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SMITHSONIAN
MISCELLANEOUS
COLLECTIONS

VOLUME 145

“EVERY MAN IS A VALUABLE MEMBER OF SOCIETY WHO, BY HIS OBSERVATIONS, RESEARCHES,
AND EXPERIMENTS, PROCURES KNOWLEDGE FOR MEN’’—JAMES SMITHSON

SMITHSONIAN INSTITUTION PRESS
CITY OF WASHINGTON
1968

SMITHSONIAN PuBLiIcaATION 4754

PORT CITY PRESS, INC.
BALTIMORE, MD., U. S. A.

ty

on

N

CONTENTS

. WETMORE, ALEXANDER. Systematic notes concerned with the

avifauna of Panama. 14 pp. June 26, 1962. (Publ. 4501.)

. Wermore, ALEXANDER. Notes on fossil and subfossil birds. 17

pp., 2 figs. June 26, 1962. (Publ. 4502.)

. FRIEDMANN, HeErBert. The problem of the Viduinae in the light

of recent publications. 10 pp. July 20, 1962. (Publ. (4506.)

. GrtocKk, WALpo S., GERMANN, PAut J., and AGERTER, SHARLENE

R. Uniformity among growth layers in three ponderosa pine.
375 pp., 13 pls., ror figs. February 21, 1963. (Publ. 4508.)

. Kier, Porter M. Tertiary echinoids from the Caloosahatchee and

Tamiami formations of Florida. 63 pp., 18 pls., 58 figs. August
2, 1963. (Publ. 4543.)

. WETMoRE, ALEXANDER. Additions to records of birds known from

the Republic of Panama. 11 pp. December 16, 1963. (Publ.
4523.)

BENNETT, CHarLes F., Jr. A phytophysiognomic reconnaissance
of Barro Colorado Island, Canal Zone. 8 pp., 1 fig. December

20, 1963. (Publ. 4527.)

. Buzas, Martin A. Foraminifera from Late Pleistocene clay near

Waterville, Maine. 30 pp., 5 pls., 4 figs. March 1, 1965. (Publ.
4590.)

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SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 145, NUMBER 1

SYSTEMATIC NOTES
CONCERINET Wibid “Erin
AVIFAUNA OF PANAMA

By
ALEXANDER WETMORE

Research Associate, Smithsonian Institution

(Pustication 4501)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JUNE 26, 1962

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SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 145, NUMBER 1

SYOLEMATIC NOTES
CONCERINE DW Iiliet hide
AVIFAUNA OF PANAMA

By
ALEXANDER WETMORE

Research Associate, Smithsonian Institution

(PusiicaTiIon 4501)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JUNE 26, 1962

PORT CITY PRESS, INC.
BALTIMORE, MD., U. S. A.

SYSTEMATIC NOTES CONCERNED WITH THE
AVIFAUNA OF PANAMA

By ALEXANDER WETMORE

Research Associate
Snuthsoman Institution

The observations and descriptions included in the following pages
have come to attention during detailed studies of the birds of the
Isthmus of Panama and their comparison with forms from other
tropical areas. They are based in the main on the collections of the
United States National Museum, with addition of specimens seen
in the course of work at other institutions.

I. GEOGRAPHIC VARIATION IN THE BLACK VULTURE,
CORAGYPS ATRATUS (BECHSTEIN)

Opinion on recognition of races in the black vulture has varied
from treatment of the entire population as undivided throughout the
vast range, from southern United States south through Central
America and South America to southern Chile and Argentina, and
recognition of two races, one northern and one tropical, based on
difference in size. The latter view has been abandoned currently in
the main since a study by Friedmann (1933, pp. 187-188) in which
he compared average size of North American birds with a small
series from southern South America to find that the wing measure-
ments appeared too similar to warrant separation. In the course of
studies during recent years I have had opportunity to examine ap-
proximately 130 black vultures in various collections in the United
States and to assemble a series of measurements that verify the
findings of Brodkorb (1944, pp. 115-121) that three size groups are
present, a smaller population in the tropical area and two larger ones
in the temperate regions to the north and south on either side.
Brodkorb was uncertain as to the nomenclature to be used since he
could not distinguish between the two larger groups, which he there-
fore treated under one name. The extensive material that I have
seen has justified the recognition of 3 races, as follows:

CORAGYPS ATRATUS ATRATUS (Bechstein)
Vultur atratus Bechstein, in John Latham’s Allg. Uebers. Vogel, Bd. 1, Anh.,

1793, p. 655. (Florida.)

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 145, NO. 1

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 145

Vultur urubu Vieillot, Hist. Nat. Ois. Amér. Sept. vol. 1, Sept. 1807, p. 23,
pl. 2. (“Carolines ... Florides et... pour patrie la Zone torride’=Caro-
lina and Florida; cf. A. O. U. Check-list North Amer. Birds, ed. 3, 1910,
Pp. 153.)

Characters.—Size large; light markings on undersurface of pri-
maries less extensive; wing <¢ (32 specimens) 414-445 (426); 9
(28 specimens) 414-438 (426).

Resident from the mountains of northeastern Sonora (Sasabe,
Moctezuma), western Texas, eastern Oklahoma, southeastern Kansas,
Missouri, southern Illinois, southern Indiana, central Ohio (northern
Licking County), eastern West Virginia, and Maryland south to the
lower Rio Grande Valley in Texas (Brownsville), Louisiana, the
Gulf coast, and Florida. The breeding range in recent years has been
extended slowly along the northern boundary, with stragglers re-
corded casually to Nebraska, the Dakotas, and southern Ontario,
eastward to southwestern Quebec, Maine, and Nova Scotia.

A male from Guadalajara, Jalisco, with the wing 415 mm., seems
to indicate that this large northern form extends south from the
international boundary through the plateau region of México and
the adjacent mountain areas. In this same connection Col. L. R. Wolfe
has pointed out to me some interesting data concerned with egg size
in these birds. Three sets comprising 6 eggs in his collection from
La Laja, in northern Veracruz, on the coastal plain about 4o miles
south of Tampico, Tamaulipas, average 73.5 X 48.4 mm., which agrees
closely with an average of 74.3 50.1 mm. for 21 eggs in 11 sets in
the U. S. National Museum from Texas and Florida. As eggs of
the subspecies of true tropical range are smaller, as will be shown
under that race, there is indication that the northern form may range
into northeastern México, though this requires check whenever skins
from that area may be available.

CORAGYPS ATRATUS BRASILIENSIS (Bonaparte)

Cathartes brasiliensis Bonaparte, Consp. Gen. Avium, vol. 1, pt. 1, 1850, p. 9.
(“ex Amer. merid. Antill.” “Brasil merid.” designated by von Berlepsch,
Nov. Zool., vol. 15, 1908, p. 289; hereby further restricted to Rio de Janeiro,
Brazil.)

? Cathartes (vultur) urbis incola “Ricord,’ Lesson, Compléments de Buffon,
ed. 2, 1838, p. 93. (Indes occidentales .. . Santo Domingo, ... bords de
l’Orenoque .. . port d’Espagne . . . Saint-Vincent, a Saint-Lucie, a la Domi-
nique et a la Santiago-de-Cuba.)

? Cathartes urbicola Des Murs, Rev. Mag. Zool., 1853, No. 4, p. 153. (Based on
Lesson, 1838, above.)

? Cathartes Ricordi Des Murs, Rev. Mag. Zool., 1853, No. 4, p. 153. (Alternate
name for C. urbicola.)

NO. I AVIFAUNA OF PANAMA—WETMORE 3

? Vultur urbis-incola “Riccord,’ Cassin, U. S. Expl. Exped. Mamm. and
Ornith., 1858, p. 81. (Based on Des Murs, 1853, above.)
Characters.—Size small; light markings on underside of primaries

more extensive and clearer white; wing ¢ (17 specimens) 386-410

(401) ; 2 (23 specimens) 388-413 (400).

Resident in the tropical zone; in México, along the Pacific coast
from southern Sonora (Camoa on the lower Rio Mayo), and on the
eastern side from southern San Luis Potosi (Bledos, Xilitla) south-
ward throughout Central America; and in South America on the
west to the coastal region of Pertti (Lima), and on the east to the
lowlands of Bolivia (Buenavista, Santa Cruz), and southern Brazil.

The southern limit from the material seen is uncertain but appears
to include most of Brazil. The small size is constant throughout this
vast range. It appears desirable to pinpoint restriction of type locality
from that proposed by von Berlepsch to Rio de Janeiro in the south-
ern part of that great country. Two specimens in the U. S. National
Museum from the Federal District are typical in small size. It is
possible that the next race may be found along the far southern
boundary since this is the form of Paraguay.

With regard to egg size in this race, Colonel Wolfe writes that
7 eggs (in 4 sets) in his collection from Trinidad average 70.1 X 49.4
mm. A set of 2 in U. S. National Museum, collected at Lagoa Santo,
Minas Gerais, by E. G. Holt, measures 70.4 x 48.4 and 68.6 xX 488
mm. These figures are definitely less than those listed above for the
typical race.

Cathartes urbicola of Des Murs (1853, p. 153), listed in the
synonymy above, is based on an account of “le Catharte citadin”
given by Ricord to Lesson and published by the latter in the second
edition of his Compléments de Buffon (Lesson, 1838, p. 93). The
bird described by Ricord is a composite based in part on the black
vulture, and in part on the turkey vulture, but with the size of a
condor as it is said to be 48 inches tall. It is described as inhabiting
the Spanish settlements in the West Indies, Trinidad, and on the
Orinoco, where it was protected by the authorities as a scavenger.
Lesson in a footnote gave it the name “Cathartes (vultur) urbis
incola, Ricord,” which was cited by Des Murs (p. 147) as “Cathartes
urbis incola (Ricord) Lesson.” Cassin (1858, p. 81) lists this under
Cathartes urbicola Des Murs as “Vultur urbis-incola, Riccord.” Des
Murs also in the original citation wrote ricordi as an alternate name,
his statement being as follows: “Jusque-la nous croyons devoir
proposer pour le nom de ce Catharte, en tant qu’on le maintiendrait

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 145

dans le série comme espéce douteuse ou a étudier, soit le nom de

Cathartes urbicola, soit encore mieux celui de Cathartes Ricordt.”
While the black vulture undoubtedly is one of the birds to which

these citations refer, the details of the accounts include the turkey

vulture and possibly the condor in such a mixture that the names
are of uncertain application. The earliest valid name is brasiliensis

Bonaparte of 1850.

CORAGYPS ATRATUS FOETENS (Lichtenstein)

Cathartes foetens Lichtenstein, Verz. Ausgest. Saug. Vogel Zool. Mus. Berlin,
1818, p. 30. (Based on “Iribu Azara’; Paraguay; restricted to Asuncion,
Paraguay, by Brodkorb, in 1944.)

Characters.—Size similar to that of C. a. atratus, but light markings
on under surface of primaries more restricted, the under wing defi-
nitely darker than in the two more northern races. Wing, male (10
specimens) 412-437 (421), female (5 specimens) 416-422 (419).

Resident in the Andes from northern Ecuador (Quito) to northern
Bolivia (Cochabamba) ; in Chile, south to Aysen; Paraguay, including
the Chaco, from the north-central section (Horqueta, 200 kilometers
west of Puerto Casado), and Argentina south to the Rio Negro;
probably through Uruguay (no specimens examined).

The large size of the southern group has been the main cause of
confusion in recognition of races in this species, since when birds of
southern South America have been examined they have not appeared
separable from northern representatives of equivalent measurements.
This misunderstanding has completely overshadowed the vast tropical
population of uniformly small dimension. The true status was clearly
outlined by Brodkorb (1944, pp. 115-121), but his analysis has been
disregarded since he applied the name of the nominate race to the
large birds of both north temperate and south temperate areas in
spite of their wide separation by another form.

With regard to eggs, Colonel Wolfe writes that a set of 2 from
Argentina in his collection measures 74.5 45.0 and 72.0X 47.0, and
that the average of 54 eggs taken in Chile, according to figures fur-
nished by A. W. Johnson, is 72.58 49.5 mm. These figures are close
to the sizes found in the typical race of the north,

Il. THE CRESTED BOBWHITE, COLINUS CRISTATUS
(LINNAEUS), OF WESTERN CHIRIQUI
The quail of Panama, described as Colinus cristatus panamensis
by Dickey and van Rossem, is locally common from western Veraguas
through Coclé to the western area of the Province of Panama. To
the south it ranges down the eastern side of the Azuero Peninsula

me

NO. I AVIFAUNA OF PANAMA—WETMORE 5

through Herrera and Los Santos, including the valley of the Rio
Tonosi. There have been few records for the Province of Chiriqui
until recently, when I found crested bobwhites in small numbers in
the coastal lowlands between Alanje and Puerto Armuelles, as well
as near Boquete. The birds of this region are an isolated population
separated by a considerable area, where no quail are known, from
the main group of the species in the Republic. The bird of Chiriqui
differs so decidedly in coloration that it requires recognition as an-
other form.

COLINUS CRISTATUS MARIAE subsp. nov.

Characters.—Similar to Colinus cristatus panamensis Dickey and
van Rossem * but decidedly darker, the markings of the upper surface
blacker, and the black areas of the lower surface more extensive in
both sexes; darker than any other population of the species.

Description—Type, U.S.N.M. No. 471174, male adult, from 7
kilometers south of Alanje, Province of Chiriqui, Panama, collected
March 8, 1960, by A. Wetmore (original number 23600). Forehead,
lores, a narrow feathered area surrounding the eye, and side of head
down to the ramal area, dull white; shorter anterior feathers of the
narrow, elongated crest pale drab-gray, the longer ones fuscous,
tipped, and spotted irregularly along the edges, with drab; a broad
superciliary extending from above the middle of the eye back to
the side of the nape behind the auricular region mikado brown, lined
irregularly with black; crown black, edged narrowly with white and
adjacent to the superciliary with bright brown; auricular area dull
olive-buff ; black of crown extended down over nape, hindneck, and
sides of neck, to extreme upper back, and spotted rather irregularly
with white; feathers of back, scapulars, tertials, wing coverts, rump,
upper tail coverts, and tail black, freckled with fine markings of fawn
color, grayish white, and white, the inner secondaries edged, and the
wings spotted more prominently, with dull white; primaries dull
mouse gray, with the edge of the outer web and the tip of both
webs varied from dull white to tilleul-buff; chin drab-gray; throat,
foreneck, and sides of upper neck, below auriculars, russet; base of
neck black, spotted with white and irregularly with russet, changing
across upper breast to sayal brown, with each feather banded basally
with white and medially with black below the brown tip to produce
an appearance of irregular spotting; abdomen pinkish buff, with
partly concealed bars of dull black, spotted with white and pinkish

1 Colinus leucotis panamensis Dickey and van Rossem, Condor, vol. 32, No. 1,
Jan. 20, 1930, p. 73. (Aguadulce, Coclé, Panama.)

