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

by Rollin H. Baker

Sterna sumatrana sumatrana Raffles

Black-naped Tern

Sterna Sumatrana Raffles, Trans. Linn. Soc. London, 13, pt. 2, 1822, p. 329. (Type locality, Sumatra.)

Sterna melanauchen Kittlitz, Obser. Zool., in Lutké, Voy. "Le Séniavine," 3, 1836, pp. 306, 308 (Guahan, Ouleai); Hartlaub and Finsch, Proc. Zool. Soc. London, 1872, pp. 90, 113 (Pelew, Uap); Gräffe, 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 (Ponapé); idem, Proc. Zool. Soc. London, 1880, p. 577 (Ruk); idem, Ibis, 1881, pp. 113, 115 (Ponapé); Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, pp. 281, 299, 330, 353 (Ponapé, Mortlock, Nukuor, Ruk); Salvadori, Ornith. Papuasia, 3, 1882, p. 444 (Pelew, Mackenzie, Ruk, Ponapé, Marshalls); Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 74 (Pelew, Uap, Ruk, Luganor, Nukuor, Ponapé, Taluit); Sanders, Cat. Birds British Mus., 25, 1896, p. 126 (Carolines, Pelews, Marshalls); Nehrkorn, Kat. Eiers., 1899, p. 222 (Palau); Hartert, Novit. Zool., 7, 1900, p. 10 (Ruk); Schnee, Zool. Jahrbücher, 20, 1904, p. 390 (Marschall-Inseln); Takatsukasa and Kuroda, Tori, 1, 1915, p. 52 (Ruk, Ponapé); Uchida, Annot. Zool. Japon., 9, 1918, pp. 483, 488 (Ponapé).

Sterna sumatrana Wetmore, in Townsend and Wetmore, Bull. Mus. Comp. Zoöl., 63, 1919, p. 186 (Arhno).

Gygisterna sumatrana Kuroda, in Momiyama, Birds Micronesia, 1922, p. 52 (Pelew, Mackenzie, Yap, Ruk, Luganor, Nukuor, Ponapé, Taluit, Arhno).

Gygisterna sumatrana sumatrana Kuroda, Avifauna Riu Kiu, 1925, p. 192 (Carolines, Pelews).

Sterna sumatrana sumatrana Yamashina, Tori, 7, 1932, p. 410 (Aruno); Hachisuka, Birds Philippines, 2, 1932, p. 335 (Caroline, Pelew); Hand-list Japanese Birds, rev., 1932, p. 195 (Palau, Guam, Saipan, Yap, Truk, Lukunor, Nukuoro, Ponapé, Jaluit, Namu, Arhno, Majuro, Aurh); Peters, Check-list Birds World, 2, 1934, p. 336 (Caroline Islands); Mayr, List New Guinea Birds, 1941, p. 36 (Micronesia); Hand-list Japanese Birds, 3d ed., 1942, p. 218 (Babelthuap, Koror, Yap, Truk, Lukunor, Nukuoro, Ponapé, 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, Malaysia, west to India, and north to the Riu Kiu Islands. In Micronesia: Palau Islands—Babelthuap, Koror, Peleliu; Caroline Islands—Yap, Ulithi, Truk, Lukunor, Nukuoro, Ponapé; 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 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 September, 1945. Coultas (field notes) obtained reports of the finding of two eggs at the Palau Islands in the period October to December, 1931.

Parasites.—Uchida (1918:483, 488) records the following Mallophaga taken at Ponapé from this tern: Docophorus albemarlensis, Colpocephalum milleri, and Colpocephalum impertunum.

Remarks.—There are no records for the Black-naped Tern from the Mariana Islands, although the species is known from the Palau, Caroline 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, these birds appeared quite unafraid 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 sumatrana are recognized by Peters (1934:336): Sterna sumatrana mathewsi known from islands of the western Indian Ocean and Sterna s. sumatrana from islands of Oceania, 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 shortest and longest tail feather.

Table 19. Measurements of Gallinula chloropus

Location	No.	Wing	Tail	Difference: Longest and shortest tail feather	Exposed culmen	Tarsus
S. s. sumatrana Micronesia	13	221	127	65	37	20.5
		211-225	117-138	54-79	35-39	20.0-21.0
Phoenix and Union	5	228	113	66	37	19.5
					36-38	18.5-20.0
Fiji, Samoa, Tonga	29	221	131	63	38	20.0
		218-229	122-142	51-74	36-41	18.0-21.0
New Caledonia Loyalty, New Hebrides	8	224	141	72	39	19.5
		221-230	135-148	68-81	37-41	18.5-20.0
Queensland, Torres Straits	4	229	142	78	38	19.5
			139-148	71-83	36-40	18.5-20.0
Solomons	52	227	144	77	36	19.0
		220-232	129-162	66-95	34.0-38.5	18.5-20.5
New Guinea, Bismarcks	10	224	143	76	34	19.5
		219-231	135-146	67-81	32.0-36.5	18.5-20.0
Malay area	49	228	141	74	34	20.0
		220-234	125-153	63-84	32.0-37.0	19.0-20.5
China coast, Riu Kiu	21	223	144	77	35	19.5
		212-234	130-151	67-85	31.5-38.0	19.0-20.0
S. s. mathewsi Micronesia	10	220	125	71	38	19.0
					35.0-40.0	18.0-20.0

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. Murphy (1938:538) has written that this phenomenon is characteristic among some species which have both continental and insular populations (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. 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 questioned. 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 certainty until additional material is obtained.

Fig. 10. Geographic variation in the average length of the exposed culmen of Sterna sumatrana sumatrana.

Sterna lunata Peale

Spectacled Tern

Sterna lunata Peale, U. S. Expl. Exped., 8, 1848, p. 277. (Type locality, Vincennes Island, Paumotu Group.)

Sterna lunata 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, pp. 90, 113 (Pelew); Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 5, 41 (Palau); Saunders, Cat. Birds British Mus., 25, 1896, p. 100 (Pelew); Takatsukasa 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); Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 76 (Pelew).

Melanosterna 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 immature male at sea south of the eastern Caroline Islands at 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).

Melanosterna 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 Micronesia: Palau Islands—exact locality not known; Marshall Islands—Bikar.

Measurements.—Four adult males from the Palau Islands have the following 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 observed 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 (Ponapé); idem, Proc. Zool. Soc. London, 1877 (1878), p. 781 (Ponapé); idem, Journ. f. Ornith., 1880, p. 295 (Ponapé); Takatsukasa and Kuroda, Tori, 1, 1915, p. 62 (Ponapé).

Onychoprion fuscata infuscata Kuroda, in Momiyama, Birds Micronesia, 1922, p. 51 (Ponapé).

Sterna fuscata nibilosa Hand-list Japanese Birds, rev., 1932, p. 195 (Ponapé); Yamashina, Tori, 10, 1940, p. 677 (Helen Reef); Hand-list Japanese Birds, 3d ed., 1942, p. 218 (Ponapé, Helen Reef).

Sterna fuscata oahuensis Mayr, Birds Southwest Pacific, 1945, p. 25 (Micronesia).

Geographic range.—Breeds from the Hawaiian, Marcus, and Bonin islands south to the Phoenix Islands and Micronesia. In Micronesia: Mariana Islands—Asuncion; Palau Islands—Helen Reef; Caroline Islands—Ponapé.

Specimens examined.—Total number, 1 unsexed from Mariana Islands, AMNH—Asuncion (Jan. 18).

Remarks.—The systematic position of the Sooty Tern in Micronesia 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 O° 90´ S and 159° 50´ E, a position south of the eastern Caroline Islands. The bird is tentatively placed in the subspecies S. f. 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 species is very abundant at Marcus Island, which is north and east of the Marianas.

Sterna albifrons sinensis Gmelin

Least Tern

Sterna sinensis 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.

Thalasseus bergii pelecanoides (King)

Crested Tern

Sterna pelecanoides King, Surv. Intertrop. 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 (Ponapé); idem, Ibis, 1880, pp. 330, 332 (Ratak Chain); idem, Journ. f. Ornith., 1880, p. 295 (Ponapé); idem, Proc. Zool. Soc. London, 1880, p. 577 (Ruk); idem, Ibis, 1881, pp. 113, 115 (Ponapé); Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, pp. 281, 299, 330, 353 (Ponapé, Mortlock, Nukuor, Ruk); Salvadori, Ornith. Papuasia, 3, 1882, p. 434 (Ruk, Ponapé, 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, Ponapé, Marshall Islands); Hartert, Novit. Zool., 7, 1900, p. 10 (Ruk); Saunders, Cat. Birds British Mus., 25, 1896, p. 89 (Ponapé, Marshalls); Takatsukasa and Kuroda, Tori, 1, 1915, p. 52 (Ponapé); Uchida, Annot. Zool. Japon., 9, 1918, pp. 483, 488 (Ponapé).

Sterna bergeri Schnee, Zool. Jahrbücher, 20, 1904, p. 390 (Marschall-Inseln).

Sterna bergii cristata Stresemann, Novit. Zool., 21, 1914, p. 58 (Truk).

Thalasseus bergii pelecanoides Oberholser, Proc. U. S. Nat. Mus., 49, 1915, p. 523 (Marshall Islands); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 51 (Luganor, Nukuor, Ruk, Ponapé, Marshall Islands); Kuroda, Avifauna Riu Kiu, 1925, p. 188 (Marshall Islands); Hand-list Japanese Birds, rev., 1932, p. 194 (Palau, Faraulep, Truk, Lukunor, Nukuoro, Ponapé, 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, Ponapé, Jaluit, Mille, Aurh, Maloelab, Ailuk).

Thalasseus bergii cristatus Peters, Check-list Birds World, 2, 1934, p. 342 (Carolines, 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 Micronesia (see [figure 11]). In Micronesia: Palau Islands—Helen Reef, Peleliu, Babelthuap; Caroline Islands—Ulithi, Truk, Faraulep, Lukunor, Nukuoro, Ponapé; 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)—Ponapé, 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 Ponapé: Docophorus albemarlensis and Colpocephalum importunum.

Remarks.—Oberholser (1915:520-526, pl. 66) lists five subspecies (T. b. cristatus, T. b. halodramus, T. b. pelecanoides, T. b. rectirostris, and T. b. poliocercus) in the region including the coast of 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.

Table 20. Measurements of Thalasseus bergii in the Pacific Area

Location	No.	Wing	Longest tail feather	Shortest tail feather	Exposed culmen	Tarsus
Thalasseus bergii pelecanoides Palaus, Carolines, Marshalls	6	343	168	82	60
		334-352	153-184	80-85	58-65
Christmas, Phoenix, Tuamotus, Society, Fiji, Loyalty, New Hebrides	48	344	170	83	58	27
		329-362	145-198	77-92	54-64	25-29
Eastern Australia	14	345	165	88	58	27
		338-349	152-174	84-92	55-63	26-29
New Guinea, Bismarck, Archipelago, Moluccas	18	342	168	81	59	27
		332-361	144-194	75-87	53-64	26-28
Totals	86	344	169	83	58	27
		329-362	144-198	75-92	53-65	25-29
Thalasseus bergii cristatus Philippines, China, Formosa, Riu Kius	18	332	162	81	58	28
		324-342	149-182	78-87	55-64	26-30
Thalasseus bergii gwendolenae Western Australia	14	354	171	86	58	27
		339-369	162-182	81-91	53-65	25-29

Measurements, as shown in [table 20], indicate a wide range of sizes but, in most series, the averages 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 lists the measurements of the wing of eight Crested Terns from the Riu Kiu Islands as 322 to 340 (average 330). The occurrence of populations with shorter wings has already been pointed out in the work of Oberholser (1915:520-526), who divided the short-winged birds into two subspecies. It seems advisable to recognize but one subspecies, T. b. cristatus, for the birds with short wings and another subspecies, T. b. pelecanoides, to include the birds with the longer wings (see [figure 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.

Fig. 11. Geographic distribution of Thalasseus bergii. (1) T. b. bergii; (2) T. b. thalassinnus; (3) T. b. velox; (4) T. b. cristatus; (5) T. b. gwendolenae; (6) T. b. pelecanoides.

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 difficult 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. 559. (Type locality, Necker Island, Hawaiian Islands.)

Procelsterna cerulea saxatilis Yamashina, 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.

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 measurements: 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 = Philippines, ex. Sonnerat.)

Sterna stolida Chamisso, in Kotzebue's Voy. "Rurick," 3, 1821, pp. 150, 157 (Marshall Islands); Kittlitz, Kupfertaf. Naturgesch. Vögel, 3, 1833, p. 27, pl. 36, fig. 1 (Mordloks-Inseln); idem, Obser. Zool., in Lutké, Voy. "Le Séniavine," 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. Ornith., 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 = Ponapé); 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 (Ponapé); idem, Proc. Zool. Soc. London, 1877 (1878), p. 781 (Ponapé); idem, Journ. f. Ornith., 1880, pp. 295, 307 (Ponapé, Ruck, Kuschai); idem, Proc. Zool. Soc. London, 1880, p. 577 (Ruk, Ponapé, Kuschai); idem, Ibis, 1881, pp. 105, 109, 115, 246, 247 (Kuschai, Ponapé); Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, pp. 299, 330, 353 (Mortlock, Nukuor, Ruk); Salvadori, Ornith. Papuasia, 3, 1882, p. 455 (Pelews, Carolines, Marshalls); Finsch, Mitth. Ornith. Ver. Wien, 1884, p. 51 (Jaluit, Ponapé); Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 76 (Pelew, Mortlock, Ruk, Nukuor, Ponapé, 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. Jahrbücher, 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 (Ponapé, Ruk); Cox, Island of Guam, 1917, p. 22 (Guam); Uchida, Annot. Zool. Japon., 9, 1918, pp. 484, 488 (Palau, Ponapé); 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," Vögel, 1865, pp. 155, 162 (Puynipet = Ponapé).

