Chapter IX: Part 9

_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

=======================================+=====+=========+=========+=======
| | | |Exposed
LOCATION | No. | Wing | Tail |culmen
---------------------------------------+-----+---------+---------+-------
_Anoüs tenuirostris melanogenys_ | | | |
Hawaiian Islands | 29 | 222 | 113 | 41
| | 210-229 | 105-120 | 41-48
| | | |
_Anoüs tenuirostris marcusi_ | | | |
Wake 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_ | 13 | 227 | 120 | 44
Christmas Island | | 220-234 | 108-128 | 41-46
| | | |
Phoenix, Howland, Union, Danger, | 9 | 229 | 119 | 46
Suvarov Islands | | 226-233 | 113-124 | 42-48
| | | |
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, | 34 | 229 | 117 | 43
Admiralty Islands, New Guinea | | 222-237 | 109-130 | 40-46
| | | |
_Anoüs tenuirostris diamesus_ | | | |
Clipperton, Cocos 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

Column headings:

A: No.
B: Wing
C: Longest tail feather
D: Shortest tail feather
E: Exposed culmen
F: Depth culmen
G: Tarsus

===================+====+=======+======+=====+=====+=======+=========
SUBSPECIES | A | B | C | D | E | F | G
-------------------+----+-------+------+-----+-----+-------+---------
_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.

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

Column headings:

A: No.
B: Wing
C: Longest tail feather
D: Shortest tail feather
E: Exposed culmen
F: Depth Culmen
G: Tarsus

======================+====+=======+=======+=====+=====+=======+=========
LOCATION | A | B | C | D | E | F | G
----------------------+----+-------+-------+-----+-----+-------+---------
_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 Islds.| 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 Islands | 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

===================+============+===============+============+===========
| | | Exposed |
SUBSPECIES | Number | Wing | 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_

=================+===============+============+==============+===========
| | | Exposed |
NUMBER | Wing | Tail | 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."

=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).