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 145

buff ; under surface of wings mouse gray, with the under wing coverts
tipped and edged lightly with dull white. Bill black; tarsus and feet
dusky neutral gray (from dried skin.)

Measurements.—Males (6 specimens), wing 92.6-95.3 (93.7), tail
46.3-52.3 (49.2), culmen from cere 12.7-13.9 (13.2), tarsus 27.9-
29.1 (28.5) mm.

Females (3 specimens), wing 92.0-96.1 (94.0), tail 45.7-50.0
(48.4), culmen from cere 12.4-13.4 (12.8), tarsus 27.2-29.4 (28.5)
mm.

Type, male, wing 93.3, tail 52.3, culmen from cere 13.0, tarsus
27.9 mm.

Range.—Western Chiriqui on the southern slopes of the Volcan
de Chiriqui near Boquete (El Salto, 1,350 meters elevation), and
Francés near El Banco; and on the coastal plain below Alanje.
Apparently restricted in distribution but fairly common when it 1s
found.

Remarks.—The crested bobwhite of Panama has been known prin-
cipally from Veraguas and Coclé, with few specimens in museum
collections from Chiriqui. Present information indicates that the
population in Chiriqui is isolated as it is known only from the western
part of the province, and there are no records of the related race
Colinus cristatus panamensis beyond a point about 10 kilometers
west of Sona in western Veraguas. From this western limit the
subspecies pananiensis is recorded in open country eastward to the
western sector of the Province of Panama (where I have found it
near Nueva Gorgona, and 10 kilometers east of Bejuco), and south
on the eastern side of the Azuero Peninsula through Herrera and
Los Santos to Pedasi and near the Rio Oria below Los Asientos ;
also in the lower Tonosi valley. It is evident that the additional race
here described is one of restricted range, since it is reported to date
only from three localities in an area between 50 and 60 kilometers
in length. As I have been long familiar with the brown subspecies
panamensis of farther east in the Republic the much darker colora-
tion of the form here described was immediately evident on my first
sight of it in life. It marks the western extension of the species
cristatus.

The new form is named for Mrs. Robert A. Terry, who as Mary E.
McLellan Davidson, through her studies in field and laboratory, has
added much to our knowledge of the bird life of the Province of
Chiriqui.

NO. I AVIFAUNA OF PANAMA—WETMORE 7

Il. AN ADDITIONAL RACE OF THE CHESTNUT-BACKED
ANTBIRD, MYRMECIZA EXSUL SCLATER

The chestnut-backed antbird, found widely through the tropical
lowlands of the Republic of Panama, is a forest-inhabiting species
that still remains in small numbers in inhabited sections since a part
of its haunt is in swampy woodlands where the land is too wet to be
available for cultivation. Here it still finds suitable habitat when
the surrounding forest has been destroyed. Three subspecies have
been recorded from Panama. A detailed study of the series now
available from the entire isthmus has indicated a fourth that requires
description.

MYRMECIZA EXSUL NIGLARUS subsp. nov.

Characters.—Similar to Myrmeciza exsul exsul,? but paler above
and below; darker than M. e. occidentalis Cherrie,* especially in the
female.

Description—Type, U.S.N.M. No. 423427, male, from the Rio
Chiman about 10 kilometers above Chiman, Province of Panama,
collected February 20, 1950, by A. Wetmore and W. M. Perrygo
(original number 15208) : Crown, sides of head, throat, and foreneck
black, shading to dark neutral gray on hindneck; rest of upper
surface Mars brown, shading to russet on rump and upper tail
coverts; rectrices fuscous, edged with russet; bend of wing black,
edged with white, the alula being fuscous-black with the outer webs
edged with white; lesser wing coverts with a shaft line of dusky
neutral gray, terminating in a slightly expanded tip of the same
color; primaries and secondaries fuscous-black, with the outer webs
Mars brown, except the outermost, which has a narrow white outer
edge; black of foreneck shading progressively to deep neutral gray
on chest, and neutral gray on lower breast; under wing coverts
dusky neutral gray, mixed scantily with white; posterior part of
sides, flanks, and under tail coverts cinnamon-brown. Bill black ; feet
fuscous-brown (from dried skin).

Measurements.—Males (15 specimens), wing 65.0-70.1 (67.1),
tail 42.5-49.7 (45.7), culmen from base 20.2-22.3 (21.4), tarsus
27.5-29.7 (28.4) mm.

2Myrmeciza exsul P. L. Sclater, Proc. Zool. Soc. London, vol. 26, 1858
(Jan.-May 1859), p. 540. (‘‘Panama,” type locality hereby restricted to near
Gatun, Canal Zone.)

8 Myrmeciza immaculata occidentalis Cherrie, Auk, vol. 8, No. 2, April 1891,
p. 191. (Pozo Azul de Pirris, Pacific slope of Province of San José, Costa
Rica.)

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 145

Females (10 specimens), wing 62.7-67.0 (64.2), tail 40.2-47.6
(43.3), culmen from base 19.9-22.2 (20.8), tarsus 26.4-29.6 (28.1)
mm.

Type, male, wing 66.8, tail 42.5, culmen from base 20.7, tarsus
28.2 mm.

Range.—Eastern Panama; on the Pacific slope from the western
end of the Cerro Azul east through the Province of Panama to
western Darién; on the Caribbean slope from the upper Chagres
Valley, above Madden Lake (Quebrada Candelaria on the Rio
Pequeni, Quebrada Peluca on the Rio Boquerén), and western
Comarca de San Blas (Mandinga) east to the Colombian boundary,
and beyond to Acandi in northernmost Chocé, Colombia.

Remarks.—The wing coverts are plain in most individuals of this
race, with the white spotting typical of M. e. cassini and M. e.
maculifer found only casually in a few. Specimens from the middle
Chucunaque Valley, near the mouth of the Rio Tuquesa, are inter-
mediate between the new form and cassini, which ranges through
the rest of the lowlands of the Tuira basin.

The name niglarus is taken from the Greek vlyAdpos, a small fife,
in allusion to the whistled calls of these birds, heard constantly as
they move through the undergrowth on the forest floor.

IV. THE GEOGRAPHIC RACES OF THE SILVER-THROATED
TANAGER, TANGARA ICTEROCEPHALA (BONAPARTE)

Specimens now available, particularly those from Costa Rica and the
western half of Panama, permit a better understanding of geographic
variation in the tanager Tangara icterocephala (Bonaparte). The
species, described from Ecuador in 1851, soon was recorded also from
Costa Rica, and from Veraguas and Chiriqui in western Panama.
Cabanis, in 1861, named the Costa Rican bird frantzu, but Ridgway
in 1902, with limited series, was not able to distinguish this as a
separate race. Hellmayr, in 1936, and others have followed Ridgway’s
treatment, though with indication by some that there may be two
forms. It is only recently that De Schauensee, in 1951, in his account
of the birds of Colombia, recognized formally that there are two
races. The uncertainty has resulted from the interesting fact that
the populations of these birds in Ecuador and in Costa Rica both
are bright in color, which obscures their differences. Specimens that
I have collected in recent years from the mountains immediately west
of the Canal Zone include another subspecies distinct from both of
the others.

Females in all three races are duller, more greenish throughout, a

NO. I AVIFAUNA OF PANAMA—WETMORE 9

fact that should be borne in mind in comparing specimens in which
the sex is not marked. Juvenile birds differ from adult females in
more greenish back, hindneck, and crown, duller-colored rump, less
definite streaking on the back, and duller yellow of the under surface.

TANGARA ICTEROCEPHALA ICTEROCEPHALA (Bonaparte)
Calliste icterocephala Bonaparte, Compt. Rend. Acad. Sci. Paris, vol. 31, No. 3

(séance du 20 janv.), 1851, p. 76. (Valley of Punta Playa, near Quito,

Ecuador. )

Characters.—Similar to T. 7. frantzii in bright coloration, but
with feathers of crown and nape somewhat greenish basally, so that
the yellow in this area appears less intense; partly concealed ring
around the base of the hindneck deeper blue; foreneck and throat
averaging faintly darker.

Measurements.—Males (12 specimens), wing 71.2-75.0 (72.7),
tail 44.2-48.5 (45.8), culmen from base 12.1-12.5 (12.3), tarsus
16.0-17.3 (16.8) mm.

Females (9 specimens), wing 66.6-72.3 (68.7), tail 41.9-46.2 (43.4),
culmen from base 12.2-14.2 (12.7), tarsus 16.2-17.5 (17.2) mm.

Range.—Mountains of eastern Darién, Panama (Cerro Tacarcuna,
Cerro Pirre) south in the western Andes through Colombia to
southern Ecuador.

TANGARA ICTEROCEPHALA ORESBIA subsp. nov.

Characters.—Decidedly duller yellow throughout than either T. 4.
frantzu or T. 1, icterocephala; partly concealed band on hindneck
more greenish blue; foreneck and throat darker; sides and flanks
darker, with a greenish-yellow cast: Female, in addition, with edging
on back feathers more green, less yellow.

DescriptionType, U.S.N.M. No. 433998, male, south face of
Cerro Campana, 850 meters elevation, western sector of the Province
of Panama, Panama, collected March 7, 1951, by A. Wetmore and
W. M. Perrygo (original number 16221). Lores, a very narrow
line around eyelids, a small spot behind the eye, a narrow line pos-
terior to the nostrils, and another from the gape across the lower
margin of the cheeks to the nape, black; crown and sides of head
slightly duller than light cadmium; nape washed with pyrite yellow;
band across hindneck bluish gray-green; back feathers distinctly
streaked, black centrally, edged broadly with sulphine yellow an-
teriorly, changing posteriorly to wax yellow; rump between light
cadmium and apricot yellow ; upper tail coverts oil green, tipped indis-
tinctly with warbler green; wings and tail black; lesser and middle
wing coverts edged with lettuce green, with a light tipping of lemon

10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 145

chrome; greater wing coverts, primaries, and secondaries edged
heavily with lettuce green; inner webs of central pair of rectrices
cedar green; outer webs of all rectrices edged with lettuce green ;
point of chin black; feathers of throat, foreneck, and upper margin
of chest dark green-blue gray, washed on throat and adjacent ramal
area with dark bluish glaucous, on foreneck with light grape green,
and on sides of neck with pinkish buff, with the darker basal color
showing through in varying amount with change in angle of the
light; center of breast and abdomen between light cadmium and
lemon chrome; sides between light cadmium and aniline yellow;
flanks and under tail coverts aniline yellow; tibia citrine; bend of
wing warbler green, stippled lightly with black; under wing coverts
light yellowish olive externally, changing to white internally, edged
lightly with cream-buff. Bill black; tarsus and toes fuscous (from
dried skin).

Measurements—Males (8 specimens), wing 72.4-77.4 (73.6),
tail 45.8-49.8 (47.3), culmen from base 12.0-14.8 (12.9), tarsus
17.5-18.8 (18.3) mm.

Females (g specimens), wing 68.7-72.4 (71.1), tail 43.5-47.6
(45.6), culmen from base 12.5-14.0 (13.2, average of 8), tarsus
17.8-18.8 (18.3) mm.

Range.—Mountain areas of west central Panama from Cerro
Campana, western Provincia de Panama, to Coclé (El Valle, Rio
Guabal).

The name of this form is taken from the Greek dpéofuos, living
on mountains.

TANGARA ICTEROCEPHALA FRANTZII (Cabanis)
Callispiza (Chrysothraupis) Frantsii Cabanis, Journ. fir Orn., vol. 9, pt. 2,

March 1861, p. 87. (Costa Rica.)

Characters.—Similar to T. 1. icterocephala, but with crown and
hindneck more yellow; partly concealed ring on base of hindneck
paler, more greenish blue; foreneck and throat paler: definitely
brighter yellow above and below than oresbia.

Measurements.—Males (14 specimens from Costa Rica), wing
72.3-78.6 (75.5), tail 44.7-49.8 (47.8), culmen from base 11.5-12.8
(12.3), tarsus 17.2-18.8 (18.0) mm.

Females (12 specimens from Costa Rica), wing 69.5-73.7 (71.7),
tail 43.0-46.3 (44.9), culmen from base 11.5-12.6 (11.9), tarsus
172-185 (17.9)-mm.

Range.—Mountains of Costa Rica and western Panama, east to
eastern Veraguas (Chitra).

NO. I AVIFAUNA OF PANAMA—WETMORE tif

V. ADDITIONS TO THE RECORDED LIST OF BIRDS FROM THE
REPUBLIC OF PANAMA

SALMON’S TIGER-BitTERN, Tigrisoma salmont SCLATER AND SALVIN:

This species, described from Medellin in the Province of Antioquia,
northwestern Colombia, with a recorded range east to Venezuela and
south through Ecuador and Pert to western Bolivia, ranges also
along the Caribbean slope of the Isthmus of Panama. It was first
noticed for this area on February 29, 1952, when I collected an adult
male on the Rio Uracillo, near the town of that name in the foothills
of the Caribbean slope of Coclé. I secured another, an immature
bird, near the Peluca Hydrographic Station on the Rio Boqueron,
Province of Colén, on February 21, 1961, and have a third, shot on
the Rio Changena, Bocas del Toro, September 9, 1961. This species
differs from the banded tiger-bittern Tigrisoma lineatum (Boddaert)
structurally in the form of the bill, which is shorter and also heavier,
less attenuate at the tip. The adult salmoni is definitely blacker, but
the immature differs only in being more extensively white on the
lower surface. With the presence of the species known I have found
several immature birds in other collections taken earlier in Darién
and the eastern Comarca de San Blas, but wrongly identified as
Tigrisoma lineatum.

