : 497. (Guatimala [sic] = Peten, (iluatemala.)
One $ (skel.), 4 km. E Papantla, 400 ft., November 9, 1947; 1 $, 7 km., NNW Cerro Gordo, October 28, 1947; 2 $ , Hacienda Potrero, 5 km. W Potrero, March 24, 1946; 1 5, 3 km. W Acultzingo, October 7, 1947; 1^,1$ (skels.), 20 km. E Jesiis Carranza, 300 ft., February 12 and 14, 1948.
This tiny bird is abundant in the Tropical Region where there are grassy fields, weedy places, and bushes along the edges of clear- ings. The overgrown milpas, or corn fields, furnish suitable habitat, as do the weeds and grasses growing along the railroad right of ways.
LowERY AND Dalquest: Birds From Veracruz, Mexico 641
Volatinia jacarina splendens (Vieillot) Blue-black Grassquit
Fringilla splendens Vieillot, Nouv. Diet. Hist. Nat., nouv. ed., 12, 1817: 173. (Cayenne = French Guiana.)
One ,J , 3 km. SW San Marcos, 200 ft., November 3, 1947; 3 $ (two skels.), 1 9 (skel.), Rio Blanco, 20 km. WNW Piedras Negras, May 14-30, 1946; 1 $ (LSUMZ), Paraje Nuevo, February 21, 1938.
The Blue-black Grassquit is abundant in weedy places on the coastal plain. Its habits closely resemble those of Passerina cyanea.
Spinus psaltria psaltria (Say) Arkansas Goldfinch
Fringilla psaltria Say, in Long, Exped. Rocky Mts., 2, 1823: 40 (note). (Arkansas River near the mountains = near Pueblo, Colorado.)
One 5,4 km. WNW Fortin, 3200 ft., March 31, 1946; 2 $ (skels.), Potrero Viejo, 1700 ft., October 22, 1947.
This goldfinch was noted several times in orange trees near Potrero and Fortin.
Loxia curvirostra stricklandi Ridgway Red Crossbill
Loxia curvirostra stricklandi Ridgway, Proc. U. S. Nat. Mus., 8, Sep- tember 2, 1885: 354. (Mexico.)
One 5 , 4 km. SE Las Vigas, 9500 ft., November 5, 1946.
The specimen listed was shot from a flock of approximately six birds. The locality is in the arid pine forest on the northeastern slope of the Cofre de Perote.
Atlapetes pileatus pileatus Wagler Rufous-capped Atlapetes
Atlapetes pileatus Wagler, Isis, 1831: 526. (Mexico = Valley of Mexico.)
One 9 4 km. W Acultzingo, 7500 ft., June 8, 1946.
This interesting sparrow was found only at high altitude in the dense brushland that caps the lip of the Mexican plateau between the states of Veracruz and Puebla.
Atlapetes brunnei-nucha brunnei-nucha (Lafresnaye) Chestnut-capped Atlapetes
Embernagra brunnei-nucha Lafresnaye, Rev. ZooL, 2, 1839: 97. (Mexico.)
One $ , 1 km. E Jalacingo, 6500 ft., November 13, 1948; 2 $ (skels.), Teocelo, 4000 ft., December 31, 1948, and January 6, 1949; 19,5 km. N Jalapa, 4500 ft., October 18, 1946; 1 5, 3 km. SE Orizaba, 5500 ft., December 21, 1946.
642 University of Kansas Publs., Mus. Nat. Hist.
The Chestnut-capped Atlapetes is a very shy and retiring species. The specimens were all taken in mouse traps. It was found only in the Upper Tropical Life-zone.
Arremonops rufivirgatus crassirostris (Ridgway)
Texas Sparrow
[Enibernagra rufivirgatal 3 crassirostris Ridgway, Proc. U. S. Nat. Mus. 1, 1878: 248. (Cordoba and Orizaba, Veracruz, Mexico.)
One unsexed (LSUMZ), Cuitlahuac, January 12, 1939; 1 9 , 5 km. S Tehuatlan, 700 ft., November 14, 1947.
Arremonops rufivirgatus rufivirgatus (Lawrence) Texas Sparrow
Embernaga rufivirgata Lawrence, Ann. Lye. Nat. Hist. New York, 5, May, 1851: p. 112, pi. 5, fig. 2. (Rio Grande in Texas = Brownsville, Texas.)
One 9 (skel.), Tampico Alto, 50 ft., January 2, 1949; 1 9 , El Cepillo, 20 ft., February 7, 1949; 1 $ (skel.), Ozuluama, 500 ft., February 9, 1949; 1 9 (skel.), Potrero Llano, 350 ft., February 15, 1949.
The specimen from El Cepillo, in northern Veracruz, is definitely assignable to rufivirgatus instead of either ridgwayi or crassirostris. As acknowledged by the describers, ridgwayi is nothing more than an intermediate population between the green-backed, large-billed birds from southern Veracruz, and adjoining areas, and the grayish, small-billed birds of southern Texas and contiguous areas. Indeed, in material from the state of San Luis Potosi, we find it difficult to distinguish most examples of ridgwayi on the basis of color, although the bills of these birds do average somewhat less massive than do the bills in examples of crassirostris. Extreme caution must be exercised in a study of geographic variation in this species because of the high degree of postmortem change that occurs in museum material. One of the most striking instances of this phenomenon that we have seen is displayed by the series of crassirostris in the United States National Museum. The splendid new series taken by Wetmore and Carriker in the Tres Zapotes area differs markedly from the older material from that general area. The upper parts of the older specimens are much more olivaceous (less grayish) and the flanks are decidedly browner.
Pipilo ocai ocai (Lawrence)
Collared Towhee
Buarremon ocai Lawrence, Ann. Lye. Nat. Hist. New York, 8, May, 1865: 126. (Jalapa, Mexico.)
One $ (LSUMZ), 7.8 mi. by highway NW Acultzingo, May 24, 1949. Charles Sibley, who recently has made an extensive study of
LowERY AND Dalquest: Birds From Veracruz, Mexico 643
geographic variation and speciation in certain Mexican popula- tions of the genus Pipilo, has kindly examined this specimen and identified it as a typical example of P. o. ocai. He was somewhat surprised to learn that the species occurs at this locality and at the comparatively low elevation of 7,500 feet. This specimen was taken by Newman and was found by him to be in breeding condi- tion and to weigh 58 grams (no fat).
Pipilo maculatus maculatus Swainson Spotted Towhee
Pipilo maculata Swainson, Philos. Mag., n. s., 1, June, 1827: 434. (Real del Monte [Hidalgo], Mexico.)
One $ (LSUMZ), 7.8 mi. by highway NW Acultzingo, May 24, 1949.
This specimen also was identified by Sibley. In referring it to the race maculatus, he points out that it shows, nevertheless, a heavy admixture of brown dorsally which indicates a trend toward a character of populations occurring farther south in Mexico. This specimen, also taken by Newman, was obtained not more than 100 feet from the place where the example of P. o. ocai was shot. The latter specimen was in a tree on the periphery of an oak wood; the specimen of P. m. maculatus was in a thicket outside of the forest. It, like the example of ocai, was in full breeding condition. The individual weighed 45.6 grams (no fat).
Pipilo fuscus Swainson Brown Towhee
Pipilo fuscus Swainson, Phil. Mag., (n. s.), 1, 1827: 434. (Temascal- tepec, Mexico.)
One $ , 6 km. WSW Zacualpilla, 6500 ft., November 6, 1948; 1 5 , 3 km. W Limon, 7500 ft., September 30, 1948.
Since the Brown Towhees are now being revised by John Davis at the Museum of Vertebrate Zoology, we sent our two specimens to him for critical study. He finds the specimen that came from six kilometers west-southwest of Zacualpilla to be a first-year male, almost through its post-ju venal molt, and showing characters of both P. /. fuscus and P. /. potosinus. He regards the specimen from three kilometers west of Limon as showing characters of both P. /. potosinus and P. /. tori. It seems, therefore, that better material than our two specimens will be required to permit subspecific identi- fications of the two populations represented.
8—3247
644 University of Kansas Publs., Mus. Nat. Hist.
Plagiospiza superciliosa (Swainson) Striped Sparrow Aimophila superciliosa Swainson, Anim. Menag., 1837: 314. (Mexico.) One S (LSUMZ), 15 km. W Las Vigas, May 30, 1949.
The specimen listed above weighed 38.7 grams and was only moderately fat.
Passerculus sandwichensis savanna (Wilson) Savannah Sparrow
Fringilla savanna Wilson, Amer. Orni., 3, 1811: p. 55, pi. 22, fig. 3. (Savannah, and Great Egg Harbor, New Jersey = Savannah, Georgia.)
One ? , 5 km. S Tehuatlan, 700 ft., November 14, 1947.
The specimen agrees closely with examples of P. s. savanna in the United States National Museum and the Louisiana State Uni- versity Museum of Zoology.
Ammodramus savannarum perpallidus (Coues) Grasshopper SpaiTow
ICoturnicAilus passerimis] var. perpallidus "Ridgway" Coues, Key N. Amer. Bds., October. 1872 : 137, in text, ("dry western regions" = An- telope Island, Great Salt Lake, Utah.)
One S , Mirador, 3500 ft., December 8, 1947.
The wing of this specimen measures 62 mm., the tail, 45.5 mm.
Pooecetes gramineus confinus (Baird) Vesper Sparrow
[Poocaetes gramineus] var. confinus Baird, in Baird. Cassin, and Law- rence, Rep. Expl. Sm-v. R. R. Pac, 9, 1858: 448. (Western U. S. = Loup Fork of the Platte River, Nebraska.)
One $ (skel.), 1 ? , 6 km. WSW Zacualpilla, 6500 ft., November 9, 1949.
The skeleton was not identified to the subspecies. The skin of the specimen taken at the same time is, however, clearly referable to P. g. confinus.
Aimophila rufescens rufescens (Swainson) Rusty Sparrow
Pipilo rufescens Swainson, Philos. Mag. (n. s.), 1, no. 6, June, 1827: 434. (Temiscaltipec [= Temascaltepec], Mexico.)
One $ (LSUMZ), Jalapa, May 27, 1949; 1 9 (LSUMZ), Potrero Viejo, 1700 ft., August 20, 1937; 1 $,l 9 , Rio Atoyac, 8 km. NW Potrero, March 20, 1946; 1 $ (skel.), 5 km. N Jalapa, 4500 ft., October 19, 1946; 1 $ (skel.), 7 km. SE Volcan San Martin, 3000 ft., January 16, 1948.
LowERY AND Dalquest: Birds From Veracruz, Mexico 645
This shy species was usually seen in pairs, along the edges of fields in the Upper Tropical Life-zone. It was generally found on or near the ground, and was quick to retire to dense brush at the slightest alarm. The specimen taken by Newman at Jalapa weighed 40 grams (no fat) .
Aimophila ruficeps boucardi (Sclater) Rock Sparrow
ZonoLrichia boucardi Sclater, Proc. Zool. Soc. London, 1867: p. 1, pi. 1. (Orizaba, Veracruz, and La Puebla, Puebia, Mexico.)
One ?, 6 km. WSW Zacualpilla, 6500 ft., November 9, 1948.
Aimophila petenica petenica (Salvin) Peten Sparrow
Ammodromus petenicus Salvin, Proc. Zool. Soc. London, 1863: 189. (Plains of JPeten, n. e. Guatemala.)
One $ , Rio Blanco, 20 km. WNW Piedras Negras, May 30, 1946.
Junco phaeonotus phaeonotus Wagler Red-backed Junco Junco phaeonotus Wagler, Isis, 1831: col. 526. (Mexico.) Three $ (one skel.), Las Vigas, 8500 ft., October 10, 11, and 19, 1948. This species is abundant in the pine forests near Las Vigas.
Spizella passerina mexicana Nelson Chipping Sparrow
Spizella socialis mexicana Nelson, Auk, 16, 1899 : 30. (San Cristobal, Chiapas, Mexico.)
Two $ (LSUMZ), Jalapa, May 27, 1949.
These two specimens weighed 14.1 and 13.1 grams, and neither possessed fat.
Spizella pusilla wortheni Ridgway Field Sparrow
Spizella wortheni Ridgway, Proc. U. S. Nat. Must., 7, 1884 : 259. (Silver City, New Mexico.)
One unsexed, 2 km. W Limon, 7500 ft., September 24, 1948.
Both in color and size, this specimen fulfills all the specifications of wortheni {cf. Burleigh and Lowery, 1942: 209-211). This is ap- parently the first record of the occurrence of either the species or the subspecies in the state of Veracruz.
646 University of Kansas Publs., Mus. Nat. Hist,
Melospiza lincolnii lincolnii (Audubon) Lincoln Sparrow
Fringilla Lincolnii Audubon, Birds Amer. [folio], 2, 1834: pi. 193. (Labrador [= near mouth of the Natashquan River, CJuebec].)
One $ (skel.), Teocelo, 45(X) ft., January 2, 1949; 1 5, Hacienda Potrero, 5 km. W Potrero, March 24, 1946; 19,4 km. WNW Fortin, 3200 ft., April 5, 1946; 1 9 (skel.), 7 km. NNW Cerro Gordo, 1500 ft., November 27, 1947.
This species was exceedingly common in the winter months and was often seen feeding in open clearings near human dwellings.
LowERY AND Dalquest: Birds From Veracruz, Mexico 647
LITERATURE CITED Allen, J. A.
1904. Mammals of southern Mexico and Central and South America. Bull. Amer. Mus. Nat. Hist., 20:29-80.
Amekican Geographical Society
1944. Index to Map of Hispanic America 1:1,000,000. Vol. 2. Geographi- cal Names in Mexico. U. S. Government Printing Office, Washing- ton. 181 pp.
American Ornithologists' Union Committee
1931. Check-list of North American birds. 4th ed. (Amer. Orni. Union, Lancaster, Pa.), xx-526 pp., October 1.
1944. Nineteenth supplement to the American Ornithologists' Union check-list of North American birds. Auk, 61 : 441-464, July 12.
