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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

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