Anous stolidus pileatus Hartert, Novit. Zool., 7, 1900, p. 9 (Ruk); Wetmore, in Townsend and Wetmore, Bull. Mus. Comp. Zoöl., 63, 1919, p. 183 (Kusaie); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 49 (Guam, Saipan, Pelew, Mortlock, Ruk, Wolea, Nukuoro, Ponapé, Kusaie, Marshalls); Hand-list Japanese Birds, rev., 1932, p. 195 (Koror, Urukthapel, Angaur, Saipan, Guam, Wolea, Truk, Mortlock, Lukunor, Nukuoro, Ponapé, 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, Ponapé, 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 unicolor? Ridgway, Bull. U. S. Nat. Mus., 50, pt. 8, 1919, p. 547 (Guam).

Table 21. Measurements of Anoüs stolidus of the Pacific Area

Location	No.	Wing	Tail	Exposed culmen
Anoüs stolidus ridgwayi Isabella, Cocos, Clipperton Islands	18	278	158	41
		260-295	147-166	38-42
Anoüs stolidus galapagensis Galapagos Islands	11	277	151	40
		274-282	142-160	38-42
Anoüs stolidus pileatus Hawaiian Islands: Nihoa to Midway	35	281	162	42
		268-299	149-176	38-40
Wake Islands	8	278	159	41
		273-285	152-170	39-43
Mariana Islands: Guam, Rota	12	280	167	41
		275-288	159-187	39-43
Palau Islands	9	278	161	41
		268-283	155-166	39-42
Caroline Islands	41	282	164	42
		270-291	150-173	39-45
Marshall Islands	3	282	164	42
		270-289	154-174	41-43
Ellice, Phoenix, Danger, Suvarov Islands	27	284	162	41
		265-295	152-174	39-44
Christmas Island	13	287	162	43
		280-292	152-174	40-46
Marquesas Islands	19	282	163	42
		275-291	155-170	40-43
Tuamotu Archipelago	38	287	165	42
		277-299	154-173	39-46
Society, Austral, Cook, Rapa Islands	16	290	164	43
		280-301	155-173	40-45
Oeno, Henderson, Ducie, Easter Islands	6	293	164	44
		285-298	154-175	41-45
Samoa, Fiji, Tonga	19	285	164	42
		277-295	153-173	39-44
Kermadecs, Norfolk	23	276	158	41
		269-289	148-173	38-43
New Hebrides, Solomons, New Guinea area	31	278	158	41
		265-287	150-172	39-44
Northwest Australia	9	263	145	40
		258-267	138-152	38-42
South China Sea area, Strait of Malacca	4	271	153	39
		262-278	148-257	37-40
Riu Kius, Japan	5	268	148	39
		259-275	143-155	37-40
Indian Ocean area: Seychelles, Aldebra, Providence, Somaliland	20	276	154	41
		270-286	146-164	39-42

Table 19. Measurements of Gallinula chloropus

Table 19. Measurements of Gallinula chloropus

Location	No.	Wing	Tail	Difference: Longest and shortest tail feather	Exposed culmen	Tarsus
S. s. sumatrana Micronesia	13	221	127	65	37	20.5
		211-225	117-138	54-79	35-39	20.0-21.0
Phoenix and Union	5	228	113	66	37	19.5
					36-38	18.5-20.0
Fiji, Samoa, Tonga	29	221	131	63	38	20.0
		218-229	122-142	51-74	36-41	18.0-21.0
New Caledonia Loyalty, New Hebrides	8	224	141	72	39	19.5
		221-230	135-148	68-81	37-41	18.5-20.0
Queensland, Torres Straits	4	229	142	78	38	19.5
			139-148	71-83	36-40	18.5-20.0
Solomons	52	227	144	77	36	19.0
		220-232	129-162	66-95	34.0-38.5	18.5-20.5
New Guinea, Bismarcks	10	224	143	76	34	19.5
		219-231	135-146	67-81	32.0-36.5	18.5-20.0
Malay area	49	228	141	74	34	20.0
		220-234	125-153	63-84	32.0-37.0	19.0-20.5
China coast, Riu Kiu	21	223	144	77	35	19.5
		212-234	130-151	67-85	31.5-38.0	19.0-20.0
S. s. mathewsi Micronesia	10	220	125	71	38	19.0
					35.0-40.0	18.0-20.0

Table 20. Measurements of Thalasseus bergii in the Pacific Area

Table 20. Measurements of Thalasseus bergii in the Pacific Area

Location	No.	Wing	Longest tail feather	Shortest tail feather	Exposed culmen	Tarsus
Thalasseus bergii pelecanoides Palaus, Carolines, Marshalls	6	343	168	82	60
		334-352	153-184	80-85	58-65
Christmas, Phoenix, Tuamotus, Society, Fiji, Loyalty, New Hebrides	48	344	170	83	58	27
		329-362	145-198	77-92	54-64	25-29
Eastern Australia	14	345	165	88	58	27
		338-349	152-174	84-92	55-63	26-29
New Guinea, Bismarck, Archipelago, Moluccas	18	342	168	81	59	27
		332-361	144-194	75-87	53-64	26-28
Totals	86	344	169	83	58	27
		329-362	144-198	75-92	53-65	25-29
Thalasseus bergii cristatus Philippines, China, Formosa, Riu Kius	18	332	162	81	58	28
		324-342	149-182	78-87	55-64	26-30
Thalasseus bergii gwendolenae Western Australia	14	354	171	86	58	27
		339-369	162-182	81-91	53-65	25-29

Table 21. Measurements of Anoüs stolidus of the Pacific Area

Table 21. Measurements of Anoüs stolidus of the Pacific Area

Location	No.	Wing	Tail	Exposed culmen
Anoüs stolidus ridgwayi Isabella, Cocos, Clipperton Islands	18	278	158	41
		260-295	147-166	38-42
Anoüs stolidus galapagensis Galapagos Islands	11	277	151	40
		274-282	142-160	38-42
Anoüs stolidus pileatus Hawaiian Islands: Nihoa to Midway	35	281	162	42
		268-299	149-176	38-40
Wake Islands	8	278	159	41
		273-285	152-170	39-43
Mariana Islands: Guam, Rota	12	280	167	41
		275-288	159-187	39-43
Palau Islands	9	278	161	41
		268-283	155-166	39-42
Caroline Islands	41	282	164	42
		270-291	150-173	39-45
Marshall Islands	3	282	164	42
		270-289	154-174	41-43
Ellice, Phoenix, Danger, Suvarov Islands	27	284	162	41
		265-295	152-174	39-44
Christmas Island	13	287	162	43
		280-292	152-174	40-46
Marquesas Islands	19	282	163	42
		275-291	155-170	40-43
Tuamotu Archipelago	38	287	165	42
		277-299	154-173	39-46
Society, Austral, Cook, Rapa Islands	16	290	164	43
		280-301	155-173	40-45
Oeno, Henderson, Ducie, Easter Islands	6	293	164	44
		285-298	154-175	41-45
Samoa, Fiji, Tonga	19	285	164	42
		277-295	153-173	39-44
Kermadecs, Norfolk	23	276	158	41
		269-289	148-173	38-43
New Hebrides, Solomons, New Guinea area	31	278	158	41
		265-287	150-172	39-44
Northwest Australia	9	263	145	40
		258-267	138-152	38-42
South China Sea area, Strait of Malacca	4	271	153	39
		262-278	148-257	37-40
Riu Kius, Japan	5	268	148	39
		259-275	143-155	37-40
Indian Ocean area: Seychelles, Aldebra, Providence, Somaliland	20	276	154	41
		270-286	146-164	39-42

Geographic range.—Islands in the Indian Ocean east to tropical parts of western and central Pacific. In Micronesia: Mariana Islands—Agrihan, Asuncion, Saipan, Rota, Guam; Palau Islands—Kayangel, Babelthuap, Koror, Urukthapel, Ngabad, Peleliu, Angaur; Caroline Islands—Ulithi, Truk, Wolea, Mortlock, Lukunor, Nukuoro, Ponapé, Kusaie; Marshall Islands—Jaluit, Mille, Aurh, Wotze, Bikini, Kwajalein.

Characters.—Adult: A large, dark-brown tern with 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 brownish, although 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, 8, 25, March 10, May 6, June 12, 13, Nov. 25, Dec. 25)—Ponapé, 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 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 obtained young birds at Kusaie in January and April and commented (field notes) that they probably nest "spasmodically at all times of the year." At Ponapé, 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 Micronesia; 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.

Parasites.—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: Neoschöngastia bougainvillensis, N. americana solomonis, N. egretta, Acariscus pluvius, and A. anous. Uchida (1918:484, 488) found the bird louse (Mallophaga), Nirmus separatus, on terms at Palau and at Ponapé he found Colpocephalum milleri on the bird. Bequaert (in litt.) has identified a fly (Hippoboscidae) as Olfersia 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). Anoüs 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 throughout the Pacific and west to Madagascar and the African coast in the Indian Ocean. These subspecies differ from one another principally 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 measurements do not appear to be significant from a taxonomic standpoint.

As shown in [table 21], measurements of length of wing for specimens from throughout most of the Pacific area are almost the same. Length of tail is correspondingly uniform. There is a gradual increase 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 indicate 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 material 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 A. s. pullus by Bangs. When specimens from the type locality of A. s. pileatus in the Philippine Islands are available, the true relationships of the populations from Micronesia and the other areas in the Pacific can be ascertained.

The tern found in the Hawaiians has the palest body and the most chalky-white forehead of any of the birds of the Pacific. 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 underparts 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 fading, 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 relatively similar conditions of plumage. In summary, the systematic position of the Common Noddy Terns of the Pacific seemingly depends on the characteristics of specimens from the type locality in the Philippines. When topotypes are available for study, 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 Ponapé, 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, where he saw approximately forty individuals in a flock with Gygis alba.

Anoüs stolidus is divided naturally into an Atlantic subspecies, which is distinguished by its browner color, and into several subspecies 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 Lutké, Voy. "Le Séniavine," 3, 1836, pp. 286, 308 (Ualan, Ouleai); idem, Denkw. Reise russ. Amer. Micron. und Kamchat., 2, 1858, p. 64 (Ualan).

Anous tenuirostris Hartlaub 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 (Ponapé, 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); Salvadori, Ornith. Papuasia, 3, 1882, p. 456 (Pelew, Ponapé, Marshalls); Finsch, Mitth. Ornith. Ver. Wien, 1884, p. 52 (Jaluit, Arno, Kuschai); Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1901 (1891), p. 77 (Pelew, Ualan, Ponapé, Nukuor, Luganor, Ruk); Hartert, Katalog Vogelsamml. Senckenb., 1891, p. 238 (Ualan); Takatsukasa and Kuroda, Tori, 1, 1915, p. 62 (Ruk); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 50 (Pelew, Ruk, Wolea, Luganor, Nukuor, Ponapé, Kusaie, Marshalls).

Anous leucocapillus Finsch, Proc. Zool. Soc. London, 1877 (1878), p. 781 (Ponapé); Nehrkorn, Journ. f. Ornith., 1879, p. 410 (Ponapé); Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, p. 281 (Ponapé); Finsch, Mitth. Ornith. Ver. Wien, 1884, p. 52 (Jaluit); Tristram, Cat. Coll. Birds, 1889, p. 10 (Pelew); Salvadori, Ornith. Papuasia, 3, 1882, p. 457 (Pelew); Wiglesworth, Abhandl. und Ber Zool. Mus. Dresden, 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, Ponapé, Kuschai, Bonham); 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. Jahrbücher, 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).

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 Australia, 2, 1912, p. 423 (Marianas?); Ridgway, Bull. U. S. Nat. Mus., 50, pt. 8, 1919, p. 553 (Mariannes?); Mathews, 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, Ponapé, 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, Ponapé, 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, Ponapé, Kusaie, Ebon, Namorik, Jaluit, Elmore, Mille, Aurh, Wotze, Ailuk).

Anous minutus marcusi Peters, Check-list Birds World, 2, 1934, p. 347 (Caroline Islands).

Anous minutus Bequaert, Mushi, 12, 1939, p. 82 (Ponapé); idem, Occ. Papers Bernice P. Bishop Mus., 16, 1941, p. 253 (Ponapé, 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: Mariana Islands—Asuncion, Saipan, Guam?; Palau Islands—Babelthuap, Koror, Peleliu; Caroline Islands—Ulithi, Truk, Ponapé, 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 eyelid with white streak, upper eyelid with white spot. Resembles A. t. melanogenys but wing and tail longer and superciliary stripe narrower. Resembles A. t. minutus but with narrower, black superciliary stripe.

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)—Ponapé, 15 (Dec. 15)—Kusaie, 17 (Jan. 10, March 10-30, April 1-10); Marshall Islands, USNM—Bikini, 4 (May 2, 14); AMNH—no locality given, 1 (Sept. 3).

Table 22. Measurements of Anoüs tenuirostris of the Pacific Area

Location	No.	Wing	Tail	Exposed culmen
Anoüs tenuirostris melanogenys Hawaiian Islands	29	222	113	41
		210-229	105-120	41-48
Anoüs tenuirostris marcusi Hawaiian Islands	8	227	118	45
		218-231	112-124	44-48
Mariana Islands	1	223	117	44
Palau Islands	3	228	122	43
		227-228	117-126	41-45
Caroline Islands	32	229	120	44
		220-240	113-127	40-47
Marshall Islands	5	224	118	44
		222-229	114-123	41-46
Anoüs tenuirostris minutus Hawaiian Islands	13	227	120	44
		220-234	108-128	41-46
Phoenix, Howland, Union, Danger, Suvarov Islands	9	229	119	46
		226-233	113-124	42-48
Anoüs tenuirostris minutus Marquesas Islands	10	226	117	45
		220-233	115-124	42-48
Tuamotu Archipelago	17	229	118	45
		222-234	112-126	42-47
Society, Cook, Austral Islands	12	230	118	46
		223-238	114-120	43-47
Samoa, Fiji, Tonga Islands	6	228	118	44
		224-231	115-121	42-47
Kermadec, Norfolk Isl'ds, New Zealand	15	226	116	44
		219-235	112-121	42-47
New Hebrides, Solomon, Bismarck, Admiralty Islands, New Guinea	34	229	117	43
		222-237	109-130	40-46
Anoüs tenuirostris diamesus Marquesas Islands	14	230	120	44
		224-237	114-127	41-47

Nesting.—Few reports have been obtained concerning the nesting of the White-capped Noddy in Micronesia. Finsch (1881b:107) recorded nests, and Nehrkorn (1899:222) reported on eggs taken at Kusaie. Yamashina (1932a:409) recorded the taking of eggs at Koror in the Palau Islands on January 19 and November 10 and in the Marshalls at Namo on October 19, and at Iringlab 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 Ponapé. 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 (Hippoboscidae), Alfersia aenescens, from the White-capped Noddy taken at Ponapé and Palau.