1945. Twentieth supplement to the American Ornithologists' Union check- list of North American birds. Auk, 62 : 436-449, July 18.
1946. Twenty-first supplement to the American Ornithologists' Union check-list of North American birds. Auk, 63: 428-432, July 24.
1947. Twenty-second supplement to the American Ornithologists' Union check-list of North American birds. Auk, 64: 445-452, July 22.
1948. Twenty-third supplement to the American Ornithologists' Union check-list of North American birds. Auk, 65: 438-443, July 27.
1949. Twenty-fourth supplement to the American Ornithologists' Union check-list of North American birds. Auk, 66: 281-285, July 22.
Baird, S. F., Brewer, T. M., and Ridgway, R.
1874. History of North American Birds. Vol. 2. Land Birds. 590 pp. plus vi, 30 pis., numerous text figs.
Bu\KE, E. R.
1949. First Mexican records of tropical American birds. Nat. Hist. Misc., Chicago Acad. Sci., 43:1-3, March 16, 1949.
Brodkorb, p.
1942. Notes on some races of the Rough-winged Swallow. Condor, 44: 214-217.
1943. Birds from the Gulf Lowlands of southern Mexico. Misc. Publ. Mus. Zool., Univ. of Michigan, 55:1-88, 1 map, January 30, 1943.
1948. Some birds from the lowlands of central Veracruz, Mexico. Quart.
Jour. Florida Acad. Sci., 10:31-38, March 4, 1948. 1948a. Taxonomic notes on the Laughing Falcon. Auk, 65:406-410, July 27,
1948.
Burleigh, T. D., and Lowery, G. H., Jr.
1942. Notes on the birds of southeastern Coahuila. Occas. Papers Mus. Zool. Louisiana State Univ., 12:185-212, 2 pis., March 4, 1942.
Chapman, F. M.
1898. Notes on birds observed at Jalapa and Las Vigas, Vera Cruz, Mexico. Bull. Amer. Mus. Nat. Hist., 10:15-43, February 24, 1898.
648 University of Kansas Publs., Mus. Nat. Hist.
CoNOVER, B. [see Hellmayr C. E., and]
Cory, C. B.
1918-1919. Catalogue of birds of the Americas and the adjacent islands . . . Field Mus. Nat. Hist., Zool. Ser., 13, Part 2, No. 1, 315 pp., 1 pL, March. Part 2, No. 2, 317-607 pp., 1 pi., Dec. 31.
Davis, W. B.
1945. Notes on Veracruzan birds. Auk, 62:272-286, April 16, 1945.
Dickey, D. R., and van Rossem, A. J.
1938. The birds of El Salvador. Field Mus. Nat. Hist., zool. ser., 23:1-609, 24 pis., 29 figs.
Friedmann H.
1941-1946. The birds of North and Middle America. Bull. U. S. Nat. Mus., 50. "By Robert Ridgway, continued by Herbert Friedmann." Part 9 (1941), ix- 254 pp., 16 text figs. Part 10 (1946), xii- 484, 28 text figs.
GiRAUD, P., Jr.
1841. A description of sixteen new species of North American birds de- scribed in the annals of the New York Lyceum of Natural History. New York, [18 pp. and 8 pis.].
Goldman, E. A.
1920. Mammals of Panama. Smithsonian Misc. Coll., 69 (5) : 1-309, 39 pis., 24 figs., 1 map.
Griscom, L.
1934. The ornithology of Guerrero, Mexico. Bull. Mus. Comp. Zool., 75: 367-422, 1 pi.
Hellmayr, C. E.
1925-1938. Catalogue of the birds of the Americas and the adjacent is- lands . . . Field Mus. Nat. Hist., Zool. Ser., 13. Part 4 (1925), iv-390 pp., 1 pi. Part 5 (1927), vi-517 pp. Part 6 (1929), v-258 pp. Part 7 (1934), vi-531 pp. Part 8 (1935), vi-541 pp. Part 9 (1936), v-458 pp. Part 10 (1937), v-228 pp. Part 11 (1938), vi-662 pp.
Hellmayr, C. E., and Conovbir, B.
1942-1949. Catalogue of birds of the Americas and the adjacent islands . . . Field Mus. Nat. Hist., Zool. Ser., 13. Part 1, No. 1 (1942), vi-636 pp., April 30. Part 1, No. 2 (1948), vii-434 pp., August 18. Part 1, No. 3 (1948), vi-383, December 16. Part 1, No. 4 (1949), vi-358.
Loetscher, F. W., Jr.
1941. Ornithology of the Mexican State of Veracruz with an annotated list of the birds. Unpublished Thesis, Cornell University.
Merriam, C. H.
1892. The geographic distribution of life in North America with special reference to the Mammaha. Proc. Biol. Soc. Washington, 7: 1-64, 1 map.
Merriam, C. H., Bailey, V., Nelson, E. W., and Preble, E. A.
1910. Zone map of North America. Publ. by Dept. of Agric, Biological Survey.
LowERY AND Dalquest: Birds From Veracruz, Mexico 649
Peters, J. L.
1931-1948. Check-list of birds of the world. (Harvard University Press, Cambridge), Vol. 1 (1931), xviii-345. Vol. 2 (1934), xvii-401. Vol. 3 (1937), xii-311. Vol. 4 (1940), xii-291. Vol. 5 (1945), xi-306. Vol. 6 (1948),xi-259.
RiDGWAY, R.
1904-1914. The birds of North and Middle America .... Bull. U. S. Nat. Mus., 50. Part 3 (1904), xx-801 pp., 19 pis. Part 4 (1907), xxii-974 pp., 34 pis. Part 5 (1911), xxiii-859 pp., 33 pis. Part 6 (1914), xx-882 pp., 36 pis.
Salvin, 0., and Godman, F. D.
1879-1904. Biologia Centrali-Americana. Aves. Taylor and Francis, London. 3 vols, of text issued in 74 dated parts. Vol. 1, xliv-512 pp. Vol. 2, [ii-] 598 pp. Vol. 3, iv-510 pp. Vol. 4, Plates 1-79, plus 4.
Secretari de la EcoNOMiA Nacional
1944. Coordenadas geograficas y alturas en metros sobre el nivel del mar, de las cabeceras municipales de la Republica. Index. Mexico.
SUMICHaVST, F.
1881. Enumeracion de las Aves observados in el territorio de la Re- publica Mexicana. La Naturaleza, 5:222-250.
Traylor, M. a., Jr.
1949. Notes on some Veracruz birds. Fieldiana, Zoology, 31:269-275.
Van Rossem, A. J.
1940. Notes on some North American birds of the genera Myiodynastes, Pitangua, and Myiochanes. Trans. San Diego Soc. Nat. Hist., 9: 79-86, April 30, 1940.
Wetmore, a.
1943. The birds of southern Veracruz. Proc. U. S. Nat. Mus., 93:215-340, 3 pis., 1 fig. (map).
1944. A collection of birds from northern Guanacaste, Costa Rica. Proc. U. S. Nat. Mus., 95 : 25-80, 6 pis.
Transmitted by authors, December 22, 1949.
n
23-3247
(Continued from inside of frcmt cover)
15. A new hylid frog from eastern Mexico. By Edward H. Taylor. Pp. 257-264, 1 figure in text. August 16, 1948.
16. A new extinct emydid turtle from the Lower Pliocene of Oklahoma. By Edwin C. Galbreath. Pp. 265-280, 1 plate. August 16, 1948.
17. Pliocene and Pleistocene records of fossil turtles from western Kan- sas and Oklahoma. By Edwin C. Galbreath. Pp. 281-284, 1 figure in text. August 16, 1948.
18. A new species of heteromyid rodent from the Middle Oligocene of northeastern Colorado with remarks on the skull. By Edwin C. Galbreath. Pp. 285-300, 2 plates. August 16, 1948.
19. Speciation in the Brazilian spiny rats (Genus Proechimys, Family Echimyidae). By Joao Moojen. Pp. 301-406, 140 figures in text. December 10, 1948.
20. Three new beavers from Utah. By Stephen D. Durant. Pp. 407- 417, 7 figures in text. December 24, 1948.
21. Two new meadow mice from Michoacan, Mexico. By E. Ray- mond Hall. Pp. 423-427, 2 figures in text. December 24, 1948.
22. An annotated check-hst of the mammals of Michoacan, Mexico. By E. Raymond Hall and Bernardo Villa-R. Pp. 431-472, 5 figures in text. December 27, 1949.
23. Subspeciation in the kangaroo rat, Dipodomys ordii. By Henry W. Setzer. Pp. 473-573, 27 figures in text, 7 tables. December 27, 1949.
24. Geographic range of the hooded skunk. Mephitis macroura, with description of a new subspecies from Mexico. By E. Raymond Hall and Walter W. Dalquest. Pp. 575-580, 1 figure in text. Janu- ary 20, 1950.
25. Pipistrellus cinnamomeus Miller 1902 referred to the Genus Myotis. By E. Raymond HaU and Walter W. Dalquest. Pp. 581-590, 5 figures in text. January 20, 1950.
26. A synopsis of the American bats of the Genus Pipistrellus. By E. Raymond Hall and Walter W. Dalquest, Pp. 591-602, 1 figure in text. January 20, 1950.
Index Pp. 605-638. Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest.
Pp. 1-444, 140 figures in text. April 9, 1948. Vol. 3. 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, 46 figures in text. June 29, 1951.
3. Phylogeny of the waxmngs and allied birds. By M. Dale Arvey. Pp. 473-530, 49 figures in text, 13 tables. October 10, 1951.
4. Birds from the State of Veracruz, Mexico. By George H. Lowery, Jr., and Walter W. Dalquest. Pp. 531-649, 7 figures in text, 2' tables. October 10, 1951.
^3
l:cr.-s? i
FEB 1 1 1952 HARVARD
LJME 3
New systematic names are in boldface type
abbas,
Tanagra, 636
Thraupis, 636 Abbot tanager, 636 Accipiter,
gularis, 104
soloensis, 104
virgatus, 104 Acrocephalus,
luscinia, 25 L 254, 256, 257
mariannae, 252
nijoi, 257
syrinx, 254
yamashinae, 256 Actitis,
hypoleucos, 143
macularia, 564 acuminata, Erolia, 152 acmninatus, Totanus, 152 acuta. Anas, 101 acutipennis, Chordeiles, 579 adolphi, Phaethornis, 582 aedon, Troglodytes, 620 aenea, Chloroceryle, 593 aeneus,
Aplonis, 297
Psarocolius, 632
Tangavius, 632 aeruginosus,
Chrysopicus, 599
Piculus, 599 aestiva,
Dcndroica, 626
Motacilla, 626 aethereus, Phaethon, 70 afBnis,
Euphonia, 635
Tanagra, 635 Aganus braccatus, 592 aglaiae, Platypsaris, 604 Agyrtrina versicolor genini, 588 Aimophila,
boucardi, 645
petenica, 645
rufescens, 644
ruficeps, 645
superciliosa, 644 Ajaia ajaja, 549 afaja,
Ajaia, 549 Platalea, 549
alba,
Crocetliia, 150
Guara, 549
Gygis, 174, 180
Scolopax, 549
Trynga, 150
Tyto, 574 albatross,
black-footed, 63
short-tailed, 64 albatnis, Diomedea, 64 albicilla,
Alcedo, 235
Halcyon, 235 albicollis,
Leucopternis, 553
Nyctidromus, 579 albifrons. Sterna, 161 iilbiejularis, Empidonax, 611 albilinea,
Iridoprocne, 616
Petrochelidon, 616 albilora, Dendroica, 627 albiventris, Aramides, 562 albonotatus, Buteo, 552 albus, Casmerodius, 547 Alcedo,
albicilla, 235
torquata, 592 alder flycatcher, 611 alexandrinus, Charadrius, 135 alpestris, Eremophila, 615 altitude, a factor in flight density,
387 amauronotus, Vireo, 625 Amazilia,
beryllina, 589
Candida, 588
cerviniventris, 589
cyanocephala, 588
genini, 588
tzacatl, 589
yucatanensis, 589 Amazona,
autumnalis, 571
farinosa, 572
guatemalae, 572
ochrocephala, 572
oratrix, 572
viridigenalis, 571 amazona, Chloroceryle, 593
— Univ. Kansas Publs., Mus. Nat. Hist., Vol. 3, 1951.
(651)
652
University of Kansas Publs., Mus. Nat. Hist.
