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Life histories of North American wood warblers, Part 1 (of 2)

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Title: Life histories of North American wood warblers, Part 1 (of 2)

Author: Arthur Cleveland Bent

Contributor: Winsor Marrett Tyler


        
Release date: June 28, 2012 [eBook #40100]
                Most recently updated: October 23, 2024

Language: English

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*** START OF THE PROJECT GUTENBERG EBOOK LIFE HISTORIES OF NORTH AMERICAN WOOD WARBLERS, PART 1 (OF 2) ***

    [Transcriber's Note: Emphasis Notation: _Italic_ and =Bold=;
       [M] and [F] represent male and female symbols respectively.
       Mathematical Notation - Whole and Fractional Part: 3-5/8;
       [=_] denotes the long sound, as in [=e]ve and [=u]se;
       [)_] denotes the short sound, as in [)e]nd and [)i]ll.]




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_Life Histories of North American Wood Warblers_


_by_


_Arthur Cleveland Bent_


_in two parts

Part 1_


_Dover Publications, Inc.
New York_




Published in the United Kingdom by Constable and Company, Limited,
10 Orange Street, London W. C. 2.


This Dover edition, first published in 1963, is an unabridged and
unaltered republication of the work first published in 1953 by the
United States Government Printing Office, as Smithsonian Institution
United States National Museum _Bulletin 203_.


_Standard Book Number: 486-21153-3_


Manufactured in the United States of America


Dover Publications, Inc.

180 Varick Street

New York 14, N. Y.




ADVERTISEMENT


The scientific publications of the National Museum include two series,
known, respectively, as _Proceedings_ and _Bulletins_.

The _Proceedings_ series, begun in 1878, is intended primarily as a
medium for the publication of original papers, based on the collections
of the National Museum, that set forth newly acquired facts in biology,
anthropology, and geology, with descriptions of new forms and revisions
of limited groups. Copies of each paper, in pamphlet form, are
distributed as published to libraries and scientific organizations and
to specialists and others interested in the different subjects. The
dates at which these separate papers are published are recorded in the
table of contents of each of the volumes.

The series of _Bulletins_, the first of which was issued in 1875,
contains separate publications comprising monographs of large zoological
groups and other general systematic treatises (occasionally in several
volumes), faunal works, reports of expeditions, catalogs of type
specimens, special collections, and other material of similar nature.
The majority of the volumes are octavo in size, but a quarto size has
been adopted in a few instances in which the larger page was regarded as
indispensable. In the _Bulletin_ series appear volumes under the heading
_Contributions from the United States National Herbarium_, in octavo
form, published by the National Museum since 1902, which contain papers
relating to the botanical collections of the Museum.

The present work forms No. 203 of the _Bulletin_ series.


REMINGTON KELLOGG,

_Director, United States National Museum_.




CONTENTS

                                                                     Page

   Introduction                                                        ix

   Order PASSERIFORMES                                                  1

   Family Parulidae: Wood warblers                                      1

     _Mniotilta varia_: Black-and-white warbler                         5
       Habits                                                           5
       Distribution                                                    14

     _Protonotaria citrea_: Prothonotary warbler                       17
       Habits                                                          17
       Distribution                                                    28

     _Limnothlypis swainsonii_: Swainson's warbler                     30
       Habits                                                          30
       Distribution                                                    37

     _Helmitheros vermivoros_: Worm-eating warbler                     38
       Habits                                                          38
       Distribution                                                    45

     _Vermivora chrysoptera_: Golden-winged warbler                    47
       Habits                                                          47
       Distribution                                                    56

     _Vermivora pinus_: Blue-winged warbler                            58
       Habits                                                          58
       Distribution                                                    65

     _Vermivora bachmanii_: Bachman's warbler                          67
       Habits                                                          67
       Distribution                                                    73

     _Vermivora peregrina_: Tennessee warbler                          75
       Habits                                                          75
       Distribution                                                    86

     _Vermivora celata celata_: Eastern orange-crowned warbler         89
       Habits                                                          89
       Distribution                                                    94

     _Vermivora celata orestera_: Rocky Mountain orange-crowned
         warbler                                                       98
       Habits                                                          98

     _Vermivora celata lutescens_: Lutescent orange-crowned warbler    99
       Habits                                                          99

     _Vermivora celata sordida_: Dusky orange-crowned warbler         103
       Habits                                                         103

     _Vermivora ruficapilla ruficapilla_: Eastern Nashville warbler   105
       Habits                                                         105
       Distribution                                                   113

     _Vermivora ruficapilla ridgwayi_: Western Nashville warbler      116
       Habits                                                         116

     _Vermivora virginiae_: Virginia's warbler                        119
       Habits                                                         119
       Distribution                                                   124

     _Vermivora crissalis_: Colima warbler                            126
       Habits                                                         126
       Distribution                                                   129

     _Vermivora luciae_: Lucy's warbler                               129
       Habits                                                         129
       Distribution                                                   134

     _Parula americana pusilla_: Northern parula warbler              135
       Habits                                                         135
       Distribution                                                   145

     _Parula americana americana_: Southern parula warbler            147
       Habits                                                         147

     _Parula pitiayumi nigrilora_: Sennett's olive-backed warbler     149
       Habits                                                         149
       Distribution                                                   152

     _Parula graysoni_: Socorro warbler                               152
       Habits                                                         152
       Distribution                                                   153

     _Peucedramus taeniatus arizonae_: Northern olive warbler         153
       Habits                                                         153
       Distribution                                                   160

     _Dendroica petechia aestiva_: Eastern yellow warbler             160
       Habits                                                         160
       Distribution                                                   178

     _Dendroica petechia amnicola_: Newfoundland yellow warbler       182
       Habits                                                         182

     _Dendroica petechia rubiginosa_: Alaska yellow warbler           184
       Habits                                                         184

     _Dendroica petechia morcomi_: Rocky Mountain yellow warbler      185
       Habits                                                         185

     _Dendroica petechia brewsteri_: California yellow warbler        186
       Habits                                                         186

     _Dendroica petechia sonorana_: Sonora yellow warbler             189
       Habits                                                         189

     _Dendroica petechia gundlachi_: Cuban yellow warbler             190
       Habits                                                         190

     _Dendroica petechia castaneiceps_: Mangrove yellow warbler       191
       Habits                                                         191

     _Dendroica magnolia_: Magnolia warbler                           195
       Habits                                                         195
       Distribution                                                   209

     _Dendroica tigrina_: Cape May warbler                            212
       Habits                                                         212
       Distribution                                                   222

     _Dendroica caerulescens caerulescens_: Northern black-throated
         blue warbler                                                 224
       Habits                                                         224
       Distribution                                                   233

     _Dendroica caerulescens cairnsi_: Cairns' warbler                237
       Habits                                                         237

     _Dendroica coronata coronata_: Eastern myrtle warbler            239
       Habits                                                         239
       Distribution 254

     _Dendroica coronata hooveri_: Alaska myrtle warbler              258
       Habits                                                         258

     _Dendroica auduboni auduboni_: Pacific Audubon's warbler         260
       Habits                                                         260
       Distribution                                                   271