Remarks.—The subspecies of Anoüs tenuirostris are well differentiated 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). Owston'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, Ponapé 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 observed small flocks of four to ten individuals flying offshore and feeding inside the reef. They were frequently observed in company with Sterna sumatrana. 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, Christmas Island.)

Gygis candida Finsch, Ibis, 1880, p. 220 (Taluit); Saunders (part), Cat. Birds British Mus., 25, 1896, p. 149 (Marshalls); Schnee, Zool. Jahrbücher, 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, Agrigan, 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 (Marianan); 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. Avium 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 Fanning 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.

Specimens examined.—Total number, 41 (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, according 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 unusual problem in that there are few characters available to distinguish the subspecies. Measurements of taxonomic value include those of the wing, tail, exposed culmen, and depth and the shape of the culmen. There appears to be no significant secondary sexual difference between males and females, 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 (Fernando 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. In G. a. alba the length of wing as well as most of the other measurements differ but slightly from those of some of the populations in the Pacific area although the slender bill of the Atlantic bird is a distinctive character, as pointed out by Murphy (1936:1166).

Table 23. Measurements of Subspecies 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	24	246	99	71	40	8.0	14.5
		239-256	93-111	68-77	35-44	7.5-9.0	13.0-16.5
Gygis alba monte	35	232	106	71	39	8.5	13.5
		224-244	98-116	64-81	37-44	8.0-8.5	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 relatively 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 examined 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 subspecies G. alba by Peters and others. On the islands of Hatutu and Motane in the Marquesas, the Whitney South Sea Expedition obtained some birds which appear to be intergrades between the two subspecies of White Terns in the area. The measurements of nine birds which show intergradation between G. a. microrhyncha and G. a. pacifica are listed in [table 24]. Probably the Marquesas population is tending toward complete reproductive isolation.

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.

Table 22. Measurements of Anoüs tenuirostris of the Pacific Area

Table 22. Measurements of Anoüs tenuirostris of the Pacific Area

Location	No.	Wing	Tail	Exposed culmen
Anoüs tenuirostris melanogenys Hawaiian Islands	29	222	113	41
		210-229	105-120	41-48
Anoüs tenuirostris marcusi Hawaiian Islands	8	227	118	45
		218-231	112-124	44-48
Mariana Islands	1	223	117	44
Palau Islands	3	228	122	43
		227-228	117-126	41-45
Caroline Islands	32	229	120	44
		220-240	113-127	40-47
Marshall Islands	5	224	118	44
		222-229	114-123	41-46
Anoüs tenuirostris minutus Hawaiian Islands	13	227	120	44
		220-234	108-128	41-46
Phoenix, Howland, Union, Danger, Suvarov Islands	9	229	119	46
		226-233	113-124	42-48
Anoüs tenuirostris minutus Marquesas Islands	10	226	117	45
		220-233	115-124	42-48
Tuamotu Archipelago	17	229	118	45
		222-234	112-126	42-47
Society, Cook, Austral Islands	12	230	118	46
		223-238	114-120	43-47
Samoa, Fiji, Tonga Islands	6	228	118	44
		224-231	115-121	42-47
Kermadec, Norfolk Isl'ds, New Zealand	15	226	116	44
		219-235	112-121	42-47
New Hebrides, Solomon, Bismarck, Admiralty Islands, New Guinea	34	229	117	43
		222-237	109-130	40-46
Anoüs tenuirostris diamesus Marquesas Islands	14	230	120	44
		224-237	114-127	41-47

Table 23. Measurements of Subspecies of Gygis alba From the Atlantic and Indian Ocean Area

Table 23. Measurements of Subspecies 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	24	246	99	71	40	8.0	14.5
		239-256	93-111	68-77	35-44	7.5-9.0	13.0-16.5
Gygis alba monte	35	232	106	71	39	8.5	13.5
		224-244	98-116	64-81	37-44	8.0-8.5	12.5-14.0

Peters (1934:348, 349) recognizes three other subspecies from the Pacific area: G. a. rothschildi Hartert from Laysan, Lisiansky, and Krusenstern islands; G. a. candida (Gmelin) from "the Carolines east to Christmas Island and south to the Tonga and Society Islands"; and G. a. royana Mathews from Norfolk and the Kermadec Islands. Birds from Revilla Gigedo, Cocos and Clipperton islands, although geographically isolated, are placed in G. a. candida. 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 Marianas east to Wake and the Hawaiian Chain and south to the Marshall, Phoenix, Christmas and the Fanning islands (see [figure 12]). 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 population occurs; it is separable as G. a. royana. The measurements of these birds are given in [table 24].

Table 24. Measurements of 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	238	109	65	36
					34-38
Mariana Islands	35	237	111	69	38	9.0	13.0
		227-246	98-120	61-75	36-41		12.0-14.0
Wake Islands	10	236	109	69	38		13.0
		232-243	101-118	64-77	37-41		13.0-14.0
Hawaiian Islands	36	235	109	68	37	8.5	13.0
		220-246	102-118	64-74	33-40	8.0-9.0	12.0-14.0
Marshall Islands	4	234	111	71	39
		231-238	107-115	70-73	38-40
Phoenix, Howland, Hull, Canton Islands	8	238	107	70	39	8.5	14.0
		237-240	101-116	64-76	37-41
Fanning, Washington, Christmas Islands	19	238	107	68	38	8.0	13.5
		227-242	97-119	65-72	37-42	7.5-9.0	12.0-15.0
Totals	116	236	109	69	38	8.5	13.0
		220-246	107-120	61-77	33-42	7.5-9.0	12.0-15.0
Gygis alba pacifica (Lesson) Caroline, Palau Islands	33	245	116	73	42	8.5	13.5
		236-253	112-125	67-76	38-44		13.0-13.5
Bismarck Arch., Solomon Islands	12	247	116	74	42
		242-256	105-129	68-78	39-45
Samoa, Wallis, Fiji, Tonga, Niue Islands	20	247	115	71	42
		239-254	110-127	67-78	39-44
Line, Danger Islands	13	245	115	73	41
		238-252	107-118	69-78	39-42
Cook, Austral Islands	29	247	114	73	42
		241-255	104-124	65-78	40-45
Society Islands	37	249	113	71	42	8.5	13.5
		241-257	107-126	62-76	40-45	8.0-9.0	12.0-14.0
Tuamotu Arch	118	245	114	72	42
		236-252	107-127	62-82	38-46
Rapa, Bass Rocks, Oeno, Henderson, Ducie, Pitcairn, Easter Islands	54	247	113	73	41
		240-255	106-126	63-84	40-45
Clipperton, Cocos Islands	10	245	115	72	40	8.5	13.5
		240-253	110-120	71-73	38-43	8.5-9.5	13.0-14.0
Totals	326	246	114	72	42	8.5	13.5
		236-257	104-129	62-84	38-46	8.0-9.5	12.0-14.0
Intergrades between G. a. microrhyncha and G. a. pacifica	9	237	105	74	38	7.5	13.0
		230-247	93-122	67-89	36-41	7.0-8.0	12.0-14.0
Gygis alba microrhyncha	55	218	78	64	36	6.5	12.0
		211-235	72-96	60-75	32-39	6.0-8.0	11.0-12.5
Gygis alba royana Mathews Norfolk Island	16	250	113	73	42
		242-257	105-124	68-79	41-44
Kermadec Islands	12	251	115	75	43
		244-255	110-121	71-81	40-46
Totals	28	250	114	74	42
		242-257	105-124	68-81	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 resemblance 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 differences 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 two islands. The occurrence of the distinct G. a. microrhyncha in the Marquesas may be accounted for by such nonmigratory behavior. Mayr (1945a:27), however, is of the opinion that White Terns found in the Bismarck Archipelago, the Solomons, Santa Cruz and New Hebrides 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 Islands, 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 Kermadecs. 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. Vögel, 3, 1833, p. 28 (Carolinen); idem, Obser. Zool., in Lutké, Voy. "Le Séniavine," 3, 1836, pp. 286, 299, 308 (Ualan, Lougounor, Ouleai).

Gygis candida Hartlaub, Archiv f. Naturgesch., 18, 1852, p. 137 (Carolinen); Hartlaub, Journ. f. Ornith., 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); Gräffe, 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 (Ponapé); idem, Proc. Zool. Soc. London, 1877 (1878), p. 782 (Ponapé); idem, Journ. f. Ornith., 1880, pp. 295, 309 (Ponapé, Kuschai); idem, Proc. Zool. Soc. London, 1880, p. 577 (Ruk); idem, Ibis, 1881, pp. 105, 106, 109, 115, 246, 247 (Kushai, Ponapé); 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, Ponapé, 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 (Carolines); Kuroda (part), Avifauna Riu Kiu, 1925, p. 193 (Carolines); Hand-list Japanese Birds (part), rev., 1932, p. 196 (Pelew, Yap, Wolea, Luganor, Ruk, Ponapé, Kusaie); Yamashina (part), Tori, 10, 1940, p. 678 (Babelthuap).

Gygis albus kittlitzi Kuroda (part), in Momiyama, Birds Micronesia, 1922, p. 50 (Pelews, Yap, Wolea, Luganor, Nukuor, Ruk, Ponapé, Kusaie).

Leucanous albus kittlitzi Mathews (part), Syst. Avium Australasianarum, 1, 1927, p. 143 (Carolines).

Gygis 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, Ponapé, 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, Ponapé, 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)—Ponapé, 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.

Remarks.—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 writer 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, Semper 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.

Columba livia Gmelin

Blue Rock Pigeon

Columba domestica [Greek: b] 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

Ptilinopus ponapensis Finsch, Proc. Zool. Soc. London, 1877 (1878), p. 779. (Type locality, Ponapé.)

Ptilinopus? fasciatus Finsch, Journ. Mus. Godeffroy, 12, 1876, pp. 18, 37 (Ponapé).

Ptilopus fasciatus Elliot, Proc. Zool. Soc. London, 1878, p. 536 (Ponapé); Tristram, Cat. Birds, 1889, p. 44 (Ponapé).

Ptilopus ponapensis Schmeltz, Verhandl. Ver. nat. Unterhaltung Hamburg, 1877 (1879), pp. 178, 179 (Ponapé); Finsch, Proc. Zool. Soc. London, 1880, p. 576 (Ruk, Ponapé); idem, Journ. f. Ornith., 1880, pp. 291, 303 (Ponapé); idem, Proc. Zool. Soc. London, 1880, p. 578 (Ruk, Ponapé); idem, Ibis, 1881, pp. 113, 115 (Ponapé); Wiglesworth, Ibis, 1891, p. 583 (Ponapé, Ruk); idem, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 50 (Ponapé, Ruk); Salvadori, Cat. Birds British Mus., 21, 1893, p. 93 (Ponapé, Ruk); Oustalet, Nouv. Arch. Mus. Hist. Nat., Paris, (3), 7, 1895, p. 222 (Ponapé); Nehrkron, Kat. Eiers., 1899, p. 180 (Ruk); Dubois, Syn. Avium, 2, 1904, p. 736 (Ruck, Ponapé); Reichenow, Die Vögel, 1, 1913, p. 354 (Ruk, Ponapé); Takatsukasa and Kuroda, Tori, 1, 1915, p. 52 (Ruk, Ponapé); Wetmore, in Townsend and Wetmore, Bull. Mus. Comp. Zoöl., 63, 1919, p. 189 (Uala, Ponapé).

Ptilinopus ponapensis Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, p. 353 (Ruk); Hartert, Novit. Zool., 7, 1900, p. 7 (Ruk, Ponapé); Seale, Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 42 (Ponapé); Matschie, Journ. f. Ornith., 1901, p. 113 (Ruck, Ponapé); Mathews, Syst. Avium Australasianarum, 1, 1927, p. 32 (Ponapé); Bequaert, Mushi, 12, 1939, pp. 81, 82 (Ponapé); Mayr. Proc. 6th Pacific Sci. Congr., 4, 1939 (1941), p. 204 (Ponapé); Bequaert, Occ. Papers Bernice P. Bishop Mus., 16, 1941, pp. 266, 290 (Ponapé).

Ptilinopus Ponapensis Christian, The Caroline Islands, 1899, p. 357 (Ponapé).

Ptilinopus ponepensis ponapensis Kuroda, in Momiyama, Birds Micronesia, 1922, p. 57 (Ponapé, Ruk); Yamashina, Tori, 7, 1932, p. 408 (Ponapé); Hand-list Japanese Birds, rev., 1932, p. 190 (Ponapé, Ruk); Peters, Check-list Birds World, 3, 1937, p. 31 (Ruk, Ponapé); Hand-list Japanese Birds, 3d ed., 1942, p. 213 (Ponapé, Truk).

Ptilinopus porphyraceus ponapensis Ripley and Birckhead, Amer. Mus. Novit., no. 1192, 1942, p. 7 (Ruk, Ponapé); Mayr, Birds Southwest Pacific, 1945, p. 289 (Truk, Ponapé); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 59 (Truk).

Geographic range.—Micronesia: Caroline Islands—Truk, Ponapé. Characters.—Adult male: A green fruit dove with forehead, anterior lores and crown near "pansy purple," faintly margined with yellow; occiput, sides of head, neck, upper breast grayish-green with bifid feathers of midbreast more 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 yellow, 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; primaries and secondaries edged on outer webs with yellowish; underwing 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 resembles adult male, but slightly smaller and duller.

Immature: Resembles adult, but entirely green with yellow edgings on feathers and lacking crimson crown and colored breast patch.

Measurements.—Measurements of subspecies of P. porphyraceus in Micronesia are presented in [table 25].