SLENDER-BILLED Kite, Helicolestes hamatus (TEMMINCK) :

The slender-billed kite is reported for Colombia on the basis of a
record by Salmon from the Rio Ité, near Remedios in Antioquia,
and is known from scattered localities from eastern Pertti to Vene-
zuela (Caicara), Surinam, and the lower Amazon. In Darién, on
February 24, 1959, as I landed from a piragua at the mouth of a
tiny stream that enters the Rio Tuira a short distance above where
the Rio Paya joins this larger river, I was interested to note shells
of an apple snail scattered along the sandy shore, and immediately
saw one of these kites perched over a shaded pool inside the forest
border. The bird was a female. Later I received another skin from
the Gorgas Memorial Laboratory, taken near the same point in the
previous year. These are the first reports of this little-known species
beyond South America.

GuAcHARO, Steatornis caripensis HUMBOLDT:

On the night of March 19, 1959, Bernard Feinstein, assistant to
Dr. Charles O. Handley, Jr., captured a female of this species in a
mist net set for bats at an elevation of 975 meters near the old
Tacarcuna village site on Cerro Tacarcuna, Darién. The guacharo

I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 145

has a wide distribution in northern South America, including Trini-
dad, but has not been found previously outside those limits.
SHORT-TAILED SwiFt, Chaetura brachyura brachyura (JARDINE) :

On September 12, 1960, Dr. Nathan Gale found one lying dead
at Corozal, Canal Zone, and brought it to the laboratory of the
Malaria Control Service. Here Eustorgio Méndez of the Gorgas
Memorial Laboratory secured it and prepared the skin, which is
now in the U. S. National Museum. The species has a wide range
in South America from the north coast to eastern Pert and central
Brazil, with populations in St. Vincent, Trinidad, and Tobago. The
Canal Zone record is the first report of it for Panama.

VI. ADDITIONS TO THE LIST OF BIRDS KNOWN
FROM COLOMBIA

Studies of the extensive collections of birds from northern Co-
lombia in the U. S. National Museum in connection with work on
the avifauna of Panama continues to add to the forms known from
that republic. Recent additions in this field are as follows:
SAVANNA Hawk, Heterospizias meridionalis rufulus (VIEILLOT) :

Examination of a considerable series of these beautiful hawks
verifies recognition of two forms on the basis of size. The southern
group that breeds from southern Paraguay and Rio Grande do Sul,
Brazil, to the provinces of Cordoba and Santa Fé, in northern Argen-
tina, ranges in wing measurement, regardless of sex, from 418 to
452 mm. During the period of southern winter part of these larger
birds move northward into the territory of the typical race. The
northern population, true mcridionalis, resident from Panama, Co-
lombia, and Venezuela to Bolivia, northern Paraguay, and south-
central Brazil, varies in wing measurement from 379 to 412 mm.

A female that I shot near Maicao in the Guajira Peninsula, north-
eastern Colombia, on April 14, 1941, with primaries worn at the
tip, has the wing still 418 mm. long, and so represents a migrant or
wanderer of the southern subspecies. Other breeding specimens taken
during the same period in the Guajira in their smaller size are
typical meridionalis.

Gray Hawk, Buteo nitidus blaket HELLMAYR AND CONOVER:

A female collected by M. A. Carriker, Jr., at Acandi in northern
Choc6, on the western side of the Gulf of Uraba, is typical of this
race of adjacent Panama. It differs from Buteo nitidus nitidus, found
elsewhere in northern Colombia, in being darker gray above, par-
ticularly on the crown and hindneck. Apparently blake: does not

NO. I AVIFAUNA OF PANAMA—WETMORE 13

extend far into Colombia since De Schauensee has reported typical
nitidus from the Rio Jurado on the Pacific slope of northern Choco,
and Carriker secured that subspecies at Nazaret, in western Cordoba,
beyond the Rio Sint.

PicEoN Hawk, Falco columbarius bendirei SWANN:

A male taken by Carriker February 26, 1946, at Manancanaca in
the higher levels (3,600 meters) of the Sierra Nevada de Santa
Marta is a well-marked adult of this migrant from western North
America. The race bendiret has not been reported previously in
South America. In this connection another specimen, a female that
W. M. Perrygo and I secured April 15, 1946, at Jaque, Darién, only
40 kilometers from the Colombian boundary, is also of interest as
the only record at present for Panama.

BANDED Woop-QualL, Rhynchortyx cinctus cinctus (SALVIN):

A series taken by Carriker at Socorro and Quebrada Salvajin,
Cordoba, near the Rio Sint, at Taraza, in northern Antioquia, near
the Rio Cauca, and at Volador in southern Bolivar, represents the
typical race, hitherto unknown outside Panama. Rhynchortyx cinctus
australis Griscom, described from the Comarca de San Blas, on the
Caribbean coast of eastern Panama, with additional material proves
not separable from typical cinctus, as the characters on which it was
based are those of individual variation,

House Wren, Troglodytes aedon inquietus Batrp:

Male and female taken January 5 and 7, 1950, by Carriker at
Acandi, Choco, on the western side of the broad entrance of the
Gulf of Uraba, are good examples of this race, which is the form
found throughout most of Panama. The occurrence at Acandi is not
surprising since this subspecies has been recorded east in the Comarca
de San Blas to Puerto Obaldia near the Colombian frontier. Carriker
secured another male inquietus February 9, 1950, at Necocli, on the
eastern shore of the Gulf of Uraba (called also Gulf of Darién),
north of Turbo, and collected two house wrens April 28 in the same
year farther south at Villa Artiaga in northwestern Antioquia that
are intermediate toward Troglodytes aedon striolatus, but nearer to
inquietus. It appears, therefore, that the form typical of most of
Panama extends around the head of the Gulf.

LITERATURE CITED

Bropkors, PIERCE.
1944. Geographical variation in the black vulture. Pap. Michigan Acad.
Sci., Arts, Letters, vol. 29 (1943), pp. 115-121, fig. I.

I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 145

/
CASSIN, JOHN.

1858. United States Exploring Expedition during the years 1838, 1839,
1840, 1841, 1842, under the command of Charles Wilkes, U.S.N.—
Mammalogy and Ornithology, pp. i-x, 1-466.

Des Murs, D.

1853. Observations sur un nom spécifique Catharte citadin Lesson—V ultur
urbis incola Ricord—Cathartcs urbis incola Lesson. Rev. Mag.
Zool., No. 4, Feb., pp. 146-154.

FRIEDMANN, HERBERT.

1933. Critical notes on American vultures. Proc. Biol. Soc. Washington,

vol. 46, Oct. 26, pp. 187-190.
Lesson, P.
1838. Le Catharte citadin. Compléments de Buffon, ed. 2, pp. 93-94.

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 145, NUMBER 2

Charles D. and Mlary Waux Halcott
Research Fund

NOTES ON FOSSIL AND SUBFOSSIL
BIRDS

By
ALEXANDER WETMORE

Research Associate, Smithsonian Institution

(Pustication 4502)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JUNE 26, 1962

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 145, NUMBER 2

Charles D. and Mary Waux CHalcott
Research Fund

NOTES ON FOSSIL AND SUBFOSSIL
BIRDS

By
ALEXANDER WETMORE

Research Associate, Smithsonian Institution

(Pustication 4502)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JUNE 26, 1962

PORT CITY PRESS, INC.
BALTIMORE, MD., U. S. A.

Charles D. and flary Waux Walcott Research Fund

NOTES ON FOSSIL AND SUBFOSSIL BIRDS
By ALEXANDER WETMORE

Research Assoctate
Smithsonian Institution

The following pages cover several studies on fossil and subfossil
birds, based mainly on material in the U.S. National Museum. The
collection from the Pleistocene of Augusta County, Va., has come
through the kindness of John E. Guilday of the Carnegie Mu-
seum. The bones from Bermuda were collected for the Smithsonian
Institution by David B. Wingate.

I. AN UPPER CRETACEOUS BIRD RELATED TO THE IBISES

In the summer of 1958, Dr. Shelton P. Applegate, now at State
College, Arkansas, collected a broken humerus of a bird in Greene
County, west-central Alabama, that appears related to the storks and
ibises of the order Ciconiiformes. According to data supplied by
Dr. Applegate, the specimen came from Hewletts farm, 3 miles
northeast of the town of Boligee, where it was found in the farther
side of a series of gullies that lie to the west of the county road,
before this reaches the farmhouse entrance. The location, in the
Mooreville formation of the Selma chalk, was about 10 feet below
the Arcola limestone.

The form of the humerus indicates a species about half the size
of the living white ibis Eudocimus albus.

PLEGADORNIS gen. nov.

Diagnosis.—A fossil storklike bird, with the distal end of the
humerus flattened, ectepicondyle long, and the brachial depression
shallow and relatively large. Characters in detail those of the only
known species, Plegadornis antecessor, the type of the genus.

PLEGADORNIS ANTECESSOR sp. nov.

Characters ——lKnown from a fragmentary left humerus that is
generally similar to living species of the suborder Ciconiae; much
smaller than the smallest of living forms of the suborder (half the

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 145, NO. 2

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 145

size of Plegadis, or less). Outline of distal end of the humerus
(fig. 1) somewhat similar to that of species of the family Threskior-

Fic. 1—Type of Plegadornis antecessor from the Upper Cretaceous of Alabama.
Natural size.

nithidae, but with the ectepicondyle elevated above the level of the
internal condyle at a slightly greater angle; attachment of anterior
articular ligament relatively much larger; internal condyle relatively
longer and slightly narrower; external condyle slightly less angular
on upper end, with more separation from the ectepicondyle on its
outer margin; the ectepicondyle much larger, so that one-fourth of
its length extends up the shaft above the level of the upper end of
the external condyle; brachial depression large and only slightly
depressed ; a slight expansion from the side of the shaft at the lower
end of the deltoid crest, below the actual articular area, like that in
modern species of Threskiornithidae, particularly of the subfamily
Plataleinae.

Transverse breadth across distal end 10.5 mm.; transverse breadth
of shaft near center 4.9 mm.

Type.—Distal end of left humerus, with part of the shaft from
the upper end, U.S.N.M. No. 22820, from the Mooreville formation
of the Selma chalk, Upper Cretaceous, 3 miles northeast of Boligee,
Greene County, Ala., collected by Shelton P. Applegate about June
20, 1958.

Remarks.—The important part of the specimen is the distal end,
which is somewhat worn, but where sufficient character is present to
allow indication of relationship. The upper segment shows a trace
of the curvature characteristic of the suborder in which it is allo-
cated, and an indication of the form at the extreme lower end of the
deltoid crest, but has lost other details. A section of the shaft between
the upper and lower portions is missing.

NO. 2 FOSSIL AND SUBFOSSIL BIRDS—-WETMORE 3

The fossil is important because of its indication, slight though that
may be, of the occurrence of ibislike birds at this early period, and
in its general similarity to species of this group that still exist. It is
the first fossil bird recorded from Alabama.

While it appears allied to species now classified in the suborder
Ciconiae, which includes the families of the hammerhead (Scopus),
the storks (family Ciconiidae), and the ibises (Threskiornithidae),
its differences, as indicated in the diagnosis, are such that it requires
a separate family, Pelagodornithidae, to be allocated in a superfamily
Pelagodornithoidea, adjacent to the superfamily Threskiornithoidea.

The generic name for this interesting species is formed from the
Greek root for Plegadis, a widely distributed modern genus of ibises,
Viz, wAnyas, ados, and dpus, bird. The specific name, the Latin word
“antecessor,” signifies a forerunner (or ancestor).

II. A RECORD OF THE COMMON LOON, GAVIA IMMER
(BRUNNICH), FROM THE PLEISTOCENE OF MARYLAND

The cranium of a loon found in December 1959 on the shore be-
tween Chesapeake Beach and Plum Point, on Chesapeake Bay, in
Calvert County, Md., has been presented to the U.S. National Mu-
seum by Miss Alice H. Howe of Arlington, Va. The specimen
(U.S.N.M. No. 22552) is stained dark brown in color and still retains
a film of fine clay silt in the deeper impressions. Its appearance, both
in color and in the clay deposit, is indication of ancient age and is
typical of the Pleistocene deposits that lie above the Miocene beds
in the earthern cliffs that line this section of Chesapeake Bay. There
is no reason therefore against listing the bone as of that age.

The bone (fig. 2) includes the upper surface of the cranium from
the base of the premaxilla to the foramen magnum, except that the
ridge immediately above the foramen is missing, and there are minor
breaks in the posterior area of the frontal. Below, the basioccipital
area has been lost.

The bone obviously is representative of an adult of a large species
of the genus Gavia. On comparison of to skulls of Gavia immer with
6 of G. adamsit, all of adult age, I find that the cranial section in the
former averages less massive in form. The angle of the anterior end
of the frontals, immediately posterior to their junction with the nasals,
in most is less abrupt, and the transverse width through the heavy
postorbital processes is less. In G. adamsii the cranium is more
massive, the anterior end of the frontals slopes more abruptly, and

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 145

Fic. 2.—Dorsal view of the cranium of a common loon, Gavia immer, from the
Pleistocene of Maryland. Natural size.

the transverse width indicated is greater. While the largest skulls of
immer are close to adamsii the smaller ones appear distinct. The
specimen under identification agrees with the medium-sized and
smaller samples of immer and is identified as that species.

The occurrence on Chesapeake Bay is an additional Pleistocene
record for Gavia immer, which has been reported previously from
deposits of that age in California and Florida.

III. THE WHOOPING CRANE, GRUS AMERICANA, IN MICHIGAN

In a recent visit to the Chicago Natural History Museum I
noted a Pleistocene bone identified tentatively as this species, which
Dr. Rainer Zangerl has kindly placed in my hands for study. The
specimen is a left tarsometatarsus of a juvenile individual which
apparently had developed the full length of this segment of the bone,
but in which the upper end was not fully ossified, as the surface of
the articulation is not completely formed. The shaft also is slender
with its outlines rounded, less angular than in adult specimens, and
the entire bone presents the slightly roughened spongy appearance
that marks an immature stage. The distal trochlea and the talon both
are broken and missing, but it is possible to ascertain the length from
the anterior end to the distal foramen, which equals that of modern
adult tarsometatarsi in the U.S. National Museum Collections. It is

NO. 2 FOSSIL AND SUBFOSSIL BIRDS—-WETMORE 5

identified, without question, as Grus americana (Linnaeus), the
whooping crane.