     _Dendroica auduboni nigrifrons_: Black-fronted Audubon's warbler 273
       Habits 273

     _Dendroica nigrescens_: Black-throated gray warbler              275
       Habits                                                         275
       Distribution                                                   281

     _Dendroica towndsendi_: Townsend's warbler                       282
       Habits                                                         282
       Distribution                                                   290

     _Dendroica virens virens_: Northern black-throated green warbler 291
       Habits                                                         291
       Distribution                                                   304

     _Dendroica virens waynei_: Wayne's black-throated green warbler  308
       Habits                                                         308

     _Dendroica chrysoparia_: Golden-cheeked warbler                  316
       Habits                                                         316
       Distribution                                                   321

     _Dendroica occidentalis_: Hermit warbler                         321
       Habits                                                         321
       Distribution                                                   328

     _Dendroica cerulea_: Cerulean warbler                            329
       Habits                                                         329
       Distribution                                                   335

     _Dendroica fusca_: Blackburnian warbler                          337
       Habits                                                         337
       Distribution                                                   347

     _Dendroica dominica dominica_: Eastern yellow-throated warbler   349
       Habits                                                         349
       Distribution                                                   358

     _Dendroica dominica albilora_: Sycamore yellow-throated warbler  359
       Habits                                                         359

     _Dendroica graciae graciae_: Northern Grace's warbler            363
       Habits                                                         363
       Distribution                                                   367



INTRODUCTION


This is the nineteenth in a series of bulletins of the United States
National Museum on the life histories of North American birds. Previous
numbers have been issued as follows:

  107. Life Histories of North American Diving Birds, August 1, 1919.

  113. Life Histories of North American Gulls and Terns, August 27,
       1921.

  121. Life Histories of North American Petrels and Pelicans and Their
       Allies, October 19, 1922.

  126. Life Histories of North American Wild Fowl (part), May 25, 1923.

  130. Life Histories of North American Wild Fowl (part), June 27, 1925.

  135. Life Histories of North American Marsh Birds, March 11, 1927.

  142. Life Histories of North American Shore Birds (pt. 1), December
       31, 1927.

  146. Life Histories of North American Shore Birds (pt. 2), March 24,
       1929.

  162. Life Histories of North American Gallinaceous Birds, May 25,
       1932.

  167. Life Histories of North American Birds of Prey (pt. 1), May 3,
       1937.

  170. Life Histories of North American Birds of Prey (pt. 2), August 8,
       1938.

  174. Life Histories of North American Woodpeckers, May 23, 1939.

  176. Life Histories of North American Cuckoos, Goatsuckers,
       Hummingbirds, and Their Allies, July 20, 1940.

  179. Life Histories of North American Flycatchers, Larks, Swallows,
       and Their Allies, May 8, 1942.

  191. Life Histories of North American Jays, Crows, and Titmice,
       January 27, 1947.

  195. Life Histories of North American Nuthatches, Wrens, Thrashers,
       and Their Allies, July 7, 1948.

  196. Life Histories of North American Thrushes, Kinglets, and Their
       Allies, June 28, 1949.

  197. Life Histories of North American Wagtails, Shrikes, Vireos, and
       Their Allies, June 21, 1950.


The paragraphs on distribution for the Colima and Kirtland's warblers
were supplied by Dr. Josselyn Van Tyne with his contributions on these
species.

All other data on distribution and migration were contributed by the
Fish and Wildlife Service under the supervision of Frederick C. Lincoln.

The same general plan has been followed as explained in previous
bulletins, and the same sources of information have been used. It does
not seem necessary to explain the plan again here. The nomenclature of
the Check-List of North American Birds (1931), with its supplements, of
the American Ornithologists' Union, has been followed. Forms not
recognized in this list have not been included.

Many who have contributed material for previous Bulletins have continued
to cooperate. Receipts of material from several hundred contributors has
been acknowledged in previous Bulletins. In addition to these, our
thanks are due to the following new contributors: G. A. Ammann, O. L.
Austin, Jr., F. S. Barkalow, Jr., Ralph Beebe, H. E. Bennett, A. J.
Berger, Virgilio Biaggi, Jr., C. H. Blake, Don Bleitz, B. J. Blincoe, L.
C. Brecher, Jeanne Broley, Maurice Broun, J. H. Buckalew, I. W. Burr, N.
K. Carpenter, May T. Cooke, H. L. Crockett, Grace Crowe, Ruby Curry, J.
V. Dennis, E. von S. Dingle, M. S. Dunlap, J. J. Elliott, A. H. Fast,
Edith K. Frey, J. E. Galley, J. H. Gerard, Lydia Getell, H. B.
Goldstein, Alan Gordon, L. I. Grinnell, Horace Groskin, F. G. Gross, G.
W. Gullion, E. M. Hall, R. H. Hansman, Katharine C. Harding, H. H.
Harrison, J. W. Hopkins, N. L. Huff, Verna R. Johnston, Malcolm Jollie,
R. S. Judd, M. B. Land, Louise de K. Lawrence, R. E. Lawrence, G. H.
Lowery, J. M. Markle, C. R. Mason, D. L. McKinley, R. J. Middleton, Lyle
Miller, A. H. Morgan, R. H. Myers, W. H. Nicholson, F. H. Orcutt, H. L.
Orians, R. A. O'Reilly, A. A. Outram, G. H. Parks, K. C. Parkes, M. M.
Peet, J. L. Peters, F. A. Pitelka, Mariana Roach, James Rooney, Jr., O.
M. Root, G. B. Saunders, James Sawders, Mary C. Shaub, Dorothy E.
Snyder, Doris Heustis Speirs, E. A. Stoner, P. B. Street, H. R. Sweet,
E. W. Teale, A. B. Williams, G. G. Williams, R. B. Williams, Mrs. T. E.
Winford, and A. M. Woodbury.

As the demand for these Bulletins is much greater than the supply, the
names of those who have not contributed to the work during recent years
will be dropped from the author's mailing list.

Dr. Winsor M. Tyler has again read and indexed for this volume a large
part of the current literature on North American birds and has
contributed four complete life histories. Dr. Alfred O. Gross has
written stories on the yellow-throats (_Geothlypis trichas_) and has
contributed three other complete life histories. Edward von S. Dingle,
Alexander Sprunt, Jr., and Dr. Josselyn Van Tyne have contributed two
complete life histories each.




INTRODUCTION


William George F. Harris has increased his valuable contribution to the
work by producing the entire paragraphs on eggs, including descriptions
of the eggs in their exact colors, assembling and averaging the
measurements, and collecting and arranging the egg dates, as they appear
under Distribution; the preparation of this last item alone required the
handling of over 5,600 records.

Clarence F. Smith has furnished references to food habits of all the
species of wood warblers. Aretas A. Saunders has contributed full and
accurate descriptions of the songs and call notes of all the species
with which he is familiar, based on his extensive musical records. Dr.
Alexander F. Skutch has sent us full accounts of all the North American
wood warblers that migrate through or spend the winter in Central
America, with dates of arrival and departure. James Lee Peters has
furnished descriptions of molts and plumages of several species and has
copied several original descriptions of subspecies from publications
that were not available to the author.