Table 25. Measurements of Ptilinopus porphyraceus in Micronesia

Subspecies	Number	Wing	Exposed culmen	Tarsus
P. p. ponapensis	12 males	137 (133-141)	14 (13-15)	25 (24-27)
	11 females	133 (126-137)	14 (13-15)	25 (24-26)
P. p. hernsheimi	6 males	134 (130-138)	13 (12-14)	25 (24-26)
	5 females	127 (125-130)	13 (12-13)	25 (24-25)
P. p. pelewensis	10 males	133 (131-134)	15 (13-15)	25 (23-26)
	4 females	133 (130-138)	15 (14-15)	24 (23-24)

Table 24. Measurements of Gygis alba From the Pacific Area

Table 24. Measurements of 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	238	109	65	36
					34-38
Mariana Islands	35	237	111	69	38	9.0	13.0
		227-246	98-120	61-75	36-41		12.0-14.0
Wake Islands	10	236	109	69	38		13.0
		232-243	101-118	64-77	37-41		13.0-14.0
Hawaiian Islands	36	235	109	68	37	8.5	13.0
		220-246	102-118	64-74	33-40	8.0-9.0	12.0-14.0
Marshall Islands	4	234	111	71	39
		231-238	107-115	70-73	38-40
Phoenix, Howland, Hull, Canton Islands	8	238	107	70	39	8.5	14.0
		237-240	101-116	64-76	37-41
Fanning, Washington, Christmas Islands	19	238	107	68	38	8.0	13.5
		227-242	97-119	65-72	37-42	7.5-9.0	12.0-15.0
Totals	116	236	109	69	38	8.5	13.0
		220-246	107-120	61-77	33-42	7.5-9.0	12.0-15.0
Gygis alba pacifica (Lesson) Caroline, Palau Islands	33	245	116	73	42	8.5	13.5
		236-253	112-125	67-76	38-44		13.0-13.5
Bismarck Arch., Solomon Islands	12	247	116	74	42
		242-256	105-129	68-78	39-45
Samoa, Wallis, Fiji, Tonga, Niue Islands	20	247	115	71	42
		239-254	110-127	67-78	39-44
Line, Danger Islands	13	245	115	73	41
		238-252	107-118	69-78	39-42
Cook, Austral Islands	29	247	114	73	42
		241-255	104-124	65-78	40-45
Society Islands	37	249	113	71	42	8.5	13.5
		241-257	107-126	62-76	40-45	8.0-9.0	12.0-14.0
Tuamotu Arch	118	245	114	72	42
		236-252	107-127	62-82	38-46
Rapa, Bass Rocks, Oeno, Henderson, Ducie, Pitcairn, Easter Islands	54	247	113	73	41
		240-255	106-126	63-84	40-45
Clipperton, Cocos Islands	10	245	115	72	40	8.5	13.5
		240-253	110-120	71-73	38-43	8.5-9.5	13.0-14.0
Totals	326	246	114	72	42	8.5	13.5
		236-257	104-129	62-84	38-46	8.0-9.5	12.0-14.0
Intergrades between G. a. microrhyncha and G. a. pacifica	9	237	105	74	38	7.5	13.0
		230-247	93-122	67-89	36-41	7.0-8.0	12.0-14.0
Gygis alba microrhyncha	55	218	78	64	36	6.5	12.0
		211-235	72-96	60-75	32-39	6.0-8.0	11.0-12.5
Gygis alba royana Mathews Norfolk Island	16	250	113	73	42
		242-257	105-124	68-79	41-44
Kermadec Islands	12	251	115	75	43
		244-255	110-121	71-81	40-46
Totals	28	250	114	74	42
		242-257	105-124	68-81	40-46

Table 25. Measurements of Ptilinopus porphyraceus in Micronesia

Table 25. Measurements of Ptilinopus porphyraceus in Micronesia

Subspecies	Number	Wing	Exposed culmen	Tarsus
P. p. ponapensis	12 males	137 (133-141)	14 (13-15)	25 (24-27)
	11 females	133 (126-137)	14 (13-15)	25 (24-26)
P. p. hernsheimi	6 males	134 (130-138)	13 (12-14)	25 (24-26)
	5 females	127 (125-130)	13 (12-13)	25 (24-25)
P. p. pelewensis	10 males	133 (131-134)	15 (13-15)	25 (23-26)
	4 females	133 (130-138)	15 (14-15)	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.)—Ponapé, 53 (Nov., Dec).

Nesting.—Yamashina (1932a:408) reports on eggs taken at Ponapé on the following dates: July 10, 12, August 1, 12, 15, 21. Only one egg was found to a nest. Hartert (1900:8) records nests containing eggs in May and June at Truk. Coultas (field notes) describes the nest as a flimsy affair. At Ponapé in November 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.

Parasites.—Bequaert (1939:81, 82, and 1941:266, 290) records the two flies (Hippoboscidae), Ornithoctona plicata and O. pusilla, from the fruit dove at Ponapé.

Remarks.—McElroy of the NAMRU2 party found the birds in mountainous areas at Truk in December, 1945. At Ponapé in November 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 the lowlands and the mountain sides. Coultas writes that the Japanese 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., Journ. 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); Salvadori, 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 Vögel, 1, 1913, p. 355 (Kuschai); Wetmore, in Townsend and Wetmore, Bull. Mus. Comp. Zoöl., 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 Momiyama, 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 (Kusaie).

Geographic range.—Micronesia: Caroline Islands—Kusaie; Marshall Islands—Ebon (extinct?).

Characters.—Adult: 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 Birckhead (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 Marshall Islands. This distribution is of particular interest because it may show the pathway by which these small fruit pigeons invaded eastern Micronesia from Polynesia.

The small fruit dove at Kusaie has apparently the same habitat requirements as others of the species. Coultas (field notes) comments that in 1931 the birds were "quite common." He found them in the high trees on the mountain sides away from the native villages and gardens.

Ptilinopus porphyraceus pelewensis Hartlaub and Finsch

Crimson-crowned Fruit Dove

Ptilinopus pelewensis Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, p. 7. (Type 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); Gräffe, Journ. Mus. Godeffroy, 1, 1873, pl. 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, Ethnogr. Abth. Mus. Godeffroy, 1881, p. 407 (Palau); Matschie, Journ. f. Ornith., 1901, p. 113 (Palau); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 56 (Pelew); Mathews, 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 Vögel, 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, Garakayo, Peleliu, Ngabad, Anguar.

Characters.—Adult male: A green fruit pigeon with anterior lores and crown purple, margined with pale yellow; forehead paler than crown; chin and midthroat 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 greenish; 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 olivaceous-brown; breast duskier. Immature resembles adult, but lacks purple crown, violet breast spot, orange abdomen and maroon under tail-coverts; upper and lower parts margined with yellow; forehead pale green; supercillary stripe pale yellow.

P. p. pelewensis 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].

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 vegetation covering over the battle areas. The bird evidently lives a solitary 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, Traité 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. Vögel, 3, 1833, p. 25, pl. 23, fig. 2 (Guahan); idem, Obser. Zool., in Lutké, Voy. "Le Séniavine," 3, 1836, p. 305 (Guahan).

Ptilinopus purpuratus Hartlaub, Journ. f. Ornith., 1854, 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, pl. 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); Oustalet, 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 (Marianne); Salvadori, Cat. Birds British Mus., 21, 1893, p. 108 (Marianne Islands); Oustalet, 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 Marianen, 1913, p. 101 (Marianen); Reichenow, Die Vögel, 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).

Kurukuru roseicapillus Prévost and Des Murs, Voy. "Venus," Oiseaux, 1855, pp. 221, 231, 257, 259, 269 (Guam).

Ptilopus roseicapilla Bonaparte, Consp. Avium, 2, 1855, p. 21 (Mariannis).

Ptilonopus roseicapillus 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 roseicapillus 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); Mathews, 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, Check-list Birds World, 3, 1937, p. 31 (Saipan, Tinian, 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 feathers 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-coverts; 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 terminal band margined with yellow; iris pale yellow; bill grass-green; legs and feet reddish-black.

Adult female: Resembles male, but slightly smaller with neck greener. Immature 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.

Specimens 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.).

Table 26. Measurements of Ptilinopus roseicapillus

Number	Wing	Tail	Exposed culmen	Tarsus
32 adult males	127 (122-133)	80 (75-84)	14 (13-15.3)	25 (24-27)
10 adult females	124 (121-130)	76 (75-79)	13 (12-13.7)	24 (22-25.5)

Nesting.—At Guam, I obtained records of nests of fruit doves on March 1, 1927, 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 post natal 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 undisturbed, 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 Ptilinopus in Micronesia.—Oceania is especially rich in species and subspecies of the genus Ptilinopus. Ripley 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 region. Within the Oceanic region there are several species of Ptilinopus 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. perousii from Samoa, Fiji, and Tonga; P. mercierii from the Marquesas; 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 northward, probably by way of the Marshall Islands. P. p. hernsheimi from Kusaie and P. p. ponapensis from Ponapé 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 (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 independent origin and subsequent convergence, it may be more advisable to continue to regard the two as separate species."

Fig. 13. Geographic distribution of Ptilinopus 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.

Ducula oceanica monacha (Momiyama)

Micronesian Pigeon

Globicera oceanica monacha Momiyama, Birds Micronesia, March, 1922, p. 4. (Type locality, Yap.)

Columba oceanica Lesson and Garnot (part), Dict. Sci. Nat., éd. Levrault, 40, 1826, p. 317 (Pelew); Lesson (part), Man. d'Ornith., 2, 1828, p. 166 (Pelew); idem (part), Voy. "La Coquille," Zool., 2, 1828, pp. 432, 709 (Pelew); idem, Compl. de Buffon, 2d ed., 2, Oiseaux, 1838, p. 292 (Pelew); Prévost and Knip, Les Pigeons, 2, 1838-43, p. 49 (Pelew).

Carpophaga oceanica Hartlaub (part), Archiv. f. Naturgesch., 18, 1852, p. 115 (Pelewinseln); idem, 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. Ornith., 1901, p. 113 (Palau); Dubois (part), Syn. Avium, 2, 1904, p. 743 (Pelew); Reichenow (part), Die Vögel, 1, 1913, p. 351 (Palau).

Globicera oceanica Bonaparte (part), Consp. Avium, 2, 1855, p. 31 (Pelew); Reichenbach (part), Tauben, 1861, p. 120 (Pelew); Salvadori (part), Cat. Birds British 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), Fauna Centralpolynesiens, 1867, p. 145 (Pelew); Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, pp. 7, 118 (Pelew); Finsch and Hartlaub, Journ. 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 (Yap, 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, Babelthuap).

Geographic range.—Micronesia: Palau Islands—Babelthuap, Koror, Garakayo, Peleliu, Angaur, Palo 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. Measurements of Ducula oceanica

Subspecies	Number	Wing	Exposed culmen	Tarsus
D. o. monacha	8 males	228 (219-233)		36 (34-37)
	6 females	221 (214-228)	22.5 (22.0-23.0)	31 (29-33)
D. o. teraokai	5 males	230 (225-237)	23.5 (23.0-25.0)	34 (33-35)
	8 females	231 (221-238)	23.0 (21.5-24.5)	34 (33-35)
D. o. townsendi	8 males	226 (211-234)	24.0 (23.5-25.0)	34 (32-35)
	5 females	226 (215-233)	24.0 (23.0-24.5)	33 (32-34)
D. o. oceanica	4 males	222 (217-228)	25.0 (24.5-26.0)	35 (34-36)
	13 females	219 (213-226)	24.0 (23.0-25.0)	32 (30-34)
D. o. ratakensis[B]	6 males	(211-217)	(25.0-27.
	3 females	(208-213)	(25.0-26.0)

^[B] 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).

Nesting.—Yamashina (1932a:408) records the finding of one egg at Yap on December 3, 1930. The NAMRU2 party obtained no evidence of breeding activity of these pigeons at the Palaus in August and September, 1945. Coultas, 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 stomachs of birds taken in September, 1945. The birds were found to be exceedingly fat at this time.

Parasites.—Uchida (1918:486, 489) records the bird lice (Mallophaga), Goniocotes carpohagae and Colopocephalum 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 compiled 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 islands. 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 remained 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 prisoners 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 island some 160 miles southeast of Peleliu. The U. S. Navy frequently 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 various kinds of birds were numerous at Pulo Anna, but he did not recall seeing the pigeon. This pigeon may occur also at Fais, 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. Vögel, 3, 1833, p. 25, pl. 33, fig. 1 (Lugunor); idem (part), Obser. Zool., in Lutké, Voy. "Le Séniavine," 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 Centralpolynesiens, 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 Salvadori (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. Ornith. Ges. Bayern, 25, 1922, p. 234 (Ualam = Truk).

Ducula oceanica teraokai 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, 1945, p. 289 (Truk).

Geographic range.—Micronesia: Caroline Islands—Truk, ?Lukunor, ?Nukuoro.

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 chestnut. 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. o. monacha and that it might be advisable to unite these under one subspecific 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

Table 26. Measurements of Ptilinopus roseicapillus

Table 26. Measurements of Ptilinopus roseicapillus

Number	Wing	Tail	Exposed culmen	Tarsus
32 adult males	127 (122-133)	80 (75-84)	14 (13-15.3)	25 (24-27)
10 adult females	124 (121-130)	76 (75-79)	13 (12-13.7)	24 (22-25.5)

Table 27. Measurements of Ducula oceanica

Table 27. Measurements of Ducula oceanica

Subspecies	Number	Wing	Exposed culmen	Tarsus
D. o. monacha	8 males	228 (219-233)		36 (34-37)
	6 females	221 (214-228)	22.5 (22.0-23.0)	31 (29-33)
D. o. teraokai	5 males	230 (225-237)	23.5 (23.0-25.0)	34 (33-35)
	8 females	231 (221-238)	23.0 (21.5-24.5)	34 (33-35)
D. o. townsendi	8 males	226 (211-234)	24.0 (23.5-25.0)	34 (32-35)
	5 females	226 (215-233)	24.0 (23.0-24.5)	33 (32-34)
D. o. oceanica	4 males	222 (217-228)	25.0 (24.5-26.0)	35 (34-36)
	13 females	219 (213-226)	24.0 (23.0-25.0)	32 (30-34)
D. o. ratakensis[B]	6 males	(211-217)	(25.0-27.
	3 females	(208-213)	(25.0-26.0)

Globicera oceanica townsendi Wetmore, in Townsend and Wetmore, Bull. Mus. Comp. Zoöl., 63, 1919, p. 191. (Type locality, Ponapé).