The specimen, Chicago Natural History Museum No. P25538,
found one-half mile northwest of Ferry, Oceana County, Mich., in
what was reported to be a Pleistocene marl, was presented to the
Museum by George W. Bowen. The record is of particular interest
since it is not only a new fossil locality for this species, but also is
the first report of this crane from the State of Michigan.

The species has been recorded previously as a fossil from the
Upper Pliocene of Idaho, and from the Pleistocene of California and
Florida.

IV. BIRDS OF LATE PLEISTOCENE AGE FROM
AUGUSTA COUNTY, VIRGINIA

Through the kindness of John E. Guilday of the Carnegie Museum
a collection of bird bones from small caves and fissures at the bases
of the rock columns known as the Natural Chimneys, a mile north
of Mount Solon, Va., has come to me for study. According to data
supplied by Mr. Guilday, the presence of bones at this site was re-
ported first in 1949 by Theodore B. Ruhoff, who has collected the
bulk of the material. Parties from the Carnegie Museum, directed
by J. LeRoy Kay, curator emeritus of the section of vertebrate paleon-
tology, also participated, until 1961. The work was possible through
the kind permission and assistance of Mr. and Mrs. Gordon E. Brown,
owners of the property.

The bird remains were associated with abundant bones of mammals
and a smaller representation of reptiles and amphibians. Most of the
specimens are of such size and condition as to indicate the probability
that the deposit was accumulated through pellets regurgitated by
ancient owls. It must be stated, however, that no bones of owls are
included. The casual intrusion of fragments of larger birds is assumed
to have come through predators that sheltered in the caves, or through
the activities of wood rats, abundantly represented among the small
mammals,

A complete report on the site prepared by Mr. Guilday (in press)
will contain a list of all the vertebrates, a detailed account of the
mammals, and a discussion of the entire fauna and its significance.
In the present account it is sufficient to state that the mammalian
remains include a number of boreal forms foreign to the area in
historic times, as well as four extinct species of the Pleistocene. These
indicate the probable age as near the end of Wisconsin time. The
birds support this assignment, as among them the spruce grouse and

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 145

the gray jay today are inhabitants of northern coniferous forests, and
the sharp-tailed grouse and the magpie also are northern and north-
western in modern distribution. None of the birds may be regarded
as typically southern since all the others identified are species that,
while found today as residents or migrants in Virginia, range widely
to the north. The presence of all at the end of the Pleistocene in
what Mr. Guilday has named the Natural Chimneys local fauna is
definitely of outstanding interest since this is the first extensive avian
fossil deposit reported for the State. The list includes 38 species,
with 2 others identified to genus. Fragmentary bits that could not be
named include several additional small passeriform species.

The bird bones are pale ivory to nearly white in color, except for
a few that are gray or blackish gray, due apparently to staining, as
none are mineralized. All are well preserved, only occasional ones
being friable or brittle. A few come from juvenile individuals, some
of them probably from young grouse, though this is not certain.

ANNOTATED LIST OF SPECIES
Family ANATIDAE: Ducks

Anas discors Linnaeus: Blue-winged Teal.

At least two individuals: Central section of right ramus of a
mandible, from the anterior end of the surangular forward to include
somewhat more than half of the dentary ; proximal ends of two right
humeri; left tarsometatarsus with the head missing.

Difference in size in the fragmentary wing bones indicates that
male and female birds may be represented. The part from the lower
leg is one with maximum development of the sculptured lines marking
the location of tendons and their attachment found in individuals
more than a year old.

This teal is recorded from several Pleistocene localities in Florida.

Bucephala albeola (Linnaeus): Bufflehead.

One individual: A left carpometacarpus, with the shaft of meta-
carpal III missing. This agrees in the details of length of the distal
symphysis, angle of anterior slope of metacarpal I, form of the facet
for articulation of the pollex, and angular compression of the inner
margin of the shaft of metacarpal III, with modern specimens.

The several Pleistocene records for the bufflehead include reports
from Oregon, California, and Florida.

Oxyura jamaicensis (Gmelin): Ruddy Duck.
One individual: Proximal two-thirds of a left humerus. The small

NO. 2 FOSSIL AND SUBFOSSIL BIRDS—WETMORE Ff

size indicates that the bird, which appears to have been adult, was a
female.

The ruddy duck has been identified in Pleistocene deposits in
Oregon, California, and Florida.

Family ACCIPITRIDAE: Hawks

Accipiter striatus Vieillot: Sharp-shinned Hawk.

One individual: A right carpometacarpus, complete, is from a
bird of small size that agrees in dimension with males.

The sharp-shinned hawk has been identified from the Pleistocene
of California and Florida, and from pre-Columbian cave deposits of
ancient but uncertain age on Great Exuma in the Bahama Islands.
Buteo jamaicensis (Gmelin) : Red-tailed Hawk.

One individual: A left femur, with broken shaft and some wear
on the proximal end.

The red-tail, widely distributed in modern time from northern
Canada to western Panama, has been found in several Pleistocene
localities in California and Florida.

Buteo lineatus (Gmelin): Red-shouldered Hawk.

One: Distal end of a right humerus, small in size.

The red-shouldered hawk, found in eastern North America from
Minnesota and southern Quebec to central México and Florida, and
west of the Rocky Mountains in California and Baja California, is
known from Pleistocene time in Florida and California.

Buteo platypterus (Vieillot): Broad-winged Hawk.

One individual: Distal third of a right tarsometatarsus, with the
trochlea intact. The specimen has the size of male birds.

Broad-wings nest in eastern North America from southern Canada
to Texas and Florida, and in the West Indies. There is one report
of the species from the Pleistocene of Florida.

Family TETRAONIDAE: Grouse

Canachites canadensis (Linnaeus): Spruce Grouse.

One individual, possibly more: Distal third of left humerus ; distal
two-thirds of left ulna; right tarsometatarsus complete. The humerus
in this species in length is similar to that of the ruffed grouse, but the
shaft is more slender, the internal condyle and the ectepicondyle are
slightly smaller, and the impression for the brachialis anticus is less
clearly outlined. The ulna is more slender, with the external condyle
smaller. The slightly shorter tarsometatarsus has the trochleae some-

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 145

what narrower, with the outer one swung more toward the center
line, so that support for the toes is narrower. Also the facet for the
articulation of the hind toe is of lesser size, and on the anterior face
the excavation below the head is smaller, with the tubercle for the
tibialis anticus shorter and less prominent.

This species definitely represents a boreal element in the fauna, as
in its modern distribution it is widely spread through the Canadian
zone forests from Alaska across Canada, south in the eastern half
of the United States only to northern Wisconsin, northern New
York, northern Vermont, northern New Hampshire, and Maine. The
present record is the first report south of these limits, as well as the
first from ancient time.
3onasa umbellus (Linnaeus): Ruffed Grouse.

Three or more individuals: Two premaxillae; proximal end of
two left humeri, and shaft and distal end of another; a left ulna; one
left coracoid, and the proximal end of another; one right carpometa-
carpus, and two others nearly complete ; distal half of a left tarsomet-
atarsus. The carpometacarpus is heavier than that of Canachites
canadensis, especially in the shaft of metacarpal III, and the inter-
metacarpal tuberosity is larger.

The ruffed grouse, common today in western Virginia, is known
from deposits of Pleistocene age in California, Tennessee, Maryland,
Pennsylvania, and Florida.

Pedioecetes phasianellus (Linnaeus): Sharp-tailed Grouse.

Four or more individuals: One partial premaxilla; a fragment
from the anterior end of a sternum; one right coracoid, somewhat
worn, head of another from the left side; heads of three left and
one right humeri, with distal ends of two from the left side, and one
from the right; one right carpometacarpus with the shaft of meta-
carpal III missing; and a fragment of the distal end of a right
tarsometatarsus. The head of the left humerus is distinctly larger
than any of the three from the right-hand side, so that it is certain
that it came from a fourth individual.

In modern time the sharp-tailed grouse has been a species of the
north and west, with a range that extends from north-central Alaska
across to central Quebec, south to eastern Oregon, in the mountains to
northern New Mexico, and east to Nebraska, Minnesota, and northern
Michigan. Formerly it ranged a little farther south to northeastern
California, western Kansas, and northern Illinois, areas from which
it has disappeared with agricultural use of the land, and increase in
hunting. The only previous report of the species east of this modern

NO. 2 FOSSIL AND SUBFOSSIL BIRDS—WETMORE 9

range is from bones of late Wisconsin age found by John I. Guilday
and his associates in Lloyd’s Rock Sinkhole in the New Paris Sink-
holes of Bedford County, western Pennsylvania. The present record,
about 120 miles to the south, is indication of a former range in the
late Pleistocene, and the period immediately following, through the
valleys of the northern Appalachian region.

The bird is known also from deposits of Pleistocene age at Fossil
Lake, Ore.

Family PHASIANIDAE: Pheasants, Quails

Colinus virgimianus (Linnaeus) : Bobwhite.

One individual: Head of a left humerus; a right femur, nearly
complete.

The bobwhite, of wide range in eastern North America, has been
found in the Pleistocene in Tennessee, and at several localities in
Florida.

Family MELEAGRIDIDAE: Turkeys

Meleagris gallopavo Linnaeus: Turkey.

Two individuals: The shaft of a left coracoid; the broken distal
end of a left tarsometatarsus. The two differ so definitely in size that
it is evident they are from separate birds.

Turkey bones have been recorded widely from Pleistocene time
in New Mexico, Illinois, Indiana, Tennessee, Arkansas, and Florida.

Family GRUIDAE: Cranes

Grus americana (Linnaeus): Whooping Crane.

One: Shaft and proximal end of a left coracoid. The bone is
fragmentary, with indications of the tooth marks of rodents, but
enough remains to indicate clearly that it is a crane, while the large
size identifies it as from the whooping crane.

This species, now much reduced in numbers, was reported in
eastern United States in the early days of European settlement from
New York, New Jersey, and South Carolina. The present record is
the first from ancient time north of Florida, where bones have been
found in Pleistocene deposits at three localities. It is also the only
report of this bird within the boundaries of present-day Virginia.

Family CHARADRIIDAE: Plovers

Charadrius vociferus Linnaeus: Killdeer.
One: Distal end of a right humerus.

IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 145

The killdeer has been recorded from the [llinoian stage of the
Pleistocene in Florida.

Family SCOLOPACIDAE: Snipe, Sandpipers

Philohela minor (Gmelin): American Woodcock.

One individual, possibly two: Proximal half of a left humerus;
a complete left tarsometatarsus. The leg bone appears to be from a
slightly smaller individual than the humerus.

The woodcock, found locally throughout Virginia, is reported from
a Pleistocene cave deposit in Florida.

Bartramia longicauda (Bechstein) : Upland Plover.

One: Right and left coracoids. These are identical in size and
color and may be from the same individual.

The upland plover, formerly common in Virginia, is now much
reduced in number. It has been found in late Pleistocene deposits in
Kansas.

Catoptrophorus semipalmatus (Gmelin): Willet.

One: Distal half of a right tarsometatarsus. The modern skeletons
at hand include a pair each of the two geographic races currently
recognized in this species. The humeri in these show the same differ-
ences in size that separate the birds in the flesh, or when preserved
as museum skins, the females in each being larger than the males.
It is significant to record that the humerus in the female of the sub-
species Catoptrophorus semipalmatus semipalmatus is appreciably
smaller than that of the male C. s. inornatus. The bone from Natural
Chimneys has the size of male inornatus and is identified as that race.
In modern times this subspecies nests through the western part of our
continent, but is common in migration and winter along the eastern
seaboard.

The only previous ancient record for the willet is from Pleisto-
cene deposits on the Newport Bay Mesa near the coast of southern
California.

Erolia minutilla (Vieillot): Least Sandpiper.

One: A complete right humerus, typical of this bird.

This is the first ancient report for the species, which now nests in
the north and spreads widely in migration, as far as Peru and central
Brazil.

Family COLUMBIDAE: Pigeons, Doves

Ectopistes migratorius (Linnaeus): Passenger Pigeon.
More than 21 individuals: 11 fragments of right humeri, and

NO. 2 FOSSIL AND SUBFOSSIL BIRDS—-WETMORE Tet

2 entire and 8 fragments of the left side; 1 entire and 3 fragmentary
ulnae from the right side, with 1 entire and 3 fragments from the
left side; 1 entire and 6 broken right carpometacarpi, with 4 frag-
ments from the left side; anterior ends of 11 sterna; 5 entire, 16 or
more fragmentary right coracoids, and 4 entire and 10 fragments
from the left side; anterior ends of 3 right and of 6 left scapulae ;
distal end of 1 right and of 2 left tibiotarsi; 1 entire and 3 partial
right tarsometatarsi, and parts of 4 from the left size.

From the abundance of these remains the passenger pigeon must
have been common and easily taken, probably from a roost, if the
deposit of bones is accepted as an accumulation from cast pellets of
night-feeding owls. All the bones are from fully adult birds which
points to a gathering outside the nesting season. This species, now
long extinct, was abundant during the period of settlement in Virginia,
with extensive roosts recorded as late as 1872. It was last reported
in the State definitely in 1890, uncertainly in 1892.

Passenger pigeon bones have been found frequently in Indian vil-
lage sites of pre-Columbian age, and are recorded from the Pleistocene
in California, Tennessee, and Florida.

Family ALCEDINIDAE: Kingfishers

Megaceryle alcyon (Linnaeus): Belted Kingfisher.
One: Proximal half of a left humerus.

There is one report of this kingfisher from the Pleistocene of
Florida.

Family PICIDAE: Woodpeckers

Colaptes auratus (Linnaeus): Yellow-shafted Flicker.

One: Distal half of a right tarsometatarsus.

The occurrence at Natural Chimneys is listed under the name of
the eastern species of the genus, following the modern geographical
ranges of these woodpeckers. But it should be noted that in available
skeletons there appear no trenchant characters on which the three
species of Colaptes of the A.O.U. Check-list may be separated.

In the eastern region of North America flickers have been reported
from three localities in the Pleistocene of Florida.

Centurus carolinus (Linnaeus): Red-bellied Woodpecker.

One: A left tarsometatarsus, complete.

The species is recorded from the Pleistocene of Florida.
Melanerpes erythrocephalus (Linnaeus): Red-headed Woodpecker.

Two individuals: A right humerus, complete, and another from the

I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 145

left side without the head; a right tarsometatarsus, complete. The
humeri are not a pair as they differ slightly in size.
There is one Pleistocene record for this species from Florida.

