North American fauna

,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 WTest 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.
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Chapman, F. M.

1896. Notes on birds observed in Yucatan. American Museum of Natural History,
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Cole, Leon J.

1906. Aves from Yucatan. Harvard University, Museum of Comparative Zoology,
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Dickey, Donald R., and A. J. van Rossem.

1938. The birds of El Salvador. Field Museum of Natural History, Zoological
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1950. Distributional check-list of the birds of Mexico, Part 1. Cooper Ornitho-
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Grant, P. R.

1965. A systematic studv of the terrestrial birds of the Tres Marias Islands,
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1932. The distribution of bird-life in Guatemala. American Museum of Natural
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Hellmayr, C. E., and B. Conover.

1942. Catalogue of birds of the Americas. Field Museum of Natural History,
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Lawrence, George N.

1874. Birds of western and northwestern Mexico, based upon collections made
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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.
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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,
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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.
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1916. Birds of North and Middle America. U. S. National Museum Bulletin 50,
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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
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Skutch, Alexander F.

1964. Life histories of Central American pigeons. Wilson Bulletin, Vol. 76, No.
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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,
p. 51-56.
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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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

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