Amazon kingfisher, 593 Amblycercus holosericeus, 632 American,
barn owl, 574
redstart, 630
robin, 621
wandering tattler, 145
whip-poor-will, 581 americana,
Chloroceryle, 593
Mycteria, 549 americanus,
Coccyzus, 573
Cuculus, 573
Daptrius, 556
Ibycter, 556 Ammodramus savannarum perpal-
lidus, 644 Ammodromus petenicus, 645 amnicola, Dendroica, 626 amurensis,
Ardea, 81
Butorides, 81 Amur green heron, 81 analis, Formicarius, 603 Anabates cervinigularis, 602 anaetheta, Sterna, 160 Anas,
acuta, 101
carolinensis, 100
clypeata, 102
crecca, 100
discors, 550
fuligula, 103
oustaleti, 94
pelewensis, 98
penelope, 102
poecilorhyncha, 98
querquedula, 100
rukensis, 98
superciliosa, 98
tzitzihoa, 101
valisineria, 103 anatum, Falco, 557 angus, Aplonis, 289 anliinga, Anhinga, 546 Anhinga,
anhinga, 546
leucogaster, 546
melanogaster, 79 ani, groove-billed, 573 annae, Psamathia, 249 Anoiis,
marcusi, 170
pileatus, 165
stolidus, 165
tenuirostris, 170 ant-eating woodpecker, 598 anthonyi,
Ardea, 546
Butorides, 546
Antliracothorax prevostii prevostii,
587 antpitta, Guatemalan, 603 antshrike, Mexican, 603 antthrush, Mexican, 603 antwren, Boucard, 603 antliracinus,
Buteogallus, 554
Falco, 554 Aphanolimnas monasa, 121 Aplomado falcon, 557 Aplonis,
aeneus, 297
angus, 289
corvinus, 301
guami, 293
harterti, 294
kvuodai, 291
orii, 292
opacus, 286, 288, 289, 291, 292, 293, 297
pelzelni, 299
ponapensis, 288 Apus pacificus, 226 aquilonis, Chrondrohierax, 552 Ara macao, 569 aracari, collared, 597 Aramides cajanea mexicana, 562 Aramus guarauna dolosus, 561 Aratinga,
astec, 570
liolochlora, 570 Archilochus colubris, 590 Ardea,
amurensis, 81
caerulea, 546
egretta, 547
flavicoUis, 94
intermedia, 82
melanolopha, 89
sacra, 84
virescens, 81, 546
violacea, 547 Ardetta eurhythma, 93 Arenaria,
interpres, 147
morinella, 149
oahuensis, 149 argentatus, Larus, 154 ariel,
Atagen, 80
Fregata, 80 Arkansas goldfinch, 641 Arremon opthalmicus, 638 Arremonops,
crassirostris, 642
rufivirgatus, 642 Artamus leucorhyncluis pelewensis,
284 Arvey, M. Dale, 473
Index to Volume 3
653
ashy,
ptilogonys, 480
starling, 302 Asiatic,
knot, 151
sparrow hawk, 104 asiatica,
Columba, 567
Zenaida, 567 Asio,
flamrneus, 217, 218
ponapensis, 218 assimilis, Turdus, 622 Astur gularis, 104 Atagen ariel, 80 Atlapetes,
brunnei-nucha, 641
pileatus, 641 atlapetes,
chestnut-capped, 641
rufous-capped, 641 atra, Fulica, 131 atratus, Coragyps, 550 atriceps,
Saltator, 638
Tanagra, 638 Atthis heloisa heloisa, 590 Audubon caracara, 556 audubonii, Polyborus, 556 Aulacorhynchus prasinus
prasinus, 596 aura,
Cathartes, 551
Vultur, 551 aurantiirostris, Catharus, 623 auricollis,
Icteria, 629
Tanagra, 629 aurifrons, Centurus, 598 aurocapilla, Motacilla, 627 aurocapillus, Seiurus, 627 austinsmithi, Oporomis, 627 Australian gray duck, 98 Automolus ochrolaemus
cervinigularis, 602 automolus, bu£E-throated, 602 autumnalis,
Amazona, 572
Psittacus, 572 axillaris, Empidonax, 611 Aythya,
fuligula, 103
valisineria, 103 Aztec parakeet, 570 azure-crown, red-billed, 588
bairdi,
Thryomanes, 619 Thryothorus, 619
Baker, Rollin H., The avifauna of Mi- cronesia, its origin, evolution and distribution, 1
Balanosphyra fonnicivora
formicivora, 598 Baltimore oriole, 633 banded crake,
Malay, 120
Philippine, 121 banded rail, 116 bannermani, Puffinus, 69 bam owl, American, 574 bam swallow, 616
eastern, 239 bartschi, CoUocalia, 222 Basileuterus,
culicivorus, 631
rufifrons, 631 baueri, Limosa, 140 beardless flycatcher, 615 becard, rose-throated, 604 becki, Pterodroma, 69 bergii, Thalasseus, 162 beryllina,
Amazilia, 589
Trochilus, 589 berylline hummingbird, 589 bewickii, Thryomanes, 619 birds,
flight densities of, 370
from the state of Veracruz, Mexico, 531
lunar observations of, 370
nocturnal migration of, 361 Bishop,
grosbeak, 639
tanager, 636 bistriatus,
Burhinus, 564
Charadrius, 564 bitorquata, Streptopelia, 198 bittern,
black, 94
Chinese least, 90
Japanese, 89
Malay, 89
Schrenk's least, 93
tiger, 548 black and white warbler, 626 black-bellied plover, 131 black-billed common tern, 155 blackbird, Sumichrast, 633 black-breasted weaver-finch, 339 black-chinned jacamar, 596 black-crowned night heron, 87 black drongo, 244 black-footed albatross, 63 black-hawk,
Mexican, 554
Ridgvi'ay, 554 black-naped tern, 156 black phoebe, 606 black-throated green warbler, 626 blatteus, Pyrocephalus, 607 bluebird, common, 623
654
University of Kansas Publs., Mus. Nat. Hist.
blue-black grassquit, 641 blue-crowned parrot, 572 blue-faced parrot-finch, 336, 337, 338 blue-gray,
gnatcatcher, 624
tern, 164 blue ground dove, 568 blue-headed vireo, 625 blue honey-creeper, 625 blue rock pigeon, 182 blue-winged teal, 550 boat-billed,
flycatcher, 608
heron, 548 bob-white, 560 Bomby cilia,
cedrorum, 482
centralasiae, 483
garrula, 482, 483
japonica, 483
pallidiceps, 483 Bombycillidae, 478 Bombycillinae, 482 Bonaparte euphonia, 635 booby,
brown, 76
masked, 75
red-footed, 75 borealis,
Nuttallomis, 610
Tyrannus, 610 Boucard,
antwren, 603
hermit, 582
tinamou, 544 boucardi,
Aimophila, 645
Crypturellus, 544
Formicivora, 603
Microrhopias, 603
Tinamus, 544
Zonotrichia, 645 braccatus,
Aganus, 592
Trogon, 592 brachyperus, Colymbus, 545 brasilianum, Glaucidium, 575 Brazilian kite, 551 brevipes,
Heteroscelus, 144
Poliolimnas, 124
Totanus, 144 brevirostris,
Cyclorhynchus, 614
Rhynchocyclus, 614 brewsteri, Empidonax, 611 bridled,
tern, 160
white-eve, 316, 318, 319, 320, 321, 322, 323 bristled-thighed curlew, 139
broadbill, Micronesian, 275, 277,
279, 280 broad-billed sandpiper, 154 broad-winged hawk, 552 brown booby, 76 brown-headed chlorospingus, 638 brown jay,
northern, 617
southern, 617 brown-throated wren, 620 brown towhee, 643 brunneicollis. Troglodytes, 620 brunnei-nucha,
Atlapetes, 641
Embemagra, 641 brunnitorques, Chaetura, 581 bryani, Ixobrychus, 90 Buarremon ocai, 642 Bubulcus ibis coromandus, 82 buff-beUied hummingbird, 589 b\ifF-f1-)roated automolus, 602 bimting,
indigo, 639
painted, 640
varied, 639 Burhinus bistriatus bistriatus, 564 burrowing owl, 575 bush-warbler, Palau, 249 Butastur indicus, 104 Buteo,
albonotatus, 552
gliiesbreghti, 553
griseocauda, 552
magnitrostris, 552
nitidus, 553
plagiatus, 553
platypterus, 552 Buteogallus anthracinus
anthracinus, 554 Butorides,
amurensis, 81
anthonyi, 546
striatus, 81
virescens, 546
cabanisi,
Lonchura, 340
Munia, 340 cachinnans, Herpetotheres, 555 Cacicus montezuma, 631 cacique, Prevost, 632 cactus wren,
banded, 619
rufous-naped, 619 caerulea,
Ardea, 546
Florida, 546
Motacilla, 624
Polioptila, 624 caerulescens,
Melanotis, 621
Orpheus, 621
Index to Volume 3
655
cajanea, Aramides, 562 caledonicus, Nycticorax, 87 Calidris tenuirostris, 151 calliope,
Luscinia, 248
Motacilla, 248 Caloenas nicobarica pelewensis, 209 Calomis kittlitzi, 286 Calothorax lucifer, 590 Campephaga,
monacha, 239
nesiotis, 241 Campephilus regius, 599 Camptostoma imberbe imberbe, 615 Campylopterus,
curvipennis, 583
excellens, 583
hemileucurus, 586 Campylorhynchus,
rufinucha, 619
zonatus, 619 Canada warbler, 630 canadensis,
Muscicapa, 630
Wilsonia, 630 Cancroma,
coromanda, 82
zeledoni, 548 cancrominus,
Platyrhynchus, 614
Platyrinchus, 614 candei,
Manacus, 605
Pipra, 605 Cande manakin, 605 Candida,
Amazilia, 588
Gygis alba, 174
Sterna, 174 canifrons,
Gallicolumba, 201
Phlegoenas, 201 canivet emerald, 588 canivetii,
Chlorostilbon, 588
Omismya, 588 canorus, Cuculus, 213 canvasback, 103 Capella gallinago delicata, 564 Caprimulgus,
indicus, 219
jotaka, 219
phalaena, 219
salvini, 580
serico-caudatus, 580
setosus, 581
vociferus, 581 caracara,
Audubon, 556
red-throated, 556 cardinal, 639
cardinal honey-eater, 304, 307, 308,
309, 312, 313 cardinalis,
Cardinalis, 639
Myzomela, 304, 307, 308, 309, 312, 313
Richmondena, 639 Cardinalis virginianus coccineus,
639 carinatum. Electron, 594 carinatus, Prionites, 594 cameipes, Cyanerpes, 625 Camifex naso, 556 Carolina, Porzana, 563 carohnensis.
Anas, 100
Columba, 567
Dumetella, 621
Muscicapa, 621
Zenaidura, 567 Carolines swiftlet, 224, 225, 226 carolinus, Rallus, 563 Carpodacus mexicanus mexicanus,
640 Caryothraustes pobogaster polio-
gaster, 639 Casmerodius albus egretta, 547 caspia,
Hydroprogne, 565
Sterna, 565 Caspian tern, 565 Cassidix mexicanus mexicanus, 633 castaneus,
Celeus, 599
Picus, 599 catbird, 621 Cathartes aura, 551 Catharus aurantiirostris melpo-
mene, 623 Catherpes mexicanus, 620 cattle egret, 82 caudacuta, Chaetura, 226 caudatus, Ptilogonys, 280 cayana, Piaya, 572 cedrorum, Bombycilla, 482 Celeus castaneus, 599 Central American squirrel cuckoo,
572 centralasiae, Bombycilla, 483 centralis,
Ciccaba, 575
Phalaenoptilus, 580 CentmTJS aurifrons veraecrucis,
598 cerulea, Procelstema, 164 cervinigularis,
Anabates, 602
Automolus, 602 cerviniventris, Amazilia, 589 chachalaca, gray-headed, 559
656
University of Kansas Publs., Mus. Nat. Hist.
Chaetura,
brunnitorques, 581
caudacuta, 226
pelagica, 581
richmondi, 581
rutila, 581
vauxi, 581 Chalcopsitta rubiginosus, 211 Chamaepelia,
inca, 567
interrupta, 568
minuta, 568
pallescens, 568
passerina, 568
rufipennis, 568 chapmani, Herpetotheres, 555 Charadrius,
alexandrinus, 135
bistriatus, 564
curonicus, 135
dubius, 135
hiaticula, 134
leschenaultii, 137
mongolus, 135
nihonensis, 135
saguineus, 135
semipalmatus, 134
stegmanni, 135
vociferus, 37, 564 chat, yellow-breasted, 629 check-list of birds of Micronesia,
16 chestnut-capped atlapetes, 641 chestnut-collared swift, 581 chestnut-colored woodpecker,
599 chestnut-sided warbler, 627 chimney swift, 581 chinensis, Coturnix, 113 Chinese blue rock thrush, 248 Chinese goshawk, 104 Chinese least bittern, 90 chipping sparrow, 645 Chlidonias leucopterus, 155 chloris, Halcyon, 233, 235, 237 Chloroceryle,
aenea, 593
amazona, 593
americana, 593
mexicana, 593 septentrionalis, 593 stictoptera, 593 chloronotus, Tyrannus, 608 chlorophonia, Mexican, 635 Chlorophonia occipitalis, 635 chloropus, Gallinula, 126, 127 chlororhynchus, PufRnus, 64 chlorospingus, brown-headed, 638 Chlorospingus ophthalmicus, 638 Chlorostilbon canivetii canivetii, 588
Chordeiles acutipennis texensis,
579 Christmas shearwater, 66 Chrondrohierax uncinatus aquilo-
nis, 552 chrysolaema, Eremophila, 615 Chrysopicus aeruginosas, 599 Chrysotis,
guatemalae, 570 viridigenaUs, 570 cicada bird, 239, 241, 242 Ciccaba virgata centralis, 575 Cinclus mexicanus mexicanus, 618 cinerea,
Drepanis, 326 Zosterops, 326, 327, 328 cineraceus, Stumus, 302 cinereus,
PoUolimnas, 123 Ptilogonys, 480, 624 cinnamomeus, Crypturellus, 545 cinnamomina. Halcyon, 227, 229, 230 cinnamon-rumped seed-eater, 640 Cinnyrorhyncha longirostra, 333 ciris,
Emberiza, 640 Passerina, 640 citreoline trogon, 591 citreolus, Trogon, 591 citrina,
Muscicapa, 629 Wilsonia, 629 clamator, Rhinoptynx, 576 clara, Erythrura, 337 Claravis pretiosa, 568 Cleptornis, marchei, 302 palauensis, 330 coccinea,
Cardinalis, 639 Richmondena, 639 Coccyzus americanus americanus, 573 cochlearius, Cochlearius, 548 Cochlearius cochlearius zeledoni, 548 coerulescens, Saltator, 638 Tanagra, 638 colchicus, Phasianus, 115 Colinus,
godmani, 560 pectoralis, 560 virginianus, 560 collared,
aracari, 597 micrastur, 556 towhee, 642 collaris, Trogon, 591 collingwoodi, Poliolimnas, 124 Collocalia,
bartschi, 222 fuciphaga, 222
Index to Volume 3
657
inexpectata, 221, 222
inquieta, 224, 225, 226
pelewensis, 221
ponapensis, 226
rukensis, 225
tachyptera, 222
ualensis, 224
vanikorensis, 226 CoUuricincla tenebrosa, 282 coloration, waxwings, 485 colubris,
Archilochus, 590
Trochilus, 590 Columba,
asiatica, 567
carolinensis, 567
flavirostris, 565
levia, 182
nigrirostris, 566
oceanica, 195
roseicapilla, 186
speciosa, 566 Columbigallina,
interrupta, 568
minuta, 568
pallescens, 568
passerina, 568
rufipennis, 568
talpacoti, 568 Colymbus,
brachypterus, 545
dominicus, 545
fulica, 563
podiceps, 545 common,
bluebird, 623
coot, 131
cuckoo, 213
mockingbird, 621
noddy, 165
sandpiper, 143
yellow-throat, 628 compressus, Picolaptes, 601 concept of flight density of birds, 370 confinus, Pooecetes, 644 connectens, Eurystomus, 238 Conopoderas luscinia yamashinae,
257 conservation of birds of
Micronesia, 58 conspicillata, Zosterops, 316, 318,
319, 320, 321, 323 conspicillatum, Dicaeum, 316 Contopus,
pertinax, 610
richardsonii, 610
sordidulus, 610 Conurus,
astec, 570
holochlorus, 570 cooperi, Myiarchus, 609 coot, common, 131
Coracias galbula, 633 Coragyps atratus, 550 cormorant, litde pied, 78 coromandus, Bubulcus, 82 coromanda, Cancroma, 82 corvina, Lamprothornis, 301 corvinus, Aplonis, 301 Corvus kubaryi, 244 Costa Rican ptilogonys, 480 Coturniculus passerinus perpal-
lidus, 644 Coturnix chinensis lineata, 113 Cotyle fulvipennis, 615 courtship, waxwings, 485 cowbird, red-eyed, 632
Malay banded, 120
Phihppine banded, 121 crassirostris,
Arremonops, 642
Embernagra, 642 Crax,
ruber, 558
rubra, 558 crecca, Anas, 100 crested,
flycatcher, Mexican, 609
tern, 162 crimson-collared tanager, 636 crimson-crowned fruit pigeon, 182,
184, 185 Crocethia alba, 150 crossbill, red, 641 Crotophaga sulcirostris sulciros-
tris, 573 Crvpturellus,
boucardi, 544
cinnamomeus, 545
salaei, 545 cuckoo,
Central American squirrel, 573
common, 213
long-tailed New Zealand, 214
oriental, 214
striped, 574
yellow-billed, 573 Cuculus,
americanus, 573
canorus, 213
horsfieldi, 214
saturatus, 214
taitensis, 214
telephonus, 213 culicivorus,
Basileuterus, 631
Sylvia, 631 cuneatus, Puflinus, 65 cunicularia, Speotyto, 575 curassow, Mexican, 558 curlew,
bristled-thighed, 139
long-billed, 140
658
University of Kansas Publs., Mus. Nat. Hist.