Carpophaga oceanica Finsch (part), Proc. Zool. Soc. London, 1877 (1878), p. 780 (Ponapé); Nehrkorn, Journ. f. Ornith., 1879, p. 407 (Ponapé); Finsch (part), Journ. f. Ornith., 1880, p. 292 (Ponapé); idem, 1881, pp. 113, 115 (Ponapé); Schmeltz and Krause (part), Ethnogr. Abth. Mus. Godeffroy, 1881, p. 281 (Ponapé); Wiglesworth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 52 (Ponapé); Matschie (part), Journ. f. Ornith., 1901, p. 113 (Guam, error = Ponapé).

Globicera oceanica Salvadori (part), Cat. Birds British Mus., 21, 1893, p. 176 (Ponapé).

Globicera oceanica townsendi Momiyama, Birds Micronesia, 1922, p. 6 (Ponapé); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 55 (Ponapé); Mathews, Syst. Avium Australasianarum, 1, 1927, p. 45 (Ponapé); Hand-list Japanese Birds, rev., 1932, p. 190 (Ponapé).

Ducula oceanica townsendi Peters, Check-list Birds World, 3, 1937, p. 44 (Ponapé); Bequaert, Mushi, vol. 12, no. 2, 1939, pp. 81, 82 (Ponapé); idem, Occ. Papers Bernice P. Bishop Mus., 16, 1941, pp. 266, 290 (Ponapé); Hand-list Japanese Birds, 3d ed., 1942, p. 212 (Ponapé); Amadon, Amer. Mus. Novit., no. 1237, 1943, p. 11 (Ponapé); Mayr, Birds Southwest Pacific, 1945, p. 289 (Ponapé).

Geographic range.—Micronesia: Caroline Islands—Ponapé.

Characters.—Adult: Resembles D. o. teraokai, but darker. Resembles. 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. townsendi and D. o. oceanica except in size.

Measurements.—Measurements are listed in [table 27].

Specimens examined.—Total number 21 (11 males, 9 females, 1 unsexed), as follows: Caroline Islands, USNM—Ponapé, 2 (Feb. 11, 12); AMNH—Ponapé, 19 (Nov. 22, 29, Dec. 1, 2, 3).

Nesting.—Coultas (field notes) writes that the pigeon at Ponapé 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.—Bequaert (1939:81, 82 and 1941:266, 290) found the flies (Hippoboscidae), Ornithoctona plicata and O. pusilla, on pigeons from Ponapé.

Remarks.—Coultas (field notes) writes that in 1930 several Japanese made a livelihood as professional hunters of pigeons at Ponapé. He notes, "Two or three years ago, 4 or 5 Japanese, each, averaged from 75 to 100 birds per day, which they sold to the inhabitants for 35 sen (17-1/2 cents) per bird.... Now these same hunters are fortunate if they obtain 4 or 5 Ducula 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 further records in his notes that the hunters used calls to attract the pigeons. In 1930, Coultas regarded the pigeon at Ponapé as a rapidly 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 Ponapé, 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 intensively by the Japanese hunting parties than ever before. Richards obtained two specimens at Ponapé in the period from August, 1947, to January, 1948.

Ducula oceanica oceanica (Lesson and Garnot)

Micronesian Pigeon

Columba oceanica Lesson and Garnot, Dict. Sci. Nat., éd., Levrault, 40, 1826, p. 316. (Type locality, Oualan = Kusaie.)

Columba oceanica Lesson (part), Voy. "La Coquille," Zool.; Atlas, 1826, pl. 41; vol. 2, 1828, pp. 432, 708 (Oualan or Strong); idem, (part), Man. d'Ornith., 11, 1828, p. 166 (Oualan); Kittlitz (part), Kupfertaf. Naturgesch. Vögel, 3, 1833, p. 25, pl. 23, fig. 1 (Ualan); idem (part), Observ. Zool., in Lutké, Voy. "Le Séniavine," 3, 1836, p. 284 (Ualan); Lesson, Compl. de Buffon, 2d ed., 2, Oiseaux, 1839, p. 292 (Oualan); Prévost and Knip (part), Les Pigeons, 2, 1838-43, p. 47, pl. 24 (Oualan); Bonaparte, Comptes Rendus Acad. Sci. Paris, 39, 1854, p. 1072 (Oualan); Kittlitz, Denkw. Reise russ. Amer. Micron. und Kamchat., 1, 1858, pp. 39, 49, 62 (Ualan).

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-Ginnasio 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, Jaluit); 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. Zoöl., 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); Takatsukasa 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 Bernice 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 Islands—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. townsendi, 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 (Hippoboscidae) 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 Marshalls at Jaluit. Takatsukasa 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 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."

Ducula 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, Voy. "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 Momiyama, Birds Micronesia, 1922, p. 55 (Arno).

Globicera oceanica ratakensis Hand-list Japanese Birds, rev., 1932, p. 190 (Arhno, 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.—Takatsukasa and Yamashina (1932:221) describe this subspecies 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 Zoölogy, 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) described this bird as new on the basis of an examination of nine skins taken at Arhno and Wotje.

Evolutionary 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 D. oceanica within a superspecies containing D. pacifica (Melanesia to Samoa and Cook Islands), D. aurorea (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 exploration; Amadon (1943:11) tentatively refers these to D. o. oceanica.

The irregular distribution of D. oceanica in the islands of Micronesia 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 undifferentiated over most of its extensive range. The pigeon of Micronesia 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 isolation. 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 differs, 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 northwest of the Marianas, but none has become established in the Marianas themselves.

Streptopelia bitorquata dusumieri (Temminck)

Philippine Turtle Dove

Columba dusumieri Temminck, Pl. 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. 266, sp. 99 (Marianis).

Columba Dussumieri Kittlitz, Obser. Zool., in Lutké, Voy. "Le Séniavine," 3, 1836, p. 305 (Guahan).

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).

Turtur (Streptopelia) Giamardi Gray, Cat. Birds Trop. Is. Pacific Ocean, 1859, p. 43 (Guam).

Turtur gaimardi Gray, Hand-list Birds, 2, 1870, p. 239 (Marian).

Turtur 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); Oustalet, 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); Safford, Osprey, 1902, p. 68 (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, Zeitschr. f. Naturwisch., 82, 1912, p. 466 (Marianen); Prowazek, Die deutschen Marianen, 1913, p. 101 (Marianen); Reichenow, Die Vögel, 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 Australasianarum, 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, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 60 (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 "vinaceous 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 whitish; 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); tarsus, 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.

Specimens examined.—Total number, 27 (16 males, 11 females), as follows: Mariana Islands, USNM—Guam, 21 (Feb. 7, May 25, 26, 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 unhatched egg was found near Mount Santa Rosa. The nest was situated approximately 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 (Rusa) 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 observed feeding on sandy beaches and tidal flats 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 remarks 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 Streptopelia 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.

Gallicolumba canifrons (Hartlaub and Finsch)

Palau Ground Dove

Phlegoenas canifrons 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, pl. 5, fig. 1 (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. 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).

Phlogoenas canifrons Sclater, Proc. Zool. Soc. London, 1877, p. 112 (Pelew); Salvadori, 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. Coll., vol. 107, no. 15, 1948, p. 62 (Garakayo, Peleliu).

Geographic range.—Micronesia: Palau Islands—Babelthuap, Koror, Garakayo, 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 habits.—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).

G. canifrons apparently came to the Palaus 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. beccarii. 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 discover 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 population 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); Garakayo—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).

Gallicolumba xanthonura xanthonura (Temminck)

White-throated Ground Dove

Columba xanthonura Temminck, Pl. col., livr. 32, 1823, pl. 190. (Type locality, Mariannes.)

Columba xanthonura Lesson, Compl. de Buffon, 2nd ed., 2, Oiseaux, 1838, p. 281 (Mariannes).

Columba Pampusan Quoy and Gaimard, Voy. "Uranie," Zool., 1824, pp. 121, 681, pl. 30 (Mariannes); Dumont, Dict. Sci. Nat., ed. Levrault, 40, 1826, p. 345 (Guam); Lesson, Traité d'Ornith., 1831, p. 471 (Mariannes); Hartlaub, Journ. f. Ornith., 1854, p. 167 (Mariannen).

Columba erythroptera Lesson, Traité d'Ornith., 1831, p. 471 (Mariannes); Kittlitz, Obser. Zool., in Lutké, Voy. "Le Séniavine," 3, 1836, p. 305 (Guahan); Hartlaub, Journ. f. Ornith., 1854, p. 167 (Mariannen).

Columba xanthura Prévost and Knip, Les Pigeons, 2, 1838-43, p. 45, pl. 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 (Type locality, Uap); Gräffe, 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. Soc. 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 Salvadori, 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 (Marianne).

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 World, 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 Vögel, 1, 1913, p. 331 (Mariannen); Bryan, 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 (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).

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.

Geographic range.—Micronesia: Mariana Islands—Asuncion, Pagan, Almagan, 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; primaries, under wing-coverts and axillaries brown; tail, lower breast and rest of underparts dark brown; bill and feet dark brown.

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; 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. Measurements of Gallicolumba xanthonura

Subspecies	Number	Wing	Tail	Culmen	Tarsus
G. x. xanthonura	43 males	146	102	22.0	32
		(139-153)	(97-111)	(21.0-23.0)	(31-33)
	31 females	136	94	20.5	30
		(131-141)	(90-98)	(20.0-21.5)	(28-32)
G. x. kubaryi	7 males	157		23.0	35
		(152-160)		(20.5-23.5)	(33-35)
	7 females	148		23.0	33
		(145-151)		(22.5-23.5)	(32-34)

Table 28. Measurements of Gallicolumba xanthonura

Table 28. Measurements of Gallicolumba xanthonura

Subspecies	Number	Wing	Tail	Culmen	Tarsus
G. x. xanthonura	43 males	146	102	22.0	32
		(139-153)	(97-111)	(21.0-23.0)	(31-33)
	31 females	136	94	20.5	30
		(131-141)	(90-98)	(20.0-21.5)	(28-32)
G. x. kubaryi	7 males	157		23.0	35
		(152-160)		(20.5-23.5)	(33-35)
	7 females	148		23.0	33
		(145-151)		(22.5-23.5)	(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 January. Nests were observed 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 observed 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 with enlarged gonads were taken in April, May, June, and July. Marche, according 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. Seale (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 conspicuous bird and was seen only occasionally in flight. Neither sex appeared to have the secretive, terrestrial habits of G. canifrons 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; Seale (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 weird uncanny 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." Seale 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 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 published regarding the status of this dove in the northern Marianas.

Gallicolumba xanthonura kubaryi (Finsch)

White-throated Ground Dove

Phlegoenas Kubaryi Finsch, Journ. f. Ornith., 1880, p. 292. (Type locality, Ruck and Ponapé.)

Phlegoenas erythroptera Bonaparte, Consp. Avium, 2, 1854, p. 89 (Carolines); Reichenbach, Tauben, 1862, p. 41 (Carolines); Finsch, Proc. Zool. Soc. London, 1877 (1878), p. 780 (Ponapé); idem, Ibis, 1881, p. 115 (Ponapé); Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, pp. 281, 353 (Ponapé, Ruk); Tristram, Cat. Birds, 1889, p. 41 (Ruk).

Phlegoenas kubaryi Reichenow and Schalow, Journ. f. Ornith., 1881, p. 75 (Ruk, Ponapé); Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 55 (Ruk, Ponapé); Hartert, Novit. Zool., 7, 1900, p. 8 (Ruk, Ponapé); Matschie, Journ. f. Ornith., 1901, p. 113 (Ruck, Ponapé); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 53 (Ruk, Ponapé).

Phlogoenas erythroptera Finsch, Proc. Zool. Soc. London, 1880, p. 576 (Ponapé, Ruk); Takatsukasa and Kuroda, Tori, 1, 1915, p. 52 (Ruk).

Phlogoenas kubaryi Salvadori, Cat. Birds British Mus., 21, 1893, p. 599 (Ruk, Ponapé); Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, p. 227 (Caroline = Truk); Bolau, Mitteil. Naturhist. Mus. Hamburg, 1898, p. 68 (Ruck); Reichenow, Die Vögel, 1, 1913, p. 331 (Karolinen).

Phlegaenas kubaryi Christian, The Caroline Islands, 1899, p. 357 (Ponapé).

Gallicolumba kubaryi Mathews, Syst. Avium Australasianarum, 1, 1927, p. 74 (Caroline Is.); Hand-list Japanese Birds, rev., 1932, p. 189 (Truk, Ponapé); Peters, Check-list Birds World, 3, 1947, p. 136 (Ruk, Ponapé); Mayr, Proc. 6th Pacific Sci. Congr., 4, 1941, p. 204 (Ponapé); Bequaert, Mushi, 12, 1939, p. 81 (Ponapé); idem, Occ. Papers Bernice P. Bishop Mus., 16, 1941, p. 266 (Ponapé); Hand-list Japanese Birds, 3d ed., 1942, p. 211 (Truk, Ponapé).

Gallicolumba xanthonura kubaryi Mayr, Birds Southwest Pacific, 1945, p. 290 (Truk, Ponapé); Baker, Smithson. Misc. Coll., vol. 107, no. 15, 1948, p. 62 (Truk).

Geographic range.—Micronesia: Caroline Islands—Truk, Ponapé.

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—Ponapé, 13 (Nov., Dec.)—Truk, 7 (Jan., Feb., May).

Nesting.—At Ponapé in November and December, Coultas obtained specimens 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. Coultas reports that one egg is laid by C. x. kubaryi.

Food habits.—Coultas (field notes) writes that the bird feeds and lives on the ground at Ponapé. He lists food as small snails, seeds, and worms.

Parasites.—Bequaert (1939:81 and 1941:266) records the fly (Hippoboscidae), Ornithoctona plicata, from the ground dove at Ponapé.

