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,KJ°"IJBLIC LIBRARY 



3 9999 06317 639 8 





SEVEN NEW WHITE -WINGED DOVES 

FROM MEXICO, CENTRAL AMERICA, AND 

SOUTHWESTERN UNITED STATES 




NUMBER 65 



UNITED STATES 

DEPARTMENT OF THE INTERIOR 

BUREAU OF SPORT FISHERIES AND WILDLIFE 



OCCIDFN PEG 

Al 



LIBRARY 



NORTH AMERICAN FAUNA 



This publication series includes monographs and other reports of scientific in- 
vestigations relating to birds, mammals, reptiles, and amphibians, for professional 
readers. It is a continuation by the Bureau of Sport Fisheries and Wildlife of the 
series begun in 1889 by the Division of Ornithology and Mammalogy (Department 
of Agriculture) and continued by succeeding bureaus— Biological Survey and Fish 
and Wildlife Service. The Bureau distributes these reports to official agencies, to 
libraries, and to researchers in fields related to the Bureau's work; additional 
copies may usually be purchased from the Division of Public Documents, U.S. 
Government Printing Office. 

Reports in North American Fauna since 1950 are as follows (an asterisk indi- 
cates that sale stock is exhausted) : 

*60. Raccoons of North and Middle America, by Edward A. Goldman. 1950. 153 p. 

*61. Fauna of the Aleutian Islands and Alaska Peninsula, by Olaus J. Murie; 
Invertebrates and Fishes Collected in the Aleutians, 1936-38, by Victor B. 
Scheffer. 1959. 406 p. 

*62. Birds of Maryland and the District of Columbia, by Robert E. Stewart and 
Chandler S. Robbins. 1958. 401 p. 

*63. The Trumpeter Swan; Its history, habits, and population in the United 
States, by Winston E. Banko. 1960. 214 p. 

*64. Pelage and Surface Topography of the Northern Fur Seal, by Victor B. 
Scheffer. 1961. 206 p. 

65. Seven New White-winged Doves From Mexico, Central America, and South- 
western United States, by George B. Saunders. 1968. 30 p. 



SEVEN NEW WHITE -WINGED DOVES 

FROM MEXICO, CENTRAL AMERICA, AND 

SOUTHWESTERN UNITED STATES 



By 

George B. Saunders 

Wildlife Biologist, Division of Wildlife Research 
BUREAU OF SPORT FISHERIES AND WILDLIFE 




NUMBER 65 



2 I 2000 



UNITED STATES 
DEPARTMENT OF THE INTERIOR 

Stewart L. Udall, Secretary 

Stanley A. Cain 
Assistant Secretary for Fish and Wildlife and Parks 

FISH AND WILDLIFE SERVICE 

Clarence F. Pautzke, Commissioner 

BUREAU OF SPORT FISHERIES AND WILDLIFE 

John S. Gottschalk, Director 




North American Fauna, Number 65 

Published by 

Bureau of Sport Fisheries and Wildlife 

May 1968 



United States Government Printing Office • Washington • 1968 



For sale by the Superintendent of Documents, U.S. Government Printing Office 
Washington, D.C. 20402 - Price 20 cents 



CONTENTS 



Page 



Abstract iv 

Introduction 1 

Methods 4 

New races 5 

Zenaida asiatica peninsulae 5 

Zenaida asiatica grandis 8 

Zenaida asiatica monticola 10 

Zenaida asiatica palustris 14 

Zenaida asiatica insularis 15 

Zenaida asiatica collina 17 

Zenaida asiatica panamensis 20 

Discussion 21 

Summary 28 

Literature cited 29 

Table— Measurements of 12 subspecies of white-winged doves . 22 

Figures 

1 . Map of breeding ranges 24 

2. Statistical comparison of wing measurements 25 

3. Statistical comparison of tail measurements 26 

4. Statistical comparison of culmen measurements 27 

Approved for publication, January 4 , 1968. 



ABSTRACT 

Seven new subspecies of Zenaida asiatica are described: Z. a. pen- 
insulae of the Yucatan Peninsula, Mexico; Z. a. grandis of the upper 
Big Bend area, central western Texas; Z. a. monticola chiefly of the 
Mexican interior plateaus and highlands; Z. a. palustris of the central 
and southern Pacific coastal plains of Mexico; Z. a. insularis of the 
Tres Marias Islands, Nayarit, Mexico; Z. a. collina of Central America, 
chiefly on the Pacific Piedmont and coastal plain from the Isthmus 
of Tehuantepec, Mexico to Costa Rica; and Z. a. panamensis of the 
northeast coast of the Azuero Peninsula, Panama. 



INTRODUCTION 

Of the white-winged dove, Zenaida asiatica, five subspecies have been 
generally recognized by taxonomists: Z. a. asiatica, Z. a. mearnsi, Z. a. 
australis, Z. a. meloda, and Z. a. alticola. Ranges of the races asiatica 
and mearnsi extend as far north as the southwestern United States, 
australis is in some of the lowlands of Central America, and meloda 
is in western South America (Peters, 1937, p. 87-88; Hellmayr and 
Conover, 1942, p. 499-503) . Later the race alticola was described 
from the Altos, the high mountain region of western Guatemala, and 
neighboring highlands (Saunders, 1951) . 

Van Rossem (1947) described a subspecies clara from the Cape 
region of Baja California, pointing out that it was paler than mearnsi, 
but clara was not generally accepted (Friedmann et al., 1950) . The 
series of 23 males and 14 females from Baja California examined dur- 
ing the present study do not show sufficient differences from mearnsi 
to justify separation from that race. The peninsular birds have longer 
wings and tail than mearnsi, but in adult males these differences 
are only 2 millimeters in average length of wings and 3 millimeters 
in average length of tail. The Baja California whitewings seem to be 
relatively sedentary— apparently they do not migrate beyond that State. 
Additional research may reveal other differences from mearnsi in 
habits and ecology. 

The present paper describes seven new subspecies. Apparently, 
ornithologists have assumed that asiatica is the resident form in much 
of Central America because their collections, made mostly in the 
autumn and winter months, include so many of this race; other white- 
wings have been lumped as asiatica with the observation that this race 
shows wide variation (Dickey and van Rossem, 1938; Griscom, 1932; 
and Ridgway, 1916) . While many asiatica winter as far south as 
Costa Rica, they do not breed in Central America (Saunders, 1959, 
1962) . It now appears that the race asiatica is much less variable in 
color and size than was formerly believed. - 

My interest in this problem began in 19-10 when some of the 
whitewing nestlings we banded in southern Texas in summer were 
reported during autumn and winter in Guatemala and El Salvador. 
In 1942 I visited these countries for the first time during winter and 
spring to study the numbers and distribution of these doves and the 
factors affecting them. 

It was during this survey that alticola was found in the Altos of 
Guatemala (Saunders, 1951), and other series of whitewings were 
collected in different parts of Guatemala and El Salvador. Further 
collecting was done there in 1946 and 1947, and in Mexico from 



2 NORTH AMERICAN FAUNA 65 

1940 through 1960. Other specimens were borrowed from the principal 
museums and from several universities and individuals. When the 
breeding specimens were sorted out and their distribution studied, 
there appeared to be several undescribed races. A statistical analysis 
of the measurements of specimens was made to determine whether 
this method would substantiate the presence of new subspecies. 

It is very significant that more than 450 Z. a. asiatica banded in 
Texas have been reported from localities in Latin America south 
of their breeding places and on wintering grounds as far away as 
Costa Rica. More than 250 banded Z. a. mearnsi from Arizona have 
been reported from western Mexico. The patterns of these recoveries 
are an invaluable aid to understanding racial distribution of this 
species. 

The extent of migration of the races that breed in Central America 
is not yet adequately known. Some subspecies appear to be mostly 
sedentary, such as alticola in the Altos of Guatemala (Saunders, 1951) . 
Skutch (1964, p. 224) reported that they nested in March and April 
in the Sierra de Tecpan, Guatemala, up to 9,000 feet above sea level, 
but after the rainy season began in mid-May they disappeared until 
late in the following November. This indicates that in part of their 
range there is a seasonal movement, perhaps mostly altitudinal. But 
we lack specimens of this race to show its distribution south of 
Guatemala. 

Members of the Bureau of Sport Fisheries and Wildlife to whom 
I am especially indebted include John W. Aldrich for advice on taxo- 
nomic questions, Ralph Andrews for field assistance on our Mexican 
survey in 1960, Richard C. Banks for advice on taxonomy, Earl Bay- 
singer for tabulations of banding and recoveries, Thomas D. Burleigh, 
formerly with the Bureau and now retired, for skins for study, Allen 
Duvall for suggestions in studies of banding, Aelred D. Geis and 
Robert G. Heath for statistical assistance, Mary W. Mann, artist, 
Bird and Mammal Laboratories, for the distribution map and figures, 
and Lester L. Short, Jr., formerly with the Bureau and now with the 
American Museum of Natural History, for help with taxonomic 
questions. 

On the surveys in Guatemala in 1947, Charles O. Handley, Jr., 
U. S. National Museum, aided in obtaining specimens, as did Clarence 
Cottam, Director, Welder Wildlife Foundation, on the 1957 survey 
in Mexico. 

I also wish to thank the following institutions and individuals for 
lending specimens essential for this study and for assistance when I 
visited and studied some of the collections: Academy of Natural 
Sciences of Philadelphia (R. M. deSchauensee and James Bond) ; 
American Museum of Natural History (Dean Amadon) ; British 
Museum (J. D. MacDonald) ; Carnegie Museum (Arthur C. Twomey 
and Kenneth C. Parkes) ; Field Museum of Natural History (E. R. 
Blake and the late Boardman Conover) ; Colorado Museum of Natural 
History (Alfred M. Bailey and Robert Niedrach) ; Louisiana State 



SEVEN NEW WHITE-WINGED DOVES 3 

University, Museum of Zoology (George H. Lowery, Jr., and Robert 
Newman) ; Texas A. & M. University, Department of Wildlife Manage- 
ment (W. B. Davis) ; U. S. National Museum (Philip S. Humphrey) ; 
University of California, Berkeley, Museum of Vertebrate Zoology 
(the late Alden H. Miller) ; University of California, Los Angeles 
(the late A. J. van Rossem) ; University of Florida, Department of 
Zoology (Pierce Brodkorb) ; University of Kansas, Museum of Natural 
History (Richard F. Johnston) ; and University of Michigan, Museum 
of Zoology (R. W. Storer and the late J. Van Tyne) . 