Eggs were measured for this volume by American Museum of Natural History
(C. K. Nichols), California Academy of Sciences (R. T. Orr), Colorado
Museum of Natural History (F. G. Brandenburg), C. E. Doe, W. E. Griffee,
W. C. Hanna, E. N. Harrison, H. L. Heaton, A. D. Henderson, Museum of
Comparative Zoology (W. G. F. Harris), and Museum of Vertebrate Zoology
(M. Jollie).

The manuscript for this Bulletin was written in 1945; only important
information could be added. If the reader fails to find in these pages
anything that he knows about the birds, he can only blame himself for
failing to send the information to--

THE AUTHOR.




LIFE HISTORIES OF NORTH AMERICAN WOOD WARBLERS




Order PASSERIFORMES: Family PARULIDAE

By ARTHUR CLEVELAND BENT

_Taunton, Mass_.



GENERAL REMARKS ON THE FAMILY PARULIDAE

CONTRIBUTED BY WINSOR MARRETT TYLER


The family of wood warblers, Parulidae, is the second largest family of
North American birds, surpassed only in number of species by the family
Fringillidae. The wood warblers occur only in the Western Hemisphere;
they are distinct from the Old World warblers, Sylviidae, although the
two families play a similar rôle in nature's economy.

The wood warblers are largely nocturnal migrants, whose long journeys in
the dark of night over sea and lake and along the coast expose them to
many perils, one being the lighthouses they strike with frequently fatal
results. Their notes are seldom heard from the night sky during their
spring migration, but on many a calm, quiet night in August and
September, as they fly overhead, their sharp, sibilant, staccato notes
punctuate the rhythmic beat of the tree-crickets singing in the
shrubbery and stand out clearly among the soft, whistled calls of the
migrating thrushes.

The length of migration varies greatly; the pine warbler withdraws in
winter only a short distance from the southern limit of its breeding
range, whereas the most northerly breeding black-polls migrate from
Alaska to the Tropics. In spring many species migrate at nearly the same
time, apparently advancing northward in intermittent waves of great
numbers during favorable nights. Flocks made up of sometimes a dozen
species together flash about in their bright plumage during the week or
two at the height of the migration and furnish days of great excitement
to ornithologists. Their return in late summer and autumn is more
leisurely and regular; in loose flocks they drift slowly by for several
weeks, their southward passage evident even in daytime. The flocking
begins early, soon after nesting is over, and to the north is apparent
early in July, if closely watched for, even before the leaves begin to
wither. The mixed fall flocks, with adults in winter plumage and young
birds in duller colors, present many fascinating problems in
identification as the birds move quietly along.

[AUTHOR'S NOTES: When I asked Dr. Tyler to contribute these remarks we
discussed Professor Cooke's (1904) theory of trans-Gulf migration, which
has been generally accepted until recently, when it was challenged by
George C. Williams (1945). This paper started a discussion in which
George H. Lowery, Jr. (1945), has taken a prominent part, and of which
we have not yet heard the last. Routes of migration from South America
to the United States are evidently well established through the West
Indies and the Bahamas to the southeastern States; across the Caribbean
to Jamaica, Cuba, and Florida; through Central America and directly
across the Gulf from Yucután to the Gulf States; through eastern Mexico
and Texas; and through western Mexico to the southwestern States.
Professor Cooke was probably correct in assuming that the majority of
wood warblers breeding in eastern North America migrate directly across
the Caribbean or the Gulf. Some species may confine themselves to only
one of the routes named, but we need more data to say just which species
uses what route.]

The literature contains descriptions of several warblers not recognized
as established species by the A. O. U. Check-List (1931). Some,
described and illustrated by older writers such as Wilson and Audubon,
cannot be identified; others are presumably hybrids; and one, _Sylvia
autumnalis_ Wilson, the autumn warbler, is clearly the black-poll in
fall plumage. The first category includes _Dendroica carbonata_
(Audubon), the carbonated warbler, of which the Check-List says "the
published plates may have been based to some extent on memory"; _D.
montana_ (Wilson), the blue mountain warbler, which is "known only from
the plates of Audubon and Wilson"; and _Wilsonia_ (?) _microcephala_
(Ridgway), the small-headed flycatcher, of which it says: "Known only
from the works of Wilson and Audubon whose specimens came from New
Jersey and Kentucky respectively. There is some question whether they
represent the same species."

In the second category is _Vermivora cincinnatiensis_ (Langdon), the
Cincinnati warbler, described in 1880. "The unique type is regarded as a
hybrid between _Vermivora pinus_ (Linnaeus) and _Oporornis formosa_
(Wilson)." Recently, in a letter dated August 3, 1948, Dr. George M.
Sutton reports to Mr. Bent the discovery of a second Cincinnati warbler,
taken in Michigan on May 28, 1948. He says: "Its bill and feet are large
for _Vermivora_ and its under tail coverts proportionately too long for
that genus. It has only a faint suggestion of wing-barring and the
merest shadow of a pattern on the outer rectrices. One of its most
interesting and beautiful characters is the gray tipping of the feathers
at the rear of the crown, as in _O. formosus_. The effect is very
unusual, for the gray-tipped feathers are yellow. It is, in short,
obviously a cross between _Vermivora_ and _Oporornis_."

The status of _Vermivora leucobronchialis_ and _V. lawrencii_ and the
relationship between them puzzled ornithologists for upward of two
generations. William Brewster (1876) described the former as a new
species, and since that time, as Walter Faxon (1911) writes, "almost
every conceivable hypothesis has been advanced by one writer or another
to fix its true status in our bird-fauna." In addition to being
considered a valid species, it has been regarded as a hybrid (Brewster,
1881), as a dichromatic phase, that is, a leucochroic phase of _V.
pinus_ (Ridgway, 1887), as a mutant (Scott, 1905), and finally as a
phase, "ancestral in character" (atavistic) of the goldenwing (C. W.
Townsend, 1908).

Lawrence's warbler is a very rare bird. The first specimen was described
in 1874 (Herold Herrick, 1874), and since that time the bird was taken
or seen infrequently, chiefly in regions where the breeding ranges of
_V. chrysoptera_ and _V. pinus_ overlap. Consensus of opinion in the
main regarded it as a hybrid between _V. chrysoptera_ and _V. pinus_, as
it combined characters of both the supposed parents. John Treadwell
Nichols (1908) some years ago brought new light to the problem. He says:

In any discussion of the status of Lawrence's and Brewster's Warblers it
is well to bear in mind the facts, including the much greater abundance
of Brewster's, are in accord with Mendel's Law of Heredity, supposing
both forms to be hybrids between _Helminthophila pinus_ and _H.
chrysoptera_. * * * All the first generation hybrids will be Brewster's
Warbler in plumage. In the next generation there will be pure
Golden-winged Warblers, pure Blue-winged Warblers, pure Brewster's
Warblers, and pure Lawrence's Warblers; also mixed birds of the first
three forms, but none of the last form, which, being recessive, comes to
light only when pure. The original hybrids then (which will be all
Brewster's in plumage) must be fertile with one another or with the
parent species for any Lawrence's to occur; and if they are perfectly
fertile Lawrence's must still remain a small minority. After the first
generation the proportion of plumages of birds with mixed parentage
should be: 9 Brewster's, 3 _chrysoptera_, 3 _pinus_, 1 Lawrence's.

