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v.39
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NATURAL HISTORY.
SURVEY

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MAY *«1968
FIELDIANA • ZOOLOgV"™

Published by
CHICAGO NATURAL HISTORY MUSEUM

Volume 39 July 25, 1961 No. 50

Variation in the Quail -Dove
Geotrygon frenata

Emmet R. Blake
Curator, Division op Birds

Gunnar Hoy1

Francisco Contino1

Over a period of several years the junior authors have sent to
Chicago Natural History Museum some hundreds of bird skins from
Argentina for critical determination, preparatory to publishing a
summary of field notes relating to the birds of Jujuy and Salta.
Among the specimens received for study by the senior author have
been a number of additions to the Argentine list, single specimens of
several probably undescribed forms, and three examples of a new
and strikingly distinct quail-dove.

The new pigeon, described below, is named in honor of Miss
Margaret Conover, sister of the late Boardman Conover, whose out-
standing collection of game birds provided much of the comparative
material used in the present study.

For the loan of additional comparative material we are indebted
to authorities of the following institutions: the American Museum
of Natural History, the British Museum (Natural History), and
Carnegie Museum.

All measurements are in millimeters and the wings were measured
flat. Names of colors when capitalized are from Ridgway's Color
Standards and Nomenclature, 1912.

Geotrygon frenata margaritae, new subspecies

Type. — Chicago Natural History Museum No. 22413 (Conover
Collection), from Yuto, Province of Jujuy, Argentina. Altitude 400
meters. Adult female collected July 7, 1957, by Francisco Contino.

1 Museo de Facultad de Ciencias Naturales, Salta, Argentina.
Library of Congress Catalog Card Number: 61-1 66S5
No. 922 567

568 FIELDIANA: ZOOLOGY, VOLUME 39

Diagnosis. — Differs from all known races of G. frenata in having
the entire crown and occiput deep reddish brown (near Liver Brown)
instead of gray or grayish slate. Nearest G. f. bourcieri of Colombia
and Ecuador but forehead and sides of head darker, near Vinaceous-
Cinnamon, and wings, lower back, upper tail coverts, and tail much
clearer olive.

Measurements. — Wing, 154; tail, 91; exposed culmen, 16.

Range. — Upper Tropical (?) and Subtropical Zones of Sierra de
Santa Barbara, southeastern Jujuy, Argentina, descending briefly
to the adjacent lowlands (Yuto, Province of Jujuy) in July and
August during periods of exceptional cold.

Habits. — According to Mr. Contino, who collected the six known
specimens, margaritae customarily lives in cloud forests of the higher
mountains of Jujuy, some 2000 meters above sea level. At that alti-
tude the vegetation is very dense and the forest composed of large
trees, of which the dominant and most conspicuous kinds are cedar
(probably Cedrela angustifolia) , walnut (Juglans australis) , and alder
(Alnus jorullensis). As the trunks and branches of these trees are
moss-covered, and laced with lianas, quail-doves are difficult to ob-
serve in their usual habitat, but examination of the six stomachs
shows that much of their diet consists of alder seeds. During ex-
tremely cold weather, usually in July or August, quail-doves some-
times visit the lowlands and may remain in cultivated areas for as
long as three weeks. At these times they lose much of their shyness
and permit one to approach within a few feet before taking alarm.
The confiding nature of margaritae while in the cultivated lowlands
has earned for it the vernacular name Paloma tonta ("foolish pigeon").
Both sexes are similar in appearance and have yellow eyes, black bills,
and reddish legs and feet. Weight of adults, 195-330 grams (aver.
244 grams).

Discussion. — The affinities of margaritae clearly lie with G. frenata
but whether or not the two actually are conspecific is at present a
matter of opinion. In color of the under parts margaritae agrees with
bourcieri of Colombia and Ecuador, geographically the most distant
race of frenata, but in color of the upper parts (exclusive of the crown
and nape) there is a closer approach to the nominate race of southern
Peru and Bolivia. Although the brown-crowned Argentine bird ap-
parently is isolated from its nearest relative (gray-crowned) by some
500 miles and might properly be considered a distinct species, we
favor a more conservative treatment until the distribution and habits
of both populations are better known.