Remarks.—Coultas (field notes) writes that in 1930 the ground dove at Ponapé was rare in the forested areas and generally found more along the sea coast and in the upland valleys. Coultas describes its call as an infrequent shrill, whistle-like call. He writes that hunting by the Japanese and natives was reducing the population of G. x. kubaryi at Ponapé 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 Ponapé 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 Gallicolumba. One invasion established G. canifrons at the Palau Islands. 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, resemble 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 relationship 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 coloring present on the head. The male and female of G. x. kubaryi 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 Ponapé by Coultas.

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 reduction of malelike plumage in the female. G. rubescens of the Marquesas 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 superspecies 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 establishing G. erythroptera in the Society and Tuamotu islands, although 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 genera 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 (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); Wiglesworth, 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 Vögel, 1, 1913, p. 328 (Palauinseln); Takatsukasa and Kuroda, Tori, 1, 1915, p. 52 (Pelew).

Caloenas nicobaricus pelewensis Kuroda, in Momiyama, Birds Micronesia, 1922, p. 53 (Pelew).

Geographic range.—Micronesia: Palau Islands-Babelthuap, Koror, Garakayo.

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 distributed 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 History 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 individuals, which I saw at Garakayo, were either perched on scrubby vegetation on high and inaccessible cliffs or were flying high overhead. In its flight overhead, the short, white tail was a particularly conspicuous mark of identification. The flight reminded me very much of that of the Black Vulture (Córagyps atrátus) 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 road to extinction unless governmental protection can be established and enforced.

Trichoglossus rubiginosus (Bonaparte)

Ponapé 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 = Ponapé.)

Chalcopsitta rubiginosus Bonaparte, Proc. Zool. Soc. London, 1850, p. 26, pl. 16 ("Ins. Barabay et Guebe," error = Ponapé); Pelzeln, Reise "Novara," Vögel, 1865, pp. 99, 162 (Puynipet); Reichenow, Journ. f. Ornith., 1881, p. 162 ("Nordwestl. Polynessische subregion Carolinen" = Ponapé); Tristram, Cat. Birds, 1889, p. 73 (Ponapé); Finsch, Deut. Verein zum Schultze der Vogelwelt, 18, 1893, p. 458 (Carolinen = Ponapé); Matschie, Journ. f. Ornith., 1901, p. 112 (Ponapé).

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).

Lorius rubiginosa Giebel, Thes. Ornith., 2, 1875, p. 502 (Senjawin = Ponapé).

Trichoglossus rubiginosus Finsch, Journ. Mus. Godeffroy, 12, 1876, pp. 17, 18 (Ponapé); idem, Proc. Zool. Soc. London, 1877 (1878), p. 778 (Ponapé); idem, Journ. f. Ornith., 1880, p. 284 (Ponapé); Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, p. 281 (Ponapé); Finsch, Ibis, 1881, pp. 110, 111, 114 (Ponapé); idem, Mitth. Ornith. Ver. Wien, 1884, p. 49 (Ponapé); Hartert, Kat. Vogelsamml. Senckenb., 1891, p. 161 (Puypinet); Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6 1890-1891 (1891), p. 8 (Ponapé); Peters, Check-list Birds World, 3, 1937, p. 151 (Ponapé); Mayr, Proc. Sixth Pac. Sci. Congr., 4, 1941, p. 204 (Ponapé); Hand-list Japanese Birds, 3d ed., 1942, p. 201 (Ponapé); Mayr, Birds Southwest Pacific, 1945, p. 291 (Ponapé).

Eos rubiginosa Salvadori, Ornith. Papuasia, 1, 1880, p. 267 (Puynipet); idem, Cat. Birds British Mus., 20, 1891, p. 29 (Ponapé); Christian, The Caroline Islands, 1899, p. 357 (Ponapé); Finsch, Notes Leyden Mus., 22, 1900, p. 142 (Ponapé); Dubois, Syn. Avium, 1902, p. 29 (Puinipet); Uchida, Annot. Zool. Japon., 9, 1918, pp. 484, 493 (Ponapé); Wetmore, in Townsend and Wetmore, Bull. Mus. Comp. Zoöl., 63, 1919, p. 192 (Ponapé).

Chalcopsittacus rubiginosus Finsch, Sammlung wissensch. Vorträge, 14th Ser., 1900, p. 639 (Ponapé).

Oenopsittacus rubiginosus Reichenow, Die Vögel, 1, 1913, p. 443 (Karolinen = Ponapé); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 58 (Ponapé); Mathews, Syst. Avium Australasianarum, 1, 1927, p. 295 (Ponapé); Hand-list Japanese Birds, rev., 1932, p. 181 (Ponapé).

Eos rubiginosus Takastukasa and Kuroda, Tori, 1, 1915, p. 53 (Ponapé).

Geographic range.—Micronesia: Caroline Islands—Ponapé.

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 toward tip; wings black with outer webs olivaceous-green; outer edges of primaries 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 grayish-white.

Immature: Resembles adult, but with narrow and more sharply pointed tail feathers.

Measurements.—Measurements are presented in [table 29].

Table 29. Measurements of Trichoglossus rubiginosus

Sex	No.	Wing	Tail	Culmenybr />from cere	Tarsus
Adult males	18	147	105	20	16
		(143-153)	(100-110)	(19-20)	(15-17)
Adult females	13	142	101	19	16
		(141-146)	(98-104)	(18-19)	(15-17)

Specimens examined.—Total number, 31 (18 males, 13 females), as follows: Caroline Islands, USNM—Ponapé, 2 (Feb. 12); AMNH—Ponapé, 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 Ponapé 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.—Uchida (1918:484, 493) found the bird lice (Mallophaga), Psittaconirmus harrisoni and Eomenopon denticulatus, on the Ponapé Lory.

Remarks.—There is little written information concerning the habits of the Ponapé 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 species; some of these unpublished notes are essentially as follows: Trichoglossus is common on Ponapé. 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 intelligent, easily tamed, and sometimes learns to repeat a few words.

Evolutionary history of Trichoglossus rubiginosus.—The Ponapé Lory is the only native parrot in Micronesia. It is an aberrant species and seemingly is of long residence on the island, as indicated by its differences from related forms to the southward and southwestward. 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 Ponapé 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 feathers 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 web of the wing, and feathers of the upper back are edged with dark coloring. T. rubiginosus may have been derived from either of these two species; however, it shows a close relationship also to the T. haematodus group from the Papuan region. In any case, the Ponapé Lory, isolated in Micronesia, has not the multicolored plumage of its relatives 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 Ponapé. Like Aplonis pelzelni, another endemic species at Ponapé, 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, Journ. f. Ornith., 1863, p. 332. (Type locality, Japan.)

Cuculus canorus Hartlaub and Finsch, Proc. Zool. Soc. London, 1872, pp. 89, 100 (Pelew); Finsch, Journ. 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 locality not given.

Remarks.—The Common Cuckoo is a straggler on winter migration to the Palau Islands.

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. Soc. London, 1872, pp. 89, 100 (Pelew); Finsch. Journ. 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 (Babelthuap, 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 and 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 taitensis Sparrman, Mus. Carls., fasc, 2, 1787, pl. 32. (No type locality = Tahiti.)

Eudynamis tahitiensis Gräffe, Journ. Mus. Godeffroy, 2, 1873, p. 123 (Yap).

Eudynamis taitiensis Finsch, Journ. Mus. Godeffroy, 8, 1875, p. 49 (Palau); idem, Journ. Mus. Godeffroy, 12, 1876, pp. 17, 20 (Ponapé); idem, Proc. Zool. Soc. London, 1877 (1878), p. 778 (Ponapé); idem, Journ. f. Ornith., 1880, pp. 284, 298 (Ponapé, Kuschai, Palaos, Marshalls); idem, Ibis, 1880, pp. 331, 332 (Taluit); idem, Ibis, 1881, pp. 104, 108, 113, 114 (Kushai, Uleai, Ponapé); Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, pp. 281, 299, 353 (Ponapé, Mortlock, Ruk); Christian, The Caroline Islands, 1899, p. 358 (Ponapé).

Urodynamis taitensis Finsch, Mitth. Ornith. Ver. Wien, 1884, p. 53 (Jaluit, Ponapé, Palau); Bogert, Amer. Mus. Novit., no. 933, 1937, p. 9 (Palau, Ruk, Kusaie, Ponapé, 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, Ponapé, Kusaie, Jaluit, Elmore, Aurh, Wotze).

Urodynamis taitiensis Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 11 (Pelew, Ualan, Ponapé, Luganor, Taluit); idem, Ibis, 1893, p. 212 (Marshalls); Hartert, Novit. Zool., 7, 1900, p. 7 (Ruk); Finsch, Notes Leyden Mus., 22, 1900, p. 120 (Ponapé, 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, Ponapé, Luganor, Ruk, Taluit); Hand-list Japanese Birds, rev., 1932, p. 180 (Palau, Kusaie, Ponapé, Luganor, Truk, Jaluit, Elmore, Aurh, Wotze).

Urdynamis taitiensis Finsch, Sammulung wissensch. Vorträge, 14th ser., 1900, p. 659 (Palau).

Eudynamis taitiensis Schnee, Zool. Jahrbücher, 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 Islands—exact locality unknown; Caroline Islands—Yap, Lukunor, Truk, Ponapé, 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 Zealand 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 Marshall and Caroline islands (see map in Bogert, 1937:3-4). There are no records 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) remarks that the cuckoo arrives at New Zealand for the breeding period 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 consequence 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 competition from resident birds and from other migrants for feed and habitat. 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, pl. 1, fig. 1 and 2 (Palau); Giebel, Thes. Ornith., 2, 1875, p. 720 (Pelew); Schmeltz and Krause, Ethnogr. 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).

Scops podargina Sharpe, Cat. Birds British Mus., 2, 1875, p. 313 (Palau); Nehrkorn, Journ. f. Ornith., 1879, p. 394 (Palau); Wiglesworth, Abhandl. und Ber. Zool. Dresden, no. 6, 1890-1891 (1891), p. 3 (Pelew); Matschie, Journ. f. Ornith., 1901, p. 112 (Palau); Dubois, Syn. Avium, 2, 1904, p. 883 (Pelew).

P[isorhina] podargina Reichenow, Die Vögel, 1913, p. 424 (Palau).

Otus podarginus Mathews, Syst. Avium Australasianarum, 1, 1927, p. 268 (Palau); Mayr. Amer. Mus. Novit., no. 1269, 1944, p. 3 (Palau); idem, Birds Southwest Pacific, 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 superciliary 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 buffy-white; lores rufous, shafts white; indistinct eye ring rufous; ear-coverts whitish with rufous tips, chin white; throat white narrowly 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, 83, 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.—Coultas (field notes) found the 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 mangrove 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, Pyrroglaux. Mayr (1944b:3) has reviewed this treatment and presents 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 O. spilocephalus, and that the characters possessed by O. podarginus are no more different or unusual 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 O. spilocephalus of Asia and Malaysia.

Asio flammeus flammeus (Pontoppidan)

Short-eared Owl

Strix Flammea Pontoppidan, Danske, Atlas, 1, 1763, p. 617, pl. 25. (Type 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).

Table 29. Measurements of Trichoglossus rubiginosus

Table 29. Measurements of Trichoglossus rubiginosus

Sex	No.	Wing	Tail	Culmenybr />from cere	Tarsus
Adult males	18	147	105	20	16
		(143-153)	(100-110)	(19-20)	(15-17)
Adult females	13	142	101	19	16
		(141-146)	(98-104)	(18-19)	(15-17)

Otus brachyotus Hartlaub, Journ. f. Ornith., 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 (Marianne); Seale, Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 44 (Mariannes); Safford, Osprey, 1902, p. 68 (Marianas); idem, Amer. 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 flammeus 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 A. f. flammeus, a migrant from Asia; possibly, however, there is an unrecorded resident population of the Short-eared Owl in the northern Marianas, which may be closely related to A. f. ponapensis of Ponapé. 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 overgrazing and burning of the grassy habitats preferred by the owl.

Asio flammeus ponapensis Mayr

Short-eared Owl

Asio flammeus ponapensis Mayr, Amer. Mus. Novit., no. 609, 1933, p. 1. (Type locality, Ponapé.)

Otus brachyotus Finsch (part), Journ. Mus. Godeffroy, 12, 1876, pp. 17, 18 (Ponapé); idem, Proc. Zool. Soc. London, 1877 (1878), p. 778 (Ponapé); idem, Journ. f. Ornith., 1880, p. 283 (Ponapé); idem, Mitth. Ornith. Ver. Wien, 1884, p. 47 (Ponapé); idem, Sammlung wissensch. Vorträge, 14 ser., 1900, p. 659 (Ponapé).

Asio brachyotus Finsch, Ibis, 1881, pp. 113, 114 (Ponapé).

Asio accipitrinus Ridgway, Proc. U. S. Nat. Mus., 4, 1882, p. 367 (Strong's Island = Kusaie); Wiglesworth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 3 (Ponapé); 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 (Ponapé); Hand-list Japanese Birds (part), rev., 1932, p. 182 (Ponapé).

Asio flammeus ponapensis Kelso, Oölogist, 1938, p. 183 (Kusaie); Peters, Check-list Birds World, 4, 1940, p. 170 (Ponapé); Hand-list Japanese Birds (part), 3d ed., 1942, p. 202 (Ponapé); Mayr, Birds Southwest Pacific, 1945, p. 291 (Ponapé).

Geographic range.—Micronesia: Caroline Islands—Ponapé, 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; forehead 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; auxilaries buffy; feathering of tibia and tarsus pale buff; bill dark slate; feet grey-brown; iris yellow.

Resembles A. f. flammeus, 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—Ponapé (Dec.).

Nesting.—Coultas (field notes) writes that the Short-eared Owl at Ponapé 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 Ponapé 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 Ponapé and apparently has somewhat the same habits as other members of the species. He estimated the 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 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 measurements 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 Ponapé closely resembles A. f. flammeus but is slightly smaller and darker. Apparently the owl came to Ponapé as a straggler on migration from Asia, and becoming acclimated and adapted to the grassy areas at Ponapé remained as a resident. The occurrence of A. f. flammeus in the Marianas on migration offers evidence as to how the bird originally reached Ponapé.