curlew sandpiper, 152 curonicus, Charadrius, 135 curvirostra, Loxia, 641 curvipennis,
Campylopterus, 583
Trochilus, 583 cyanea,
Passerina, 639
Tanagra, 639 Cyanerpes cyaneus carneipes, 625 cyaneus, Cyanerpes, 625 cyanocephala, Amazilia, 588 cyanocephalus, Ornismya, 588 Cyclorhynchus brevirostris, 614 Cynanthus lucifer, 590 Cypselus inquietiis, 224
dactylatra, Sula, 75
Dalquest, Walter W., George H.
Lowery, Jr., and, Birds from
the state of Veracruz, Mexico,
531 Daptrius americanus guatemalen-
sis, 556 Delattre saber-wing, 586 delawarensis, Larus, 565 delicata,
Capella, 564
Gallinago, 37
Scolopax, 564 Demigretta sacra sacra, 84 Dendrocopos,
jardinii, 600
ridgwayi, 600
scalaris, 600
villosus, 600 Dendroica,
aestiva, 626
albilora, 627
amnicola, 626
dominica, 627
magnolia, 626
occidentalis, 627
pensylvanica, 627
petechia, 626
virens, 626 deppei, Polioptila, 624 derby flycatcher, 609 diaconus,
Tanagra, 636
Thraupis, 636 Dicaeum conspicillatum, 316 dichrous, Puffinus, 66 Dicrurus macrocercus harterti,
244 Diomedea,
albatrus, 64
nigripes, 63 Diplopterus excellens, 574 dipper, Mexican, 618 discors. Anas, 550
dives.
Dives, 633
Icterus, 633 Dives dives dives, 633 doliatus, Thamnophilus, 603 dollar bird, 238 dolosus,
Aramus, 561 domesticus,
Fringilla, 631
Passer, 631 dominica,
Dendroica, 627
Pluviahs, 132 dominicus,
Colymbus, 545
Dulus, 484 dorotheae, Phaethon, 72 dotterel,
large sand, 137
Mongohan, 135 dove,
blue ground, 568
crimson-crowned fruit, 182, 184, 185
ground, 568
Inca, 567
Marianas fruit, 186
mourning, 567
Palau ground, 201
Phihppine turtle, 198
plain-breasted ground, 568
plumbeous-headed, 569
ruddy ground, 568
Verreaux, 569
white-throated ground, 203, 207
white-winged, 567 dowitcher, long-billed, 564 Drepanis cinerea, 326 drongo, black, 244 Dryobates scalaris ridgwayi, 600 Dryocopus,
ebumeirostris, 601
lineatus, 599
similis, 599 dubius,
Charadrius, 135
Melanerpes, 598 duck,
Australian gray, 98
tufted, 103 duck hawk, 557 Ducula,
monacha, 190
oceanica, 190, 193, 194, 195, 197
ratakensis, 197
teraokai, 193
townsendi, 194 Dulinae, 484 Dulus,
dominicus, 484
oviedo, 484
Index to Volume 3
659
Dumetella carolinensis, 621 Dupetor flavicollis, 94 dusky,
shearwater, 66
thrush, 248 dusky-capped flycatcher, 610 dusumieri,
Columba, 198
Streptopelia, 198
eagle, harpy, 554 eastern meadowlark, 634 ebumeirostris,
Dryocopus, 601
Xiphorhynchus, 601 edible nest swiftlet, 221, 222 Edolisoma,
insperatum, 242
monachum, 239
nesiotis, 241
tenuirostre, 239, 241, 242 egret,
cattle, 82
plumed, 82 Egretta intermedia intermedia, 82 egretta,
Ardea, 547
Casmerodius, 547 Elaenia,
flavogaster, 614
placens, 615
subpagana, 614
viridicata, 615 elaenia,
placid, 614
yellow-bellied, 614 Elanus leucurus majusculus, 551 Electron carinatum, 594 Emberiza ciris, 640 Embemagra,
brunnei-nucha, 641
crassirostris, 642
rufivirgata, 642 emerald,
canivet, 588
white-bellied, 588 emerald toucanet, 596 Empidonax,
albigularis, 611
axillaris, 611
brewsteri, 611
flaviventris, 610
fulvifrons, 611
minimus, 611
rubicundus, 611
traillii, 611 English sparrow, 631 episcopus,
Sialia, 623
Thraupis. 636 Erator inquisitor fraserii, 605
Eremopliila alpestris chrysolaema,
615 Ergaticus ruber, 630 Erolia,
acuminata, 152
ferruginea, 153
ferrugineus, 153
melanotos, 152
minuta, 151
minutilla, 564
ruficollis, 151
subminuta, 152 erytlirogaster, Hirundo, 616 erythrops, Myiagra, 275 erythrothorax, Synallaxis, 601 Erythrura,
clara, 337
pelewensis, 338
trichroa, 336, 337, 338 Eudynamis,
tahitiensis, 214
taitensis, 214 Eupherusa eximia nelsoni, 589 Euphonia,
affinis, 635
occipitalis, 635 euphonia,
Bonaparte, 635
Lesson, 635 eurhythma, Ardetta, 93 eurhythmus, Ixobrychus, 93 eurizonoides,
Gallinula, 121
Rallina, 121 Eurystomus,
connectens, 238
orientalis, 238
pacificus, 238 excellens,
Campylopterus, 583
Diplopterus, 574
Pampa, 583
Tapera, 574 eximia, Eupherusa, 589
falcinellus, Llinicola, 154 Falco,
anatum, 557
femoralis, 557
fruitii, 106
I'aponensis, 105
palliatus, 552
paulus, 557
peregrinus, 105, 557
septentrionalis, 557
soloensis, 104
sparverius, 557 falcon,
Aplomado, 557
laughing, 555
peregrine, 105
660
University of Kansas Publs., Mus. Nat. Hist.
fantail,
Palau, 266
rufous-fronted, 261, 262, 263, 264, 265 farinosa, Amazona, 572 fasciata, Rallina, 120 fasciatus, Rallus, 120 femoralis, Falco, 557 ferruginea,
Calidris, 153
Erolia, 153 ferrugineus, Tringa, 153 ferruginous pigmy owl, 575 field sparrow, 645 finch, house, 640 finfoot, 563 finschii,
Tephras, 328
Zosterops, 328 flamingo, 549 flammea, Strix, 217 flammeus, Asio, 217, 218 flat-bill,
Mexican, 614
short-billed, 614 flavicoUis,
Ardea, 94
Dupetor, 94 flavigaster, Xiphorhynchus, 600,
601 flavirostris, Columba, 565 flaviventris,
Empidonax, 610
Tyrannula, 610 flavogaster, Elaenia, 614 flavoviridis,
Vireo, 625
Vireosylvia, 625 flight density,
definition of, 385
of birds, 370 Florida caerulea, 546 Florisnga mellivora mellivora, 587 flycatcher,
alder, 611
beardless, 615
boat-billed, 608
Chinese gray-spotted, 282
derby, 609
dusky-capped, 610
forked-tailed, 608
fulvous, 611
least, 611
Mexican crested, 609
narcissus, 282
olive-sided, 610
scissortailed, 607
social, 609
striped, 608
sulphur-bellied, 608
sulphur-rumped, 614
tufted, 613
vermilion, 607
white-throated, 611
yellow-bellied, 610 food, waxwings, 493 forbesi, Rhinoptynx clamator, 576 forficata,
Muscicapa, 607
Muscivora, 607 forked-tailed flycatcher, 608 Formicarius analis moniliger, 603 Formicivora boucardi, 603 formicivora, Balanosphyra, 598 formicivorus, Picus, 598 formosa, Sylvia, 627 formosus, Oporomis, 627 fowl, red jungle, 114 fraserii,
Erator, 605
Psaris, 605 Fregata,
ariel, 80
minor, 79 fresh-water birds of Micronesia, 42 freycineti, Myiagra oceanica, 277 Fringilla,
domestica, 631
hncolnii, 646
mexicana, 640
psaltria, 641
rubra, 637
savanna, 644
splendens, 641 fruitii, Falco peregrinus, 106 fuciphaga, Collocalia, 222, 225 Fulica,
atra, 131
spinosa, 563 fulica,
Colymbus, 563
Heliomis, 563 fulicarius, Phalaropus, 37 fulva, Pluvialis, 132 fulvifrons, Empidonax, 611 fulvipennis,
Cotyle, 615
Stelgidopteryx, 615 fulviventris,
Leptoptila, 569
Leptotila, 569 fulvous flycatcher, 611 fumigatus, Veniliornis, 600 furtiva, Synallaxis, 601 fuscata. Sterna, 161 fuscus, Pipilo, 643
gadfly petrel, stout-billed, 70 gaimardi, Streptopelia, 199 Galbula melanogenia, 596 glabula,
Coracias, 633
Icterus, 633
Index to Volume 3
661
Gallicolumba,
canifrons, 201
kubaryi, 207
xanthonura, 203, 207
yapensis, 204 Gallinago,
delicata, 37
gallinago, 150
megala, 149 gallinago,
Gallinago, 150
Capella, 564 Gallinula,
chloropus, 126, 127
eurizonoides, 121
guami, 127
indica, 128
lozanoi, 128
orientalis, 126
sandvicensis, 128 gallinule, 126, 127 Gallus,
gallus, 114
micronesiae, 114 gallus, Phasianus, 114 gartered trogon, 592 garrula, Bombycilla, 482, 483 gazetteer of localities,
Micronesia, 11
Veracruz, 538 genini,
Agyrtrina, 588
Amazilia, 588 Geothlypis,
nelsoni, 628
occidentalis, 628
trichas, 628
typhicola, 628 Geranospiza nigra nigra, 555 ghiesbreghti,
Buteo, 553
Leucoptemis, 553 gilvus, Vireo, 625 glareola, Tringa, 142 Glaucidium brasilianum
ridgwayi, 575 Globicera,
momiyamai, 191
monacha, 190
oceanica, 190, 193, 194, 197
ratakensis, 197
teraokai, 193
townsendi, 194 Glyphorynchus spirurus pectoralis,
601 gnatcatcher, blue-gray, 624 godeffroyi, Monarcha, 272 godmani. Colinus, 560 godwit, Pacific, 140 goisagi,
Gorsachius, 89
Nycticorax, 89
golden-cheeked parrot, 572 goldfinch, Arkansas, 641 golden-fronted woodpecker, 598 golden honey-eater, 302 golden plover. Pacific, 132 Gorsachius,
goisagi, 89
kutteri, 90
melanolophys, 89 goshawk,
Chinese, 104
Mexican, 553 grackle,
great-tailed, 633
Mexican rice, 632 Grallaria guatimalensis guatimal-
ensis, 603 gramineus, Pooecetes, 644 grandis,
Saltator, 638
Tanagra, 638 grasshopper sparrow, 644 grassquit,
blue-backed, 641
yellow-faced, 640 gray-breasted wood wren, 620 gray-headed chachalaca, 559 grayi, Turdus, 622 gray robin, 621 gray-tailed,
hawk, 552
tattler, 145 greater white-eye,
Palau, 330
Ponape, 333
Truk, 332
Yap, 331 greater yellow-legs, 142 great-tailed grackle, 633 great tinamou, 543 grebe,
least, 545
pied-billed, 545
sun, 563 green,
heron, 546
jay, 618
kingfisher, 593
parakeet, 570 green heron, Amur, 81 greenshank, 141 griseicapillus, Sittasomus, 601 griseisticta,
Hemichelidon, 282
Muscicapa, 282 griseocauda,
Buteo, 552
Rupomis, 552 griseus,
Nyctibeus, 578
Vireo, 625 groove-billed ani, 573
662
University of Kansas Publs., Mus. Nat. Hist.