This explanation removed the stumbling block, long believed to
be insurmountable, that a black-throated bird, mating with a
yellow-throated bird, could produce progeny having a white throat.
Under Mendel's Law the dominant color (white) of _chrysoptera_ would
appear by the suppression of the recessive black throat.

Fortunately, Walter Faxon (1913) not long afterward found a female
blue-winged warbler mated with a goldenwing and was successful in
following the resulting brood of young birds until they had acquired
their first winter plumage when, fulfilling Mendel's Law, they were all
in "the garb of _Helminthophila leucobronchialis_," thus establishing
beyond a doubt the hybrid nature of the bird. At the end of his paper,
Walter Faxon (1913) relates a bit of interesting ancient history
regarding these three species of _Vermivora_. He says:

In my paper published in 1911, after stating the different hypotheses
proposed in order to explain the relations existing among the
Golden-winged, Blue-winged, Brewster's, and Lawrence's Warblers I added,
half in jest, that the only hypothesis left for a new-comer in the field
was this: that the Golden-winged and the Blue-winged Warblers themselves
were merely two forms of one species. Curiously enough, not long after
this I found that this very opinion had been expressed, and in a most
unexpected quarter: in a letter dated Edinburgh, Sept. 15, 1835, Audubon
wrote to Bachman that he suspected the golden-winged warbler and the
blue-winged warbler were one species! That Audubon at that early date,
ignorant (as he was assumed to be) of the existence of Brewster's and
Lawrence's warblers, and but superficially acquainted with the
golden-wing, should suspect that two birds so diverse as the blue-wing
and the golden-wing were one species seemed incomprehensible, and in the
light of what we now know about these birds, his surmise seemed to
presuppose an almost superhuman faculty of prevision.

As a possible explanation of Audubon's letter I have only this to offer:
in the winter of 1876-77 Dr. Spencer Trotter discovered in the
collection of the Academy of Natural Sciences of Philadelphia a specimen
of Brewster's warbler without a label, the third specimen known up to
that time; on the bottom of the stand was written in the autograph of
John Cassin, "J. C., 20 October, 1862," and also a badly blurred legend
"Not from Bell." An appeal to J. G. Bell elicited the response that he
remembered shooting a peculiar warbler in Rockland Co., N. Y., about the
year 1832--a warbler something like a golden-wing, but lacking, although
in high plumage, the black throat of that species; a great many years
afterward, he sold this specimen in Philadelphia but knew nothing of its
ultimate fate. Dr. Trotter justly inferred that the Philadelphia Academy
specimen was in all probability the very bird shot by Bell.

Now as Audubon was intimately associated with Bell, is it not possible
that he had examined this example of Brewster's warbler? In that case,
seeing that this bird's characters were in part those of the blue-wing,
in part those of the golden-wing, he may have inferred the interbreeding
of these two birds, and so (rather unwarrantably, it is true) their
identity. If this be not the explanation of the passage in Audubon's
letter to Bachman I have no other to suggest.

When Audubon came to publish his account of the Golden-winged Warbler in
1839 (Ornithological Biography, 1839, 5, p. 154) he said not a word
about its connection with the Blue-winged Warbler.

Recently Karl W. Haller (1940) described "a new wood warbler from West
Virginia" from two specimens, male and female, which he collected on May
30 and June 1, 1939, respectively, at points 18 miles apart, and
proposed for it the new name _Dendroica potomac_, Sutton's warbler.
These birds resemble the yellow-throated warbler in plumage but lack
streaks on the sides. They also suggest the parula warbler in having a
faint yellowish wash on the back and, in the male, "an almost
imperceptible hint of raw sienna" on the upper breast. The male sang a
song much like the parula's, but doubled by repetition.

Two more Sutton's warblers have been carefully observed in the field:
one at the point where the type was collected on May 21, 1942, by
Maurice Brooks and Bayard H. Christy (1942); the second about 18 miles
to the westward on June 21, 1944, by George H. Breiding and Lawrence E.
Hicks (1945). Another aberrant warbler has been described by Stanley G.
Jewett (1944), who examined four specimens which show a curious
intermingling of the plumage characters of the hermit and Townsend
warblers.

[AUTHOR'S NOTE: Since the above was written, Kenneth C. Parkes (1951)
has published a study of the genetics of the golden-winged--blue-winged
warbler complex, to which the reader is referred.]




MNIOTILTA VARIA (Linnaeus)

BLACK-AND-WHITE WARBLER


CONTRIBUTED BY WINSOR MARRETT TYLER

PLATES 1-3


HABITS

The black-and-white warbler is one of the earliest spring warblers to
reach its breeding-ground in the Transition Zone. Most of the other
members of this family arrive in or pass through the region in mid-May
or somewhat later, according to the season, when the oaks are in bloom
and the opening flowers attract swarms of insects.

The black-and-white warbler, however, owing to its peculiar habit of
feeding on the trunks and the large limbs of the trees, does not have to
wait for the bounty supplied by the oaks but finds its special
feeding-ground well stocked with food long before the oaks blossom or
their leaves unfold. It comes with the yellow palm warbler late in
April, when many of the trees are nearly bare, and not long after the
pine warbler.

_Mniotilta_ is a neat little bird, dressed in modest colors, at this
season singing its simple but sprightly song as it scrambles over the
bark--the black-and-white creeper, Alexander Wilson calls it.

Milton B. Trautman (1940), speaking of the spring migration at Buckeye
Lake, Ohio, shows that the male birds are preponderant in the earliest
flights. He says: "The first spring arrivals, chiefly males, were noted
between April 16 and 30, and between May 1 and 5, 2 to 15 birds, mostly
males, could be daily noted. The peak of migration usually lasted from
May 6 to May 18, and then from 3 to 42 individuals, consisting of a few
old males and the remainder females and young males, were daily
observed. On May 18 or shortly thereafter a decided lessening in numbers
occurred, and by May 23 all except an occasional straggler had left."

_Courtship._--Forbush (1929) gives this hint of courtship, which
resembles the activities of most warblers at this season: "When the
females arrive there is much agitation, and often a long-continued
intermittent pursuit, with much song and fluttering of black and white
plumage, and much interference from rival males before the happy pair
are united and begin nesting."

_Nesting._--The black-and-white warbler usually builds its nest on the
ground, tucking it away against a shrub or tree, or even under the
shelter of an overhanging stone or bank. The nest is generally concealed
among an accumulation of dead leaves which, arching over it, hides it
from above. It is made, according to A. C. Bent (MS.), "of dry leaves,
coarse grass, strips of inner bark, pine needles and rootlets, and is
lined with finer grasses and rootlets and horsehair." I have seen a nest
made chiefly of pine needles on a base of dry leaves.