Others to whom I am grateful for specimens or information are 
Rollin H. Baker, Michigan State University; Alvaro Collado M., San 
Jose, Costa Rica; Robert W. Dickerman, then with the Oficina San- 
itaria Pan, Americana, Mexico, D. F., and now with the Cornell 
University Medical College, New York, N. Y.; Herbert Friedmann, 
then Curator of Birds, U. S. National Museum, and now Director, 
Los Angeles County Museum; Roland W. Hawkins, Pittsburgh, Pa.; 
Hugh C. Land, Northwestern State College, Natchitoches, La.; D. B. 
Legters, Merida, Yucatan; the late Luis Macias A., Chief, 
Department of Game, Mexico, D. F.; Burt L. Monroe, Jr., University 
of Louisville; Allan R. Phillips, Instituto de Biologia, Universidad 
Nacional Autonoma de Mexico, D. F.; Albert Schwartz, Miami, 
Fla.; Gilbert Shaw and William Stone, U.S.D.A., Laboratorio 
Entomilogico, Mexico, D. F.; Alexander F. Skutch, San Isidro del 
General, Costa Rica; Austin P. Smith, Zarcero, Costa Rica; and 
Helmuth Wagner, Ubersee Museum, Bremen, West Germany. 

Several ornithologists and other scientists now deceased gave me 
helpful information or other assistance in earlier years: Lee Arnold 
and Frederick C. Lincoln, Bureau of Sport Fisheries & Wildlife; Wil- 
fred H. Osgood and Karl P. Schmidt, Field Museum of Natural 
History; James L. Peters, Museum of Comparative Zoology; Charles 
Plummer, U. S. Department of Agriculture, Mexico, D.F.; and John 
T. Zimmer, American Museum of Nautral History. My thanks go 
also to my fellow biologists and other field men of the States of 
Arizona and Texas, who have banded many thousands of white- 
winged doves since 1940 to obtain information on migration, mortal- 
ity, and other important subjects. Without so many band recoveries 
of asiatica and mearnsi to help in clarifying the distribution of these 
birds in migration and on the wintering grounds in Latin America, 
the relationships of several races would continue to be much more 
puzzling. 

To Alexander Wetmore I am much indebted for his advice on 
taxonomic and distributional problems, for his generosity in giving 
me access to field journals of his 1948 and 1963 collecting trips in 
Panama, and for the privilege of describing the subspecies panamensis. 

To my wife, Dorothy Chapman Saunders, I owe the principal 
acknowledgment for her assistance in collecting and preparing speci- 
mens, making color comparisons, statistical calculations, and editorial 
suggestions. 



NORTH AMERICAN FAUNA 65 



METHODS 

Since differences in coloration, especially of females, are often 
subtle, some of the races can generally be separated most accurately 
on the basis of body size and dimensions. Differences in ecology, time 
of breeding, and extent of migration also help in characterizing some 
subspecies. 

When field observations and study of laboratory skins suggested 
differences in a particular population, a statistical analysis was made 
to determine the possible significance of morphological differences. 
Mayr considers the conventional level of subspecies difference to be 
90 percent or more (Mayr, Linsley, and Usinger, 1953) . The cri- 
terion used here in determining the validity of races was whether 95 
percent or more of the specimens of a population were separable 
from 95 percent or more of the specimens of the adjacent race or 
races. To determine the degree of difference and the percentage of 
joint nonoverlap between characters of races, the standard error was 
calculated for each mean. Confidence limits were determined for the 
.05 probability level as the mean ± "t" times the standard error. 
The means, confidence limits, and ranges are given in the table and 
in figures 2 to 4. When the confidence limits do not overlap, 
statistically significant differences are indicated. 

In this review, 463 adult specimens and several juveniles were 
studied. They represent most of the known populations from Arizona 
and Texas south to Ecuador and Peru. The largest series were those 
of the Fish and Wildlife Service collection in the U. S. National 
Museum that I had obtained in Mexico and Central America. Speci- 
mens from the W T est Indies were not included. Birds from Jamaica, 
Cuba, Haiti, Dominican Republic, the Bahamas, and Old Providence 
Island were examined, but additional breeding specimens are needed 
before some puzzling questions can be answered and an adequate 
appraisal made of their taxonomy. The specimens of asiatica in- 
cluded in the present tabulation of measurements were from breeding 
grounds in southern Texas and northeastern Mexico. 

Measurements used in this study include length of wing (chord) , 
tail, culmen, and tarsus, although the tarsus is not included in most 
of the comparisons as it is not of significant diagnostic value. Statistical 
differences in dimensions are shown in the table and in figures 2 to 
4 (see pages 22-27) . Plumage colors were studied under natural light 
in most instances. A few were determined under special lights which 
approximate daylight in the Bird Division, U.S. Museum of Natural 
History. Color names are from Ridgway (1912) . 

Since migrant subspecies may mingle with resident birds during 
the winter, it is essential that full information, particularly condition 
of gonads and amount of fat, be recorded in the field. In Mexico and 
Central America it was usually possible, from February to April, to 
separate the resident and migrant forms by the greater development 



SEVEN NEW WHITE-WINGED DOVES 5 

of the gonads and the much smaller amount of fat in the residents. 

In the following comparisons of specimens, males are referred 
to unless females are specified. In general, males show more racial 
differences in dimensions, as well as color, than do females, and 
consequently are of greater value for taxonomic study. 

In the original determination of taxonomic differences in any 
migratory animal it is essential to use specimens that are represent- 
ative of breeding populations. It would be desirable to have more 
specimens in fresh plumage for comparison, but by the time the 
postnuptial molt is completed, considerable migration away from the 
breeding ground may have occurred, and specimens taken together 
at that time may represent two or more subspecies. Therefore, the 
descriptions of these seven new races are based on specimens in 
breeding condition, with due consideration of adventitious effects of 
wear and fading. In contrast, the type specimen of australis, taken at 
Cerro Santa Maria, Costa Rica, January 9, 1908, (Peters, 1913) 
is in fine, fresh plumage, so it is not directly comparable widi 
specimens in breeding plumage. It should be noted that no breeding 
white-winged doves have been reported from that locality. This is an 
example of doubtful situations which can arise from using wintering 
or freshly molted specimens in original descriptions. 

NEW RACES 

The series of white-winged doves from the Mexican States of Yucatan, 
Campeche, and Quintana Roo examined in this study exhibits con- 
siderable variation. It includes some wintering birds with dimensions 
and coloration of typical asiatica and others that were smaller and 
mostly paler. Fifteen of these birds which were collected by Gaumer 
have no date, and many of them have no locality other than "Yuca- 
tan." However, the general season during which they were obtained 
could be determined by molt and wear of the plumage. When the 
specimens were sorted according to season, it was found that almost 
all of those in worn breeding plumage had shorter wings and tails than 
typical specimens of asiatica in comparable plumage. Field study 
and this review of specimens have shown. that the breeding population 
of the above area is a distinct subspecies which may be called: 

Zenaida asiatica peninsulae, new subspecies 
Yucatan White-winged Dove 

CHARACTERS 

Nearest to Z. a. asiatica but with shorter wings and tail. In breed- 
ing plumage the back averages slightly paler and grayer than that 
of most specimens of asiatica seen. The crown of the male is paler 
and a lighter purple, and in some specimens is more suffused with 



6 NORTH AMERICAN FAUNA 65 

brown than in asiatica. The latter has more extensive purple that 
extends farther back on the hindneck. In most of the specimens of 
peninsulae seen the underparts are slightly paler, and some have a 
cinnamon tone to the throat and upper breast that occurs in rel- 
atively few asiatica. 

DESCRIPTION 

Type, Field Mus. Nat. Hist. No. 13223, adult male, San Felipe, 
near the mouth of the Rio Lagartos, northeastern Yucatan, collected 
June 6, 1893, by W. W. Brown. Crown and hindneck vinaceous drab; 
back drab; tertiaries buffy brown; middle rectrices olive brown; throat 
wood brown basally with cinnamon tips; breast light drab; abdomen 
pearl gray; and flanks pale olive gray. 

MEASUREMENTS 

Males (14 specimens): wing 148.5-156.3 mm. (av. 151.5), tail 
97.1-108.0 (103.5), and culmen 18.0-21.3 (20.2). Females (18 speci- 
mens): wing 143.0-155.0 mm. (av. 148.6), tail 93.0-104.0 (99.6), and 
culmen 18.4-21.7 (20.0). Most of these specimens were taken during 
the breeding season. 

Ridgway (1916, p. 379) gave the average measurements of wing, tail, 
and culmen of 9 males from Yucatan as 156.4, 103.6, and 19.7 mm., 
and of 7 females as 149.6, 99.4, and 20.2 mm. Judging from the large 
average wing length there was no sorting of these specimens accord- 
ing to season, and apparently several wintering asiatica males were 
included. 

RANGE 

The only breeding specimens seen were from areas of Mexico includ- 
ing the northern half of Yucatan, coastal localities in Quintana Roo, 
the adjacent islands of Cozumel and Mujeres, and northern Campeche. 
A specimen from Jaina, Campeche, taken on June 15, 1900, by 
Nelson and Goldman, is referable to this form. The only breeder 
taken by Paynter (1955, p. 118) was a male collected at Vigia Chico, 
Quintana Roo, March 30, 1949. That locality is adjacent to Bahia de 
la Ascension. Additional specimens from southern Yucatan and Quin- 
tana Roo are needed for clarification of the extent of the breeding 
range. 