grosbeak,
Bishop, 839
rose-breasted, 639 ground dove,
blue, 568
Palau, 201
plain-breasted, 568
ruddy, 568
white-throated, 203, 207 guami,
Aplonis, 293
Gallinula, 127 guan, purplish, 558 Guara alba, 549 guarauna, Aramus, 561 guatemalae,
Amazona, 572
Clirysotis, 572
Siaha, 623 Guatemalan,
antpitta, 603
ivory-billed woodpecker, 599
sclerurus, 602 guatemalensis,
Daptrius, 556
Ibycter, 556
Phloeoceastes, 599
Sclerurus, 602 guatimalensis, Grallaria, 603 gularis,
Accipiter, 104
Astur, 104
Icterus, 634 Gulf of Mexico, migration of birds
over, 438 gull,
herring, 154
ring-billed, 565 guttata, Ortyx, 561 guttatus, Odontophorus, 561 gutturalis,
Habia, 637
Hirundo, 239 ■Gygis,
alba, 174, 180
Candida, 174
pacifica, 180 Gymnostinops montezuma, 631
Habia,
gutturalis, 637
rubica, 637
rubicoides, 637
salvini, 637 hairy woodpecker, 600 Halcyon,
albicilla, 235
chloris, 233, 235, 237
cinnamomina, 227, 229
mediocris, 230
orii, 235
owstoni, 237
pelewensis, 229
reichenbachii, 230
rufigularis, 227
teraokai, 233 haliaetus, Pandion, 105 Harpia harpyja, 554 harpy eagle, 554 harpyja,
Harpia, 554
Vultur, 554 harterti,
Aplonis, 294
Dicrurus, 244 hawk,
Asiatic sparrow, 104
blackish crane, 555
broad-winged, 552
duck, 557
gray-tailed, 552
Mexican black, 554
Ridgway black, 554
sparrow, 557
white snake, 553
zone-tailed, 552 Heliomaster longirostris pallidiceps,
589 Hehomis fulica, 563 heloisa,
Arismya, 590
Atthis, 590 Heloise hummingbird, 590 hemileucurus,
Campylopterus, 586
Trochilus, 586 Henicorhina,
leucophrys, 620
leucosticta, 620
mexicana, 620
prostheleuca, 620 hermit,
Boucard, 582
white-browed, 583 hermit warbler, 627 hemsheimi, Ptilinopus, 184 heron,
Amur green, 81
black-crowned night, 87
boat-billed, 548
green, 546
httle blue, 546
reef, 84
rufous night, 87
yellow-crowned night, 547 Herpetotheres cachinnans
chapmani, 555 herring gull, 154 Heterocnus mexicanus
mexicanus, 548 Heteroscelus,
brevipes, 144
incanus, 145 hiaticula, Charadrius, 134
Index to Volume 3
663
hidalgensis, Mitrephanes, 613 Himantopus himantopus
knudsoni, 33 Hirundo,
erythrogaster, 616
gutturalis, 239
pelagica, 581
rustica, 239, 616 hirundo, Sterna, 155 hoactii, Nycticorax, 49 holerythra, Rhytiptema, 604 holerythrus, Lipaugus, 604 holochlorus,
Aratinga, 570
Conurus, 570 holosericeus,
Amblycercus, 632
Sturnus, 632 honey-creeper, blue, 625 honey-eater,
cardinal, 304, 307, 308, 309, 312, 313
golden, 302 hooded warbler, 629 hooked-billed kite, 551 homed lark, 615 horsfieldi, Cuculus, 214 house,
finch, 640
wren, 620 hummingbird,
bervlline, 589
buff-bellied, 589
Heloise, 590
Jacobin, 587
Lucifer, 590
Rieffer, 589
ruby-throated, 550
striped-tail, 589 hunsteini, Munia, 339 Hydrocorax melanoleucus, 78 Hydroprogne caspia, 565 Hylocichla,
mustelina, 622
ustulata, 623 Hylomanes momotula momotvJa,
594 hj'polais, Zosterops, 323 hypoleuca,
Oestrelata, 70
Pterodroma, 70 hypoleucos,
Actitis, 143
Tringa, 143 Hypotaenidia philippensis pelewen-
sis, 116 hypugaea,
Speotyto, 575
Strix, 575
ibis, Bubulcus, 82
ibis,
white, 549
wood, 549 Ibycter americanus guatemalen-
sis, 556 Icteria virens auricoUis, 629 Icterus,
dives, 633
galbula, 633
gularis, 634
mesomelas, 634
spurius, 633
tamaulipensis, 634 imberbe, Camptostoma, 615 impacifus, Psomocolax, 632 Inca dove, 567 inca,
Cliamaepeha, 567
Scardafella, 567 indica, Gallinula, 128 indicus,
Butastur, 104
Caprimulgus, 219 indigo bunting, 639 inexpectata, Collocalia, 221, 222 inquieta, Collocalia, 224, 225, 226 inquietus, Cypselus, 224 inquisitive tityra, 605 inquisitor, Erator, 605 insignis,
Lepidocolaptes, 601
Picolaptes, 601 insperata, Volvocivora, 242 insperatum, Edolisoma, 242 intennedia,
Ardea, 82
Egretta, 82 intermedins, Thamnophilus, 603 interpres,
Arenaria, 147
Tringa, 147 interrupta,
Chaemepelia, 568
Columbigallina, 568 Iridoprocne albilinea, 616 Ischnosceles niger, 555 ivory-billed woodhewer, 600 Ixobrychus,
bryani, 90
eurhytlimys, 93
sinensis, 90
yapensis, 91
jacamar, black-chinned, 596 facana, Mexican, 563 Jacana spinosa spinosa, 563 jacarina, Volatinia, 641 Jacobin hummingbird, 587 Jalapa,
robin, 621
trogon, 591
664
University of Kansas Publs., Mus. Nat. Hist.
jamaicensis, Nyctibeus, 578 Japanese bittern, 89 japonensis, Falco, 105 japonica, Bombycilla, 483 jardinii,
Dendrocopos, 600
Picus, 600 iay,
green, 618
northern brown, 617
southern brown, 617 jotaka, Caprimulgus, 219 Junco phaeonotus, 645 junco, red-backed, 645 jungle nightjar, 219
keel-billed,
motmot, 594
toucan, 597 Kentish plover, 135 Kentucky warbler, 627 killdeer, 564
kingbird, melancholy, 608 kingfisher,
Amazon, 593
green, 593
least, 593
Micronesian, 227, 229, 230
ringed, 259
white-collared, 233 king vulture, 550 kite,
Brazilian, 552
hooked-billed, 552
white-tailed, 551 kittlitzi, Calomis, 286 knot, Asiatic, 151 knudsoni, Himantopus, 33 kobayashii, Myzomela, 313 kubaryi,
Corvus, 244
Gallicolumba, 207
Rhipidura, 265 kurodai,
Aplonis, 291
Myzomela, 312 Kusaie,
black rail, 121
mountain starUng, 301 kutteri, Gorsachius, 90
ladder-backed woodpecker, 600 Lamprothomis,
corvina, 301
opaca, 286 land birds of Micronesia, 42 Lanius ludovicianus mexicanus, 624 laperouse, Megapodius, 106, 109 lapponica. Limosa, 140 large sand dotterel, 137 lark, horned, 615
Larus,
argentatus, 154
delawarensis, 565
ridibundus, 154
vegae, 154 laughing falcon, 555 lauta, Tanagra, 635 lawrenceii, Muscicapa, GIO least,
bittern, 90, 93
flycatcher, 611
kingfisher, 593
man-o'-war, 80
sandpiper, 152, 564
tern, 80 least bittern,
Chinese, 90
Schrenck's, 93 Legatus leucophaius variegatus, 608 lepida, Phainopepla, 481 Lepidocolaptes souleyetii
insignis, 601 Leptoptila,
fulviventris, 569
plumbeiceps, 569 Leptotila,
fulviventris, 569
plumbeiceps, 569
verreauxi, 569 lepturus, Phaethon, 72 leschenaultii, Charadrius, 137 Lesson,
euphonia, 635
motmot, 595 lessonii, Momotus, 595 leucogaster,
Anhinga, 546
Plotus, 546
Sula, 76 leucophaius, Legatus, 608 leucophrys, Henicorhina, 620 leucoptera,
Piranga, 637
Pyranga, 637
Sterna, 155 Leucoptemis albicollis ghiesbreghti,
553 leucopterus,
Chlidonias, 155
Mimus, 621
Orpheus, 621 leucorhynchus, Artamus, 284 leucosticta, Henicorhina, 620 leucurus, Elanus, 551 Iherminieri, Puffinus, 66, 69 Lichtenstein,
oriole, 634
warbler, 631
woodpecker, 599 life-zones, Veracruz, 535 Limicola falcinellus sibirica, 154
Index to Volume 3
665
limicola, Rallus, 562 Limnodromus scolopaceus, 564 Limosa,
baueri, 140
lapponica, 140
scolopacea, 546 limpkin, 561 lincolnii,
Fringilla, 646
Melospiza, 646 Lincoln sparrow, 646 lineata, Coturnix, 113 lineated woodpecker, 599 lineatus,
Dryocopus, 599
Oriolus, 113 Lipaugus holerythrus, 604 lipida, Rhipidura, 266 little,
blue heron, 546
pied cormorant, 78
stint, 151 livia, Columba, 182 lobata, Tringa, 154 lobatus, Phalaropus, 154 loggerhead shrike, 624 Lonchura,
cabanisi, 340
minor, 339
nigerrima, 339
punctulata, 340 long-billed,
curlew, 140
dowitcher, 564 longipennis. Sterna, 155 longirostra, Cinnyrorhyncha, 333 longirostris, Heliomaster, 589 long-tailed.
New Zealand cuckoo, 214
saber-wing, 583 longuemareus, Phaethomis, 582 lory, Ponape, 211 Louisiana water-thrush, 627 Lowery, George H., Jr.,
A quantitative study of the noc- turnal migration of birds, 361 Lowery, George H., Jr., and Walter
W. Dalquest, Birds from the state
of Veracruz, Me.xico, 531 Loxia,
curvirostra, 641
ludoviciana, 639
stricklandi, 641 lozanoi, Gallinula chloropus, 128 Lucifer hummingbird, 590 lucifer,
Calothorax, 590
Cynanthus, 590 ludoviciana, Loxia, 639 ludovicianus,
Lanius, 624
Pheucticus, 639
lunar observations of birds, 370 lunata, Sterna, 160 Luscinia calliope, 248 luscinia, Acrocephalus, 251, 254 luscinius, Thryotliorus, 251 luteiventris, Myiodynastes, 608 luxuosa, Xanthoura, 618
macao,
Ara, 569
Psittacus, 569 macaw, red and blue, 569 Macgillivray warbler, 627 macrocercus, Dicrurus, 244 macroura, Zenaidura, 567 macularia,
Actitis, 564
Tringa, 564 maculatus, Pipilo, 643 madagascariensis,
Numenius, 140
Scolopax, 140 magna, Sturnella, 634 magnitrostris,
Buteo, 552
Rupornis, 552 magnolia,
Dendroica, 626
Sylvia, 626 magnolia warbler, 626 major,
Myzomela, 308
Tinamus, 543 majusculus, Elanus, 551 Malay banded crake, 120 mallard, Marianas, 94 Manacus candei, 605 manakin,
Cande, 605
yellow-thighed, 605 mango, Prevost, 587 mangrove swallow, 616 mannikin, Philippine nutmeg, 340 man-'o-war,
least. 80
Pacific, 79 marchei,
Cloptomis, 302
Ptilotis, 302 Marianas,
fruit dove, 186
mallard, 94 mariae, Rhipidura, 263 mariannae, Acrocephalus, 252 marsh snipe, 149 masked booby, 75 massena trogon, 590 massena, Trogon, 590 meadowlark, eastern, 634 mediocris, Halcyon, 230 Megaceryle torquata torquata, 592 megala, Gallinago, 149
666
University of Kansas Publs., Mus. Nat. Hist.
inegapode, Micronesian, 106, 109 Megapodius,
laperouse, 106, 109
senex, 106 Megarhynchus pitangua mexi-
canus, 608 Megazosterops palauensis, 330 melancholicus, Tyrannus, 608 melancholy kingbird, 608 Melanerpes dubius veraecrucis,
598 melanocephalus, Trogon, 591 melanogaster, Anhinga, 79 melanogenia, Galbula, 596 melanoieuca,
Scolopax, 142
Tringa, 142 melanoleucus,
Hydrocorax, 78
Phalacrocorax, 78 melanolopha, Ardea, 89 melanolophys, Gorsachius, 89 Melanotis caerulescens, 621 melanotus, Phorphyrio, 129 melanotos,
Erolia, 152
Tringa, 152 melanoxantha, Phainoptila, 479 mellivora,
Florisuga, 587
Trochilus, 587 Melospiza lincolnii, 646 melpomene,
Catliarus, 623
Turdus, 623 melvillensis, Pandion, 105 mentalis, Pipra, 605 merrilli, Nyctidromus, 579 mesomelas,
Icterus, 634
Psarocolius, 634 mesonauta, Phaethon, 70 Metabolus rugensis, 269 Mexican,
antshrike, 603
antthrush, 603
black hawk, 554
chlorophonia, 635
crested flycatcher, 609
curassow, 558
dipper, 618
flat-bill, 614
goshawk, 553
jacana, 563
ptilogonys, 624
rice grackle, 632
sclerurus, 602
sittasomiis, 601
tityra, 605 mexicana,
Aramides, 562
Chloroceryle, 593
Fringilla, 640
Henicorhina, 620
Spizella, 645
Sturnella, 634
Tigrisoma, 548 mexicanus,
Aramides, 562
Carpodacus, 640
Cassidix, 633
Catlierpes, 620
Cinclus, 618
Corvus, 633
Heterocnus, 548
Lanius, 624
Megarhynchus, 608
Nyctibeus, 578
Psilorhinus, 617
Scaphorhynchus, 608
Sclerurus, 602
Thryothorus, 620 Micrastur semitorquatus naso,
556 micrastur, collared, 556 Micronesia,
avifauna of, 1
check-list of birds of, 16
climate of, 5
conservation of avifauna of, 58
fresh-water birds of, 42
gazetteer of islands of, 11
land birds of, 42
migratory shore birds of, 32
oceanic birds of, 28
ornithological exploration of, 16
soils of, 8
speciation of birds of, 48
surface water of, 9
vegetation of, 9 micronesiae.