Caprimulgus indicus jotaka Temminck and Schlegel

Jungle Nightjar

Caprimulgus jotaka Temminck and Schlegel, in Siebold's Fauna Japonica, Aves, 1847, p. 37, pl. 12, 13. (Type locality, Japan.)

Caprimulgus 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 December 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

Caprimulgus phalaena Hartlaub and Finsch, Proc. Zool. Soc. London, 1872, p. 91. (Type locality, Pelew.)

Caprimulgus phalaena Finsch, Journ. Mus. Godeffroy, 8, 1875, pp. 4, 13, pl. 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, Syn. Avium 1, 1902, p. 124 (Pelew); Reichenow, Die Vögel, 2, 1914, p. 154 (Palau); Mathews, Syst. Avium. Australasianarum, 1, 1927, p. 396 (Pelew); Hachisuka, Birds Philippines, 2, 1934, p. 120 (Pelew).

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, Garakayo.

Characters.—Adult male: "Above grayish-brown, very finely vermiculated, more rufous on the back, with large longitudinal streaks and a few cross markings; 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 buff 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.

Measurements.—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, 8 (4 males, 4 females), as follows: Palau Islands, USNM—Koror, 3 (Nov. 3, 20, 29); AMNH—exact locality not given, 5 (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 (field 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 species which reached the Palaus without expanding their ranges farther 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. Soc. 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. 111 (Pelew); Wiglesworth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 18 (Pelew); Matschie (part), Journ. f. Ornith., 1901, p. 112 (Palau).

Collocalia vanikorensis Gray, Hand-list Birds, 1, 1869, p. 66 (Pelew); Giebel, Thes. Ornith., 1, 1872, p. 737 (Pelew).

Collocalia fuciphaga Oustalet (part), Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, p. 189 (Palaos); Reichenow, Die Vögel, 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, Garakayo, Peleliu, Angaur.

Characters.—Adult, 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 glossy green; white loral spot inconspicuous."

Measurements.—Measurements are listed in [table 30].

Table 30. Measurements of Collocalia inexpectata in Micronesia

Subspecies	No.	Wing	Tail
C. i. pelewensis	14	111 (109-113)	50 (47-51)
C. i. bartschi	13	108 (105-108)	54 (52-57)

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 found the swiftlet to be numerous on islands in the southern Palaus in 1945. The birds were observed 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 Lutké., Voy. "Le Séniavine," 3, 1836, p. 304 (Guahan); idem (part), Denkw. Reise russ. Amer. Micron. und Kamchat., 2, 1858, p. 26 (Guahan).

Collocalia nidifica Gray (part), Ann. Mag. Nat. Hist., (3), 17, 1866, p. 125 (Marianne); idem (part), Hand-list Birds, 1, 1869, p. 65 (Marianne).

Collocalia vanicorensis Finsch (part), Journ. Mus. Godeffroy, 12, 1876, p. 24 (Marianen); 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 Bernice 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 (Marianne).

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.

Characters.—Resembles C. i. pelewensis, 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 subspecies 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 islands 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 which 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 subspecies 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 rearrange these species and subspecies, it appears to me that the Micronesian swiftlets fall into the two natural groups (C. inexpectata 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 concentrated at cliff areas, flying about in large groups. Away from the cliffs fewer were seen and singles were observed in woodland openings, over fields, and in the coconut groves. On May 18, 1945, a colony of nesting birds was found approximately two miles east of Agaña 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, sheltered 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.

Collocalia inquieta inquieta (Kittlitz)

Carolines Swiftlet

Cypselus inquietus Kittlitz, Obser. Zool., in Lutké, Voy. "Le Séniavine," 3, 1836, p. 285. (Type locality, Ualan.)

Cypselus inquietus Kittlitz (part), Denkw. Reise russ. Amer. Micron. und Kamchat., 2, 1858, p. 26 (Ualan).

Collocalia ualensis Streubel, Isis, 1848, p. 368 (no type locality = Kusaie?).

Collocalia nidifica ualensis Gray, Ann. Nat. Hist., 17, 1866, p. 123 (Caroline Islands = Kusaie?).

Collocalia vanicorensis Finsch (part), Journ. Mus. Godeffroy, 12, 1876, p. 24 (Ualan); idem (part), Proc. Zool. Soc. London, 1880, p. 575 (Kuschai); idem (part), Journ. f. Ornith., 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), Journ. f. Ornith., 1901, p. 112 (Ualan).

Collocalia 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).

Collocalia fuciphaga fuciphaga Oberholser (part), Proc. Acad. Nat. Sci. Phila., 1906, p. 186 (Ualan).

Collocalia fuciphaga vanikorensis Oberholser (part), Proc. U. S. Nat. Mus., 42, 1912, p. 20 (Kusaie).

Collocalia fuciphaga inquieta Stresemann, Verhandl. Ornith. 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).

Collocalia 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).

Collocalia 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.—Kittlitz obtained this swiftlet when he visited Kusaie from December 8, 1827, to January 1, 1828. In 1931, Coultas found the bird common at Kusaie. The name Collocalia ualensis, published 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 Caroline Islands.

Table 31. Measurements of Collocalia inquieta

Subspecies	No.	Wing
Collocalia i. inquieta	11	119 (116-125)
Collocalia i. ponapensis	10	110 (107-114)
Collocalia i. rukensis	10	(112-119.5)[C]

^[C] (Mayr, 1935:3).

Collocalia inquieta rukensis Kuroda

Carolines Swiftlet

Collocalia fuciphaga rukensis Kuroda, Tori, 1, 1915, pp. 58, 59, pl. 3, fig. 1. (Type locality, Ruk.)

Collocalia vanicorensis Finsch (part), Proc. Zool. London, 1880, p. 575 (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. 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 Australasianarum, 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 Mount Tonáchian 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, Ponapé.)

Collocalia vanicorensis Finsch (part), Journ. Mus. Godeffroy, 12, 1876, pp. 17, 23 (Ponapé); idem, Proc. Zool. Soc. London, 1877 (1878), p. 778 (Ponapé); idem (part), Journ. f. Ornith., 1880, p. 285 (Ponapé); idem, Ibis, 1881, p. 115 (Ponapé); Wiglesworth (part), Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 18 (Ponapé); Matschie (part), Journ. f. Ornith., 1901, p. 112 (Ponapé).

Collocalia fuciphaga Hartert, Cat. Birds British Mus., 16, 1892, p. 498 (Ponapé).

Collocalia fuciphaga vanikorensis Takatsukasa and Kuroda, Tori, 1, 1915, p. 53 (Ponapé).

Collocalia fuciphaga inquieta Kuroda (part), in Momiyama, Birds Micronesia, 1922, p. 62 (Ponapé).

Collocalia vanikorensis ponapensis Mayr, Amer. Mus. Novit., no. 828, 1936, p. 12 (Ponapé); Hand-list Japanese Birds, 3d ed., 1942, p. 198 (Ponapé).

Collocalia inquieta ponapensis Mayr, Amer. Novit., no. 915, 1937, p. 11 (Ponapé); Peters, Check-list Birds World, 4, 1940, p. 225 (Ponapé); Mayr, Birds Southwest Pacific, 1945, p. 292 (Ponapé).

Collocalia inquieta Mayr, Proc. 6th Pac. Sci. Congr., 4, 1941, p. 204 (Ponapé).

Geographic range.—Micronesia: Caroline Islands—Ponapé.

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 examined.—Total number, 37 (19 males, 18 females) from Caroline Islands, AMNH—Ponapé (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. vanikorensis in Northern Melanesia. The birds found in New Guinea and the Solomons are similar in size to the birds in the Carolines, while those in the Moluccas, Admiralties and Lihir are larger. Color differences 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.

Halcyon cinnamomina cinnamomina Swainson

Micronesian Kingfisher

Halcyon cinnamomina Swainson, Zool. Illustr., 2, 1821, text to pl. 67. (No type locality = Guam.)

Halcyon cinnamomina Hartlaub, Journ. f. Ornith., 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, pl. 80 (Guam); Gray, Hand-list Birds, 1, 1869, p. 93 (Mariannes = Guam); Oustalet, Le Nat., 1889, p. 260 (Mariannes = 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, Journ. 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 ruficeps Dumont, Dict. Sci. Nat., 29, 1823, p. 273 (Mariannes = Guam); Pucheran, Rev. et Mag. de Zool., 1853, p. 387 (Mariannes = Guam); Hartlaub, Journ. f. Ornith., 1855, p. 423 (Mariannen = Guam).

Dacela ruficeps Lesson, Traité d'Ornith., 1831, p. 247 (Mariannes = Guam).

Halcyon cinnamomeus Kittlitz, Obser. Zool., in Lutké, Voy. "Le Séniavine," 3, 1836, p. 304 (Guahan).

Dacelo cinnamomina Kittlitz, Denkw. Reise russ. Amer. Micron. und Kamchat., 2, 1858, p. 131 (Guahan); 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 cinnamominus 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 Vögel, 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).

Table 30. Measurements of Collocalia inexpectata in Micronesia

Table 30. Measurements of Collocalia inexpectata in Micronesia

Subspecies	No.	Wing	Tail
C. i. pelewensis	14	111 (109-113)	50 (47-51)
C. i. bartschi	13	108 (105-108)	54 (52-57)

Table 31. Measurements of Collocalia inquieta

Table 31. Measurements of Collocalia inquieta

Subspecies	No.	Wing
Collocalia i. inquieta	11	119 (116-125)
Collocalia i. ponapensis	10	110 (107-114)
Collocalia i. rukensis	10	(112-119.5)[C]

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 Southwest 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 surface 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; 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 culmen	Tarsus
H. c. cinnamomina	31 males	102 (96-105)	77 (73-83)	37 (35-39)	15 (14-17)
	25 females	102 (99-106)	79 (74-84)	38 (35-38)	15 (14-17)
H. c. pelewensis	5 males	89 (88-89)	61 (58-64)	39 (38-40)	14 (13-14)
	4 females	88 (88-89)	64 (61-67)	39 (38-39)	14 (13-14)
H. c. reichenbachii	14 males	99 (95-101)	74 (72-77)	41 (39-43)	16 (16-17)
	15 females	100 (96-102)	74 (71-76)	41 (39-42)	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 Islands, 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 nestling 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 irregular 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 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 à têterouse." Kittlitz recorded the bird in March, 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 frequenting 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, pl. 30 (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 Vögel, 2, 1914, p. 116 (Palau).

Halcyon cinnamominus Finsch (part), Journ. Mus. Godeffroy, 12, 1876, pp. 17, 20 (Palau).

Sauropatis cinnamomina Salvadori (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).

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 (Palau); 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. Coultas (field notes) writes that foods of this bird consist of grubs and ants.

Parasites.—Uchida (1918:483) found the bird louse (Mallophaga), Docophorus alatoclypeatus, 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. McElroy of the NAMRU2 party saw a kingfisher with cinnamon underparts 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, Ponapé.)

Halcyon cinnamominus Finsch (part), Journ. Mus. Godeffroy, 12, 1876, pp. 17, 19 (Ponapé); idem, Proc. Zool. Soc. London, 1877 (1878), p. 778 (Ponapé); idem, Journ. f. Ornith., 1880, p. 285 (Ponapé); idem, Journ. f. Ornith., 1880, p. 285 (Ponapé); idem, Mitth. Ornith. Ver. Wien, 1884, p. 47 (Ponapé).

Sauropatis cinnamomina Salvadori (part), Ornith. Papuasia, 1, 1880, p. 481 (Ponapé).

Halcyon cinnamomina Finsch, Ibis, 1881, pp. 112, 114 (Ponapé); Tristram (part), Cat. Birds, 1889, p. 92 (Ponapé).

Halcyon mediocris Sharpe, Cat. Birds British Mus., 17, 1892, p. 260 (Type locality, Ponapé); Wiglesworth, Abhandl. and Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 16 (Ponapé); Oustalet, Nouv. Arch. Mus. Hist. Nat. Paris, (3), 7, 1895, pp. 177, 180, 181, 184, 185, 186 (Ponapi); Reichenow, Die Vögel, 2, 1914, p. 116 (Ponapé).

Halcyon reichenbachi Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 15 (Ponapé); 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 (Ponapé); Matschie, Journ. f. Ornith., 1901, pp. 112, 113 (Ponapé); Mayr, Proc. 6th Pacific Sci. Congr., 4, 1941, p. 204 (Ponapé).

Halcyon cinnamominus var. reichenbachi Dubois, Syn. Avium, 1, 1902, p. 108 (Ponapé).

Halcyon cinnamominus var. mediocris Dubois, Syn. Avium, 1, 1902, p. 108 (Ponapé).

Halcyon reichenbachii Takatsukasa and Kuroda, Tori, 1, 1915, p. 53 (Ponapé).

Sauropatis mediocris Wetmore, in Townsend and Wetmore, Bull. Mus. Comp. Zoöl., 63, 1919, p. 195 (Ponapé).

Sauropatis reichenbachii reichenbachii Kuroda, in Momiyama, Birds Micronesia, 1922, p. 60 (Ponapé).

Hyposyma cinnamomina reichenbachii Mathews, Syst. Avium Australasianarum, 1, 1927, p. 384 (Ponapé).

Halycyon cinnamomina reichenbachii Hand-list Japanese Birds, rev., 1932, p. 180 (Ponapé); Hand-list Japanese Birds, 3d ed., 1942, p. 200 (Ponapé); Mayr, Birds Southwest Pacific, 1945, p. 293 (Ponapé); Peters, Check-list Birds World, 5, 1945, p. 206 (Ponapé).

Halcyon cinnamomina reichenbachii Bequaert, Mushi, 12, 1939, p. 82 (Ponapé); idem, Occ. Papers Bernice P. Bishop Mus., 16, 1941, p. 290 (Ponapé).

Geographic range.—Micronesia: Caroline Islands—Ponapé.

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 creamy, 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 Islands, USNM—Ponapé, 1 (Feb. 12); AMNH—Ponapé, 48 (Nov., Dec).

Molt.—Most of the specimens taken by Coultas in November and December are either worn or in molt.

Parasites.—Bequaert (1939:82 and 1941:290) records a fly (Hippoboscidae), Ornithoica pusilla, from the Micronesian Kingfisher at Ponapé.