Dendrocopos pubescens (Linnaeus): Downy Woodpecker.
One: A right humerus with the distal end missing.
This is the first ancient record for this species.

Family TYRANNIDAE: Tyrant Flycatchers

Sayornis phoebe (Latham): Eastern Phoebe.

Three individuals: Two right humeri, and another from the left
side, all complete. Slight differences in size indicate that each bone
comes from a separate individual. The occurrence of this species is
one that would be expected from its habit of placing its nest on
sheltered projections on rock faces.

The record is the first one for this bird in ancient time.

Contopus virens (Linnaeus): Eastern Wood Pewee.

One: A complete left humerus. This agrees with the wood pewees,
and is listed as above on geographic grounds.

It is the first report of this group in prehistoric time.

Family HIRUNDINIDAE: Swallows

Petrochelidon pyrrhonota (Vieillot): Cliff Swallow.

Eight or more individuals: A series of humeri that includes
two complete and two fragments from the right side, and four com-
plete and three additional segments from the left.

The humerus in this species is approached in size among our
smaller swallows by the tree swallow, but has the head slightly larger
and the shaft heavier. The other species concerned are all distinctly
smaller.

The relative abundance of bones of this species compared to those
of other of the small birds indicates a nesting colony, a supposition
that appears to be verified by one bone with the porous structure of
the head typical of immature individuals not fully grown.

Cliff swallow bones are reported from the Pleistocene of California.

Family CORVIDAE: Jays, Magpies, Crows

Perisoreus canadensis (Linnaeus): Gray Jay.

One: A right tarsometatarsus with the trochlea for the fourth
digit missing, but otherwise complete. More slender form, greater
outward slant of the external face of the talon, relatively smaller

NO. 2 FOSSIL AND SUBFOSSIL BIRDS—-WETMORE 13

trochleae, and more widely open groove on the anterior face of the
head between the external and internal cotylae, identify this bone
in the gray jays from species of similar size of the genera Cyanocitta
and Aphelocoma.

This is another bird that is found in modern times in the coniferous
forests of the north and northwest, with extension southward only
along the higher mountains of the west. In much of this area it
ranges in the same regions as the spruce grouse, also its companion
in ancient Virginia.

The present record is the first report of the gray jay in the
prehistoric period.

Cyanocitta cristata (Linnaeus): Blue Jay.

One: A complete left humerus.

The widely ranging eastern blue jay is reported from the Pleisto-
cene of Florida.

Pica pica (Linnaeus): Black-billed Magpie.

One: Proximal half of a left humerus.

This record is one of particular interest since, though the magpie
in the Old World is spread from western Europe across northern
Siberia, in North America it has been restricted to the western half
of the continent. The find in Virginia indicates an early distribution
to the eastward, with subsequent withdrawal westward, a circum-
stance without apparent explanation. Many magpie bones have been
found in caves and other ancient deposits throughout Europe, but
the present find is the first report from America, since Dr. Brodkorb
informs me that a record for it from the lower Pleistocene of Randall
County, Tex., refers to another species.

Family SITTIDAE: Nuthatches

Sitta canadensis Linnaeus: Red-breasted Nuthatch.

One: A left humerus, complete.

This nuthatch is present in Virginia now as a breeding species
wherever spruce forest remains on the higher mountains, and as a
winter visitor from the north.

It is recorded from deposits of late Pleistocene age in California.

Family MIMIDAE: Mockingbirds, Thrashers

Toxostoma rufum (Linnaeus): Brown Thrasher.
One: A complete right humerus.
This is the first report of this bird in the prehistoric period.

I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 145

Family TURDIDAE: Thrushes, Bluebirds

Turdus migratorius Linnaeus: Robin.

One or more: A premaxilla; a complete right humerus, and one
from the left side with the head missing. The wing bones are of
maximum size for this species.

The only other ancient record for the robin is from the late Pleisto-
cene of California.

Hylocichla sp.: Thrush.

One: A left humerus complete. This comes from one of the
smaller species of this group. It is not the wood thrush, which is
larger, but except for this, it is not practicable to indicate relationship,
since the related species may not be separated from one another on
the basis of this single bone.

Family ICTERIDAE: Meadowlarks, Blackbirds, Orioles

Agelaius phoeniceus (Linnaeus): Red-winged Blackbird.

Two individuals: Right and left humeri with the heads broken.
These differ in size so that they come from two individuals.

The species is known from the Pleistocene of Ontario and Florida.

Molothrus ater (Boddaert): Brown-headed Cowbird.

One or more: Right and left humeri of such similar size that they
may be a pair.

This is the first ancient record for the species.

Family FRINGILLIDAE: Grosbeaks, Finches, Sparrows, Buntings

Junco sp.: Junco.

One: A complete right humerus.

While this agrees with the slate-colored junco it is not practicable
to make a specific identification among the several species of similar
size in this genus.

Zonotrichia albicollis (Gmelin) : White-throated Sparrow.

One: A complete left humerus.

This is the first ancient record for this species.
Passerella iliaca (Merrem): Fox Sparrow.

One: The symphysis of a lower mandible. This agrees in full
detail with modern skeletons. The form of the thickened inner margin
of the anterior end of the dentary, smooth and rounded when viewed
from above, and shelflike when seen from below, is characteristic of
this species. The bone is similar to that of the small-billed eastern
subspecies.

NO. 2 FOSSIL AND SUBFOSSIL BIRDS—-WET MORE I5

Fox sparrows have been recorded from two Pleistocene localities
in California.
Melospiza melodia (Wilson): Song Sparrow.

One: A right humerus.

The widely ranging song sparrow is reported from the Pleistocene
of California.

V. BONES OF BIRDS FROM COCKROACH ISLAND, BERMUDA

In November 1958, David B. Wingate forwarded a considerable
collection of bones from Bermuda, collected on Cockroach Island,
located in Harrington Sound off the base of Abbott’s Cliff. Most of
these specimens were dug from about 4 cubic feet of sandy soil and
rubble, some of them from near the surface where they were among
roots of plants. Many are of young birds, ranging from nearly adult
to half or even one-third grown indicating a breeding colony. In
careful digging no associated skeletons were encountered, so that
the site was one where separate bones had accumulated.

While the age of these specimens is unknown, the material prob-
ably is Recent, though, with one exception from the pre-Columbian
period. The few remains of the white-tailed tropicbird obviously are
of modern age. The uniform pale brownish-white cast in all the other
material indicates a deposit of some antiquity, though whether this
is of hundreds of years or of a longer period remains uncertain.
A few molluscan shells that accompanied the bones have been identi-
fied by Dr. J. P. E. Morrison of the National Museum as Poectlo-
sonites bermudensis Pfeiffer, a living species that in time ranges back
to deposits of Pleistocene age.

There have been several reports of bones of birds from caves in
the Bermudas but usually without identification, the earliest account
that I have seen being that of Nelson (1840, p. 113). In view of
the small amount of definite information on such deposits in Ber-
muda, I have prepared the brief account of the collection made by
Mr. Wingate which follows.

Family PROCELLARIIDAE: Shearwaters, Fulmars
PUFFINUS LHERMINIERI Lesson: Audubon’s Shearwater

Puffinus [sic] lherminiert Lesson, Rev. Zool., vol. 2, No. 3, April (May), 18309,
p. 102. (Guadeloupe, Lesser Antilles.)

Puffinus parvus Shufeldt, Ibis, ser. 10, vol. 4, No. 2, Oct. 2, 1961, p. 632. (Recent
deposits in the bone caves of Bermuda.)

The few bones of this species include humeri, radii, ulnae, meta-

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 145

carpals, coracoids, a femur, tibiotarsus, tarsometatarsi, a sternum,
and parts of a skull, that probably represent half a dozen individuals.
While the wing and leg bones may be sorted in two groups one of
which is slightly smaller than the other, it is seen on close scrutiny
that the specimens of lesser size all are obviously immature, some
of them quite young. It is my opinion therefore that the smaller size
in these is due to their not having attained full growth.

Shufeldt (1916, p. 632) in study of a collection of cave bones from
Bermuda noted two apparent size groups and named the smaller one
Puffinus parvus. While I have not had opportunity as yet to examine
his material, the plates that he published in a later account (Shufeldt,
1922) do not appear to substantiate his claims, particularly since at
the time he had available only one skeleton of Puffinus lhermintert
in the U. S. National Museum for comparison. This individual is
near the maximum size for the species. His smaller specimens as
illustrated show no differences in size from the range of variation
found in the series now available, particularly when it is under-
stood that all Shufeldt’s illustrations are not natural size, though so
indicated in the legends. I regard parvus, therefore, as a synonym of
lherminiert.

Puffinus mcgalli Shufeldt (1916, p. 630; 1922, p. 354), based on
a nearly complete sternum, appears to be an example of Puffinus
puffinus, as the figures agree exactly with a sternum of a female
Puffinus puffinus puffinus, No. 227465 in the U. S. National Museum
collections.

PTERODROMA CAHOW (Nichols and Mowbray): Bermuda Petrel

Aestrelata cahow Nichols and Mowbray, Auk, vol. 33, No. 2, April (March 31),

1916, p. 194. (Southeast side of Castle Island, Bermuda.)

Aestrelata vociferans Shufeldt, Ibis, ser. 10, vol. 4, No. 4, Oct. 2, 1916, p. 633.

(Bermuda. )

The greater part of the bones in the present collection are those
of this species, including abundant representation of wing and leg
bones, parts of 12 skulls, 12 sterna, 23 furculae, several coracoids,
scapulae and parts of more than 14 pelves. The indication is that
more than 25 individual birds are represented. About one-half come
from young birds that range from one-third grown to full size, but
the latter with the ends of some of the long bones still spongy. The
indication is clear that the site where the bones were found was a
breeding colony of this petrel, formerly abundant in Bermuda.

The adult bones all agree in detail with the modern skeletons of
the cahow in the U. S. National Museum.

NO. 2 FOSSIL AND SUBFOSSIL BIRDS—-WET MORE 17

Family PHAETHONTIDAE: Tropicbirds
PHAETHON LEPTURUS Daudin: White-tailed Tropicbird

Phaéton lepturus Daudin, in Buffon, Hist. Nat., ed. Didot, Quadrupédes, vol. 14,
1802, p. 319. (Mauritius. )

The right and left ulna, right and left radius, carpometacarpus,
and scapula that represent this species are obviously modern in ap-
pearance, and are believed to represent an intrusion in the older
deposit. It is probable that all come from one individual as the
duplicate elements are paired.

LITERATURE CITED

NE son, R. J.
1840. On the geology of the Bermudas. Trans. Geol. Soc. London, ser. 2,
vol. 5 (1837), pp. 103-123, figs. 1-16, map.
SHUFELDT, R. W.
1916. The bird-caves of the Bermudas and their former inhabitants. Ibis,
ser. 10, vol. 4, No. 2, Oct. 2, pp. 623-635, pl. 20.
1922. A comparative study of some subfossil remains of birds from Ber-
muda, including the “cahow.” Ann. Carnegie Mus., vol. 13, Nos.
3-4, Apr. 25, pp. 333-418, pls. 16-31.

DIVISION OF BIRDS
SMITHSONIAN INSTITUTION

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 145, NUMBER 3

THE PROBLEM: OF THE, VIDUINAE IN
THE LIGHT? /OP- RECENT PUBLICATIONS

By
HERBERT FRIEDMANN

Director, Los Angeles County Museum

an Rate IN res
f RE Ao

(GTN

(Pustication 4506)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JULY 20, 1962

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 145, NUMBER 3

THE PROBLEM OF THE VIDUINAE IN
THE LIGHT OF RECENT PUBLICATIONS

By
HERBERT FRIEDMANN

Director, Los Angeles County Museum

©#2eee06e°

(Pustication 4506)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JULY 20, 1962

PORT CITY PRESS, INC.
BALTIMORE, MD., U. S. A.

THE PROBLEM OF THE VIDUINAE IN THE
LIiGH POP RECENT: PUBEICATIONS

By HeErsert FRIEDMANN

Director, Los Angeles County Museum

While my study of the parasitic weaverbirds (1960) was in press,
an important paper by Steiner (1960) appeared. Although his at-
tention was centered largely upon the waxbills and their allies (the
spermestids of his paper; estrildids of mine), he briefly discussed
the systematic position of the Viduinae and their relationships with
the waxbills and came to conclusions different from my own. Inas-
much as Steiner’s experience and thinking concerning the waxbills
were both prolonged and extensive, it is necessary to consider his com-
ments carefully and objectively, even though I am still of the
opinion that to accept them poses more difficulties than it solves.

The various recommendations made by Steiner and others prior
to 1959 were reviewed in my account (1960, pp. 3-9), where a
consideration of their not altogether harmonious contents led me to
conclude that it was more nearly correct and acceptable to keep the
waxbills and their allies in the Ploceidae than to erect a separate
family for them. It was recognized that there were substantial argu-
ments for recognizing a separate family for the estrildines, but there
were equally suggestive ones for keeping them as a subfamily of the
Ploceidae. One could not lightly overlook the conclusion that they
constitute a distinct family arrived at by two of their most careful
investigators, Steiner and Nicolai, under conditions of aviculture.
On the other hand, Chapin’s very extensive field acquaintance with
many of the included genera and species and his interest in the classi-
fication of the whole assemblage caused him to consider them as one
family. In his last extensive treatment of a good portion of the
whole weaverbird complex, Chapin (1954, pp. 286-287) has this
to say:

The three most highly specialized subfamilies are believed to be the Pas-
serinae, Ploceinae, and Estrildinae. The most primitive group of all is the
Bubalornithinae, which at one time I believed should be treated as a distinct

family. In 1925 Peter Sushkin convinced me that the Plocepasserinae are dis-
tinctly intermediate between the buffalo-weavers and the sparrows, and he

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 145, NO. 3

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 145

regarded Sporopipes as fairly close to the ancestral line of both Ploceinae and
Estrildinae.

I still find it difficult to visualize a possible common ancestor for these two
subfamilies. Sushkin considered the Vidua group to be fairly close to the
Estrildinae, yet showing some rather primitive characters in their anatomy. I
have always felt that the Viduinae, now commonly raised to subfamily rank, are
closely allied to the Estrildinae, of which they appear to be always nest parasites.
They share the curious mouth markings and gape wattles of nestlings, and these
were not acquired independently, in my opinion, by the Viduinae through
mimicry.