Callus, 114
Poliolimnas, 123 Micronesian,
broadbill, 275, 277, 279, 280
dusky white-eve, 326, 327, 328
kingfisher, 227, 229, 230
megapode, 106, 109
pigeon, 190, 193, 194, 195, 197
starling, 286, 288, 289, 291, 292, 293, 297 Microrhopias quixensis boucardi,
603 migration,
in relation to topography, 424
of birds, nocturnal, 361
of birds over Gulf of Mexico, 438 migratorius, Turdus, 621 migratory birds of Micronesia, 32 Mimus polyglottos leucopterus, 621 mindanensis, Rattus, 74 miniata, Setophaga, 630 miniatus, Myioborus, 630
Index to Volume 3
667
minima, Tyrannula, 611 minimus, Empidonax, 611 minor,
Fregata, 79
Lonchmra, 339
Munia, 339
Pelecanus, 79
Phainoptila, 479
Puffinus, 66, 68 minuta,
Chaemaepelia, 568
Colmnbigallina, 568
Erolia, 151 minutilla,
Erolia, 564
Tringa, 564 Mitrephanes phaeocercus hidal-
gensis, 613 Mniotilta varia, 626 mockingbird, common, 621 molochinus, Myioborus, 630 molybdophanes, Ptilogonys, 480 momiyamai, Globicera, 191 momota, Momotus, 595 momotula, Hylomanes, 594 Momotus momota lessonii, 595 monacha,
Campephaga, 239
Ducula, 190
Globicera, 190 monachum, Edolisoma, 239 monachus,
Muscivora, 608
Tyrannus, 608 monarch,
Tinian, 274
Truk, 269
Yap, 272 Monarcha,
godeffroyi, 272
takatsukasae, 274 Monarcharses takatsukasae, 274 monasa,
Aphanolimnas, 121
Rallus, 121 Mongolian dotterel, 135 mongolus, Charadrius, 135 moniliger, Formicarius, 603 Montezuma oropendola, 631 montezuma,
Cacicus, 631
Gymnostinops, 631 Monticola solitaria philippensis,
248 morelleti,
Spermophila, 640
Sporophila, 640 morinella, Arenaria, 149 morio,
Pica, 617
Psilorhinus, 617 morning bird, Palau, 282
Motacilla,
aestiva, 626
aurocapilla, 627
caerulea, 624
calliope, 248
pensylvanica, 627
philippensis, 302
pileolata, 629
tricolor, 630
varia, 626
virens, 626 motacilla,
Seiujus, 627
Turdus, 627 motmot,
keel-billed, 594
Lesson, 595
tody, 594 mourner, rufous, 604 mourning dove, 567 Munia,
cabanisi, 340
hunsteini, 339
minor, 339 Muscadivora oceanica winkleri,
191 Muscicapa,
canadensis, 630
carolinensis, 621
citrina, 629
forficata, 607
griseisticta, 282
lawrenceii, 610
narcissina, 282
noveboracensis, 625
pusilla, 629
rugensis, 269
saya, 606
solitaria, 625
texensis, 609 Muscivora,
forficata, 607
monachus, 608
tyrannus, 608 muscovy, 550 mustelina, Hylocichia, 622 mustelinus, Turdus, 622 Mycteria americana, 549 Myiagra,
erythrops, 275
freycineti 277
oceanica, '275, 277, 279
pluto, 280 Myiarchus,
cooperi, 609
lawrenceii, 610
tuberculifer, 610
tyrannulus, 609 Myiobius sulphureipygius, 614 Myioborus,
miniatus, 630
molochinus, 630
668
University of Kansas Publs., Mus. Nat, Hist.
Myiodynastes luteiventris lutei-
ventris, 608 Myiozetetes similis texensis, 609 Myzomela,
cardinalis, 304, 307, 308, 309, 312, 313
dichromata, 307
kobayashii, 313
kurodai, 312
major, 308
rubratra, 304
safiFordi, 309
naevia, Tapera, 574 narcissina, Muscicapa, 282 narcissus flycatcher, 282 naso,
Carnifex, 556
Micrastur, 556 nativitatis, PufEnus, 66 nebularia, Tringa, 141 nebularis, Scolopax, 141 nelsoni,
Eupherusa, 589
Scolopax, 141 Nelson yellow-throat, 628 nesiotis,
Campephaga, 241
Edolisoma, 241 nest building, waxwings, 491 net trend density, 386 nicobarica, Caloenas, 209 Nicobar pigeon, 209 nigerrima, Lonchura, 339 night heron,
black-crowned, 87
rufous, 87
yellow-crowned, 547 nightingale reed-warbler, 251,
254, 256, 257 nightingale tlirush, orange-billed,
623 nightjar, jungle, 219 nigra,
Geranospiza, 555
Ischnosceles, 555 nigricans,
Sayomis, 606
Tyrannula, 606 nigripes, Diomedea, 63 nigrirostris, Columba, 566 nihonensis, Charadrius, 135 nitens, Phainopepla, 481 nitidus, Buteo, 553 nocturnal migration of birds, 361,
408 noddy,
common, 165
white-capped, 170 northern brown jay, 617 Nothocernus salaei, 545
noveboracensis,
Muscicapa, 625
Vireo, 625 Numenius,
madagascariensis, 140
phaeopus, 137
tahitiensis, 139
variegatus, 137 nuttallii, Phalaenoptilus, 580 Nuttallornis borealis, 610 Nyctanassa violacea violacea, 547 Nyctibeus,
griseus, 578
jamaicensis, 578
mexicanus, 578 Nycticorax,
caledonicus, 87
goisagi, 89
hoactH, 49
nycticorax, 49, 87
pelewensis, 87 nycticorax,
Ardea, 87
Nycticorax, 49, 87 Nyctidromus,
albicoUis, 579
merrilli, 579
yucatanensis, 579
oahuensis,
Arenaria, 149
Sterna, 161 obscurus, Turdus, 248 obsoletus, Salpinctes, 620 ocai,
Buarremon, 642
Pipilo, 642 occidentalis,
Dendroica, 627
Geothlypis, 628
Sylvia, 627 occipitalis,
Chlorophonia, 635
Euphonia, 635 oceanica,
Columba, 195
Ducula, 190, 193, 194, 195, 197
Globicera, 190, 193, 194
Muscadivora, 191
Myiagra, 275, 277, 279, 280 oceanic birds of Micronesia, 28 ochrocephala, Amazona, 572 ochrolaemus, Automolus, 602 Odontophorus guttatus, 561 Odontriorchis palliatus, 552 Oestrelata hypoleuca, 70 oleaginea,
Rukia, 331
Zosterops, 331 oleaginous woodpecker, 600
Index to Volume 3
669
oleaginus.
Picus, 600
Veniliornis, 600 olivacea,
Sylvia, 626
Tiaris, 640 olivaceus,
Peucedramus, 626
Vireo, 625 olive-sided flycatcher, 610 olive warbler, 626 opaca, Lamprothornis, 286 opacus, Aplonis, 286, 288, 289,
291, 292, 293, 297 opthalmicus,
Arremon, 638
Cholorospingus, 638 opisthomelas, PufRnus, 66, 68 Oporornis,
austinsmithi, 627
formosus, 627
tolmiei, 627 orange-billed nightingale thrush,
623 oratrix, Amazona, 572 orchard oriole, 633 orientalis,
Eurystomus, 238
Gallinula, 126 oriental cuckoo, 214 origin of species, waxwings, 519 orii,
Aplonis, 292
Halcyon, 235 oriole,
Baltimore, 633
Lichtenstein, 634
orchard, 633
yellow-tailed, 633 Oriolus spurius, 633 Omismya,
canivetii, 588
cyanocephalus, 588
heloisa, 590 oropendola, Montezuma, 631 Orpheus,
caerulescens, 621
leucopterus, 621 Ortalis vetula vetula, 599 Ortyx,
guttata, 561
pectoralis, 561 oryzivorus, Psomocolax, 632 osprey, 105 Otus podarginus, 215 oustaleti. Anas, 94 oven-bird, 627 oviedo, Dulus, 484 owl,
American barn, 574
burrowing, 575
ferruginous pigmy, 575
Palau scops, 215 short-eared, 217, 218 spectacled, 574 striped, 575 wood, 575 owstoni,
Halcyon, 237 Rallus, 118 Zosterops, 321
Pacific,
godwit, 140
golden plover, 132
man-o'-war, 79 pacifica,
Gygis, 180
Sterna, 180 pacificus,
Apus, 226
Eurystomus. 238
PuflBnus, 64 painted,
bunting, 640
quail, 113 Palau,
bush-warbler, 249
fantail, 266
greater white-eye, 330
ground dove, 201
morning bird, 282
scops owl, 215 palauensis,
Cleptornis, 330
Megazosterops, 330
Rukia, 330 pale-crowned star-throat, 589 pallescens,
Chamaepelia, 568
Columbigallina, 568 palliatus,
Falco, 552
Odontriorchis, 552 pallidiceps,
Bombycilla, 483
Heliomaster, 589 palm-chat, 484 Pampa pampa excellens, 586 Pandion haliaetus melvillensis, 105 papa,
Sarcoramphus, 550
Vultur, 550 parakeet, Aztec, 570 green, 570 parkmanii. Troglodytes, 620 parrot,
blue-crowned, 572
golden-cheeked, 572
red-crowned, 571
white-crowned, 571
yellow-headed, 572
670
University of Kansas Publs., Mus. Nat. Hist.
parrot-finch, blue-faced, 336, 337,
338 Passer domesticus, 631 Passerculus sandwichensis savanna,
644 Passerina,
ciris, 640
cyanea, 639
versicolor, 639 passerina,
Chamaepelia, 568
Columbigallina, 568
Spizella, 645 passerinus, Cotumiculus, 644 paulus,
Cerchneis, 557
Falco, 557 pauraque, 579 pectoralis,
Colinus, 560
Glyphorhynchus, 601
Ortyx, 560
Rhipidura, 261 pectoral sandpiper, 152 pelagica,
Chaetura, 581
Hirundo, 581 Pelecanus,
minor, 79
plotus, 76 pelecanoides.
Sterna, 162
Thalasseus, 162 pelewensis,
Anas, 98
Artamus, 284
Caloenas, 209
CoUocalia, 221
Erythrura, 338
Halcyon, 229
Hypotaenidia, 116
Nycticorax, 87
Phorphyrio, 129
Ptilinopus, 185
Rallus, 116 pelzelni, Aplonis, 299 Penelope,
purpurascens, 558
vetula, 559 pensylvanica,
Dendroica, 627
Motacilla, 627 peregrine falcon, 105 peregrinus, Falco, 105, 557 Peristera pretiosa, 568 perpallidus,
Ammodramus, 644
Coturniculus, 644 personata,
Sula, 75
Tityra, 605
perspicillata, Pulsatrix, 574 pertinax, Contopus, 610 petechia, Dendroica, 626 Peten sparrow, 645 petenica, Aimophila, 645 petenicus, Ammodromus, 645 petrel,
stout-billed gadfly, 70 Tahiti, 69 Petrochelidon albilinea, 616 Peucedramus olivaceus olivaceus,
626 pewee,
Swainson, 610
western wood, 610 phaeocercus, Mitrephanes, 613 phaenotus, Junco, 645 Phaethon,
aethereus, 70
dorotlieae, 72
lepturus, 72
mesonauta, 70
rothschildi, 71
rubicauda, 71 Phaethornis,
adolphi, 582
longuemareus, 582
superciliosus, 582
veraecrucis, 582 phaincipepla, 481 Phainopepla,
lepida, 481
nitens. 481 phainoptila, 479 Phainoptila,
melanoxantha, 479
minor, 479 Phalacrocorax melanoleucus, 78 phalaena, Caprimulgus, 219 Phalaenoptilus nuttallii centralis,
580 Phalaropus,
fulicarius, 37
lobatus, 154 Phasianus,
c'olchicus, 115
gallus, 114 Pheucticus ludovicianus, 639 philippensis,
Hypotaenidia, 116
Monticola, 248
Motacilla, 302
Rallus, 116
Sturnus, 302
Turdus, 248 Philippine,
banded crake, 121
nutmeg mannikin, 340
turtle dove, 198 phillipsi, Turdus, 621
Index to Volume 3
671
Phlegoenas,
canifrons, 201
kubaryi, 207
yapensis, 203 Phloeoceastes guatemalensis
regius, 599 Phlogothraupis sanguinolenta
sanguinolenta, 636 phoebe,
black, 606
Say, 606 Phoenicopterus ruber, 549 Phoenicothraupis salvini, 637 Phorphyrio,
melanotus, 129
pelewensis, 129 Phylogeny of the waxwings and
allied birds, 473 Piaya cayana thermophila, 572 Pica morio, 617 Picolaptes compressus insignis,
601 pictus, Aramus, 561 Piculus rubiginosus aeruginosus,
599 Picus,
castaneus, 599
formicivorus, 598
jardinii, 600
oleaginus, 600
similis, 599 pigeon,
blue rock, 182
Micronesian, 190, 193, 194,
195, 197
Nicobar, 209
scaled, 566
short-billed, 566 pigmy owl, ferruginous, 575 pileata. Sterna, 165 pileatus,
Anoiis, 165
Atlapetes, 641 pileolata,
Motacilla, 629
Wilsonia, 629 pintail, 101
Pionus senilis senihs, 570 Pipilo,
fuscus, 643
maculatus, 643
ocai, 642
rufescens, 644 Pipra,
candei, 605
mentalis, 605 Piranga,
leucoptera, 637
rubra, 637 pitangua, Megarhynchus, 608 Pitangus sulfuratus texanus, 609
Pitylus poliogaster, 639 placens, Elaenia, 615 placid elaenia, 614 plagiatus, Buteo, 553 Plagiospiza superciliosa, 644 plain-breasted ground dove, 568 Platalea ajaja, 549 Platypsaris aglaiae sumichrasti,
604 platypterus,
Buteo, 552
Sparvius, 552 Platyrhynchus
cancrominus, 614 Platyrinchus cancrominus, 614 Plotus leucogaster, 546 plotus,
Pelecanus, 76
Sula, 76 plover,
black-bellied, 131
Kentish, 135
Pacific golden, 132
ring-necked, 135
semipalmated, 134 plumbeiceps,
Leptoptila, 569
Leptotila, 569 plumed egret, 82 pluto, Myiagra, 280 Pluvialis dominica fulva, 132 podargina,
Noctua, 214
Pyrroglaux, 216 podarginus, Otus, 215 podiceps,
Colymbus, 545
Podilymbus, 545 Podilvinbus podiceps podiceps,
545 poecilorhyncha. Anas, 98 poliogaster,
Caryothraustes, 639
Pitylus, 639 Poliolimnas,
brevipes, 124
cinereus, 123
collingwoodi, 124
micronesiae, 123 Polio tila,
caerulea, 624
deppei, 624 Polvborus cheriway audubonii,
556 polyglottos, Mimus, 621 polynesiae, PuflBnus, 69 Ponap6,
greater white-eye, 333
lory, 211
mountain starling, 299
672
University of Kansas Publs., Mus. Nat. Hist.