Based on the relative scarcity of this race in northern Yucatan dur- 
ing the winter, I believe that, most of these birds winter farther 
south in the State, and possibly also in arid interior valleys of eastern 
Guatemala and Honduras. If so, peninsulae would be associated in 
some localities with a subspecies to be described further on, as well 
as with australis. 

REMARKS 

The white-winged dove was listed for Mujeres and Cozumel Islands, 
and was considered a well-known species on the mainland by Salvin 



SEVEN NEW WHITE-WINGED DOVES 7 

(1889). Cole (1906) was at Chichen Itza in March 1904 and collected 
three specimens March 10-12. 

Paynter (1955, p. 118) reported the habitat to be chiefly in coastal 
scrub and deciduous forest, but occasionally in clearings within the 
rain forest zone. 

Peters (1913, p. 372) took two females on March 16 and 17, 1912, 
at Camp Mengel, on the Rio Hondo, 36 miles southwest of Chetumal, 
Quintana Roo. I examined one of these specimens, MCZ No. 60754; 
it is a small bird, typical of peninsulae. Its rectrices are narrow, and 
its measurements are wing 145.0, tail 96.0, tarsus 22.5, and culmen 
18.5 mm. 

D. B. Legters, Merida, who lived and hunted in Yucatan for many 
years, wrote me in 1961 that he found great numbers of white-winged 
doves on the northern coast of Yucatan between Dzilam and Telchac 
in April, May, and June, nesting among the coconut palms and 
mangroves. A very few remained through the year, mostly in the 
coconut groves. Chapman (1896) found whitewings in large numbers 
in the old cornfields near Chichen Itza in March. 

Dr. Allan R. Phillips wrote me in 1965 that he saw no whitewings 
in Mexico on Isla Mujeres, January 15-18, only one on Isla Cozumel, 
January 19-23, and very few anywhere in Yucatan, or on the pen- 
insula north or east of Isla del Carmen, where they should have 
been common. He found none on the Isla Cozumel during exten- 
sive daily field collecting, November 3-18, 1965. 

During my field work in northwestern Yucatan in January 1960, 
the only white- winged doves seen or heard were several in the dry 
woodland south of Uman near the aguada (watering place) Xcamal, 
and one near the boundary with Campeche, a few miles north of 
Bolonchen de Rejon, both localities on the Merida-Campeche highway. 

Indians who live near the aguada Xcamal and who had learned a 
good deal about the "zac pakal," as the Mayas call this dove, said 
that they were more common during the nesting season, which 
begins in late March and extends through May, and that few were 
present the rest of the year. They added that nests were found in 
densely foliaged, thorny trees near the aguada, some of them placed 
quite low, and that only one brood was raised. I heard two sing 
briefly, and their songs were weaker in volume than typical asiatica. 
When flushed they flew low through the trees, more in the manner 
of white-fronted doves, rather than above the trees as whitewings 
usually do. 

Three were collected and 20 seen at a watering place 16 miles east 
of the city of Campeche on January 31, 1960. One specimen was 
an adult male, the second an adult female, and the third an immature 
female with two juvenal primaries; all were peninsulae. Two birds 
there sang in the same low volume that characterized those noted at 
the aguada Xcamal. Two specimens from San Jose - Carpizo, Campeche 



8 NORTH AMERICAN FAUNA 65 

(Storer, 1961) , approximately 27 miles south of the capital, are im- 
mature females taken November 1 and 24, 1946. They are probably 
peninsulae, judging from their measurements. 

A juvenile male of peninsulae less than 5 weeks old was collected 
at Santa Clara, Yucatan, September 2, 1950 (Yale U. No. 14384) . It 
differs from juveniles of asiatica in having conspicuous buffy edging 
on many of the lesser coverts. The color of the underparts is slightly 
grayer, and the tips of the breast feathers are more tawny than in 
asiatica. 

Additional proof that asiatica from the north occurs on the Pen- 
insula in winter is found in two white-winged doves banded in Tamau- 
lipas during the breeding season which were shot during winter in 
Tabasco and Campeche, and two banded in southern Texas which 
were recovered in Yucatan and Campeche. 

SPECIMENS EXAMINED 

Mexico: Yucatan: Izamal, Santa Clara, Chichen Itza, Sisal, San 
Felipe, and Xocempich. Some of those collected by Gaumer in Yucatan 
were not marked as to sex, locality, and date. Campeche: Isla del Car- 
men near Puerto Real, Jaina, Champoton, and 16 miles east of the 
city of Campeche. Quintana Roo: Vigia Chico, Camp Mengel, Chet- 
umal, and Isla Cozumel. 

Field investigations in the Big Bend sector of Texas and the review 
of specimens collected there and elsewhere in the Southwest and in 
Mexico proved that the white-winged dove of the Chinati Mountains 
and the adjacent valley of the Rio Grande in the upper Big Bend 
region of Texas is a distinct geographic race. As it is larger than the 
other known North American species, it is named: 

Zenaida asiatica grandis, new subspecies 
Upper Big Bend White-winged Dove 

CHARACTERS 

It has longer wings, tail, and tarsus than mearnsi, monticola, and 
asiatica, but its culmen is shorter than that of mearnsi. It is grayer 
above and paler on the breast than asiatica, and slightly grayer tfian 
most mearnsi. The few specimens seen of monticola from the Chisos 
Mountains and northern Mexico average very slightly browner on the 
back than grandis, with the purple crown of the male slightly less 
bright and more veiled with brown. The underparts are somewhat 
paler in grandis than in asiatica. It is similar to Z. a. meloda of 
South America in length of wing, tail, tarsus, and culmen, but it is 
much browner. The race meloda has a thicker bill, gray instead of 
white tips to the rectrices, and other differences in color of plumage. 



SEVEN NEW WHITE-WINGED DOVES 9 

DESCRIPTION 

Type, U. S. Nat. Mus. (Fish and Wildlife Service Collection) No. 
481592, adult male, breeding, active milk glands, near Ruidosa, 
Presidio County, Texas, altitude about 3,000 feet, May 25, 1957, by 
George B. Saunders, collector's number 2662. 

Crown vinaceous drab; hindneck light purple drab; back and terti- 
aries drab; middle rectrices nearest olive brown; throat deep olive 
buff to avellaneous; breast light drab; abdomen pale smoke gray to 
pale olive gray; and flanks pale mouse gray. 

MEASUREMENTS 

Males (10 specimens) : wing 166.0-175.0 mm. (av. 172.0), tail 120.5- 
133.0 (126.6), tarsus 26.0-28.0 (26.5), culmen 20.0-23.0 (21.5), length 
(2) 317-340 (332) , extent (2) 518-538 (526) . Females (5 specimens) : 
wing 162.0-164.9 mm. (av. 163.5), tail 112.9-121.3 (116.5), tarsus 
25.0-26.9 (25.8) , culmen 20.7-22.3 (21.5) , length (2) 304-306 (305), 
extent (2) 494-498 (496). 

RANGE 

Breeds in the Chinati Mountains and adjacent parts of the Rio 
Grande bottomland from near Presidio, Presidio County, north to 
Indian Hot Springs, Hudspeth County, Texas. It probably also occurs 
in the Sierra Vieja, as white-winged doves were seen flying to the 
bottomland near Porvenir from the direction of those mountains. 

A few are reported to winter in the Big Bend region of Texas, but 
the majority journey farther south into Mexico. How far they go at 
that season is not known, but a specimen taken in February near 
Presa Calles, Aguascalientes, Mexico, at 7,000 feet, has the character- 
istics of this race. 

REMARKS 

When male specimens from the lower Big Bend (Lajitas, Castolon, 
Chisos Mountains) are compared with those from the Chinati Moun- 
tains and adjacent valley of the Rio Grande, in the upper Big Bend, 
it is apparent that the latter have significantly longer wings and tail. 
This is surprising since the two ranges are separated by less than 60 
miles. In the collecting done from Chinati north to Porvenir, 9 of 
10 males taken in the breeding season were typical grandis and the 
tenth was intermediate between grandis and monticola. The ecologi- 
cal differences between their habitats are believed to be the principal 
basis for the separation of the two populations. 

Along the Rio Grande Valley the principal physical barriers between 
these two populations are the Colorado Canyon, 3.1 miles in length, 
and the Grand Canyon of Santa Elena, 7 miles in length. These are 
very narrow, deep canyons, with vertical rocky walls rising at each 
side of the Rio Grande, and no trees or shrubs bordering the river. 



10 NORTH AMERICAN FAUNA 65 

The upland on each side of the canyon walls is barren, with gravelly 
hills and no cover for whitewings. The only habitat between these 
two canyons is in small, isolated coves in the river bottom and at the 
mouths of tributary arroyos or washes where there are scattered 
clumps of tamarisk (Tamarix) , mesquite (Prosopis) , and associated 
trees. 

Formerly there was much more nesting habitat in the river bottom- 
land from Presidio to El Paso. From Presidio northward almost all 
river bottom woodland that was suitable for nesting. has been cleared 
for agricultural crops for a distance of approximately 25 miles, except 
for an occasional thin fringe near the river. From near Chinati 
north to Ruidosa and locally beyond as far as Indian Hot Springs 
there are occasional coves of woodland, chiefly of dense tamarisk and 
mesquite, that offer nesting cover for whitewings. When this part of 
the Rio Grande was scouted by plane in 1949 no adequate cover 
for white-winged doves was seen north of Esperanza to El Paso. 

Although most of the specimens of grandis were collected near 
Ruidosa where the eastern edge of the river bottomland meets the 
foothills, the doves flew in from the direction of the Chinati Moun- 
tains. This range is to the east, and its highest elevation, Chinati Peak, 
is 7,730 feet. Local hunters said the whitewings nested in the oak 
woodland of the mountains. How many grandis nest in the oak wood- 
land of the Chinati Mountains and how many utilize other plant 
associations remains to be determined. 

SPECIMENS EXAMINED 

United States: Texas (Presidio County) : Chinati, Ruidosa, and 
near Porvenir. 