Henry Mousley (1916), writing of Hatley, Quebec, mentions moss as a
component part of the nest, and says of three nests that they were all
"heavily lined with long black and white horse hairs," a peculiarity of
coloration mentioned in one of Mr. Bent's nests. Thomas D. Burleigh
(1927b) speaks of a nest in Pennsylvania "built of dead leaves and
rhododendron berry stems, lined with fine black rootlets and a few white
hairs." H. H. Brimley (1941) describes an exceptional nest. He says:
"There was no particular departure from normal in its construction
except for the fact that it was lined with a mixture of fine rootlets
and very fine copper wire, such as is used in telephone cables.
Fragments of such cable, discarded by repair men, were found nearby
where a telephone line ran through the woods."

Cordelia J. Stanwood (1910c) speaks of a nest "built in a depression
full of leaves, behind a flat rock. * * * The cavity was shaped on a
slant, the upper wall forming a partial roof. * * * It looked not unlike
a small-sized nest of an Oven-bird. On the inside, the length was 2-1/2
inches, width 1-1/2 inches, depth 2 inches. On the outside, length 3-1/2
inches, width 2-1/2 inches, depth 2-1/2 inches. Thickness of wall at the
top of nest, 1 inch; at the bottom, 1/2 inch." Henry Mousley (1916)
gives the average dimensions of three nests as "outside diameter 3-3/4,
inside 1-3/4 inches; outside depth 2-1/4, inside 1-1/2 inches."

F. A. E. Starr (MS.) writes to A. C. Bent from Toronto, Ontario, that
all the nests he has found have been in broken-off stumps in low woods.
"The cavity in the top of the stump," he says, "is filled with old
leaves, and the nest proper is made chiefly of strips of bark with grass
and fiber." Guy H. Briggs (1900) reports a nest "in a decayed hemlock
stump, fifteen inches from the ground." In such cases, of course, while
the nest is well above the ground level, it rests on a firm foundation.

Audubon (1841) says: "In Louisiana, its nest is usually placed in some
small hole in a tree," but he quotes a letter to him from Dr. T. M.
Brewer on the subject, thus: "This bird, which you speak of as breeding
in the hollows of trees, with us always builds its nest on the ground. I
say always, because I never knew it to lay anywhere else. I have by me a
nest brought to me by Mr. Appleton from Batternits, New York, which was
found in the drain of the house in which he resided."

Minot (1877) speaks of two nests found near Boston, Mass., well above
the ground. He says: "The first was in a pine grove, in the cavity of a
tree rent by lightning, and about five feet from the ground, and the
other on the top of a low birch stump, which stood in a grove of white
oaks."

Gordon Boit Wellman (1905) states: "Toward the last of the incubation
time one of the birds was constantly on the nest. I found the male
sitting usually at about dusk, but I think the female sat on the eggs
over night."

_Eggs._--[AUTHOR'S NOTE: The black-and-white warbler usually lays 4 or 5
eggs to a set, normally 5, seldom fewer or more. These are ovate to
short ovate and slightly glossy. The ground color is white or creamy
white. Some are finely sprinkled over the entire surface with
"cinnamon-brown," "Mars brown," and "dark purplish drab"; others are
boldly spotted and blotched with "russet" and "Vandyke brown," with
underlying spots of "brownish drab," "light brownish drab," and "light
vinaceous-drab." Speckled eggs are commoner than the more boldly
blotched type. The markings are usually concentrated at the large end,
and on some of the heavily spotted eggs there is a solid wreath of
different shades of russet and drab. The measurements of 50 eggs average
17.2 by 13.3 millimeters; the eggs showing the four extremes measure
=18.8= by 13.7, 17.9 by =14.7=, =15.7= by 12.7, and 16.3 by =12.2=
millimeters (Harris).]

_Young._--Cordelia J. Stanwood (MS.) speaks of the nestlings a few days
from the egg as "very dark gray, much like young juncos and Nashville
warblers." But when they leave the nest they are clearly recognizable as
young black-and-white warblers, although they are slightly tinged with
brownish. By mid-July, here in New England, they assume their first
winter plumage, and, as both sexes of the young birds have whitish
cheeks, they resemble very closely their female parent.

Unlike the young of some of the other warblers which remain near the
ground for many days, the young black-and-white warblers shortly ascend
to the branches of trees where they are fed by the old birds.

I find no definite record of the length of the incubation period, but in
a nest I watched in 1914 it was close to 10 days. Burns (1921) gives the
period of nestling life as 8 to 12 days.

_Plumages._--[AUTHOR'S NOTE: Dr. Dwight (1900) calls the natal down
mouse gray, and describes the juvenal plumage as follows: "Above,
wood-brown streaked with dull olive-brown, the upper tail coverts dusky;
median crown and superciliary stripe dingy white. Wings and tail dull
black, edged chiefly with ashy gray, the tertiaries (except the proximal
which is entirely black) broadly edged with white, buff tinged on the
middle one. Two buffy white wing bands at tips of greater and median
wing coverts. The outer two rectrices with terminal white blotches of
variable extent on the inner webs. Below, dull white, washed on the
throat and sides with wood-brown, obscurely streaked on throat, breast,
sides and crissum with dull grayish black."

A postjuvenal molt begins early in July, involving everything but the
flight feathers; this produces in the young male a first winter plumage
which is similar to the juvenal, but whiter and more definitely
streaked. "Above, striped in black and white, the upper tail coverts
black broadly edged with white; median crown and superciliary stripe
pure white. The wing bands white. Below, pure white streaked with bluish
black on sides of breast, flanks and crissum, the black veiled by
overlapping white edgings; the chin, throat, breast and abdomen
unmarked. Postocular stripe black; the white feathers of the sides of
the head tipped with black."

The first nuptial plumage is acquired by a partial prenuptial molt in
late winter, which involves a large part of the body plumage, but not
the wings or the tail. "The black streaks of the chin and throat are
acquired, veiled with white, and the loral, subocular and auricular
regions become jet-black. The brown primary coverts distinguish young
birds and the chin is less often solidly black than in adults."

The adult winter plumage is acquired by a complete postnuptial molt,
beginning early in July. It differs from the first winter dress "in
having the chin and throat heavily streaked with irregular chains of
black spots veiled with white edgings, the wings and tail blacker and
the edgings a brighter gray. * * * The female has corresponding plumages
and moults, the first prenuptial moult often very limited or suppressed.
In juvenal dress the wings and tail are usually browner with duller
edgings and the streaking below obscure. In first winter plumage the
streakings are dull and obscure everywhere, a brown wash conspicuous on
the flanks and sides of the throat. The first nuptial plumage is gained
chiefly by wear through which the brown tints are largely lost, the
general color becoming whiter and the streaks more distinct. The adult
winter plumage is rather less brown than the female first winter, the
streaking less obscure and the wings and tail darker. The adult nuptial
plumage, acquired partly by moult, is indistinguishable with certainty
from the first nuptial."]