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 Hartlaub in 1852 for a kingfisher with a white abdomen in the Dresden Museum, which had been figured by Reichenbach (Synopsis Avium, Alcedineae, 1851) and called Todiramphus 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 Ponapé were taken, Hartlaub's bird was found to be identical with them; thus the name H. reichenbachii could be restricted to the bird at Ponapé, and Wiglesworth supplied the new name H. pelewensis for the population at Palau. H. mediocris was used by Sharpe to designate the cinnamon-breasted birds at Ponapé, 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 cinnamomina.—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 Heinroth 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 populations of H. cinnamomina.

In H. c. pelewensis and H. c. reichenbachii, the adult birds resemble 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 Ponapé (eastern Carolines) and spread to Guam and Palau. He states further (1942b:181, 182) that the presence of H. cinnamomina and H. chloris as reproductively isolated groups in the Palaus may not indicate that they are distinct species, but that they represent the overlap 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, pl. 3, fig. 3. (Type locality, Pelew.)

Halcyon albicilla Hartlaub, Proc. Zool. Soc. 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, Journ. 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), Thes. 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, Garakayo, 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.

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 culmen	Tarsus
H. c. teraokai	17	113 (110-116)	76 (72-81)	45 (41-52)	14 (13-16)
H. c. orii	9	111 (109-116)	80 (78-83)	44 (42-45)	16 (15-16)
H. c. albicilla	17	116 (109-119)	81 (78-84)	46 (42-49)	16 (14-17)
H. c. owstoni	3	115 (114-116)	81 (80-82)	44 (42-45)	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, obtain 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), Docophorus alatoclypeatus, from this bird at Palau. Bequaert (1939:82 and 1941:290) lists the fly (Hippoboscidae), Ornithoica 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 overlap that of the secretive and inconspicuous H. cinnamomina pelewensis, 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 numerous 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.

Halcyon chloris orii Takatsukasa and Yamashina

White-collared Kingfisher

Halcyon chloris orii Takatsukasa and Yamashina, Dobutsu. Zasshi, 43, 1931, p. 484. (Type locality, Rota.)

Halcyon albicillus Sharpe (part), Cat. Birds British Mus., 17, 1892, p. 249 (Marianne = 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 albicillus 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 Southwest 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.—Measurements 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, Dict. Sci. Nat., éd. 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).

Halcyon albicilla Hartlaub, Journ. f. Ornith., 1854, p. 167 (Mariannen = Tinian); Gray, Cat. Birds Trop. Is. Pacific Ocean, 1859, p. 5 (Ladrone or Marian Islands = Tinian); Gray (part), Hand-list Birds, 1, 1869, p. 93 (Mariannes = Tinian); Oustalet, 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. Ornith., 1901, pp. 112, 113, 114 (Saipan); Seale, Occ. Papers Bernice P. Bishop Mus., 1, 1901, p. 45 (Saipan).

Dacelo albicilla Giebel (part), Thes. Ornith., 2, 1875, p. 1 (Marianne = Tinian).

Sauropatis albicilla Salvadori, Ornith. Papuasia, 1, 1880, p. 470 (Marianne = Tinian).

Halcyon albicillus Sharpe (part), Cat. Birds British Mus., 17, 1892, p. 249 (Marianne = Saipan).

Halcyon saurophagus Schnee, Zeitschr. f. Naturwisch., 82, 1912, p. 463 (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, Check-list 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 Islands, 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. albicilla closely resembles that in H. s. saurophaga Gould of Melanesia. These two species resemble 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 greenish. 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 locality, Asuncion.)

Halcyon albicillus Sharpe (part), Cat. Birds British Mus., 17, 1892, p. 249 (Marianne = 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).

Sauropatis chloris owstoni Kuroda, in Momiyama, Birds Micronesia, 1922, p. 59 (Asuncion).

Table 32. Measurements of Halcyon cinnamomina

Table 32. Measurements of Halcyon cinnamomina

Subspecies	Number	Wing	Tail	Exposed culmen	Tarsus
H. c. cinnamomina	31 males	102 (96-105)	77 (73-83)	37 (35-39)	15 (14-17)
	25 females	102 (99-106)	79 (74-84)	38 (35-38)	15 (14-17)
H. c. pelewensis	5 males	89 (88-89)	61 (58-64)	39 (38-40)	14 (13-14)
	4 females	88 (88-89)	64 (61-67)	39 (38-39)	14 (13-14)
H. c. reichenbachii	14 males	99 (95-101)	74 (72-77)	41 (39-43)	16 (16-17)
	15 females	100 (96-102)	74 (71-76)	41 (39-42)	16 (15-17)

Table 33. Measurements of Halcyon chloris in Micronesia

Table 33. Measurements of Halcyon chloris in Micronesia

Subspecies	No.	Wing	Tail	Exposed culmen	Tarsus
H. c. teraokai	17	113 (110-116)	76 (72-81)	45 (41-52)	14 (13-16)
H. c. orii	9	111 (109-116)	80 (78-83)	44 (42-45)	16 (15-16)
H. c. albicilla	17	116 (109-119)	81 (78-84)	46 (42-49)	16 (14-17)
H. c. owstoni	3	115 (114-116)	81 (80-82)	44 (42-45)	17 (16-17)

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. 180 (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, Checklist Birds World, 5, 1945, p. 209 (Asuncion, Pagan, Alamagan); Borror, Auk, 64, 1947, p. 417 (Agrigan).

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 buffy 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: Mariana 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 February, 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 Malaysia, 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. 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 structure. The species did not succeed in establishing itself in the Carolines or at Guam, but did so in the Marianas at Rota and northward. In comparison with other subspecies of H. chloris those in the Marianas 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 Micronesia 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 Micronesia via the Palaus and that of H. cinnamomina via Ponapé (eastern 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, Halmahera and Batchian. Mayr (1045a:302) refers this visitor to Palau to E. o. pacificus (Latham); this form is migratory and may fly 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 rustica Gräffe, 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 Pacific, 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, Peleliu, 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 Micronesia 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 Agaña 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, pl. 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); Tristram, 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 (Palau); Dubois, Syn. Avium., 1, 1902, p. 303 (Pelew); Reichenow, Die Vögel, 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 (Pelew).

Edolisoma tenuirostre monacha Stresemann, Ornith. 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).

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, innermost 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. Measurements of Edolisoma tenuirostre in Micronesia

Subspecies	No.	Wing	Tail	Exposed culmen	Tarsus
E. t. monachum	10	98	80	21.0	23.0
		96-103	76-83	20.0-22.5	22.5-24.0
E. t. insperatum	35	109	86	23.0	24.0
		107-112	82-91	22.0-24.0	23.0-25.0

Molt.—Molt in this bird appears to take place in the 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, especially 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. 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.)

Campehaga nesiotis Gräffe, 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); Wiglesworth, 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); Reichenow, Die Vögel, 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 (Mackenzie group); Hand-list Japanese Birds, rev., 1932, p. 174 (Yap); Hand-list Japanese 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).

Geographic 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 Stresemann (1939:126) arranged it as a subspecies of Edolisoma tenuirostre. The type specimen is an immature, and the adult is unknown. The presence of rufous coloring shows a relationship with E. t. insperatum of Ponapé, 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 barring 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.

Edolisoma tenuirostre insperatum (Finsch)

Cicada Bird

Volvocivora inseperata Finsch, Proc. Zool. Soc. London, 1875, (1876), p. 644. (Type locality, Ponapé.)

Volvocivora insperata Finsch, Journ. Mus. Godeffroy, 12, 1876, pp. 17, 27 (Ponapé); idem, Proc. Zool. Soc. London, 1877 (1878), p. 779 (Ponapé); idem, Ibis, 1881, pp. 110, 112, 115 (Ponapé); Schmeltz and Krause, Ethnogr. Abth. Mus. Godeffroy, 1881, p. 281 (Ponapé).

Volvozivora insperata Finsch, Journ. f. Ornith., 1880, p. 289 (Ponapé).

Lalage insperata Sharpe, Cat. Birds British Mus., 4, 1879, p. 108 (Ponapé); Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 25 (Ponapé); Bolau, Mitteil. Naturhist. Mus. Hamburg, 1898, p. 53 (Ponapé); Matschie, Journ. f. Ornith., 1901, pp. 112, 113 (Ponapé); Reichenow, Die Vögel, 2, 1914, p. 276 (Karolinen = Ponapé); Kuroda, in Momiyama, Birds Micronesia, 1922, p. 68 (Ponapé); Hand-list Japanese Birds, rev., 1932, p. 174 (Ponapé); Hand-list Japanese Birds, 3d ed., 1942, p. 194 (Ponapé).

Lisomada insperata Mathews, Novit. Zool., 24, 1928, p. 372 (new generic name); idem, Syst. Avium Australasianarum, 2, 1930, p. 545 (Ponapé).

Edolisoma tenuirostre insperata Stresemann, Ornith. Monatsber., 47, 1939, p. 126 (Ponapé); Mayr, Birds Southwest Pacific, 1945, p. 294 (Ponapé).

Edolisoma tenuirostre Mayr, Proc. 6th Pacific Sci. Congr., 4, 1941, p. 204 (Ponapé).

Geographic range.—Micronesia: Caroline Islands—Ponapé.

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; forehead 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 of 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—Ponapé (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 season had just been completed in November and December (the time of his visit to Ponapé), 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 Ponapé.

Examination of the large series of birds taken by Coultas at Ponapé 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 present two plumages, which, if Mayr's arrangement is followed, may be interpreted as a more primitive or "retarded" type 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 Ponapé 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 relationship between the cicada birds of Micronesia and Edolisoma tenuirostre 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 differences 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 coloring of the primaries and secondaries. E. t. nesiotis may have arrived 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 Ponapé bird in color; however, in size it probably more closely approaches the latter subspecies.

The Ponapé 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 subspecies in Micronesia may represent remnants of a single colonization, 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).

Specimens 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 Kohatsu Kabushiki Kaisha) apparently in 1935. An illustrated booklet, 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 cultivated 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

Corvus Kubaryi Reichenow, Journ. f. Ornith., 1885, p. 110. (Type locality, Palau, error = Guam.)

Corvus solitarius Kittlitz, Obser. Zool., in Lutké, Voy. "Le Séniavine," 3, 1836, p. 305 (Guahan); Bonaparte, Comptes Rendus Acad. Sci. Paris, 37, 1853, p. 830 (Mariannes); 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 kubaryi 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 Bernice P. Bishop Mus., 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 deutschen Marianen, 1913, pp. 87, 102 (Marianen); Reichenow, Die Vögel, 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); Bryan, 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, 1946, 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 greenish-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 extending over nostrils and base of culmen; 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. Measurements of Corvus kubaryi

Location	Number and sex	Wing	Tail	Full culmen	Tarsus
Guam	9 males	236 (229-244)	165 (158-170)	55 (51-57)	51 (49-52)
	19 females	227 (222-241)	151 (143-166)	50 (47-54)	50 (46-54)
Rota	3 males	235 (233-236)	167 (166-169)	54 (53-56)	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 nesting 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 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 September. 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 contained both plant and animal food. Both Seale (1901:55) and Safford (1905:79) comment on the damage which the crow does to the corn 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 singles 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 roadway counts made. Coultas (field notes) noted the birds at the northern 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.

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 Bryan (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 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. Winters 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 (Müller)

Chinese Blue Rock Thrush

Turdus philippensis Müller, 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 infrequent 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 Momiyama, Birds Micronesia, 1922, p. 66 (Pelew).

Table 34. Measurements of Edolisoma tenuirostre in Micronesia

Table 34. Measurements of Edolisoma tenuirostre in Micronesia

Subspecies	No.	Wing	Tail	Exposed culmen	Tarsus
E. t. monachum	10	98	80	21.0	23.0
		96-103	76-83	20.0-22.5	22.5-24.0
E. t. insperatum	35	109	86	23.0	24.0
		107-112	82-91	22.0-24.0	23.0-25.0

Table 35. Measurements of Corvus kubaryi

Table 35. Measurements of Corvus kubaryi

Location	Number and sex	Wing	Tail	Full culmen	Tarsus
Guam	9 males	236 (229-244)	165 (158-170)	55 (51-57)	51 (49-52)
	19 females	227 (222-241)	151 (143-166)	50 (47-54)	50 (46-54)
Rota	3 males	235 (233-236)	167 (166-169)	54 (53-56)	50 (49-51)

Merula obscura Wiglesworth, Abhandl. und Ber. Zool. Mus. Dresden, no. 6, 1890-1891 (1891), p. 39 (Pelew).

Turdus obscuras obscuras 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.—Breeds in northeastern Asia. Winters south to Malaysia. 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).

Psamathia annae Hartlaub and Finsch

Palau Bush-warbler

Psamathia annae Hartlaub and Finsch, Proc. Zool. Soc. London, 1868, p. 5, pl. 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, Journ. Mus. Godeffroy, 8, 1875, pp. 5, 22 (Palau); Nehrkorn, Journ. f. Ornith., 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 Vögel, 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, Garakayo, Peleliu, Ngabad.

Characters.—Adult: A medium-sized 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-buff; 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: Resembles 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	74	64	21.0	28.5
		(72-77)	(62-68)	(19.5-22.5)	(27.0-30.0)
Adult females	11	69	58	21.0	26.5
		(65-74)	(55-61)	(19.5-22.0)	(25.0-29.0)

Specimens 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 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 movements; 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 ecologic 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 Psamathia, there seems to be a great resemblance to the genus Arundinax of Blyth, a form with which it is not in my power to compare." The genera Calamoherpe and Tatare are now included in Acrocephalus; 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 "Aberrant 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 pattern of evolution which characterizes some of the other island birds in that the bill and legs are 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 having 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 marchei 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 appearance and structure of the feet, tail, wing, body and bill are the same.

Table 36. Measurements of Psamathia annae

Table 36. Measurements of Psamathia annae

Sex	No.	Wing	Tail	Exposed culmen	Tarsus
Adult males	7	74	64	21.0	28.5
		(72-77)	(62-68)	(19.5-22.5)	(27.0-30.0)
Adult females	11	69	58	21.0	26.5
		(65-74)	(55-61)	(19.5-22.0)	(25.0-29.0)