All white-winged doves in the interior highlands of Mexico have 
been referred to the race mearnsi (Friedmann et al., 1950) , but breed- 
ing specimens collected there have longer wings and tail, shorter 
bill, and average slightly grayer plumage than Arizona mearnsi. Birds 
from the more northern highlands, as in Nayarit and Durango, are 
slightly darker than those of Oaxaca and Puebla, but they too have 
longer wings and tail than mearnsi. Study of these highland birds 
confirms that they are a distinct subspecies which may be named: 

Zenaida asiatica monticola, new subspecies 
Mexican Highland White-winged Dove 

CHARACTERS 

It has shorter wings and tail than grandis. Birds from the highlands 
of Nayarit, Durango, and more northern States average browner 
than grandis, but those from Puebla and Oaxaca average slightly 



SEVEN NEW WHITE-WINGED DOVES 11 

grayer. It is larger and longer in wing and tail than asiatica and 
mearnsi, and has a shorter bill than the latter. Similar in dimensions 
to an undescribed race on the Tres Marias Islands, Mexico, but with 
paler underparts; similar also to Z. a. alticola of the highlands of 
Guatemala (Saunders, 1951) but much paler above and below. 

DESCRIPTION 

Type, U. S. Nat. Mus. (Fish and Wildlife Service Collection) No. 
481589, adult male, breeding, singing on territory, testes 6 x 12 mm., 
11 miles south of Acatlan, Puebla, Mexico, April 28, 1957, collected 
by George B. Saunders, collector's number 2648. Crown brownish drab; 
hindneck light brownish drab; back hair-brown; tertiaries buffy brown; 
middle rectrices clove brown; throat and breast light drab; abdomen 
pale smoke gray; and flanks light quaker drab. 

MEASUREMENTS 

Males (44 specimens, all seasons) : wing 161.3-177.0 mm. (av. 
167.2), tail 114.0-127.5 (120.4), and culmen 18.9-22.9 (21.1). Females 
(22 specimens, all seasons): wing 156.0-171.0 mm. (a v. 162.8), tail 
111.0-124.4 (116.2), and culmen 19.2-23.0 (21.0). 

RANGE 

Interior plateau and some of the mountains from Oaxaca north 
of the Isthmus of Tehuantepec, northward in the mesquite and 
guamachil associations, thorn forest, tropical deciduous forest, and 
in some localities in oak-pine woodland, to northern Chihuahua, Coa- 
huila, and Nuevo Leon of Mexico, and the Chisos Mountains and 
lower Big Bend of central western Texas. Most were observed at 
elevations of 4,000 to 8,000 feet. They have also been found during 
summer in Hidalgo County, southwestern New Mexico, where 3 of 
15 specimens examined were identified as monticola: 2 were collected 
in 1892, and 1 in 1933. The other 12 were nearer mearnsi. Northern 
Chihuahua and the southwestern corner of New Mexico may be a 
zone of intergradation between these two races, but this is an area 
where whitewings are scarce and local in distribution. 

REMARKS 

Although monticola is widely distributed in the highlands, there 
are many localities where it is absent. Most were observed in dry 
woodlands or thorn forest, but some were in agricultural valleys 
where large guamachiles (Pithecrllobium dnlce) and mesquites of- 
fered nesting cover and food, or in pecan groves of some of the 
valleys, and villages. A few others were seen in higher oak and pine 
woodland. In many localities their absence was due to a lack of 
suitable habitat, but conversely many places with what appeared to 
be a good habitat lacked whitewings. In field work during 1950, 1952, 
1957, and 1960 they were observed in the highlands of every interior 



12 NORTH AMERICAN FAUNA 65 

State of Mexico, and they were collected in Oaxaca, Puebla, Guerrero, 
Morelos, Jalisco, Michoacan, Nayarit, and Aguascalientes during this 
study. 

This race was breeding in Oaxaca and Puebla during the first 
week in February, although the altitudes were from 4,000 to 7,000 
feet. Also, they were breeding in the mountains of Guerrero and 
Nayarit in March when mearnsi, still heavy with winter fat, were 
in flocks there in the foothills, and on the coastal plain of these States. 

Most monticola are believed to winter in or near their breeding 
range. Some at the northern end of the range move south for the 
winter for an undetermined distance, but there are winter flocks as 
far north as Coahuila and Durango. No specimens of monticola are 
known to have been collected from coastal areas or any locality south 
of the Isthmus of Tehuantepec. 

Several individuals of asiatica banded in Texas and Tamaulipas, 
Mexico, and recovered in the Mexican highlands, chiefly in the States 
of Morelos and Oaxaca, prove that some asiatica migrate through or 
winter in the range of monticola. Other winter specimens of asiatica 
have been collected in these southern highlands. Recoveries in the 
western highlands, and especially in the States of Michoacan and 
Jalisco, of many Arizona-banded mearnsi prove that many of this sub- 
species winter there with the resident monticola. If most of the 
museum specimens of white-winged doves were obtained in winter, it 
is not surprising that mearnsi was for so long thought to be the resi- 
dent subspecies throughout the western highlands. 

Peters (1937, p. 87) , as well as Hellmayr and Conover (1942, p. 
500) , gave the range of mearnsi as extending southward and eastward 
in Mexico to Puebla. The present study shows that much of this 
area is within the range of the new race monticola. Specimens from 
the southern highlands of Mexico in Guerrero, D. F., Morelos, and 
San Luis Potosi, considered by Pitelka (1948) to be intermediate 
between asiatica and mearnsi, included some wintering mearnsi and 
asiatica, as well as summer and autumn monticola. 

Several specimens taken in winter in Oaxaca and Puebla were un- 
usually large, and may represent a different race that breeds in the 
higher mountains of those States and winters at lower elevations with 
monticola, or they may be grandis which wintered south of the prin- 
cipal range. The inclusion of their measurements with those of 
monticola is responsible for the upper limits of the wing and tail 
measurements of monticola exceeding those of grandis. Further field 
study of breeding populations in different parts of Oaxaca and Puebla 
is needed to give information on this subject. 

Another interesting discovery concerns a population in the moun- 
tainous part of Durango, where dimensions of the sexes are about 
equal. The several females were sexed and labeled in different years 



SEVEN NEW WHITE-WINGED DOVES 13 

by two experienced collectors. They also may represent a separate 
race, for this characteristic is not true of any other known population 
of white-winged doves. 

The white-winged doves from the interior highlands of Sonora at 
Tecoripa, Cerro Blanco, Opodepe, and Moctezuma probably are re- 
ferable to monticola, judging from their dimensions, but inclusion 
in this race should await further study of specimens and ecology. 

Several whitewings and a nest with a single nestling were found 
in northern Coahuila at Noria de Gilberto, by Walter P. Taylor 
and Clifford C. Presnall, April 9, 1945 (personal correspondence, 
1945) . These birds probably were of the race monticola, since it 
occurs north of this area in Brewster County, Texas (Van Tyne and 
Sutton, 1937). 

The differences in habitats occupied by monticola in the lower Big 
Bend and by asiatica at the northwestern corner of its breeding range 
in Val Verde County, Texas, apparently serve as ecological barriers 
and seem to be effective in maintaining the separateness of these 
populations. In addition, the three long, narrow canyons of the 
Rio Grande and the barren hills between the breeding ranges of 
monticola and asiatica probably also assist in separating these races. 
Although a strong flier like the white-winged dove could easily fly 
this distance, no flights have been reported along this route. No 
specimen of either monticola or grandis has been taken near Del 
Rio, to my knowledge, nor do I know of a specimen of asiatica re- 
ported from the Big Bend. 



SPECIMENS EXAMINED 

Mexico: Aguascalientes: Presa Calles. Chihuahua: Meoqui, Rio 
Conchos, Rio San Pedro. Coahuila: Las Delicias, Piedra Blanca. 
Distrito Federal: Pedregal. Durango: La Boquilla, Las Bocas, Rio 
Sestin, San Juan (C. Lerdo) . Guerrero: Chilpancingo, Colotlipa, and 
Iguala. Hidalgo: Zimapan. Jalisco: Autlan, Bolanos, Hacienda La Ven- 
ta, La Cienega, Santa Cruz, Tizapan el Alto (west of L. Chapala) , Villa 
Corona, Zapotlan. Michoacan: Zamora. Morelos: Cuernavaca, Puente 
de Ixtla, Temilpa. Nayarit: Amatlan de Canas, Hacienda de Ambas 
Aguas, and Tepic. Oaxaca: Cuilapan (near Oaxaca) and La Compania 
(near Ejutla de Crespo) . Puebla: Acatlan, Atotonilco, Chila, Huejotz- 
ingo, Matamoros, Tecomatlan. San Luis Potosi: Hacienda Capulin, 
Salinas Reg, and Santo Domingo. United States: Texas: Lower Big 
Bend (Black Gap, Boquillas, Castolon, Lajitas, Pine Canyon, Still- 
well Crossing, and Wade Canyon. New Mexico: Guadalupe Canyon 
and Animas Mountains, Hidalgo County. All seasons are represented 
by this series. 



14 NORTH AMERICAN FAUNA 65 

Series of white-winged doves were collected in Mexico on March 
28-30 near San Bias, Nayarit, and on April 1, 1960, south of 
Escuinapa, Sinaloa. Study of these specimens, in addition to field 
observations on their distribution, habitat, and habits, showed clearly 
that at least two different populations were present. Most of the 
collecting was done in or near the mangrove swamps, and most of the 
specimens were of a darker population that was breeding in that 
habitat. The others seen were in drier upland habitats, were in flocks, 
were fat, and their gonads were little, if any, enlarged. The latter 
birds were obviously winter residents or migrants, and were identified 
as mearnsi. The darker breeding race has not been reported previously, 
so it is described as: 

Zenaida asiatica palustris, new subspecies 
San Bias White-winged Dove 

CHARACTERS 

Nearest to Z. a. mearnsi (Ridgway, 1915) in size, but darker than 
that race, and with a shorter bill. It is darker in coloration and has 
shorter wings and tail than the Tres Marias Islands population 
described beyond. 