None of the characters that have been cited for the recognition
of the Estrildidae is completely trenchant, and none is wholly con-
stant. While it is true that the estrildines show no seasonal plumage
dimorphism, which many of the ploceids do have, there are numbers
of the latter group that agree in this respect with the waxbills.
Among such examples may be cited such genera as Amblyospiza,
Bubalornis, Dinemellia, Histurgops, Malimbus, Passer, Petronia,
Philetairus, Phormoplectes, Plocepasser, Ploceus (many _ species,
especially of the subgenera Heteryphantes, Hyphanturgus, Icteropsis,
Melanoploceus, Melanopteryx, Otyphantes, and Xanthoploceus, al-
though many other species have marked seasonal plumages in the
adult males), Sorella, Sporoptpes, and Symplectes. As shown in my
1960 summary, the presumed behavioral differences are also not con-
stant and therefore they cannot be looked upon as trenchant systematic
criteria. It may seem that the point at issue is a very minor one—
whether we have two closely related families or two subfamilies of
one family—but the difference in the status of the two is supposed
to reflect something of the closeness or remoteness of their relation-
ship, and this is important.

The recognition of a separate family Estrildidae, based on ad-
mittedly “average,” nontrenchant characters, would result in either
of two unfortunate situations. If the viduines were to be included
as a specialized subfamily of the waxbills, the supposed criteria of
the family would break down completely. If the viduines were not
included, but were left as a subfamily of the Ploceidae, they would
then be separated systematically from the birds to which they seem
most closely allied. The closeness of their affinity to the waxbills
appears to be agreed upon by most students of the viduines—Chapin,
Delacour, Friedmann, Sushkin, and others. For that matter, Steiner,
who places them as a subfamily of the Ploceidae and recognizes a
separate family for the waxbills and their relatives, admits that the
widowbirds developed reflection globules and buccal patterns essen-

NO. 3 THE PROBLEM OF THE VIDUINAE—FRIEDMANN 3
tially similar to their estrildine hosts, “. . . auf Grund wirklicher
Verwandtschaft ... ,”’ and he considers the Viduinae as the sec-
tion of Ploceidae nearest to the Estrildidae.

The immediate problem of uppermost concern to me was, and still
is, how to interpret most cautiously and most accurately the parasitic
breeding habit of the viduines, and it was obvious that to do so
entailed an appreciation of the degree of their phylogenetic affinity
to their chief hosts, the waxbills forming the estrildine group.

If the two groups, Viduinae and Estrildinae, were considered as
closely related and as stemming from a common ancestral stock, the
striking similarity in the mouth markings and reflection globules of
their nestlings could be interpreted readily as something retained by
both from the stock from which the two groups bifurcated. If, how-
ever, the two groups were looked upon as not so closely related
and as not derived from a common ancestry, this important feature
of their young would have to be treated as a parallel development,
and quite probably as an adaptive one on the part of the parasitic
Viduinae. This is, in fact, what Steiner concludes when he writes
(translation mine) that “in the viduines, as a specialized small sub-
family of the ploceines, we have nothing else but a case of true
mimicry, which, in the imitation of the mouth markings, is not more
astonishing than are other known examples in insects, snakes, and
other creatures, and which have developed in the viduines in place
of the complicated reflex behavior of nestlings of other brood para-
sites . . . ,” such as the evicting behavior of young cuckoos of some
species, and the deliberate and usually lethal attacks by newly hatched
Indicators on their nest mates. Steiner expressly calls the mouth
markings a “spermestid character” in the viduines, and he considers
that in any evaluation of them a decisive role would have to be as-
signed to the thought that the viduines obtained or developed
“through true relationship, in their 6 or 7 species, various distinct
mouth-markings similar to those of their similarly distinguishable
host species—Pytilias, Granatinas, Lagonostictas, and Estrildas. This
would presume that each of their species had developed with its co-
ordinated host species from a primitive form, which, in retrospect,
must be assumed to have had a disclosed value for each presumed
parasite-host pair of species.” As I pointed out in my account, this
point of view has also been stated by Southern (1954), who accepted
the opinion that the viduines were extremely specialized brood para-
sites, each species being practically an obligate parasite of a single
species of estrildine host to which it was thought to be permanently

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 145
attached by virtue of a “very complicated form of mimicry. .. .”

The great difficulty in accepting this appraisal of the host-parasite
situation lies in the fact that the several species of Vidua are not each
rigidly restricted to single species of hosts. Of some of these birds
our knowledge is still very scanty (or even wanting), but of others,
such as . macroura with 18 recorded kinds of hosts, V. regia with 7,
V. chalybeata with 2, and Steganura with 9g, the available data cer-
tainly contradict any postulated rigid host specificity. To account for
the development of nestling mouth markings similar to those of the
host species would necessitate, as Steiner himself outlined, a strictly
limited host—parasite specificity, and this we do not find to be the
case. It is true that each of the species of viduines does appear to
have a single most-favored host, but the percentage of deviates from
it is too great to ignore. Thus, of the best known species, Vidua
macroura, I was able to assemble data on 77 records with 18 species
of hosts, and of these more than three-quarters were of IO species
of waxbills of the genus Estrilda and more than half were of the
races of a single species, Estrilda astrild. However, the different
species of waxbills differ as much in their mouth markings among
themselves as do the species of Vidua. If, as Steiner implies, the
mimetic similarity of buccal patterns of each species of parasite and
its normal host can only be looked upon as having an importantly
selective survival value, we would expect a considerably higher ad-
herence to the specific host relationship it is supposed to serve.

It might be considered that there may have been such a rigid host
selection originally and that subsequently the parasites broadened
their range of fosterers, but this would imply a subsequent denial
of an original, and ostensibly a continuing, selective force. In view
of the inconstant nature of the differences tabulated in support of
familial rank for the waxbills, and in view of the great difficulties
such an arrangement would make in interpreting the breeding biology
of the widowbirds, I still think it better to keep them all in one
systematic family group.

It has occurred to me that the above argument may make it seem
that the conclusions arrived at may imply something akin to a manipu-
lation of classification to simplify or to eliminate what would other-
wise be a perplexing problem, rather than to maintain a systematic
arrangement based purely on traditional characters, and to let the tan-
gential problem continue to perplex us if need be. This is not the
case, as the characters advanced by the proponents of familial rank
for the waxbills are not constant, on the one hand, and the mouth

NO. 3 THE PROBLEM OF THE VIDUINAE—FRIEDMANN 5

markings of the nestlings are also valid morphological characters in
themselves. The fact that these buccal patterns may be functional as
well as morphological, and hence to some extent possibly subject to
the pressure of natural selection, need not rule out the possibility,
the probability even, that they are also phylogenetically stable charac-
ters, useful as indicators of relationship. This idea is by no means
novel at this point, nor was it in my 1960 discussion, where (p. 24)
I pointed out that Morris (1957, p. 199) concluded that these mouth
markings were conservative taxonomic characters and as such were
useful aids to understanding the evolution of the birds that have
them.

Nicolai (1961) has recently published in abbreviated form the re-
sults of a study of the vocalization of several species of Vidua
under aviary conditions. He studied with a tape recording the sounds
produced by V. macroura, V. regia, V. chalybeata, and Steganura
paradisaea and reported that part of the notes of each was a fairly
accurate copy of the song of their host species. He stated that the
viduine sounds comprised a “weaverbird-like” series of notes, scarcely
distinguishable in the four species, and a series of loud notes and
songs of the respective host species (various species and races of
Estrilda). Nicolai found in the ploceids and estrildids closest to the
viduines all songs and notes to be consistently innate and nonvaria-
ble, and he concluded that probably the notes of the viduines were
similarly somewhat “‘fixed.”’ He went on to speculate that the young
Viduinae probably acquired their vocabulary from their foster parents
during their period of dependency in and out of the nest. Only in
this way did he think the exclusive reproduction of the vocabulary
of the particular host species could have been made possible. Fur-
thermore, he pointed out that in the case of V. macroura, which is
known to parasitize a number of species of Estrilda, each male had
invariably only the notes of one host species. There were no cases
of mixed songs, a fact which he considered in agreement with his
premise as to how the imitative process could have taken place. On
the other hand, Nicolai further contended that the “whispering nest
notes” of the male, which appear in the vocalization of V. regia
and V.. chalybeata, were learned somewhat later, after the birds had
become self-sufficient and no longer were in constant contact with
their fosterers, when the latter began preparing to breed again and
began nest building anew.

Nicolai further concluded that whereas, at the close of the period
of parental dependency, the young of other, self-breeding, passerines

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I45

might go through what seemed like playing at nest building or play-
ing at heterosexual pursuit, the young parasitic widowbirds were
interested in watching the breeding preparations of their fosterers.
The precise observations they made and the degree to which they
seemed to incorporate these impressions were thought to become im-
portant later in their lives in helping to synchronize their reproduc-
tive cycles and activities with those of their hosts, and so to be-
come significant in the breeding success of the widowbirds.

Inasmuch as Nicolai’s work has not yet been published with suffi-
ciently detailed documentation, it is somewhat difficult to appraise
and to criticize his conclusions. The following comments must be
read with this in mind, and some doubts that are raised here may prove
to be baseless. I must stress that the observations, surprising as
they seem to me, merit serious and respectful consideration. Their
interpretation seems to be less certain.

For one thing, in a state of captivity birds may sometimes do
things they would have little chance of doing or, as far as we know,
do not do, in a wild state. I do not know whether Nicolai’s birds
had the presumed fosterers with them in the cage or in nearby cages
where they could hear them. If they were not actually raised in
captivity by these fosterers, one wonders how Nicolai could know
which was the foster parent species in each instance, unless he as-
sumed the most likely one from the total recorded literature (as
was brought together in my book), or unless he assumed the iden-
tity of the host from the vocabulary of the parasite. The latter
would be a matter of circular reasoning which would hardly be con-
vineing, and which I cannot believe was done. Yet this was the way
in which some of Neunzig’s original (1929) conclusions seem to have
been achieved.

I am wholly convinced that it is possible to learn many things,
including vocalizations, from captive birds that it would be very
difficult to learn in the free state, but I am still surprised that no
one ever reported any constant and marked specific differences in
the notes of the various species of Vidua in Africa. Although my
own fieldwork is now many years past, and I do not pretend to re-
member accurately the songs and calls of these birds, I can find no
mention in my journals of any marked differences between them,
and I have found no published observations of others to this effect.
This suggests that the differences noted in aviary birds are not suffi-
ciently striking to be obvious in the field but require close-up ob-
servation for their discrimination. As a matter of fact, the vocali-

NO. 3 THE PROBLEM OF THE VIDUINAE—-FRIEDMANN 7

zations of the various host species of the genus Estrilda, as de-
scribed in the literature, are all quite similar, or at least their
specific patterns vary only slightly among themselves. This does not
mean that the differences are less real, but I cannot dispell the
thought that these portions of the songs resembling the notes of
the presumed host species may have been due to the limiting condi-
tions of the aviary, whereas the “weaverbird-like” notes common to
all four species agree with what is known of their calls in the state
of nature.

The very abbreviated form in which Nicolai’s data were reported
caused them to appear to imply further evidence for a definite host—
parasite relationship, but this is not actually implicit in them. We
are not informed how many individuals of each species of viduine
were observed or under what conditions. Thorpe’s (1958) work on
the learning of song patterns by small passerine birds, especially the
chaffinch, has indicated that the learned, as opposed to the innate,
pattern of song is restricted to the “first 13 months of life and towards
the end of this time there is a peak period of learning activity of a
few weeks during which a young Chaffinch may learn, as a result
of singing in a territory, the fine details of as many as six different
songs.” If Nicolai’s assumption is correct, that the young parasitic
widowbirds learn the utterances of their foster parents during the
first two or three weeks of life, they are apparently more precocious
than chaffinches in this respect. Furthermore, we may recall that
in the case of parasitic cowbirds and cuckoos there is no sign what-
ever of the young learning any of the vocalisms of their fosterers.
This cannot be looked upon as meaning that the same situation
necessarily is true for the parasitic weavers, but judgment must be
delayed until evidence is forthcoming. If eventual fuller publication
of Nicolai’s work should convince us that the viduines enhance their
reproductive potential even very slightly by vocal mimicry of their
common hosts, we would have to admit an unexpected uniqueness in
these birds.

Another study that appeared too late for me to discuss in my ac-
count was Ziswiler’s (1959) paper on features of the ontogenetic
development of the waxbills. While presenting some data on the
relative lack of sensitivity of later developmental stages to increas-
ingly long interruptions of brooding, and also some data on the
postembryonic (i.e., nestling) growth curves of several species,
Ziswiler does not concern himself with the viduines at all; he does not
even mention them. His paper therefore gives us no opinions to eval-

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 145

uate in the present connection. He does consider the waxbills a sys-
tematic family, but he gives no arguments or data either supporting or
contradicting this treatment. The data he does present are not given as
systematic criteria and show nothing peculiar to the ‘““Spermestidae.”

The problem as to which of the numerous described species or
races of the combassous are really valid still awaits an answer based
on much more extensive and more complete knowledge of them in
the field. From my own field studies of many years ago and from
much more recent examination of large numbers of museum speci-
mens I arrived at the arrangement given in my 1960 publication.
However, almost simultaneously, Wolters (1960) proposed a some-
what different treatment, based in part on observations of aviary
birds. These differences are not particularly important, as no one
has the data on which to formulate a completely convincing and
wholly satisfying classification, but they do point out that until such
information is assembled, all our judgments can have only limited
validity. In our understanding of the combassous, as contrasted with
the present knowledge of the long-tailed viduas, we are still con-
fronted with the species of the systematists rather than the species
of the naturalists. This is bound to continue until the living birds
are studied much more thoroughly, as further examination of their
preserved corpses will only lead to divergent and inconclusive
arrangements.

Still more recently, Wolters (1961) has published an arrangement
of the viduines in which the short-tailed species (subgenus
“Hypochera’) are placed at the top, whereas I put them at the base
of the group. Wolters considers the absence of elongated rectrices
in the breeding plumage of adult males to be a secondarily arrived
at condition, and that the long-tailed species (subgenus Vidua
proper) are to be looked upon as representing the original, ancestral
character of the group. Also, he suggests that Steganura is the basic
or primitive member of the viduines, whereas I placed it at the apex
of the assemblage. While it is obvious that each of us came to our
respective conclusions on the basis of all the evidence we could
muster, it now becomes clear that, in the absence of any really con-
clusive data, these alternate, and, in fact, opposite, arrangements can
only be looked upon as interpretations of the purely circumstantial
evidence afforded by the appearance and the habits of the existing
species. Actually the two classifications agree closely in the relative
placement of the included species and genera, but differ in their over-
all orientation.