BLAKE, HOY AND CONTINO: VARIATION IN QUAIL-DOVE 569

Characters of the several described races of G. frenata have been
conveniently summarized by Hellmayr and Conover (1942, footnotes,
pp. 618-620). Unfortunately, individual variation within these pop-
ulations has received scant attention and we find no reference to an
apparently significant cline in color of the upper parts that becomes
evident on comparison of representative specimens in adequate series.
The following remarks reflect the results of a study of such series, and
critical examination of the unique type of loricata Todd and of two
examples (1 adult, 1 immature) of the exceedingly rare and contro-
versial Ecuador form, erythropareia Salvadori.

The specimens now assembled show a wide range of individual
variation and when arranged by subspecies it becomes obvious that
the accepted racial characters are themselves often unreliable in the
allocation of individuals. Fortuitous variation is especially evident
in the extent and shade of the gray and tawny coloring of the head,
and in the coloring of the under parts, whether essentially brown or
grayish, dark or pale. As some specimens representing each of the
recognized gray-crowned populations are to all appearances identical,
any subspecies based on only one or two specimens is suspect and
requires authentication. Nevertheless, there is clear evidence in
G. frenata of two plumage trends, in effect color clines, that indi-
cate significant geographic variation. This is expressed in the pro-
gressive loss, from north to south, of rufescence on the upper parts,
and in the darkening of the under parts in peripheral populations.
Although the clarity of both clines is somewhat clouded by the ran-
dom occurrence of individual variants, average differences in color
of plumage serve to distinguish the following additional populations.

Geotrygon frenata bourcieri Bonaparte

Geotrygon bourcieri Bonaparte, 1855, Compt. Rend. Acad. Sci. Paris, 40, no. 3,
p. 101 — Valley of Lloa, Ecuador.

Geotrygon erythropareia Salvadori, 1893, Cat. Bds. Brit. Mus., 21 : 565 (in key),
578 — Jima and Rosario, Ecuador.

Characters. — Similar to G. f. subgrisea but somewhat more rufes-
cent above and, on the average, decidedly darker and browner (less
grayish) below.

Distribution. — Upper Tropical and Subtropical Zones (870-2500
meters) of Colombia and Ecuador; north in the western Andes to
Rio San Juan. In Colombia reported east of the Andes only in south-
eastern Narino (Cerro Pax).

570 FIELDIANA: ZOOLOGY, VOLUME 39

Discussion. — In series this peripheral race averages much darker
below than either subgrisea or frenata, and in this respect is identical
with both margaritae and the type of "loricata." Pale-bellied individ-
uals similar in all respects to darker examples of subgrisea occur at
random both in Colombia (Cauca) and Ecuador, but these variants
are in the minority and do not alter the reality of a separable, dark-
bellied northern form. According to Chapman (1926, p. 173), bour-
cieri intergrades with subgrisea in extreme southwestern Ecuador
(Zaruma) .

The status of "erythropareia," which is known from but three
specimens (2 adults, 1 immature) in the British Museum, remains
somewhat equivocal. The type, as depicted in Sclater and Salvin
(1869, pi. 40) closely resembles several of the darker, more richly
colored examples of bourcieri at our disposal, but on direct compari-
son both Chapman (loc. cit.) and Hellmayr and Conover (loc. cit.)
found it to differ in several minor respects from any specimens of
the latter seen by them. Our examination of the other adult (from
Jima) has verified their observations except as to the distinctive in-
tensity of rufescence on the upper parts. In this connection it is
interesting to note that the plumage of the immature bird (from
Rosario), while not definitely diagnostic, strongly suggests that of
typical bourcieri, especially in color of the forehead, malar region,
hind neck, and belly. Although the exact plumage of "erythropareia"
as now known has not yet been found among unquestioned examples
of bourcieri, the pronounced variability of the latter strongly suggests
that the former is only a fortuitous and presumably rare color phase.
As the supposed characters of "erythropareia" are relatively incon-
spicuous at best, and its range is apparently covered by that of
bourcieri, it seems best to combine the two provisionally.

Geotrygon frenata subgrisea (Chapman)

Oreopelia (sic) bourcieri subgrisea Chapman, 1922, Amer. Mus. Nov., 31: 2 —
Alamor, 4350 feet, Province of Loja, Ecuador.

Characters. — Similar to G. f. frenata but on the average slightly
more rufescent above, and the gray of the crown, whether pale or
dark, without a suggestion of vinaceous coloring.