_Food._--McAtee (1926) summarizes the food of the species thus:

In its excursions over the trunks and larger limbs of trees the Black
and White Creeper is certainly not looking for vegetable food, and only
a trace of such matter has been found in the stomachs examined. The
food is chiefly insects but considerable numbers of spiders and
daddy-long-legs also are eaten. Beetles, caterpillars, and ants are the
larger classes of insect food, but moths, flies, bugs, and a few
hymenoptera also are eaten. Among forest enemies that have been found in
stomachs of this species are round-headed wood borers, leaf beetles,
flea beetles, weevils, bark beetles, leaf hoppers, and jumping plant
lice. The hackberry caterpillar, the hackberry psyllid, an oak leaf
beetle _Xanthonia 10-notata_, and the willow flea beetle, are forms
specifically identified. Observers have reported this warbler to feed
also upon ordinary plant lice, and upon larvae of the gypsy moth.

Forbush (1929) adds the following observation: "The food of this bird
consists mostly of the enemies of trees, such as plant-lice, scale-lice,
caterpillars, both hairy and hairless, among them such destructive
enemies of orchard, shade and forest trees as the canker-worm and the
gipsy, brown-tail, tent and forest tent caterpillars. Wood-boring and
bark-boring insects, click beetles, curculios and many other winged
insects are taken. Sometimes when the quick-moving insects escape its
sharp bill, it pursues them on the wing but most of its attention is
devoted to those on the trees."

H. H. Tuttle (1919), speaking of the male parent feeding the young
birds, says: "The fare which he provided was composed entirely of small
green caterpillars, cut up into half-lengths."

_Behavior._--The black-and-white warbler seems set apart from others of
the group, perhaps because of its marked propensity for clambering over
the trunks of trees and their larger branches. Although, like other
warblers, it seems at home among the smaller twigs, it spends a large
part of its time on upright surfaces over which it moves easily and
quickly, upward, downward, and spirally, with great agility and sureness
of footing, constantly changing direction, and not using the tail for
support. As it scrambles over the bark, it switches from side to side as
if at each hop it placed one foot and then the other in advance, and
even on slim branches it hops in the same way, the tail alternately
appearing first on one side of the branch and then on the other; it
reminds us of a little schoolgirl swishing her skirt from side to side
as she walks down the street. The bird is alert and watchful, and if it
starts an insect from the bark, or sees one flying near, it may pursue
it and catch it in the air.

H. H. Tuttle (1919) describes an extreme example of behavior simulating
a wounded bird. He says: "She struck the leaves with a slight thud and
turned over on her side, while the toes of one up-stretched leg
clutched at the air and her tail spread slowly into a pointed fan. * * *
Deceived for a moment then, I turned a step in her direction. She lay
quite still except for a quivering wing. I reached out toward her with a
small stick and touched her side; she screamed pitifully; I stretched
out my hand to pick her up, but with a last effort she righted herself,
and by kicking desperately with one leg, succeeded in pushing forward a
few inches."

We associate this warbler with dry, rocky hillsides where the ground is
strewn with dead leaves, but the bird may breed also in the dry portions
of shady, wooded swamps.

_Voice._--The black-and-white is one of the high-voiced singers. Its
song is made up of a series of squeaky couplets given with a
back-and-forth rhythm, a seesawing effect, like the ovenbird's song
played on a fine, delicate instrument. It may be suggested by
pronouncing the syllables _we see_ rapidly four or five times in a
whispered voice. In the distance the song has a sibilant quality; when
heard near at hand a high, clear whistle may be detected in the notes.
The final note in the song is the accented _see_.

Albert R. Brand (1938), in his mechanically recorded songs of warblers,
placed the black-and-white's song as the fourth highest in pitch in his
last of 16 species, the black-poll, blue-winged, and the Blackburnian
being higher. He gives the approximate mean (vibrations per second) of
the black and white as 6,900 and of the blackpool as 8,900.

Aretas A. Saunders (MS.) says: "The pitch of the songs varies, according
to my records, from B´´´ to E´´´´, a range of three and a half tones
more than an octave. A single song, however, does not vary more than
three and a half tones."

A second song, not heard, I think, until the bird has been on its
breeding ground for some time, is rather more pleasing, less monotonous,
than the first. It is longer, somewhat faster, more lively, and is
modulated in pitch. Francis H. Allen (MS.) speaks of it thus: "Later in
the season a more elaborate song is very commonly heard. I have been
accustomed to syllabify it as _weesy, weesy, weesy, weesy, woosy, woosy,
weesy, weesy_. The notes indicated by _woosy_ really differ from the
others only by being pitched lower."

Occasionally we hear aberrant songs which prove puzzling until we can
see the singer. Allen remarks that he has heard several such songs, and
I remember hearing one in which the lower note of each couplet was
reduplicated, thereby strongly suggested one of the songs of the
Blackburnian warbler. Sometimes _Mniotilta_ sings during flight. I once
heard a song from a bird flying within a few feet of me--at this range a
sound of piercing sharpness.

Of the minor notes Andrew Allison (1907) says: "I know of no other
warbler except the Chat that can produce so great a variety of sounds;
and since nearly all of the notes resemble those of other warblers, this
is a most confusing bird to deal with during the busy season of
'waves'."

The call note often has a buzzing quality, and often runs into a long
chatter (also characteristic of the young bird), but it may be given so
sharply enunciated that it suggests the _chip_ of the black-poll. Allen
(MS.) writes it _chi_, "like pebbles struck together," and Cordelia J.
Stanwood (1910) renders it _sptz_, saying "the sound resembled the noise
made by a drop of syrup sputtering on a hot stove."

_Field marks._--The black-poll, in its spring plumage, and the
black-and-white warbler resemble each other in coloration, but the
latter bird may be readily distinguished by its white stripe down the
center of the crown and the white line over the eye. The contrast in the
behavior of the two birds separates them at a glance.

_Enemies._--Like other birds which build on the ground, the
black-and-white is subject, during the nesting season, to attacks by
snakes and predatory mammals. A. D. DuBois (MS.) cites a case in which
maggots destroyed a nestful of young birds.

Harold S. Peters (1936) reports that a fly, _Ornithoica confluens_ Say,
and a louse, _Myrsidea incerta_ (Kellogg), have been found in the
plumage of the black-and-white warbler.

Herbert Friedmann (1929) says: "This aberrant warbler is a rather
uncommon victim of the Cowbird, only a couple dozen definite instances
having come to my notice. * * * The largest number of Cowbirds' eggs
found in a single nest of this Warbler is five, together with three eggs
of the owner." George W. Byers (1950) reports a nest of this warbler, in
Michigan, that held two eggs of the warbler and eight of the cowbird, on
which the warbler was incubating. His photograph of the eggs suggests
that they were probably laid by four different cowbirds.

_Fall._--Several of the warblers show a tendency to stray from their
breeding grounds soon after their young are able to care for themselves,
perhaps even before the postnuptial molt is completed and long before
the birds gather into the mixed autumn flocks. Among these early
wandering birds the black-and-white warbler is a very conspicuous
species, perhaps because it is one of our commoner birds or, more
probably, because of its habit of feeding in plain sight on the trunks
and low branches of dead or dying trees and shrubs instead of hiding,
like other warblers, high up in the foliage. It may be that the warblers
we see at some distance from their breeding grounds thus early in the
season have already begun their migration toward the south: they often
appear to be migrating.