NO. 3 THE PROBLEM OF THE VIDUINAE—FRIEDMANN 9

In defense of the arrangement proposed in my book I can only
repeat here what I outlined there, namely, that inasmuch as rectricial
elongation in male nuptial plumage is a character that has developed
wholly independently in two of the main groups or sections of the
family, it seems probable that within each of these groups the short-
tailed species are nearer the stock from which they evolved than are
their long-tailed relatives. There is nothing in the life histories of
the short-tailed species to suggest that they are in any way more ad-
vanced than their congeners with elongated rectrices; in fact, the
reverse is more in keeping with our still all too incomplete informa-
tion. The courtship antics of the combassous are simpler, less in-
volved, apparently more primitive than are those of the long-tailed
species. All the viduines are quite similar in their vocalisms and, ex-
cept for size (in Steganura), in the appearance of their eggs. It is
perhaps a necessary commentary on so much of our present avian
systematics to end this discussion with the observation that the one
point of agreement in all these attempts is that we need to know
more about the birds themselves.

LITERATURE. CITED

CHAPIN, JAMES P.
1954. The birds of the Belgian Congo. Part IV. Amer. Mus. Nat. Hist.
Bull. 75B, 846 pp.
FRIEDMANN, HERBERT.
1960. The parasitic weaverbirds. U. S. Nat. Mus. Bull. 223, 196 pp., 16 pls.
Morris, DESMOND.
1957. The reproductive behaviour of the bronze mannikin, Lonchura cucul-
lata. Behaviour, vol. 11, pp. 156-201.
NEUNZIG, RUDOLF.
1929. Zum Brutparasitismus der Viduinen. Journ. fiir Orn., vol. 77, pp.
I-21, 2 pls.
NIcoal, J.
1961. Die Stimmen einiger Viduinen. Journ. fiir Orn., vol. 102, pp. 213-
214.
SouTHERN, H. N.
1954. Mimicry in cuckoos’ eggs. In Evolution as a process, ed. J. Huxley,
A. C. Hardy, and E. B. Ford, pp. 219-232.
STEINER, Hans.
1960. Klassifikation der Prachtfinken, Spermestidae, auf Grund der
Rachenzeichnungen iherer Nestlinge. Journ. fiir Orn., vol. ror,
Pp. 92-112.
TuorpPe, Wn. H.
1958. The learning of song pattern by birds, with especial reference to
the song of the chaffinch, Fringilla coelebs. Ibis, vol. 100, pp. 535-

570.

10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 145

Wotters, H. E.
1960. Zur Systematik der Atlasfinken (Untergattung Hypochera der
Gattung Vidua, Viduinae, Estrildidae, Aves). Bonner Zool. Beitr.,
vol. 1, pp. 19-25.
1961. Uber die verwandschaftlichen Beziehungen der Arten der Viduinae.
Vogelring, vol. 30, pp. 12-15.
ZISWILER, VINZENZ.
1959. Besonderheiten in der Ontogenese der Prachtfinken (Spermestidae).
Festschr. Steiner, Vierteljahrsschr. Naturf. Ges. Zurich, vol. 104,
pp. 222-226.

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 145, NUMBER 6

ADDITIONS TO RECORDS OF BIRDS
KNOWN FROM THE REPUBLIC
OF PANAMA

By
ALEXANDER WETMORE

Research Associate
Smithsonian Institution

(Pusiication 4523)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
DECEMBER 16, 1963

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 145, NUMBER 6

ADDITIONS TO RECORDS OF BIRDS
KNOWN FROM THE REPUBLIC
OF PANAMA

By
ALEXANDER WETMORE

Research Associate
Smithsonian Institution

iy ‘2
5ST SON ~
INGTON

2500000

(Pusiication 4523)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
DECEMBER 16, 1963

PORT CITY PRESS, INC.
BALTIMORE, MD., U. S. A.

ADDITIONS TO RECORDS OF BIRDS KNOWN
FROM THE REPUBLIC OF PANAMA

By ALEXANDER WETMORE
Research Associate, Smithsonian Institution

THE notes that follow, pertaining to recent studies on avian collec-
tions made in Panama, include descriptions of two species and two
geographic races not known previously. The two named from
Darién are based on specimens received from the Gorgas Memorial
Laboratory in Panama and derive from recent field work directed
by Dr. Pedro Galindo. Included with these are further records of
birds from the little-known island of Escudo de Veraguas, located
18 kilometers at sea off the base of the Valiente Peninsula, Bocas del
Toro, and report of two North American migrants not found
previously in the republic.

I. ADDITIONAL RECORDS FROM ISLA ESCUDO DE
VERAGUAS, WITH DESCRIPTION OF A NEW
SPECIES OF HUMMINGBIRD

In the course of a visit to Isla Escudo de Veraguas early in
March 1958, I collected a thick-spined rat (genus Hoplomys), the
first island record for this group, and a race that proved to be new
to science (Handley, 1959, pp. 9-10). Following its description,
Dr. C. O. Handley, Jr., of the U. S. National Museum, through
cooperation of the U. S. Army, came to the island in 1962 and lived
there in a shore camp from March 20 to 24. In addition to a series
of the rat, and many bats, he preserved in formalin a number of birds
caught in mist nets, and prepared a few others, shot for specimens,
as study skins. The 41 birds collected have added considerably to
earlier information on the avifauna, available from my own brief
visit four years earlier (Wetmore, 1959, pp. 1-27).

Migrants recorded by Dr. Handley include several that had not
been listed from the island previously. A belted kingfisher, taken
March 21, is the eastern subspecies Megaceryle alcyon alcyon.
Several eastern wood peewees (Contopus virens) were present and

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 145, NO. 6

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 145

in song on March 23, when one was taken for a skin. Several smail
groups of barn swallows passed on this same day, moving toward
the north. On March 21 single purple martins were reported at
intervals during the day in northward flight off shore. One was
recorded on March 19 on the airstrip at Fort Sherman, Canal Zone,
and another was observed March 20 at sea about 15 kilometers off the
mouth of Rio Belén, on the boundary between the provinces of
Colon and Veraguas. Swainson’s thrushes (Hylocichla ustulata),
taken March 21 and 24, and a red-eyed vireo (Vireo olivaceus), on
March 23, are identified to species only, as they were placed in
formalin. Other migrants, all in formalin, include the black-and-
white warbler (Mniotilta varia), on March 22, and the prothonotary
warbler (Protonotaria citrea), worm-eating warbler (Helmitheros
vermivorus), ovenbird (Seiurus aurocapillus), and northern water-
thrush (Seiurus noveboracensis), taken on March 21. A male
summer tanager (Piranga rubra rubra) in full breeding plumage,
prepared as a skin, was collected March 21, and another was re-
corded on the day following.

An immature yellow-crowned night heron, another addition to the
island list, appears to be the resident race of Panama, Nyctanassa
violacea caliginis, while a single green heron (Butorides virescens),
seen March 20 and 21, was believed to be a migrant. A pair of
pygmy kingfishers (Chloroceryle aenea aenea) caught in mist nets
set up near the lagoon March 24, form an interesting addition to
the island residents. Men with me in 1958, and those with Handley,
saw a small rail that was not collected, but from the description it
may have been the white-throated rail (Laterallus albigularis) which
is common on the mainland. George Barratt, with Handley, also
reported a night bird with batlike flight that probably was a species
of goatsucker.

Specimens in formalin of the endemic races of the manakin,
Manacus vitellinus amitinus, bay wren, Thryothorus nigricapillus
odicus, and blue-gray tanager, Thraupis virens caesitia, all show
clearly the decidedly larger size of the first two, and the heavier bill
found in the tanager when compared to birds of the adjacent main-
land. The same character of larger dimension is present in the
island form of the thick-spined rat, described by Dr. Handley, and is
in much greater evidence in the hummingbird, whose description
follows.

No. 6 RECORDS OF BIRDS FROM PANAMA—WETMORE 3

AMAZILIA HANDLEYI, new species T

Characters—In general appearance similar to Amazilia tzacail
tzacatl* but much larger, and darker in color; bill decidedly heavier ;
feet larger; brown of tail, upper and under tail coverts, and lores,
darker ; back and wing coverts decidedly darker and duller.

Description—tType, ¢ ad., U. S. Nat. Mus. 477282, from Isla
Escudo de Veraguas, collected March 22, 1962, by C. O. Handley,
Jr., and F. M. Greenwell (orig. no. 1188). Crown, hindneck, back,
and wing coverts (except the primary coverts) deep green, with a
sheen of dull bronze that changes on lower back and rump to a
darker shade with an iridescence of dull russet; upper tail coverts
liver brown; tail chocolate, edged and tipped with dull black; pri-
maries, secondaries, and primary coverts dull black with a faint
sheen of violet; a narrow line of chocolate on the lores; foreneck,
sides of neck, upper breast, and sides clear bright green, with some
of the throat feathers edged narrowly with dull white; a small tuft
of white feathers on the upper line of the sides near center; lower
breast and upper abdomen hair brown; lower abdomen and tibial
tufts white; under tail coverts walnut brown; edge of wing lined
narrowly with chocolate. Tips and sides of maxilla and tip of
mandible dull black; rest of bill dull reddish brown; bare lower
end of tarsus, toes, and claws dull black. (From dried skin.)

Measurements—Males (4 specimens), wing 67.5-68.7 (68.1),
tail 40.0-41.5 (40.6), culmen from base 24.4-27.6 (24.5) mm.

Female (one specimen), wing 67.1, tail 41.1, culmen from base
25.8 mm.

Type, male, wing 67.5, tail 40.0, culmen from base 27.6 mm.

Range.—Confined to Isla Escudo de Veraguas, off the base of
Peninsula Valiente, Bocas del Toro, Panama.

Remarks——During my visit to Escudo de Veraguas early in
March 1958 I had brief glimpses of hummingbirds at flowers among
low bushes back of the beach, but none came sufficiently near to
allow me to shoot any for specimens. As they turned in flight I had
brief glimpses of brown in the tail as in Rieffer’s hummingbird
(Amazilia t. tzacatl) common on the mainland, and these Escudo
birds were so identified and recorded (Wetmore, 1959, p. 6). During
the work of Dr. Handley five were captured in mist nets set for bats,
and with these in hand it was obvious immediately that while they

1Trochilus Tzacatl De la Llave, Registro Trimestre, vol. 2, no. 5, Jan. 1833,
p. 48. ( México.)

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 145

resembled Rieffer’s hummingbird in color pattern, they were so
much larger, especially in bulk of body and total length, and also
so much darker colored, that they were examples of an unknown
form. From careful examination the differences are of such a nature
that they must be considered as representative of a distinct species.
This is named for Charles O. Handley, Jr., in recognition of his
continuing interest in the avifauna during his field work concerned
with the mammals of Panama.

The five birds taken by Dr. Handley were preserved in formalin
and were prepared as skins by Mrs. Roxie Laybourne on their arrival
at the U. S. National Museum. Their much greater size was obvious,
but to make certain that the color differences were not due to the
preservative I placed a recently taken study skin of Amazilia tzacatl
tsacatl in the same fluid in which the hummingbirds from Isla
Escudo de Veraguas had been received. When dried after a month
of such immersion this specimen showed no change of any kind.
It is interesting to record that in this skin, and in the larger relative
here described, the feathers along the side of the neck when wet
were metallic reddish purple, a color that disappeared completely as
the specimens dried.

Amasilia tzacatl as a species maintains uniform size, within the
usual limits of individual variation, throughout a vast area from
eastern México, Central America, and Colombia to western Ecuador
and western Venezuela. The only variation apparent is in a buffy
wash on the abdomen in that part of the population found in south-
western Colombia and Ecuador on which birds of that section are
separated as a geographic race under the name A. ft. jucunda. It has
been noted above that the bird of Isla Escudo de Veraguas com-
pared with tzacatl differs in decidedly darker coloration and in much
larger size. It is clearly evident that the island group is of a stock
similar to that of the mainland, so that on first consideration it
would appear that they should be related as subspecies. The color
differences, while considerable, would not militate against this. But
the size difference in terms of bulk of body of the island bird is
so much greater—over 50 percent more than that of the mainland
group—with its complete isolation, make it reasonable to regard
handleyi as a separate species.

As stated in my earlier paper (Wetmore, 1959, pp. 3-4) it is
probable that the island had connection with the mainland during the
fluctuations in sea level of Pleistocene time so that the present
inhabitants among birds and mammals may have come to it during

No. 6 RECORDS OF BIRDS FROM PANAMA—-WETMORE 5

such periods. It is interesting that the hummingbird, the manakin,
the wren, and the rat all differ from present-day mainland relatives
in definitely greater size. In the tanager this distinction is also evident
but is restricted to the bill. Perhaps this species has come to the
island more recently than the others.

II. DESCRIPTIONS OF A WOOD-QUAIL AND A TYRANT
FLYCATCHER FROM THE SERRANIA DEL DARIEN

During part of June and July 1963, the Gorgas Memorial Labora-
tory, under arrangements directed by Dr. Pedro Galindo, established
three camps in the Serrania del Darién, in the vicinity of Cerro
Tacarcuna, to serve as bases for the investigation of this little-known
area. The birds collected included specimens of a beautiful wood-
quail, related to Andean mountain forms to the south but unlike any
of those known, and a flycatcher of a South American species not
recorded before from Panama. Descriptions of these follow.

Family PHASIANIDAE

ODONTOPHORUS DIALEUCOS, new species

Characters.—Generally similar to Odontophorus strophium (Gould)?
but with crown black; back and scapulars without white shaft lines ;
entire upper surface olive, with rufous only as a band on the hindneck ;
foreneck similar in the two white bands above and below, with the
space between mixed black and dull rufous; rest of lower surface
olive rather than rufous and cinnamon, without shaft lines or a black
collar below the lower white band; breast, sides, and flanks dull olive-
buff, finely barred and mottled with slaty black.