ponapensis,
Aplonis, 288
Asio, 218
Collocalia, 226
Ptilinopus, 182
Zosterops, 327 Pooecetes gramineus confiniis, 644 poor-will, 580 porphyraceus, Ptilinopus, 182, 184,
185 Porzana Carolina, 563 potoo, 578 prasinus,
Aulacorhynchus, 596
Pteroglossus, 596 pratincola,
Strix, 574
Tyto, 574 pretiosa,
Claravis, 568
Peristera, 568 Prevost,
cacique, 632
mango, 587 prevostii,
Anthracotliorax, 587
Trochilus, 587 Prionites carinatus, 594 Procellaria,
rostrata, 69
tenuirostris, 66 Procelsterna cerulea saxatilis, 164 prostheleuca, Henicorhina, 620 prostlieleucus, Scytalopus, 620 psaltria,
Fringilla, 641
Spinus, 641 Psamathia annae, 249 Psarocolius,
aeneus, 632
mesomelas, 634 Psilorhinus,
mexicanus, 617
morio, 617 Psittacus,
autumnalis, 570
macao, 569
senilis, 570 Psomocolax oryzivorus impacifuj;,
632 Pterodroma,
becki, 69
hypoleuca, 70
rostrata, 69 Pteroglossus,
prasinus, 596
torquatus, 597 Ptilinopus,
hernsheimi, 184
pelewensis, 185
ponapensis, 182
porphyraceus, 182, 184, 185
roseicapillus, 186 Ptilogonatinae, 478 Ptilogonys,
caudatus, 480
cinereus, 480, 624
molybdophanes, 480 ptilogonys,
ashy, 480
Costa Rican, 480
Mexican, 624 Ptilotis marchei, 302 puella, Trogon, 591 PuflBnus,
bannermani, 69
chlororhynchus, 64
cuneatiis, 65
dichrous, 66
Iherminieri, 66
nativitatis, 66
opisthomelas, 66, 68
pacificus, 64, 65
polynesiae, 69
tenuirostris, 66 Pulsatrix perspicillata saturata, 574 punctulata, Lonchura, 340 purple swamphen, 129 purplish guan, 558 purpurascens, Penelope, 559 pusilla,
Muscicapa, 629
Spizella, 645
Tiaris, 640
Wilsonia, 629 Pyranga leucoptera, 637 Pyrocephalus rubinus blatteus, 607 Pyrroglaux podargina, 216
quail,
painted, 113
spotted wood, 561 Quantitative study of the nocturnal
migration of birds, 361 querquedula. Anas, 100 quixensis, Microrhopias, 603
rail,
banded, 116
Guam, 118
Kusaie black, 121
Virginia, 562
white-browed, 123
wood, 562 Rallina,
eurizonoides, 121
fasciata, 120 Rallus,
carolinus, 563
fasciatus, 120
limicola, 562
monasa, 12]
Index to Volume 3
673
owstoni, 118
pelewensis, 116
philippensis, 116 Ramphastos,
sulfuratus, 597
torquatus, 597 ratakensis,
Ducula, 197
Globicera, 197 Rattus,
mindanensis, 74
rattus, 4 rattus, Rattus, 74 Rectes tenebrosus, 282 red,
crossbill, 641
jungle fowl, 114
warbler, 630 red and blue macaw, 569 red-backed junco, 645 red-bellied redstart, 630 red-billed,
azure-crown, 588
pigeon, 565
tropic-bird, 70 red-crowned parrot, 570 red-eyed,
cowbird, 632
vireo, 625 red-footed booby, 75 red-tailed tropic bird, 71 red-throated caracara, 556 redstart,
American, 630
red-bellied, 630 reed-warbler, nightingale, 251
254, 256, 257 reef heron, 84 regius,
Campephilus, 599
Phloeoceastes, 599 reichenbachii.
Halcyon, 230
Todirhamphus, 230 Rhamphozosterops sanfordi, 333 Rhinoptynx clamator forbesi, 576 Rhipidura,
kubaryi, 265
lepida, 266
mariae, 263
rufifrons, 261, 262, 263, 264, 265
saipanensis, 262
uraniae, 261
versicolor, 264 Rhynchocyclus brevirostris, 614 Rhvtiptema holerythra holerythra,
'604 richardsonii, Contopus, 610 Richmondena cardinalis,
coccinea, 639
richmondi, Chaetura, 581 Ridgway black hawk, 554 ridgwayi,
Dendrocopos, 600
Dryobates, 600
Glaucidium, 575
Hypomorphnus, 554
Urubitinga, 554 ridibundus, Larus, 154 Rieffer hummingbird, 589 ring-billed gull, 565 ring-necked,
pheasant, 115
plover, 135 ringed kingfisher, 592 robin,
American, 621
gray, 621
Jalapa, 621 robustus, Tinamus, 543 rock sparrow, 645 rock thrush, Chinese blue, 248 roseate spoonbill, 549 roseicapilla, Columba, 186 roseicapillus, Ptilinopus, 186 rose-breasted grosbeak, 639 rose-throated becard, 604 rostrata,
Procellaria, 69
Pterodroma, 69 rotensis, Zosterops, 319 rothschildi,
Phaethon, 71
Scaeophaethon, 71 roui^li.winged swallow, 615 ruber,
Crax, 558
Ergaticus, 630
Phoenicopterus, 549
Setophaga, 630 rubica, Habia, 637 rubicoides,
Habia. 637
Saltator, 637 rubicundus, Empidonax, 611 rubiginosus,
Chalcopsitta, 211
Piculus, 599
Trichoglossus, 211 rubinus, Pyrocephalus, 607 rubra,
Crax, 558
Fringilla, 637
Piranga, 637 rubratra,
Cinnyris, 304
Myzomela, 304 rubricauda,
Phaethon, 71
Scaeophaetlion, 71 rubripes, Sula, 75
674
University of Kansas Publs., Mus. Nat. Hist.
rubythroat, Siberian, 248 ruby-tliroated hummingbird, 590 niddy ground dove, 568 rufescens,
Aimophila, 644
Pipilo, 644 ruficeps, Aimophila, 645 nificoUis,
EroHa,151
Stelgidopteryx, 615
Trynga, 151 rufifrons,
Basileuterus, 631
Setophaga, 631 rufigularis, Halcyon, 227 rufinucha,
Campylorhynchus, 619
Picolaptes, 619 rufipennis,
Chamaepelia, 568
Columbigallina, 568 rufivirgata, Embernagra, 642 rufivirgatus, Arremonops, 642 rufous-breasted synallaxis, 601 rufous-capped,
atlapetes, 641
warbler, 631 rufous-fronted fantail, 261, 262,
263, 264 rufous mourner, 604 rugensis,
Metabolus, 269
Muscicapa, 269 rukensis.
Anas, 98
Collocalia, 225 ruki,
Rukia, 332
Tephras, 332 Rukia,
oleaginea, 331
palauensis, 330
ruki, 332
sanfordi, 333 russet-backed thrush, 623 rustica, Hirundo, 239, 616 rusty sparrow, 644 ruticilla, Setophaga, 630 rutila, Chaetura, 581 Rupornis magnitrostris griseocauda,
552
saber-wing,
curved-winged, 583
Delattre, 586
long-tailed, 583 sacra,
Ardea, 84
Demigretta, 84 saffordi, Myzomela, 309 saipanensis, Rhipidura, 262
sallaei,
Crypturellus, 545
Notliocemus, 545 Salpinctes obsoletus, 620 Saltator,
atriceps, 638
coerulescens, 638
grandis, 638 saltator,
black-headed, 638
grayish, 638 saltuarius, Xiphorhynchus, 600 Salvin whip-poor-will, 580 salvini,
Caprimulgus, 580
Habia, 637
Phoenicotliraupis, 637 sanderling, 150 sandpiper,
broad-billed, 154
common, 143
curlew, 153
least, 152, 564
pectoral, 152
sharp-tailed, 152
spotted, 564
wood, 142 sandvicensis, Gallinula, 128 sandwichensis, Passerculus, 644 sanfordi,
Rhamphozosterops, 333
Rukia, 333 sanguineus, Charadrius, 135 sanguinolenta, Phlogothraupis, 636 sanguinolentus,
Tachyphonus, 636
Tanagra, 636 Sarcoramphus papa, 550 sflturata, Pulsatrix, 574 saturatus, Cuculus, 214 savanna,
Fringilla, 644
Passerculus, 644 savannah sparrow, 644 savannarum, Ammodramus, 644 saxatilis, Procelsterna, 164 Say phoebe, 606 saya,
Muscicapa, 606
Sayornis, 606 Sayomis,
nigricans, 606
saya, 606 saypani, Zosterops, 318 Scaeophaethon rubricauda roths-
childi, 71 scalaris,
Dendrocopos, 600
Dryobates, 600 Scaphorliynchus mexicanus, 608 Scardafella squammata inca, 567
Index to Volume 3
675
Schrenck's least bittern, 93 scissor-tailed flycatcher, 607 Sclerurus,
guatemalensis, 602
mexicanus, 602 scerurus,
Guatemalan, 602
Mexican, 602 scolopacea, Limosa, 564 scolopaceus, Limnodromus, 564 Scolopax,
alba, 549
delicata, 564
gallinago, 150
incana, 145
melanoleuca, 142
nebularis, 141
taliitiensis, 139 Scytalopus prostheleucus, 620 sector density, 385 seed-eater, cinnamon-rumped, 640 Seiurus,
aurocapillus, 627
motacilla, 627 semifasciata, Tityra, 605 semipalmated plover, 134 semipalmatus, Charadrius, 134 semitorquatus, Micrastur, 556 semperi, Zosterops, 320 senex, Megapodius, 106 senilis,
Pionus, 570
Psittacus, 570 septentrionalis,
Chloroceryle, 593
Falco, 557 serico-caudatus, Caprimulgus, 580 Setophaga,
miniata, 630
rubra, 630
rufifrons, 631
ruticilla, 630
tricolora, 630 setosus, Caprimulgus, 581 sharp-tailed sandpiper, 152 shearwater,
Christmas, 66
dusky, 66
short-tailed, 66
wed^e-tailed, 64, 65 short-billed flat bill, 614 short-eared owl, 217, 218 short-tailed,
albatross, 64
shearwater, 66 shoveller, 102 shrike, loggerhead, 624 Sialia,
episcopus, 623
guatemalae, 623
sialis, 623
sialis, Sialia, 623 Siberian rubythroat, 248 sibirica, Limicola, 154 similis,
Dryocopus, 599
Myiozetetes, 609
Picus, 599 sinensis,
Ardea, 90
Ixobrychus, 90
Sterna, 161 Sittasomus griseicapillus sylviodes,
601 sittasomus, Mexican, 601 skeleton, waxwings, 494 snipe,
common, 150
marsh, 149
Wilson, 564 social flycatcher, 609 socialis, Spizella, 645 solitaria,
Monticola, 248
Vireo, 625 solitarius, Vireo, 625 soloensis,
Accipiter, 104
Falco, 104 sooty tern, 161 sora, 563
sordidulus, Contopus, 610 souleyetii, Lepidocolaptes, 601 southern browTi jay, 617 sparrow,
chipping, 645
English, 631
field, 645
grasshopper, 644
Lincoln, 646
Peten, 644
rock, 644
rusty, 644
savannah, 644
striped, 644
Texas, 642
vesper, 644 sparrow hawk, 557 sparrow hawk, Asiatic, 104 sparverius, Falco, 557 speciation of birds of Micronesia,
48 speciosa, Columba, 566 spectacled tern, 160 Spermophila morelleti, 640 Speotvto cunicularia hypugaea,
575 spinosa,
Fulica, 563
Jacana, 563 Spinus psaltria psaltria, 641 spirurus, Glyphorhynchus, 601
676
University of Kansas Publs., Mus. Nat. Hist.
Spiza versicolor, 639 Spizella,
mexicana, 645
passerina, 645
pusilla, 6^45
socialis, 645
wortheni, 645 splendens,
Fringilla, 641
Volatinia, 641 spoonbill, roseate, 549 Sporophila torqueola morelleti, 640 spotted,
sandpiper, 564
towhee, 643
wood quail, 561 spurius.