DESCRIPTION 

Type, U. S. Nat. Mus. (Fish and Wildlife Service collection) No. 
481591, adult male, breeding, near San Bias, Nayarit, Mexico, March 
29, 1960, collected by George B. Saunders, collector's number 2672. 
Crown and nape between vinaceous drab and dark vinaceous drab; 
back Prout's brown; tertiaries Saccardo's umber to cinnamon brown; 
middle rectrices between Prout's brown and mummy brown; breast 
nearest Saccardo's umber; belly pale ecru drab to smoke gray; and 
flanks light quaker drab. 

Principal differences of diagnostic value are: palustris males average 
a shorter bill than mearnsi, but differentiation of these races is chiefly 
on the basis of the darker color of both sexes of palustris. In compar- 
ison with the Tres Marias Islands population described beyond, 
palustris has a shorter wing and is darker in color. 

MEASUREMENTS 

Males (22 specimens, mostly breeding) : wing 158.0-169.0 mm. (av. 
163.2), tail 109.-123.6 (117.2), culmen 19.0-23.0 (20.7). Females 
(20 specimens, mostly breeding): wing 153.5-166.0 mm. (av. 159.5), 
tail 107.0-117.0 (112.2) , culmen 19.8-23.0 (21.4). 

RANGE 

The specimens of palustris examined in this study were from the 
mangrove swamps near San Bias, Nayarit, and northward to near 



SEVEN NEW WHITE-WINGED DOVES 15 

Mazatlan, Sinaloa. This race also occurs in some mangrove swamps 
and adjacent woodlands southeast to localities in Guerrero, but further 
study of museum collections and additional field work will be needed 
to determine distribution in more detail. There are several specimens 
of palnstris in museum collections from near Acapulco, Guerrero, and 
six in my series from there. Two of the latter were taken in the 
breeding season (February 6 and 11, 1949), and four in August and 
September, 1965 (from A. R. Phillips collection) . Most of the speci- 
mens I have seen from Sinaloa, Nayarit, Colima (Schaldach, 1963) , and 
Guerrero were taken in winter and were chiefly migrant and winter- 
ing mearnsi. 

The northern limit of the range of palustris may extend to about 
Culiacan, Sinaloa. A male collected at Providencia, 15 miles west of 
Culiacan, April 11, 1963, by A. R. Phillips, is palustris both in color- 
ation and dimensions, although its back is slightly paler than average. 
There is little mangrove swamp north of Culiacan, and no speci- 
mens of palustris have been seen beyond there. 

SPECIMENS EXAMINED 

Mexico: Guerrero: Laguna Coyuca, Laguna Cayaco, and Laguna 
Tres Palos, all near Acapulco; Ciruelar and Tuncingo. Nayarit: San 
Bias and Quimeche River (Rio Acaponeta) . Sinaloa: Escuinapa, 
Mazatlan, and Providencia (La Palma) . Most of these were taken 
during the breeding season. 

A review of specimens of white-winged doves from the Tres Marias 
Islands, Nayarit, Mexico, and comparison of them with series taken on 
the mainland show that they are a separate race. The name proposed 
is: 

Zenaida asiatica insularis, new subspecies 
Tres Marias White-winged Dove 

CHARACTERS 

Similar to palustris of the adjoining mainland of Nayarit and 
Sinaloa, but with paler plumage and longer wings. It likewise has 
longer wings than mearnsi of Arizona and Sonora, but is slightly darker. 
It is similar in dimensions to Z. a. monticola of the Mexican highlands, 
but has darker underparts. 

DESCRIPTION 

Type, Acad. Nat. Sci. Phila. No. 150095, adult male, Maria Madre, 
Tres Marias Islands, Nayarit, Mexico, July 12, 1941, collected by 
Dawson Feathers, Fifth George Vanderbilt Expedition. Crown deep 
brownish drab; hindneck brownish drab; back olive brown; tertiaries 
Saccardo's umber; middle rectrices sepia; throat wood brown to 



16 NORTH AMERICAN FAUNA 65 

sayal brown; breast wood brown; abdomen pale smoke gray to smoke 
gray; and flanks light mouse gray to pale mouse gray. 

Two specimens collected on Maria Madre, May 7, 1897, by Nelson 
and Goldman are slightly paler than the type; the back is Saccardo's 
umber and the breast avellaneous. 

MEASUREMENTS 

Males (9 specimens): wing 163.0-176.0 mm. (av. 170.3), tail 114.0- 
126.0 (121.4), tarsus 25.0-27.0 (25.7), culmen 19.2-21.0 (20.5). 
Females (4 specimens): wing 161.8-165.0 mm. (av. 163.3), tail 108.0- 
113.00 (110.0), tarsus 24.0-26.0 (24.6), culmen 20.0-22.0 (20.6). 

RANGE 

The Tres Marias Islands, Nayarit, Mexico, 60-75 miles west of 
San Bias, Nayarit. 

REMARKS 

Nelson (1899) was the first to record this species for the Tres 
Marias. He found white-winged doves rather common residents on 
both Maria Madre and Maria Magdalena, and a few were seen on 
Maria Cleofas, breeding in the last half of May. The fact that Gray- 
son did not report this species from the Tres Marias on his trips 
there in 1865, 1866, and 1867 (Lawrence, 1874) led Nelson (1899) 
to think that these birds were recent residents on these islands. 
However, white-winged doves have been overlooked in many other 
places. They can be very local in distribution, especially under 
adverse ecological conditions. 

McLellan (1927) found these doves fairly common at all places 
visited in Sinaloa and Nayarit, including Maria Madre, in the fall 
of 1925. The collection includes a male and female taken on Maria 
Madre, October 23, 1925. 

The Fifth George Vanderbilt Expedition of the Academy of Natural 
Sciences of Philadelphia collected a good series including the type of 
insularis on Maria Madre from July 10 to 16, 1941 (Bond and de 
Schauensee, 1944) . 

Stager (1957) found Zenaida in considerable numbers on all islands 
of the Tres Marias group, although they were outnumbered by 
Leptotila and Columbigallina. Zenaida was likewise the least abund- 
ant on Maria Cleofas, but could always be found in the forest margin 
directly behind the beaches on the eastern side of that island. 

Grant (1965) , in his taxonomic study of the birds of the Tres 
Marias, examined a series of 10 male and 5 female white- winged 
doves from the islands and 9 males and 12 females from a nearby 
area on the mainland. He concluded that the two groups do not 
differ sufficiently to warrant taxonomic recognition. He probably 
reached this conclusion largely because some of his mainland speci- 
mens were mearnsi migrants and winter visitants from farther north, 



SEVEN NEW WHITE-WINGED DOVES 17 

and not the breeding subspecies of the Nayarit mainland. In dis- 
cussing the disparity in male bill lengths between some of the main- 
land and the island birds, Grant correctly diagnosed the reason for 
this difference when he inferred that it might be due to the presence 
of migrants in the mainland sample. 

The nearest breeding population on the mainland is near San Bias, 
Nayarit, mostly in or near the mangrove swamps. The 22 adult males 
I examined from this part of the mainland were breeders I collected 
mostly in that locality and near Escuinapa, Sinaloa, in April 1960. 
Their bills average almost 2 mm. shorter than those of mearnsi. In 
late autumn, winter, and early spring, the population of mearnsi in 
drier woodlands and fields of this coastal plain and foothills of these 
States, many of them from Arizona as proved by band recoveries, 
greatly outnumbers that of palustris, which is much more local in 
distribution. 

A series of 25 specimens from the Mexican mainland of Sinaloa 
and Nayarit nearest the Tres Marias includes no individuals that 
have the principal characteristics of insularis. The latter are distinctly 
different although these islands are only 65 miles offshore, west of San 
Bias, Nayarit. 

SPECIMENS EXAMINED 

Mexico: Nayarit: Tres Marias Islands, Maria Madre. 

Field studies in Guatemala in 1942, 1946, and 1947 provided 
many observations on the biology of white-winged doves there. A 
series of specimens from the dry woodlands was first recorded as 
mearnsi (Saunders et al., 1950) . Later study showed that these repre- 
sented a new race whose principal range is in the dry woodland of 
the Pacific piedmont hills of Central America. This new race may be 
called: 

Zenaida asiatica collina, new subspecies 
Piedmont White-winded Dove 



CHARACTERS 

Compared with alticola, collina is paler and smaller and has shorter 
wings and tail. It has shorter wings and tail than monticola. It is 
slightly larger than australis, averages paler on the underparts, and 
lacks the cinnamon brown tone to the breast and upperparts which 
usually characterizes that race. Also, it is grayer on the back and 
rump, and usually more purple on the crown than australis. Com- 
pared with panamensis, collina has longer wings and averages darker 
on the breast and upperparts. Compared with asiatica, collina is 
slightly larger, and its wings and tail are longer. Compared with 



18 NORTH AMERICAN FAUNA 65 

mearnsi from Arizona, it averages browner, the wing of the male is 
longer and the culmen shorter; in the female the tail and culmen 
are shorter. 

DESCRIPTION 

Type, U. S. Nat. Mus. (Fish and Wildlife Service Collection) No. 
481590, adult male, breeding, near Progreso, Department of Jutiapa, 
Guatemala, elevation approximately 3,100 feet, March 13, 1942, col- 
lected by George B. Saunders, collector's number 1622. Crown dark 
vinaceous drab; hindneck vinaceous drab; back olive brown; tertiaries 
Prout's brown; middle rectrices mummy brown; throat and breast 
nearest buffy brown; abdomen pale drab gray; and flanks pale quaker 
drab. 

MEASUREMENTS 

Males (52 specimens) : wing 158.0-169.9 mm. (av. 162.6) , tail 108.0- 
121.9 (113.0), culmen 17.5-22.0 (20.0). Females (22 specimens): 
wing 151.0-165.6 mm. (av. 157.4), tail 103.7-111.0 (107.0), and cul- 
men 18.1-22.1 (20.1) . 