Behind the house in Lexington, Mass., where I lived for years, there was
a little hill, sparsely covered with locust trees, to the southward from
my dooryard. This hill was a favorite resort for warblers in late
summer. No warbler bred within a mile of the spot, except the summer
yellowbird, to use the old name, yet soon after the first of July the
black-and-white warblers began to assemble there. Not infrequently I
have seen a single bird come to the hill, flying in from the north
across Lexington Common, and join others there. The small company might
remain for an hour or more, frequently singing (evidently adult males)
as the birds fed in the locust trees.

Later in the season, as August advances, migration appears more evident.
The birds now gather in larger numbers, sometimes as many as eight or
ten; they pause in the locust trees for a shorter time before flying
off; they are no longer in song; and the majority of the birds have
white cheeks, most of them presumably young birds. Although they are
almost silent as they climb about feeding, if you stand quietly in the
midst of a company of four or five, now and then you may hear a faint
note, and at once the note comes from all sides, each bird apparently
reporting its whereabouts--a sound which calls to mind the south-bound
migrants as they roam through the quiet autumn woods. Other warblers,
unquestionably migrants, visit this hillside in August, notably the
Tennessee, an early arrival who has already traveled a long way.

The fall migration of the black-and-white is long-drawn-out. The bird
does not depend, like many of the warblers, on finding food among the
foliage, so it may linger long after the trees are bare of leaves,
sometimes, here in New England, well into October. I saw a bird in
eastern Massachusetts on October 23, 1940, a very late date.

_Winter._--Dr. Alexander F. Skutch (MS.) sent to A. C. Bent the
following comprehensive account of the bird on its winter quarters:
"None of our warblers is more catholic in its choice of a winter home
than the black-and-white. Upon its departure from its nesting range, it
spreads over a vast area from the Gulf States south to Ecuador and
Venezuela, from the Pacific coast of Mexico and Central America eastward
through the Antilles. And in the mountainous regions of its winter range
it does not, like so many members of the family, restrict itself to a
particular altitudinal zone, but on the contrary scatters from sea level
high up into the mountains. As a result of this wide dispersion,
latitudinal and altitudinal, it appears to be nowhere abundant in
Central America during the winter months, yet it has been recorded from
more widely scattered localities than most other winter visitants. On
the southern coast of Jamaica, in December 1930, I found a greater
concentration of individuals than I have ever seen in Central America
during midwinter.

"Wintering throughout the length of Central America, from near sea level
up to 9,000 feet and rarely higher, the black-and-white warbler is
somewhat more abundant in that portion of its altitudinal range
comprised between 2,000 or 3,000 and 7,000 or 8,000 feet above sea
level. It is found in the heavy forest, in the more open types of
woodland, among the shade trees of the coffee plantations, and even amid
low second-growth with scattered trees. It creeps along the branches in
exactly the same fashion in its winter as in its summer home. Solitary
in its disposition, two of the kind are almost never seen together. The
only time I have heard this warbler sing in Central America was also one
of the very few occasions when I found two together. Early on the bright
morning of September 1, 1933, when the warblers were arriving from the
north, I heard the black-and-white's weak little song repeated several
times among the trees at the edge of an oak wood, at an altitude of
8,500 feet in the Guatemalan highlands. Looking into the tree tops, I
saw two of these birds together. Apparently they were singing in
rivalry, as red-faced warblers, Kaup's redstarts, yellow warblers, and
other members of the family solitary during the winter months will sing
in the face of another of their kind, at seasons when they are usually
silent. Often such songs lead to a pursuit or even a fight; but I have
never seen black-and-white warblers actually engaged in a conflict in
their winter home.

"Although intolerant of their own kind, the black-and-white warblers are
not entirely hermits; for often a single one will attach itself to a
mixed flock of small birds. In the Guatemalan highlands, during the
winter months, such flocks are composed chiefly of Townsend's warblers;
and each flock, in addition to numbers of the truly gregarious birds,
will contain single representatives of various species of more solitary
disposition, among them often a lone black-and-white, so different in
appearance and habits from any of its associates.

"This warbler arrives and departs early. It has been recorded during the
first week of August in Guatemala, and by the latter part of the month
in Costa Rica and Panamá. In Costa Rica, it appears not to linger beyond
the middle or more rarely the end of March; while for northern Central
America my latest date is April 22.

"Early dates of fall arrival in Central America are: Guatemala--passim
(Griscom), August 3; Sierra de Tecpán, August 23, 1933; Santa María de
Jesús, August 6, 1934; Huehuetenango, August 14, 1934. Honduras--Tela,
August 19, 1930. Costa Rica--San José (Cherrie), August 20; Carrillo
(Carriker), September 1; San Isidro de Coronado, September 8, 1935;
Basin of El General, September 19, 1936; Vara Blanca, September 5, 1937;
Murcia, September 11, 1941. Panamá--Canal Zone (Arbib and Loetscher),
August 24, 1933, and August 29, 1934. Ecuador--Pastaza Valley, below
Baños, October 17, 1939.

"Late dates of spring departure from Central America are: Costa
Rica--Basin of El General, February 23, 1936, March 10, 1939, March 26,
1940, March 3, 1942, March 18, 1943; Vara Blanca, March 13, 1938;
Guayabo (Carriker), March 30; Juan Viñas (Carriker), March 21.
Honduras--Tela, April 22, 1930. Guatemala--Motagua Valley, near Los
Amates, April 17, 1932; Sierra de Tecpán, February 20, 1933."

The bird has a wide winter range, as shown above. Dr. Thomas Barbour
(1943) speaks of it thus in Cuba: "Common in woods and thickets. A few
arrive in August, and by September they are very abundant, especially in
the overgrown jungles about the Ciénaga."

Edward S. Dingle (MS.) has sent to A. C. Bent a remarkable winter record
of a black-and-white warbler seen on Middleburg plantation, Huger, S.
C., on January 13, 1944.


DISTRIBUTION

_Range._--Canada to northern South America.

_Breeding range._--The black-and-white warbler breeds =north= to
southwestern Mackenzie, rarely (Simpson and Providence; has been
collected at Norman); northern Alberta (Chipewyan and McMurray); central
Saskatchewan (Flotten Lake, probably Grand Rapids, and Cumberland
House); southern Manitoba (Duck Mountain, Lake St. Martin, Winnipeg, and
Indian Bay); central Ontario (Kenora, Pagwachuan River mouth, and Lake
Abitibi; has occurred at Piscapecassy Creek on James Bay, and at Moose
Factory); southern Quebec (Lake Tamiskaming, Blue Sea Lake, Quebec,
Mingan, and Mascanin; has occurred at Sandwich Bay, Labrador); and
central Newfoundland (Deer Lake, Nicholsville, Lewisport, and Fogo
Island). =East= to Newfoundland (Fogo Island and White Bear River); Nova
Scotia (Halifax and Yarmouth); the Atlantic coast to northern New Jersey
(Elizabeth and Morristown); eastern Pennsylvania (Berwyn); Maryland
(Baltimore and Cambridge); eastern Virginia (Ashland and Lawrenceville);
North Carolina (Raleigh and Charlotte); South Carolina (Columbia and
Aiken); and central Georgia (Augusta and Milledgeville). =South= to
central Georgia (Milledgeville); south central Alabama (Autaugaville);
north-central Mississippi (Starkville and Legion Lake); northern
Louisiana (Monroe; rarely to southern Louisiana, Bayou Sora); and
northeastern and south-central Texas (Marshall, Dallas, Classen,
Kerrville, and Junction). =West= to central Texas (Junction and Palo
Dura Canyon); central Kansas (Clearwater); central-northern Nebraska
(Valentine); possibly eastern Montana (Glasgow); central Alberta
(Camrose, Glenevis, and Lesser Slave Lake); to southwestern Mackenzie
(Simpson). There is a single record of its occurrence in June at Gautay,
Baja California, 25 miles south of the international border.