Distribution. — Upper Tropical and Subtropical Zones (1200-3000
meters) of extreme southwestern Ecuador (Province of Loja) and
northwestern Peru, eastward at least to the lower Maranon Valley
(Santa Rosa).

BLAKE, HOY AND CONTINO: VARIATION IN QUAIL-DOVE 571

Discussion. — An intermediate form, having the clear gray crown
of bourcieri and pale under parts of frenata. The darkest individuals
of subgrisea are indistinguishable from the palest examples of bour-
cieri, but in series the two populations can be separated by average
differences. Color of the forehead, considered important by Hellmayr
and Conover, proves to be highly variable and of no diagnostic value.

Geotrygon frenata frenata (Tschudi)

Columba frenata Tschudi, 1843, Arch. Naturg., 9, (1), p. 386 — eastern slopes
of the Andes of Peru (= probably Chanchamayo, Department of Junfn).

Oreopeleia erythropareia loricata Todd, 1947, Proc. Biol. Soc. Wash., 60: 67 —
Buena Vista (Rfo Yapacani), Bolivia.

Characters. — Similar to G. f. subgrisea but gray of the crown,
whether dark or pale, usually more or less conspicuously tinged with
violaceous.

Distribution. — Upper Tropical and Subtropical Zones (800-3000
meters) of southern Peru (north to Junin) and northern Bolivia in De-
partments of La Paz, Cochabamba, and Santa Cruz (Rio Yapacani) .

Discussion. — The distinction between frenata and subgrisea is
rather finely drawn but the diagnostic character of the former shows
quite well in pale-crowned birds. Both races are identical in color
of the under parts, and individuals of each are in this respect virtually
indistinguishable from the palest examples of bourcieri.

In describing "loricata" from a single specimen Todd evidently
was unaware of the wide range of variation to be found even in the
adjacent population (frenata) and unfortunately compared his bird
only with specimens from Incachaca (Bolivia), and with "erythro-
pareia" as portrayed by Sclater and Salvin (1869, pi. 40).

The unique type of "loricata" is, in fact, notably different from
the three Incachaca birds seen by us but there is little to distinguish
it from several examples of both frenata (Cuzco) and subgrisea, and
it is virtually identical with numerous typical examples of bourcieri.
Furthermore, there is no certain evidence that the type of "loricata,"
an August (=cold season) specimen, represents a lowland form. This
assumption is suspect by virtue of the fact that no other race of the
species is restricted to the highlands, and at least one (margaritae)
customarily descends to lower altitudes at intervals in July and
August. In our opinion the Santa Cruz bird has no present claim
to recognition.

572 FIELDIANA: ZOOLOGY, VOLUME 39

SPECIMENS EXAMINED

G. f. bourcieri. — Colombia: La Costa, 2; Munchique, 6; San An-
tonio, 1; Cocal, west of Popayan, 1. — Ecuador: Chical, 3; Tagua-
quiri, 2; Ramos- Urco, 1; San Cadoo-Mindo, 1; Montes Anagumba, 4;
Pacto, 1; Cerro San Jos£, 1; Rio Tigre, 1; Paramba, 1; Monte Chuga,
1; Gualea, 1; Chimiplayas, 1; Condeja, 1; Baeza, 1; San Jose" de
Sumaco, 3; Jima, 1; Rosario, 1.

G. /. subgrisea. — Ecuador: Alamor, 2; Celicia, 1. — Peru: Uscho, 1;
La Lejia, 1; San Pedro, 1; Santa Rosa, 1.

G. f. frenata. — Peru: Chanchamayo, 7; Marcapata, 3; Oconeque,
1. — Bolivia: Incachaca, 3; Rio Yapacani, 1 (type of "loricata") .

G.f. margaritae. — Argentina: Yuto, 2; Sierra de Santa Barbara, 1.

REFERENCES

Chapman, Frank M.

1926. The distribution of bird life in Ecuador. Bull. Amer. Mus. Nat. Hist.,
55, xiii+784 pp.

Hellmayr, Charles E., and Conover, Boardman

1942. Catalogue of birds of the Americas. Field Mus. Nat. Hist., Zool. Ser.,
13, pt. 1, no. 1, vi+636 pp.

Sclater, Philip Lutley, and Salvin, Osbert
1869. Exotic Ornithology, vi+204 pp. London.

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