RANGE 

Southernmost Mexico from southeast of the Isthmus of Tehuantepec 
(Oaxaca and Chiapas) through Central America to the Guanacaste 
district of Costa Rica (Carriker, 1910) , chiefly on the Pacific slope. 

REMARKS 

Field studies and specimens indicate that collina occurs in the dry 
woodland and thorn forests of die coastal plain foothills and lower 
mountains of the Pacific slope, and in many of the arid interior valleys, 
including some in the Caribbean drainage. In some places, as at Punta 
Piedra, Costa Rica, on the Gulf of Nicoya, collina breeds locally in the 
coastal lowlands. It is not known whether it also nests in mangrove 
swamps there. 

In some localities on the Pacific slope of Guatemala and El Salvador 
during winter months every white-winged dove I collected was asiatica; 
in other places they were in equal numbers with collina, and in yet 
other habitats only a few miles away I found only collina. In some 
instances collina was the only race present in the thorn forest, and 
asiatica was often more common in valleys that had extensive weed 
fields and farms with grain. 

One specimen of collina, labeled Panama, is probably from Guate- 
mala. Ridgway (1916, p. 380) wrote, "There is a specimen in the 
collection of the Carnegie Museum labeled Nata, Code, Panama (no. 
20777; Heyde and Lux, collectors) ; but this is evidently referable to 
the larger and grayer form from western Mexico, and if really from 



SEVEN NEW WHITE-WINGED DOVES 19 

Panama must have been a cage bird; indeed its appearance suggests 
its having been kept in confinement." This adult male has its primaries 
and rectrices moderately worn, but no more so than many other 
breeding white-winged doves. Nothing about the appearance of this 
specimen indicates that it had been caged; its plumage is not soiled, 
nor are feathers broken or fault-barred. The date, May 20, 1889, was 
within the breeding season. It is comparable in size and color to 
specimens of collina from the Pacific coast and piedmont from Guate- 
mala to Costa Rica. According to Alexander Wetmore, with whom 
this specimen was discussed, the collectors Heyde and Lux obtained 
a large number of "trade skins," many of them from Guatemala. 
There are several instances in which Guatemalan birds in their col- 
lections were mislabeled "Panama." For the present it seems inadvis- 
able to accept this specimen as proof that collina occurs there. 
Collina breeds and winters as far south as the Guanacaste region 
of Costa Rica, so a few of them may have populated arid woodlands of 
southwestern Panama, but if so it is strange that no specimens of 
this race have been collected or reported there since 1889. 



SPECIMENS EXAMINED 

Mexico: Oaxaca: La Ventosa; Chiapas: Berriozabal, Chicomuselo, 
Cd. Cuauhtemoc, Entronque Santa Isabel, Esperanza, Hacienda Mon- 
serrate, Mazapa, Pinuela, Rezo de Oro, San Bartolo, San Jose (near 
Comitan) , Tuxtla Gutierrez. Guatemala: Antigua, Chanquejelve, 
Chiquimula, El Rancho, Lake Atescatempa, Progreso, Sacapulas, San 
Jose de Arada, Usumatlan, Zacapa. El Salvador: Laguna de las Ranas, 
Puerto El Triunfo, Rio Goascoran, Rio Lempa (near Puente Cuscat- 
lan) , San Miguel, Sonsonate. Honduras: Comayagua, Coyoles, El 
Hatillo, La Hor Archaga, Monte Redondo, Rio Hondo, Siguatepeque, 
Subirana. Nicaragua: Calabasas, San Rafael del Norte. Costa Rica: 
Hacienda El Pelon, Las Canas, La Palma de Nicoya, Miravalles, 
Punta Piedra, Tenorio. All seasons are represented by these specimens, 
but the majority are of spring and winter months. 



Only 11 specimens of white-winged doves from Panama were avail- 
able for this study. A twelfth specimen was not sufficiently authenti- 
cated to be considered a satisfactory record. The 11 birds were from 
the coastal mangrove swamps of southwestern Panama, and eight of 
the nine males were collected during the breeding season. They are 
different from other resident populations farther north in Central 
America and are described as a distinct subspecies which may be called: 



20 NORTH AMERICAN FAUNA 65 

Zenaida asiatica panamensis, new subspecies 
Panamanian White-winged Dove 

CHARACTERS 

Smaller than collina, especially in wing and tail length, and aver- 
aging paler brown on the back and breast. Its undertail coverts are 
paler than those of collina which average darker gray. It lacks the 
rich cinnamon brown tones of the back and breast characteristic 
of most specimens of australis. 

DESCRIPTION 

Type, U. S. Nat. Mus. No. 476630, adult male, breeding, Rio Pocri, 
Puerto Aguadulce, Code, Panama, March 12, 1962, collected by A. 
Wetmore. Crown dark vinaceous drab; hindneck vinaceous drab; back 
and tertiaries buffy brown; middle rectrices mummy brown; throat 
slightly paler than sayal brown; breast drab; abdomen pearl gray; and 
flanks pale quaker drab. 

Several of the males from Aguadulce are slightly paler on the 
breast and upper parts than the type specimen. Five of the eight have 
the middle pair of rectrices marked with a more or less visible 
terminal band of grayer or paler brown. 

The female, No. 477593, is slightly paler and less vinaceous brown 
on the breast, and slightly grayer brown on the upper parts. Her 
crown and nape are a paler vinaceous drab than those of the males. 

Compared with collina, most specimens of panamensis are paler 
and grayer, especially on the back and tertiaries; the chin is grayer and 
less brownish; the middle rectrices are lighter brown or grayer and 
often show a paler terminal bar, whereas in collina they are usually 
more uniformly brown. 

Z. a. australis is darker, with a cinnamon brown tone to the breast, 
and this color usually extends farther down on the under parts than 
in panamensis, The back, scapulars, tertiaries, wing coverts, and middle 
rectrices are a darker brown in australis, and its middle rectrices 
usually are uniformly colored and lack the paler terminal band so 
common in panamensis. Both male and female panamensis have sig- 
nificantly shorter wings than collina and australis. 

MEASUREMENTS 

Males (9 specimens): wing 150.0-158.0 mm. (av. 154.8), tail 106.5- 
114.0 (110.3), tarsus 23.5-26.0 (24.7), and culmen 19.0-20.5 (19.8). 
Females (2 specimens): wing 150.0-151.5 mm. (av. 150.8), tail 104.0- 
105.0 (104.5) , tarsus 23.8-24.0 (23.0) , and culmen 20.0. 

RANGE 

Resident and fairly common in the mangrove swamps around the 
shores of the Gulf of Parita on the northeastern coast of the Azuero 
Peninsula, from the lower Rio Parita (Monagrillo) , Herrera, to the 



SEVEN NEW WHITE-WINGED DOVES 21 

Rio Pocri and the Rio Anton, Code. This race is believed to be non- 
migratory. 

The breeding habitat in the mangrove swamp woodland apparently 
is characteristic of this race. None of these birds was observed in 
other habitat types in the localities visited. 

SPECIMENS EXAMINED 

Panama: La Isleta and Rio Pocri in Aquadulce, Province of 
Code; and Rio Parita in Monagrillo, Province of Herrera. 



DISCUSSION 

Of the 12 subspecies mentioned, their grouping in the following 
table, map, and figures is based on their breeding distribution in 
three zones, (1) the Gulf and Caribbean, (2) the interior high- 
lands, and (3) the Pacific Coast. Within these groups the arrangement 
is from north to south. Those of group 1 breed in lowlands of the 
eastern coast of Mexico and Central America and in the West Indies, 
and include asiatica, peninsulae, and australis. They are of medium 
to small sizes for this species, have shorter wings and tail, and live 
chiefly in tropical and subtropical lowlands. 

Group 2 of the interior highlands includes grandis, monticola, and 
alticola. They are large for whitewings, have longer wings and tail, 
and do not occur in lowland localities. 

Group 3, Pacific Coast, includes mearnsi, palustris, insularis, collina, 
panamensis, and meloda. The races in this last group are of medium 
to large size, and the ranges of several extend from the lowlands in- 
land to higher elevations. Their wings and tail are of medium length 
to long except in the mangrove-dwelling race panamensis, in which 
these dimensions are smaller. 

Figure 1 shows the tentative boundaries of the breeding ranges of 
the various subspecies, but further information will undoubtedly 
result in many changes in this map. In the case of australis, for ex- 
ample, the breeding range is without doubt more extensive than 
shown, but in the series of birds examined, only the localities in north- 
ern Honduras were represented by breeding specimens. Their winter- 
ing ranges are not mapped because the overlapping of asiatica and 
mearnsi on the ranges of resident races is so complex. In some places 
three different races may be wintering in the same locality. 

There are many unanswered questions in white-winged dove tax- 
onomy and distribution, especially in areas where insufficient field work 
and collecting have been done. To determine the ecological distribu- 
tion of distinct populations, additional specimens should be taken 
during the breeding season from Mexico south to Costa Rica. Likewise, 
there are considerable gaps in our knowledge of the distribution and 
taxonomy of this species in South America and the West Indies. 