Icterus, 633
Oriolus, 633 squammata, Scardafella, 567 Squatarola squatarola, 131 squatarola,
Squatarola, 131
Tringa, 131 squirrel cuckoo. Central American,
572 starling,
ashy, 302
Kusaie mountain, 301
Micronesian, 286, 288, 289, 291, 292, 293, 297
Ponape mountain, 299
violet-backed, 302 station density, 386 star-throat, pale-crowned, 589 stegmanni, Charadrius, 135 Stelgidopteryx ruficoUis
fulvipennis, 615 Sterna,
albifrons, 161
anaetheta, 160
Candida, 174
fuscata, 161
hirundo, 155
leucoptera, 155
longipennis, 155
lunata, 160
oahuensis, 161
pelecanoides, 162
pileata, 165
sinensis, 161 sumatrana, 155 stictoptera, Chloroceryle, 593 stint, httle, 151 stolidus, Anoiis, 165 Streptopelia,
bitorquata, 198 dnsumieri, 198 gaimardi, 199 strinhis, Butnrides, 81 stricklandi, Loxia, 641
striped,
cuckoo, 574 flycatcher, 608 owl, 576 sparrow, 644 striped-tail hummingbird, 589 Strix,
flammea, 217 hypugaea, 575 pratincola, 574 Sturnella magna mexicana, 634 Stumus,
cineraceus, 302 holosericeus, 632 philippensis, 302 subminuta, Erolia, 152 Tringa, 152 subpagana, Elaenia, 614 Sula,
dactylatra, 75 leucogaster, 76 personata, 75 plotus, 76 rubripes, 75 sula, 75 sula, Sula, 75
sulcirostris, Crotophaga, 573 sulphur-bellied flycatcher, 608 sulphur-rumped flycatcher, 614 sulphuratus, Pitangus, 609 Ramphastos, 597 sulphureipygius, Myiobius, 614 Tyrannula, 614 sumatrana. Sterna, 156 Sumichrast blackbird, 633 sumichrasti, Platypsaris, 604 sun grebe, 563 superciliosa,
Aimophila, 644 Plagiospiza, 644 superciliosus, Phaethomis, 582 Swainson pewee, 610 swallow, bam, 616 eastern bam, 239 mangrove, 616 rough-winged, 615 swamphen, purple, 129 swift,
chestnut-collared, 581 chimney, 581 Vaux, 581 swiftlet,
Carolines, 224, 225, 226 edible nest, 221, 222 Synallaxis erythrothorax furtiva, 601 synallaxis, mfous-breasted, 601
Index TO Volume 3
677
Sylvia,
culicivora, 631
formosa, 627
magnolia, 626
occidentalis, 627
olivacea, 626
syrinx, 254 sylviodes, Sittasomus, 601 syrinx,
Acrocephalus, 254
Sylvia, 254
Tachyphonus sanguinolentus, 636 tach>T)tera, Collocalia, 222 tahitiensis,
Eudynamis, 214
Numenius, 139
Scolopax, 139 taitensis,
Cuculus, 214
Eudynamis, 214 takatsukasae,
Monarcha, 274
Monarcharses, 274 takatsukasai, Zosterops, 322 talpacoti, Columbigallina, 568 tamaulipensis. Icterus, 634 tanager,
Abbot, 636
Bishop, 636
crimson-collared, 636
red ant, 637
Sclater ant, 637
summer, 637
white-winged, 637 Tanagra,
abbas, 636
affinis, 635
atriceps, 638
auricollis, 629
cyanea, 639
diaconus, 636
grandis, 638
lauta, 635
sanguinolentus, 636 Tangavius aeneus aeneus, 632 Tantalus variegatus. 137 Tapera naevia excellens, 574 tattler,
American wandering, 145
gray-tailed, 144 teal,
blue-winged, 550
European, 100
Garganey, 100
green-vi'inged, 100 telephonus, Cuculus, 213 telescope, in study of bird
migrations, 372 tenebrosa, Collurincincla, 282 tenebrosus, Rectes, 282 tenuirostre, Edolisoma, 239, 241
tenuirostris,
Calidris, 151
Procellaria, 66
Puffinus, 66
Totanus, 151 Tephras,
finschii, 328
ruki, 332 teraokai,
Ducula, 193
Globicera, 193
Halcyon, 233 tern,
black -billed common, 155
black-naped, 156
blue-gray, 164
bridled, 160
Caspian, 565
crested, 162
least, 161
sooty, 161
spectacled, 160
white, 174, 180
white-winged black, 155 texanus, Pitangus, 609 Texas sparrow, 642 texensis,
Chordeiles, 579
Muscicapa, 609
Myiozetetes, 609 Thamnophilus dohatus intermedius,
603 thermophila, Piaya, 572 thick-knee, 564 Thraupis,
abbas, 636
diaconus, 636
episcopus, 636 thrush,
Chinese blue rock, 248
dusky, 248
orange-billed nightingale, 623
russet-backed, 623
wood, 622 Thryomanes bewdcldi bairdi, 619 Thryothonis,
bairdi, 619
luscinius, 251
mexicanus, 620 Tiaris olivacea pusilla, 640 tiger bittern, 548 Tigrisoma mexicana, 548 time patterns in bird migrations, 414 tinamou,
Boucard, 544
cinnamomeous, 545
great, 543
Tinamus,
boucardi, 544 m^.jor, 543 robustus, 543
678
University of Kansas Publs., Mus. Nat. Hist.
Tinian monarch, 274
Tityra semifasciata personata, 605
tityra,
inquisitive, 605
Mexican, 605 tody motmot, 594 toliniei, Oporomis, 627 topography, in relation to migration,
424 torquata,
Alcedo, 592
Megaceryle, 592 torquatus,
Pteroglossus, 597
Ramphastos, 597 torqueola, Sporophila, 640 Totanus,
acuminatus, 152
brevipes, 144
tenuirostris, 151 toucanet, emerald, 596 towhee,
brown, 643
collared, 642
spotted, 643 townsendi,
Ducula, 194
Globicera, 194 traillii, Empidonax, 611 trichas,
Geothlypis, 628
Turdus, 628 Trichoglossus, rubiginosus, 211 trichroa,
Erythrura, 336, 337, 338
Fringilla, 336 tricolora,
Motacilla, 630
Setophaga, 630 Tringa,
glareola, 142
hypoleucos, 143
interpres, 147
lobata, 154
melanoleuca, 142
minutilla, 564
nebularia, 141
squatarola, 131 Trochilus,
beryllinus, 589
colubris, 590
curvipennis, 583
hemileucurus, 586
mellivorus, 587
prevostii, 587
tzacati, 589 Troglodytes,
aedon, 620
brunneicollis, 620
obsoleta, 620
parkmanii, 620
Trogon,
braccatus, 591
citreolus, 591
collaris, 591
massena, 590
melanocephalus, 591
puella, 591
violaceus, 591 trogon,
citreohne, 591
gartered, 591
Jalapa, 591
massena, 590 tropic-bird,
red-billed, 70
red-tailed, 71
white-tailed, 72 Truk,
greater white-eye, 332
monarch, 269 Trynga alba, 150 tufted,
duck, 10;5
flycatcher, 613 Turdus,
assimihs, 622
cestulatus, 623
grayi, 622
melpomene, 623
migratorius, 621
motacilla, 627
mustelinus, 622
obscurus, 248
philippensis, 248
phillipsi, 621
trichas, 628
ustulatus, 623 turkey vulture, 551 tumstone, 147 typhicola, Geothlypis, 628 Tyrannula,
flaviventris, 610
minima, 610
nigricans, 606
sulphureipygia, 614 tvrannulus, Myiarchus, 609 Tyrannus,
borealis, 610
chloronotus, 608
melancholicus, 608
monachus, 608 tvrannus, Muscivora, 608 Tyto alba pratincola, 574 tzacati,
Amazilia, 589
Trochilus, 589 tzitzdhoa. Anas, 101
ualensis, Collocalia, 224 uncinatus, Chrondrohierax, 552 uraniae, Rhipidura, 261
Index to Volume 3
679
Urubitinga ridgwayi, 554 urubitinga, Hypomorphnus, 554 ustulata, Hylocichla, 623 iistulatus, Turdus, 623
vanikorensis, CoUocalia, 226 varia,
Mniotilta, 626
Motacilla, 626 varied bunting, 639 variegata, Elaenia, 608 variegatus,
Legatus, 608
Numenius, 137
Tantalus, 137 Vaux swift, 581 vauxi, Chaetura, 581 vegae, Larus, 154
Veniliomis fumigatus oleaginus, 600 Veracruz,
Birds from the state of, 531
gazetteer of localities, 538
life-zones, 535 veraecrucis,
Centurus, 598
Melanerpes, 598
Phaethomis, 582 vermilion flycatcher, 607 verreauxi, Leptotila, 569 versicolor,
Agyrtrina, 588
Passerina, 639
Rhipidura, 264
Spiza, 639 vesper sparrow, 644 vetula,
Ortalis, 559
Penelope, 559 villosus, Dendrocopos, 600 violacea, Nyctanassa, 547 violaceus, Trogon, 592 violet-backed starling, 302 virens,
Dendroica, 626
Icteria, 629
Motacilla, 626 Vireo,
amauronotus, 625
flavoviridis, 625
gilvus, 625
griseus, 625
noveboracensis, 625
olivaceus, 625
solitarius, 625 vireo,
blue-headed, 625
red-eved, 625
warbling, 625
white-eyed, 625 Vireosvlvia flavoviridis, 625 virescens,
Ardea, 546
Butorides, 546
virgata, Ciccaba, 575 virgatus, Accipiter, 104 Virginia rail, 562 virginianus, Colinus, 560 viridicata, Elaenia, 615 viridigenalis,
Amazona, 570
Chrysotis, 570 vivida, Xanthoura, 618 vociferus,
Caprimulgus, 581
Charadrius, 37, 564 Volatinia jacarina splendens, 641 Volvocivora insperata, 242 Vultur,
atratus, 550
aura, 551
harpyja, 554
papa, 550 vulture,
black, 550
king, 550
turkey, 551
warbler,
black and white, 626
black-throated green, 626
Canada, 630
chestnut-sided, 627
hermit, 627
hooded, 629
Kentucky, 627
Lichtenstein, 631
Macgillivrav, 627
magnolia, 626
olive, 626
red, 630
rufous-capped, 631
Wilson, 629
yellow, 626
yellow-throated, 627 warbling vireo, 625 water-thrush, Louisiana, 627 water-turkey, 546 waxwing,
Bohemian, 482
cedar, 482
coloration, 485
food, 493
Japanese, 483
nest building, 491
origin of species, 519
phylogeny, 473
skeleton, 494 weaver-finch, black-breasted, 339 wedgebill, 601
wedge-tailed shearwater, 64, 65 western wood pewee, 610 whimbrel, 137 whip-poor-will,
American, 581
Salvin, 580
680
University of Kansas Publs., Mus. Nat. Hist.
white,
ibis, 549
snake hawk, 553 white-beUied emerald, 588 white-breasted,
wood-swallow, 248
wood wren, 620 white-browed,
hermit, 582
rail, 123 white-capped noddy, 170 white-collared kingfisher, 233, 235,
237 white-crowned parrot, 570 white-eye,
bridled, 316, 318, 319, 320, 321
Micronesian dusky, 326, 327, 328
Palau greater, 330
Ponape greater, 333
Tnik greater, 332
Yap greater, 331 white-eyed vireo, 625 white-tailed,
kite, 551
tropic-bird, 72 white-throated,
flycatcher, 611
ground dove, 203, 207 white-winged,
dove, 567
black tern, 155 widgeon, 102 Wilson,
snipe, 564
warbler, 629 Wilsonia,
canadensis, 630
citrina, 629
pileolata, 629
pusilla, 629 winkleri, Muscadivora, 191 wood,
ibis, 549
owl, 575
rail, 562
sandpiper, 142
thrush, 622 wren,
gray-breasted, 620
white-breasted, 620 woodhewer,
ivory-billed, 601
streaked-headed, 601 woodpecker,
ant-eating, 598
chestnut-colored, 599
golden-fronted, 598
Guatemalan ivory-billed, 599
hairy, 600
ladder-backed, 600
Lichtenstein, 599
lineated, 599
oleaginous, 600 wood-swallow, white-breasted, 284 wortheni, Spizella, 645 wren,
banded cactus, 619
Bewick, 619
brown-throated, 620
canyon, 620
common rock, 620
house, 620
rufous-naped cactus, 619
white-breasted wood, 620
xanthonura, Gallicolumba, 203, 207 Xanthoura,
luxuosa, 618
vivida, 618
yncas, 618 Xiphorhynchus,
eburneirostris, 601
flavigaster, 600, 601
saltuarius, 600
yamashinae,
Acrocephalus, 256
Conopoderas, 257 yapensis,
Ixobrychus, 91
Phlegoenas, 203 Yap,
greater white-eye, 331
monarch, 272 yellow warbler, 626 yellow-bellied,
elaenia, 614
flycatcher, 610 yellow-billed cuckoo, 573 yellow-breasted chat, 629 yellow-crowned night heron, 547 yellow-faced grassquit, 640 yellow-headed parrot, 572 yellow-legs, greater, 142 yellow-tailed oriole, 634 yellow-thighed manakin, 605 yellow-throat,
common, 628
Nelson, 628 yellow-throated warbler, 627 yncas, Xanthoura, 618 yucatanensis,
Amazilia, 589
Nyctidromus, 579
zeledoni,
Cancroma, 548
Cochlearius, 548 Zenaida asiatica, 567 Zenaidura macroura carolinensis, 567 zonatus,
Campvlorhvnchus, 619
Picolaptes, 619
Index to Volume 3 681
zone-tailed hawk, 552 oleaginea, 331
Zonotrichia boucardi, 645 owstoni, 321
Zosterops, ponapensis, 327
cinerea, 326, 327, 328 rotensis, 319
conspicillata, 316, 318, 319, 320, saypani, 318
321, 322, 323 semperi, 320
finschii, 328 takatsukasai, 322
h^TDolais, 323
PRINTED BY
FERC VOILAND. JR.. STATE PRINTER
TOPEKA. KANSAS
1951
24-1735
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