Description—tType, 3, U.S. Nat. Mus. 483327, from 1,450 meters
elevation, 64 kilometers west of the summit of Cerro Mali, Darién,
Panama, taken June 7, 1963, by Pedro Galindo (orig. no., Gorgas
Mem. Lab. 4-00384) : Crown black with slight, partly concealed mot-
tling of dull rufous, and tiny spots of white; a prominent white super-
ciliary streak that extends back of the eye; a band of hazel mottled
and lined with sooty black on the hindneck that laterally becomes
cinnamon-buff as it extends around to meet the posterior end of the
white superciliary ; back, rump, and upper tail coverts brownish olive,
finely barred and mottled with sooty black, with scattered faint spots
and indistinct bars of cinnamon; wing coverts, inner secondaries, and

2 Ortyx (Odontophorus) strophium Gould, Proc. Zool. Soc. London, vol. 11,
1843 (March, 1944), p. 134. (Bogota, Colombia.)

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 145

tertials snuff brown, barred and spotted finely with sooty black, and
lined and spotted sparingly with small, irregular marks of buffy white;
tertials with heavy, irregular markings of black, inner secondaries
barred broadly with black ; primaries fuscous, finely mottled with dull
cinnamon-buff on outer webs; a band of white across the upper fore-
neck extending at either side over the malar region beneath the eye,
and on the lower eyelid; a broad band of black mixed with Mars
brown and russet extending between the two white bands from the
lower cheeks across the middle foreneck and upper throat, changing
to dull black over the ear coverts; a broad band of white across the
lower foreneck; rest of lower surface dull buffy brown to tawny-olive,
heavily mottled with sooty black, spotted sparingly and indistinctly
with buffy white, becoming Saccardo’s umber, with slightly heavier
markings of black and cinnamon-buff on the flanks and under tail
coverts ; under wing coverts fuscous, sparingly and indistinctly spotted
with dull Saccardo’s umber. Bill, tarsi, and toes black (in dried skin).

Measurements.—Male (type), wing 129.5, tail 44.3, culmen from
base 19.8, tarsus 45.2 mm.

Female, wing 131.0, tail 46.7, culmen from base 19.6, tarsus 47.5
mim.

Remarks.—The male and female from which this bird is described
were taken together. The adult female is very slightly browner than
the male. This specimen has the lores and the superciliary area black
like the crown, with only a fine spotting of white. The chin also seems
to have had the white band considerably reduced by black (though
this can not be ascertained clearly as some of the feathers of this area
are missing.) The line of the culmen and the tip of the maxilla in this
bird are partly brown.

The discovery of this beautiful wood-quail, isolated in the higher
levels of the Serrania del Darién, adds another form to populations
of this genus with prominent markings of white on the head and neck.
It is most like Odontophorus strophium of the Bogota region of Co-
lombia, which has the foreneck similar, with a black center bordered
broadly with white above and below. This species differs, however,
in the presence of a narrow black collar on the neck below the border
of the lower white band. Also strophium is rufous and cinnamon on
the breast and sides, with prominent white shaft lines and spots, has
the crown fuscous-brown, and the whole upper surface rufescent
rather than olive, with heavier, more prominent markings. Odonto-
phorus columbianus (Gould) of the subtropical zone of the moun-
tains of northern Venezuela in general resembles strophium but has

No. 6 RECORDS OF BIRDS FROM PANAMA——-WETMORE 7

the entire foreneck white above the narrow basal black collar. Also it
is spotted along the sides with black, and heavily with white on the
breast. Odontophorus parambae Rothschild, found in the tropical zone
from west-central Colombia south to Ecuador, has a single white band
across the lower area of the black foreneck. Odontophorus leucolae-
mus, more remote, in Costa Rica and western Panama, has the entire
upper foreneck white, with the lower area and upper breast jet black.
And finally there may be noted Odontophorus atrifrons of the Andes
of northern Colombia and O. erythrops melanotis found on Cerro
Pirre and Cerro Azul in Panama, in which the foreneck is solid black.

All these are similar in size, form, and, so far as known, in habits,
so that it is reasonable to postulate common ancestry. Their present-
day differences in pattern of markings, coupled with variations in color,
may unite them in a super species, but these distinctions appear so
fixed and so definite that to group them as subspecies under one specific
name would conceal their interesting divergences.

The name for the species here described is from the Greek dialeukos,
marked with white.

Family TyRANNIDAE
ELAENIA CANICEPS ABSITA, new subspecies

Characters —Male, similar to that of Elaenta caniceps parambae
(Hellmayr)* but lighter, clearer gray above and across the breast;
whiter on throat and abdomen; partly concealed white area of center
of crown larger ; white edgings on lesser wing coverts more extensive.
Female, with pileum darker gray (around the white center) ; breast,
sides, and abdomen decidedly paler, less deeply yellow.

Description.—Type, ¢, U.S. Nat. Mus. 483342, from the old Tacar-
cuna Village site, headwaters of the Rio Pucro, 950 meters elevation,
on the base of Cerro Mali, Serrania del Darién, collected by Pedro
Galindo, July 4, 1963 (orig. no., Gorgas Mem. Lab. 3-00329). Crown
deep neutral gray, with an extensive, partly concealed central area in
which the basal two-thirds of each feather is pure white; a narrow
line of grayish white across forehead and upper edge of lores; back,
rump, and upper tail coverts neutral gray ; wings black, with the wing
coverts tipped, and the inner primaries, secondaries, and tertials
broadly edged with white; tail feathers mouse gray edged with neu-
tral gray, mainly toward base, and tipped narrowly with grayish white ;

3 Serpophaga parambae Hellmayr, Bull. Brit. Orn. Club, vol. 14, Feb. 27,
1904, p. 54. (Paramba, elevation 3,500 feet, Provincia de Esmeraldas, Ecuador.)

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 145

a very narrow line of white on the edge of both eyelids ; lores, side of
head below eye, and anterior auricular feathers neutral gray at the
tips and more or less white at the base; throat and upper foreneck
very pale grayish white ; lower foreneck, upper breast, and sides pallid
neutral gray ; flanks, abdomen, and under tail coverts pure white; edge
of wing white with a slight spotting of neutral gray; inner wing
coverts and edge of inner webs of primaries white; outermost wing
coverts white mixed with neutral gray. Bill dull black, with the base of
the gonys whitish; tarsus, toes, and claws black. (From dried skin.)

Measurements.—Male, type, wing 58.0, tail 49.4, culmen from base
10.5, tarsus 15,5 mm.

Female, wing 52.5, tail 42.8, culmen from base 10.2, tarsus 15.6 mm.

Range.—Known only from the upper Rio Pucro, at 950 meters
elevation on the base of Cerro Mali, Serrania del Darién, Panama.

Remarks.—A female, U.S. Nat. Mus. 483341, was taken with the
male at the same location, on July 4, 1963 (Gorgas Mem. Lab. no.
3-00328). This bird has the following colors: Crown slightly darker
than in the male, with the same partly hidden white center; upper
surface Kronberg’s green; tail feathers blacker than in the male,
edged lightly with dull green; light edgings on wing chartreuse
yellow ; side of head as in male; throat and upper foreneck duller
white; lower foreneck, breast, and sides washed lightly with vetiver
green; abdomen sea-foam green; under tail coverts chalcedony yel-
low ; lighter part of under wing coverts, and inner webs of basal part
of primaries like abdomen.

The male has been compared with the type of Elaema c. parambae
in the American Museum of Natural History. Through the kindness
of James Bond I have examined a female and two males of that race
in immature dress in the Academy of Natural Sciences of Philadelphia,
taken on the Rio Jurubida, inland from Nuqui near the central coast
of the Department of Choco, northwestern Colombia. The two marked
male in color are like the female. Compared with the female from
Cerro Mali the three from Nuqui are very slightly clearer green on
the back, with the base color of the crown faintly lighter gray. Below
they differ decidedly as the throat and upper foreneck are duller,
grayer, the lower foreneck, breast, and sides are much greener, and
the rest of the under surface is decidedly deeper yellow.

The specimens from the Cerro Tacarcuna massif in Darién mark an
interesting addition to the flycatchers known from Panama. As a
species, Elaenia caniceps ranges from Colombia and southern Vene-
zuela to Bolivia, northern Argentina, and southern Brazil. With the
present description four subspecies are recognized in this area.

NO. 6 RECORDS OF BIRDS FROM PANAMA—WETMORE 9

The name of the present race, the most northern population known,
is taken from the Latin absitus, in the sense of one living remote or
distant from its relatives.

III. A WESTERN SUBSPECIES OF THE PLAIN-COLORED
TANAGER

The plain-colored tanager, in its subspecies Tangara inornata lan-
guens Bangs and Barbour, is a common bird of the tropical zone in
Panama from the central lowlands eastward into Colombia. Through
recent work of the Gorgas Memorial Laboratory at a field station in
Bocas del Toro, I have received from Eustorgio Méndez three speci-
mens from Almirante that mark a considerable extension of range.
These prove to represent an undescribed race.

TANGARA INORNATA RAVA, new subspecies

Characters.—Similar to Tangara inornata languens Bangs and Bar-
bour,* but with throat, lower breast, abdomen, and under tail coverts
light buff to pinkish buff; a faint wash of the same color on lower
rump and upper tail coverts ; lesser wing coverts decidedly darker blue.

Description—Type, 3, U.S. Nat. Mus. 483344, from Almirante
(Milla 2), Bocas del Toro, Panama, collected August 23, 1963, by
Eustorgio Méndez (orig. no., Gorgas Mem. Lab. 6936). Dorsal sur-
face from crown to upper tail coverts neutral gray; wings and tail
sooty black ; lesser wing coverts methyl blue to Paris blue, with a me-
tallic sheen ; sides of head like crown, with the feathers of the ear cov-
erts with faint grayish white shaft lines; chin sooty gray; foreneck
pale olive-buff; chest and sides pale neutral gray; center of breast,
abdomen, under tail coverts, and axillars pale pinkish buff to pinkish
buff. Bill, tarsus, toes, and claws black. (From dried skin.)

Measurements.—Male (one, the type), wing 69.7, tail 45.4, culmen
from base 10.9, tarsus 17.8 mm.

Female (two specimens), wing 65.4, 68.0, tail 43.6, 43.7, culmen
from base 10.9, 11.0, tarsus 17.1, 17.2 mm.

Range.—Western area of the Province of Bocas del Toro in the
Caribbean lowlands of Panama; probably extending on the Caribbean
slope in Costa Rica.

Remarks.—The well-known race Tangara inornata languens of this
tanager has been recorded in Panama on the Pacific slope west through

4 Tangara inornata languens Bangs and Barbour, Bull. Mus. Comp. Zodl.,
vol. 65, Sept. 1922, p. 227. (Lion Hill, Canal Zone, Panama.)

IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 145

the Canal Zone to Chorrera in the western sector of the Province of
Panama. On the Caribbean side I have taken it in the valley of the
Rio Indio west to El Uracillo, in northern Coclé, and Chilar in west-
ern Colén. In the American Museum of Natural History there is one
from “Cascajal, Coclé” collected February 5, 1889. No collector is
indicated, but the label and writing are those of Heyde and Lux,
whose locality is believed to have been on the Caribbean slope on the
Rio Cascajal, a tributary of the Rio Coclé del Norte.

The first specimen from Bocas del Toro, an immature female, taken
in a mist net and prepared by Rudolfo Hinds, December 16, 1960,
marked a considerable extension of range. The prominent buff of the
under surface of this bird, which attracted immediate attention, was
attributed at the time with some uncertainty to the age of the speci-
men, though this color did not agree with that found in other young
birds of this species that I had seen. A second skin from Almirante,
an adult female, taken September 25, 1962, was as deep buff as
the first one, and with the receipt of an adult male secured August
23, 1963, it was apparent that a racial difference was indicated.

Approximately 75 skins of the race languens and 40 of T. 1. tnornata
from central and eastern Colombia have been available for comparison.
A faint wash of pale pinkish buff on the center of the abdomen and
the under tail coverts is found in a number of these specimens, but in
none is this color prominent as it is in the skins from Bocas del Toro.
It is most evident in a few skins taken a hundred years ago by Mc-
Leannan in which this color seems due in part to discoloration from
age as museum specimens. In all individuals in these long series the
lesser wing coverts are light blue, with no approach to the darker color
of this area in the birds from Bocas del Toro.

It is probable that the race described here ranges beyond the inter-
national boundary in the lowlands of the Caribbean slope of Costa
Rica. While no specimens are available at this time, Dr. Paul Slud
informs me that he has a few records of Tangara inornata (which he
will publish in detail later) from that area.

The name is from the Latin adjective ravus, in the sense of tawny.

IV. ADDITIONS TO THE RECORDED LIST OF BIRDS FROM
THE REPUBLIC OF PANAMA

Knot, Calidris canutus rufa (Wilson) : Two immature males taken
at Puerto Obaldia, San Blas, on September 12 and 22, 1934, by Hasso
von Wedel are the only present report for this species. The specimens,
originally in the Herbert Brandt collection at the Museum of the Uni-

No. 6 RECORDS OF BIRDS FROM PANAMA—WETMORE II

versity of Cincinnati, are now in the U.S. National Museum. In the
period of northern winter the knot, which nests in the far north, is
found from eastern United States south to Tierra del Fuego. There
are, however, few records of it in Central America.

Caspian tern, Hydroprogne caspia (Pallas): In the files of the U.S.
Fish and Wildlife Service there is record of one banded by L. Tyler
on South Limestone Island, in Georgian Bay, Lake Huron, Ontario,
on June 11, 1955, that was found wounded at Aligandi, San Blas, on
the evening of November 12 of that year. According to the report,
forwarded by Dr. Alcibiades Iglésias, the bird died the following day.
The occurrence on the San Blas coast is one to be expected, as this
tern is reported as a migrant to the Caribbean coast of Colombia from
Cartagena to the lower Rio Magdalena.

LITERATURE CITED

HANDLEY, CHARLEs O., Jr.

1959. A review of the genus Hoplomys (thick-spined rats), with descrip-
tion of a new form from Isla Escudo de Veraguas, Panama. Smith-
sonian Misc. Coll., vol. 139, no. 4, July 3, pp. 1-10, 1 fig.

WETMORE, ALEXANDER.

1959. The birds of Isla Escudo de Veraguas, Panama. Smithsonian Misc.

Coll., vol. 139, no. 2, July &, pp. 1-27, 1 pl., 3 figs.

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