22 



NORTH AMERICAN FAUNA 65 



Table.— Measurements of 12 subspecies of white-winged doves 
[In millimeters] 



Subspecies 
and sex 



Number 
in sample 



Range 



Standard Standard Mean ± Confi- 
deviation error dence limits .05 



Wing measurements: 

asiatica: 

males 32 

females 20 

peninsulae: 

males 14 

females 18 

australis: 

males 10 

females 11 

grandis: 

males 10 

females 5 

monticola: 

males 44 

females 22 

alticola: 

males 8 

females 6 

mearnsi: 

males 38 

females 27 

palustris: 

males 22 

females 20 

insularis: 

males 9 

females 4 

collina: 

males 52 

females 22 

panamensis: 

males 9 

females 2 

meloda: 

males 11 

females 10 

Tail measurements: 

asiatica: 

males 32 

females 20 

peninsulae: 

males 14 

females 18 

australis: 

males 10 

females 11 

grandis: 

males 9 

females 5 

monticola: 

males 44 

females 22 

alticola: 

males 8 

females 6 



151.0-164.0 
146.0-158.9 


2.93 
3.13 


0.52 
0.70 


157.3 + 1.1 
153.5 ±1.5 


148.5-156.3 
143.0-155.0 


2.75 
3.80 


0.76 
0.90 


151.5 + 1.6 

148.6 ±1.9 


154.0-161.0 
150.8-158.2 


2.08 
2.59 


0.66 
0.78 


158.5 + 1.5 
154.1 ±1.7 


166.0-175.0 
162.0-164.9 


2.90 
1.37 


0.92 
0.61 


172.0 + 2.1 
163.5 ±1.7 


161.3-177.0 
156.0-171.0 


3.41 
4.12 


0.51 

0.85 


167.2 + 1.0 
162.8 ±1.8 


164.5-173.0 
153.0-162.0 


2.70 
3.83 


0.95 
1.56 


167.9 + 2.3 
156.7 ±4.0 


155.0-170.4 
148.0-164.0 


3.08 
3.65 


0.50 
0.70 


161.6 + 1.0 
157.9 ±1.4 


158.0-169.0 
153.5-166.0 


2.74 
0.74 


0.58 
0.74 


163.2 + 1.2 
159.5 ±1.6 


163.0-176.0 
161.8 165.0 


4.33 
1.45 


1.44 
0.73 


170.3 + 3.3 
163.3 ±2.3 


158.0-169.9 
151.0-165.6 


2.38 
3.65 


0.33 

0.78 


162.6 + 0.7 
157.4±1.6 


150.5-158.0 
150.0-151.5 


2.58 
1.06 


0.86 
0.75 


154.8 + 2.0 
150.8 ± - 


164.0-175.5 
159.4-169.0 


4.14 

3.86 


1.25 
1.22 


168.3 + 2.8 
164.5 ±2.8 


100.7-115.0 
101.0-112.6 


3.30 
2.68 


0.58 
0.60 


108.8 + 1.2 
107 .2 ±1.2 


97.1-108.0 
93.0-104.0 


2.72 
3.22 


0.72 
0.76 


103.5 + 1.5 
99.6 ±1.6 


108.0-114.0 
103.0-112.0 


2.43 
2.29 


0.77 
0.69 


111.3 + 1.7 
106.2 ±1.5 


120.5-133.0 
112.9-121.3 


3.60 

3.87 


1.2 
1.7 


126.6 + 2.8 
116.5 ±4.7 


114.0-127.5 
111.0-124.4 


3.52 
3.67 


0.55 
0.71 


120.4+1.1 
116.2 ±1.5 


116.6-129.5 
104.0-113.3 


3.89 
3.58 


1.37 
1.46 


122.0 + 3.3 
109.9 + 3.8 



SEVEN NEW WHITE-WINGED DOVES 



23 



Table.— Measurements of 12 subspecies of white-winged doves— Continued 

[In millimeters] 



Subspecies Number 

and sex in sample 

Tail measurements— Cont. 

mearnsi: 

males 38 

females 27 

palustris: 

males 22 

females 20 

insularis: 

males 8 

females 4 

collina: 

males 52 

females 22 

panamensis: 

males 9 

females 2 

meloda: 

males 10 

females 10 

Culmen measurements: 

asiatica: 

males 21 

females 19 

peninsulae: 

males 13 

females 18 

australis: 

males 10 

females 10 

grandis: 

males 10 

females 5 

monticola: 

males 41 

females 21 

alticola: 

males 8 

females 6 

mearnsi: 

males 35 

females 23 

palustris: 

males 22 

females 20 

insularis: 

males 9 

females 3 

collina: 

males 49 

females 18 

panamensis: 

males 9 

females 2 

meloda: 

males 11 

females 10 



Range 



Standard Standard Mean ± Confi- 
deviation error dence limits .05 



106.7-123.5 
103.8-116.0 


3.80 
2.69 


0.63 
0.52 


116.3 + 1.3 
110.1 ±1.1 


109.0-123.6 
107.0-117.0 


3.75 
3.42' 


0.79 
0.72 


117.2 + 1.6 
112.2 + 1.5 


114.0-126.0 
108.0-113.0 


3.47 
2.16 


1.24 
1.08 


121.4 + 2.9 
110.0 ±3.4 


108.0-121.9 
103.7-111.0 


2.63 
2.53 


0.36 
0.54 


113.0 + 0.7 
107.0 ±1.1 


106.5-114.0 
104.0-105.0 


2.82 
0.71 


0.94 
0.50 


110.3 + 2.2 
104.5 ± - 


120.0-133.0 
113.3-125.0 


3.95 
4.69 


1.19 
1.48 


125.2 + 2.7 
118.9 ±3.4 


19.0-21.3 
17.0-21.2 


0.50 
1.16 


0.11 
0.27 


20.1+0.2 
19.5 ±0.6 


18.0-21.3 
18.4-21.7 


1.04 
0.97 


0.29 
0.22 


20.2 + 0.6 
20.0 ±0.5 


18.0-20.9 
18.8-21.0 


1.12 
0.85 


0.35 
0.27 


19.4 + 0.8 
19.9 ±0.6 


20.0-23.0 
20.7-22.3 


0.85 
0.67 


0.26 
0.3 


21.5 + 0.6 
21.5 ±0.8 


18.9-22.9 
19.2-23.0 


0.91 
1.07 


0.17 
0.24 


21.1+0.3 
21.0 ±0.5 


19.0-21.9 
18.5-20.0 


1.23 
0.51 


0.43 
0.21 


20.2 + 1.0 
19.2 ±0.5 


20.8-24.8 
20.7-25.4 


0.96 
1.16 


0.16 
0.24 


22.5 + 0.3 
22.1 ±0.5 


19.0-23.0 
19.8-23.0 


1.15 
1.02 


0.25 
0.23 


20.7 + 0.5 
21.4 ±0.5 


19.2-21.0 
20.0-22.0 


0.64 
1.16 


0.21 

0.58 


20.5 + 0.5 

20.6 ± 2.5 


17.5-22.2 
18.1-22.1 


0.98 
1.32 


0.14 
0.31 


20.0 + 0.3 

20.1 ±0.7 


19.0-20.5 
20.0-20.0 


0.52 
0.00 


0.17 


19.8 + 0.4 
20.0 ± - 


20.5-23.2 
20.5-22.5 


1.00 
0.66 


0.30 
0.21 


21.6 + 0.7 
21. 4 ±0.5 



24 



NORTH AMERICAN FAUNA 65 



6 £ S g a I 
©©©©©© 




1 


1 


I 




1 

1 ~" < 


*l 








/- 


rS\Sb 




i / O ^SV'i.^' 








p — vy 


1 




\ 


o Jl 


b 







SEVEN NEW WHITE-WINGED DOVES 



25 



140 142 144 146 148 ISO 152 154 156 158 160 162 164 166 168 170 172 174 176 178 MM 

1 1 1 1 1 1 1 1 I I 1 



-i 1 1 r 



MALES 
osiotico (32) 
E=!=3 



peninsuloo(l4) 



oustrolis(IO) 

EEE^E^ 



monticolo (44) 

4=±3 



meornsl (381 



olticola (8) 

I I I 



polustris(22) 

r~\—i 



panomensis (9) 



Colli no (52) 
-E*3 



insular is ($) 

r I 



me I o do (ll) 



osiotico (20) 

E=£=3 

pen insuloe (18) 

i I ' 

oustrolis(ll) 

— r~ 1 1 — 



monticolo (22) 



otticolo (6) 



meornsi (27) 



ooljttris (201 



col Una (22) 

- ' I ' 



panomensi s(2 1 



I I I L 



l I I I I I 1 U 



J I L. 



Figure 2.— Statistical comparison of wing measurements of subspecies 

of white-winged doves. 



The present studies indicate that asiatica does not occur as a breeder 
anywhere on the mainland south of the Isthmus of Tehuantepec, 
and mearnsi may not breed south of northern Sinaloa. 

Morphological differences between populations are not great, and 
in several races the general trends do not conform to the classical 
rules of morphological variation correlated with climate. In general, 
long-winged birds are characteristic of the higher altitudes and more 
temperate areas, and short-winged birds are typical of the tropical 
lowlands of the Gulf and Caribbean. However, some of the subspecies 
having the longest wings and tails are those in tropical lowlands of 
some Pacific coastal localities from Mexico south to northern Chile. 



26 NORTH AMERICAN FAUNA 65 



92 94 96 98 100 102 104 106 108 110 112 114 116 118 120 122 124 126 128 130 132 134 MM 

I 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 

MALES 
OS lot loo (32) 



peninsulas (14) 

r^ — i 



austral is (10! 



moat: col a (44) 



all I col a (8 1 



meornst(38) 

I . . I 



col I in o(5Z) 

-m — 



insula r is (8) 

I I 3- 



ponamensis (9) 

i I i 



eloOo(lo) 
1 ' 



pentnsuloe (IB) 

I 1 I 



oslotlco!20) 
I I I 



aostralis(ll) 

I 1 1 



olllcolo (6) 

I I 



monticola (22) 

^A — i 



paluatris (20) 



panomensis (2) 



col Una (22) 



melodo(IO) 

I I 



-I I I 1 1 I I 1 I I I I 1 J 



Figure 3.— Statistical comparison of tail measurements of subspecies 
of white-winged doves. 



The various subspecies of white-winged doves do not seem to 
follow Allen's rules in terms of bill length. All of the races in the 
hottest year-round habitats, in the tropical lowlands of Central 
America, and on the Pacific coast of Mexico have short bills. The 
race with the longest bill is mearnsi, yet its Arizona range is at the 
greatest latitude of any of the whitewings. Although Arizona desert 
summers are hot, the more tropical habitats in Latin America are 
hotter. 

The subspecies of whitewings show more agreement with Bergman's 
rule, but there are exceptions. The largest birds in body size are 
those in the highlands of Mexico and Central America. They live at 



SEVEN NEW WHITE-WINGED DOVES 27 



MALES 

os i an co (21) 
-E*3- 



peninsular fi3> 



a a strolls I/O) 



grand/ s CO I 
I I I 



moot i cola (41) 

F=t=» - 



meornsi (35) 



insulonsl 9) 

1 I I 

colli no (49) 
F=l=» 



caluslrls (221 
^=3 



ponomensis(9) 

meloda(ll) 

1 | I 



FEMALES 



os lo 1 1 co (l 9/ 
1 I ■ 



penmsjloe (18) 
auttrollt(IO) 



all I c ol o (6) 
I I 3— 



monticolo (21 1 

1 1 I 



grand it (S) 



meornsi (231 

3 



poluslris (201 

' 1 ' 



col lino (181 



ponamentls (2) 



meloda(io) 

1 I ' 



Figure 4.— Statistical comparison of culmen measurements of sub- 
species of white-winged doves. 

higher altitudes where the climate is cooler, but meloda, which also 
is large, lives in Pacific coastal and piedmont areas of tropical west- 
ern South America. The climate of the range of meloda is modified 
somewhat by the proximity of the Humboldt Current. The smallest 
whitewings in body size are those in the hot, tropical lowlands of 
Yucatan and Panama. 

In conformance with Gloger's rule, dark pigmentation appears to 
be associated with the more humid habitats and paler hues with 
the more arid areas. The darkest are those of the wooded highlands 
of Guatemala and of the mangrove swamps of the Pacific lowlands of 
Mexico from southern Sinaloa to Guerrero. The palest are those of 



28 NORTH AMERICAN FAUNA 65 

the deserts of Arizona, Baja California, the highlands of Mexico, 
and the Pacific coast of South America, especially if breeding speci- 
mens are compared. The summer (breeding) plumage shows the 
greatest contrast between some of the subspecies as birds of arid, 
rocky habitats show much more fading and wear of plumage than 
those in more moist woodlands. The plumage of most white-winged 
doves is darkest and richest in color when in fresh condition after the 
postnuptial molt. Combinations of these environmentally related char- 
acters of dimensions and plumage color are the basis for the differ- 
ences found in the several races described in this paper. 



SUMMARY 

In a study of the distribution and taxonomy of the white-winged 
dove, Zenaida asiatica, it was found that the subspecies Z. a. asiatica 
of Texas and northeastern Mexico and Z. a. mearnsi of Arizona are 
strongly migratory. The former winters chiefly in Central America and 
the latter in western Mexico. With the clarification of their ranges 
and the study of breeding populations in Mexico and Central America 
it became apparent that several undescribed races were resident in 
these countries, Systematic collecting in many localities, the review 
of museum specimens, and field studies of ecological differences 
among populations indicated the presence of at least seven undescribed 
subspecies which are described in this paper. 

These subspecies are Z. a. peninsulae of the Yucatan peninsula, 
Z. a. grandis of central western Texas, Z. a. monticola of the Mexican 
highlands, Z. a. palustris of the central and southern Pacific coastal 
plain of Mexico, Z. a. insularis of the Tres Marias Islands, Nayarit, 
Mexico, Z. a. collina of Central America, chiefly on the Pacific pied- 
mont and coastal plain from Chiapas, Mexico, to Costa Rica, and Z. a. 
panamensis of the northeast coast of the Azuero peninsula, Panama. 
Measurements of specimens, with figures presenting a statistical analysis 
of these measurements, are given, together with a map showing the 
breeding ranges. 



SEVEN NEW WHITE-WINGED DOVES 29 



LITERATURE CITED 



Bond, James, and R. M. de Schaufnsee. 

1944. Results of the Fifth George Vanderbilt Expedition. The Birds. Academy 
of Natural Sciences of Philadelphia, Monograph 6. p. 7-56. 
Carriker, M. A., Jr. 

1910. An annotated list of the birds of Costa Rica including Cocos Island. 
Carnegie Institute, Annals of the Carnegie Museum, Vol. 6, p. 314-915. 
Chapman, F. M. 

1896. Notes on birds observed in Yucatan. American Museum of Natural History, 
Bulletin 8, p. 271-290. 
Cole, Leon J. 

1906. Aves from Yucatan. Harvard University, Museum of Comparative Zoology, 
Bulletin 50, p. 109-146. 
Dickey, Donald R., and A. J. van Rossem. 

1938. The birds of El Salvador. Field Museum of Natural History, Zoological 
Series 23, Publication No. 406. 609 p. 
Friedmann, Herbert, Ludlow Griscom, and Robert T. Moore. 

1950. Distributional check-list of the birds of Mexico, Part 1. Cooper Ornitho- 
logical Club, Pacific Coast Avifauna No. 29. 202 p. 

Grant, P. R. 

1965. A systematic studv of the terrestrial birds of the Tres Marias Islands, 
Mexico. Yale University, Peabody Museum of Natural History, Postilla, No. 
90. 106 p. 
Griscom, Ludlow. 

1932. The distribution of bird-life in Guatemala. American Museum of Natural 
History, Bulletin 64. 425 p. 

Hellmayr, C. E., and B. Conover. 

1942. Catalogue of birds of the Americas. Field Museum of Natural History, 
Zoological Series 13, Publication No. 514. 636 p. 

Lawrence, George N. 

1874. Birds of western and northwestern Mexico, based upon collections made 
by Col. A. J. Grayson, Capt. J. Xantus and Ferd. Bischoff, now in the 
Museum of the Smithsonian Inst, at Washington, D. C. Boston Society of 
Natural History, Memoirs, Vol. 2, No. 30, p. 265-319. 

Mayr, E., E. G. Linsley, and R. L. Usinoer. 

1953. Methods and principles of svstematic zoology. McGraw-Hill, New York. 
797 p. 

McLellan, M. E. 

1927. Notes on the birds of Sinaloa and Nayarit, Mexico, in the fall of 1925. 
California Academy of Science, Proceedings, Vol. 16, No. 1, p. 1-51. 

Nelson, E. W. 

1899. Birds of the Tres Marias Islands, western Mexico. U. S. Biological Survey, 
North American Fauna, No. 14, p. 7-62. 

Paynter, Raymond A., Jr. 

1955. The ornithogeography of the Yucatan Peninsula. Yale University, Peabody 
Museum of Natural History, Bulletin 9. 347 p. 

Peters, James L. 

1913. List of birds collected in the Territory of Quintana Roo. Auk, Vol. 30, 

p. 367-380. 
1937. Check-list of the birds of the world. Harvard University Press, Cambridge. 
Vol. 3, 311 p. 



30 NORTH AMERICAN FAUNA 65 

Pitelka, Frank A. 

1948. Notes on the distribution and taxonomy of Mexican game birds. Condor, 
Vol. 50, p. 121-122. 
Ridgway, Robert. 

1912. Color standards and color nomenclature. Published by the author, Wash- 
ington, D. C, 43 p., 53 color plates. 

1915. Descriptions of some new forms of American cuckoos, parrots, and pigeons. 
Biological Society of Washington, Proceedings, Vol. 28, p. 105-107. 

1916. Birds of North and Middle America. U. S. National Museum Bulletin 50, 
Part 7. 543 p. 

Salvin, Osbert. 

1889. A list of the birds of the islands of the coast of Yucatan and of the Bay 
of Honduras.. Ibis, Vol. 31, p. 377. 

and F. D. Godman. 

1902. Biologia Centrali-Americana. Aves, Vol. 3, p. 245-247. 
Saunders, George B. 

1951. A new white-winged dove from Guatemala. Biological Society of Wash- 
ington, Proceedings, Vol. 64, P. 83-87. 
1959. La paloma de alas blancas en las Americas. Memoria de la Segunda 
Convencion Nacional Forestal, Departamento de Divulgacion y Propaganda 
de la Subsecretaria de Recursos Forestales, Mexico, D. F., 1959, p. 414-^122. 

1962. The white-winged doves of the Americas. U. S. Fish and Wildlife Service, 
mimeographed, 10 p. (Based on a translation of the paper, "La paloma de 
alas blancas en las Americas," Memoria de la Segunda Convencion Nacional 
Forestal, Mexico, 1959.) 

, C. O. Hanuley, Jr., and A. D. Hoixoway. 

1950. A fish and wildlife survey of Guatemala. U. S. Fish and Wildlife Service, 
Special Scientific Report— Wildlife, No. 5. 162 p. 

SCHALDACH, W. J., Jr. 

1963. The avifauna of Colima and adjacent Jalisco, Mexico. Western Foundation 
of Vertebrate Zoology, Proceedings, Vol. 1, No. 1. 100 p. 

Skutch, Alexander F. 

1964. Life histories of Central American pigeons. Wilson Bulletin, Vol. 76, No. 
3, p. 211-247. 

Stager, Kenneth E. 

1957. The avifauna of the Ties Marias Islands, Mexico. Auk, Vol. 74, No. 4, 
p. 413^32. 
Storer, Robert W. 

1961. Two collections of birds from Campeche, Mexico. Occasional Papers of 
the Museum of Zoology, University of Michigan, No. 621, p. 1-20. 
van Rossem, A. J. 

1947. Comment on certain birds of Baja California, including descriptions of 
three new races. Biological Society of Washington, Proceedings, Vol. 60, 
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Van Tyne, Josselyn, and George M. Sutton. 

1937. The birds of Brewster County, Texas. Miscellaneous Publications of the 
Museum of Zoology, University of Michigan, No. 37, 119 p. 



a U.S. GOVERNMENT PRINTING OFFICE : 1968 O— 310-811 



I 49.30:65 docus 

Seven new white-winged doves /Saunders, 




3 5043 00332 4984 



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servation, concerned with management, conservation, and development of the 
Nation's water, wildlife, fish, mineral, forest, and park and recreational resources. 
It has major responsibilities also for Indian and Territorial affairs. 

As America's principal conservation agency, the Department works to assure 
that nc - uu -°crM,r™>* aw developed and used wisely, that park and rec- 

reation /able resources 



make 

United 



DATE DUE 



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