_Winter range._--In winter the black-and-white warbler is found =north=
to southern Texas (Cameron County, occasionally Cove, and Texarkana);
central Mississippi, occasionally (Clinton); accidental in winter at
Nashville, Tenn.; southern Alabama (Fairfield); southern Georgia (Lumber
City, occasionally Milledgeville, and Athens); and rarely to
central-eastern South Carolina (Edisto Island and Charleston). =East= to
the coast of South Carolina, occasionally (Charleston); Georgia
(Blackbeard Island); Florida (St. Augustine, New Smyrna, and Miami); the
Bahamas (Abaco, Watling, and Great Abaco Islands); Dominican Republic
(Samaná); Puerto Rico; Virgin Islands and the Lesser Antilles to
Dominica; and eastern Venezuela (Paria Peninsula). =South= to northern
Venezuela (Paria Peninsula, Rancho Grande, and Mérida); west-central
Colombia (Bogotá); and central Ecuador (Pastazo Valley). =West= to
central and western Ecuador (Pastazo Valley and Quito); western Colombia
(Pueblo Rico); western Panamá (Dvala); El Salvador (Mount Cacaguatique);
western Guatemala (Mazatenango); Guerrero (Acapulco and Coyuca); Colima
(Manzanillo); northwestern Pueblo (Metlatayuca); western Nuevo León
(Monterey); and southern Texas (Cameron County). It also occurs casually
in the Cape region of Baja California and in southern California (Dehesa
and Carpenteria). There are also several records in migration from
California and from western Sinaloa.

_Migration._--Late dates of spring departure from the winter home are:
Venezuela--Yacua, Paria Peninsula, March 20. Colombia--Santa Marta
region, March 12. Panamá--Gatún, March 26. Costa Rica--El General, April
9. Honduras--Tola, April 22. Guatemala--Quiriguá, April 17. Veracruz--El
Conejo, May 15. Puerto Rico--Algonobo, April 27. Haiti--Île à Vache, May
6. Cuba--Habana, May 25. Bahamas--Abaco, May 6. Florida--Orlando, May
21. Georgia--Cumberland, May 26. Louisiana--Avery Island, April 27.

Early dates of spring arrival are: South Carolina--Clemson College,
March 20. North Carolina--Weaverville, March 3. Virginia--Lawrenceville,
March 23. District of Columbia--Washington, March 30. New York--Corning,
April 18. Massachusetts--Stockbridge, April 16. Vermont--St. Johnsbury,
April 19. Maine--Lewiston, April 27. Quebec--Montreal, April 26. Nova
Scotia--Wolfville, April 29. Mississippi--Deer Island, March 4.
Louisiana--Schriever, March 8. Arkansas--March 12. Tennessee--Nashville,
March 20. Illinois--Chicago, April 17. Michigan--Ann Arbor, April 6.
Ohio--Toledo, April 7. Ontario--Guelph, April 22. Missouri--Marionville,
April 3. Iowa--Grinnell, April 16. Wisconsin--Milwaukee, April 20.
Minnesota--Lanesboro, April 23. Kansas--Independence, April 1.
Omaha--April 21. North Dakota--April 28. Manitoba--Winnipeg, April 28.
Alberta--Edmonton, May 6; McMurray, May 15. Mackenzie--Simpson, May 22.

Late dates of fall departure are: Alberta--Athabaska Landing, September
11. Manitoba--Aweme, September 22. North Dakota--Argusville, October 2.
Minnesota--Minneapolis, October 10. Iowa--Davenport, October 1.
Missouri--Columbia, October 24. Wisconsin--Madison, October 7.
Illinois--Port Byron, October 15. Ontario--Hamilton, October 3.
Michigan--Detroit, October 15. Ohio--Youngstown, October 15.
Kentucky--Danville, October 14. Tennessee--Athens, October 17.
Arkansas--Winslow, October 17. Louisiana--New Orleans, October 25.
Mississippi--Gulfport, November 19. Quebec--Quebec, September 18. New
Brunswick--St. John, September 19. Nova Scotia--Yarmouth, September 23.
Maine--Portland, October 17. New Hampshire--Ossipee, October 18.
Massachusetts--Cambridge, October 15. New York--New York, October 6.
Pennsylvania--Atglen, October 29. District of Columbia--Washington,
October 18. Virginia--Charlottesville, October 18. North
Carolina--Raleigh, October 29. South Carolina--Charleston, November 15.
Georgia--Savannah, October 29.

Early dates of fall arrival are: South Carolina--Charleston, July 19.
Florida--Pensacola, July 12. Cuba--Artemisa, Pinar del Río, August 1.
Dominican Republic--Ciudad Trujillo, September 27. Puerto
Rico--Mayagüez, October 9. Louisiana--New Orleans, July 21.
Mississippi--Bay St. Louis, July 4. Michoacán--Tancitaro, August 7.
Guatemala--Huehuetenango, August 14. Honduras--Cantarranas, August 7.
Costa Rica--San José, August 20. Panamá--Tapia, Canal Zone, August 24.
Colombia--Bonda, Santa Marta region, August 21. Ecuador--Pastaza Valley,
October 17. Venezuela--Estado Carabobo Las Trincheras, October 9.

_Banding._--A single banding recovery is of considerable interest: A
black-and-white banded at Manchester, N. H., on August 31, 1944, was
found on March 17, 1945, at Friendship P. O., Westmoreland, Jamaica.

_Casual records._--This warbler is casual in migration or winters in
Bermuda, having been recorded in six different years from October to
May.

At Tingwall, Shetland Islands, north of Scotland one was picked up on
November 28, 1936. This is almost as far north as the northernmost
record of occurrence in North America and later than it is normally
found in the United States.

A specimen was collected near Pullman, Wash., on August 15, 1948, the
first record for the State.

_Egg dates._--Massachusetts: 31 records, May 18 to June 14; 17 records,
May 25 to June 3, indicating the height of the season.

New Jersey: 7 records, May 18 to June 8.

Tennessee: 3 records, May 1 to 17.

North Carolina: 6 records, April 20 to 28.

West Virginia: 7 records, May 6 to 29 (Harris).




PROTONOTARIA CITREA (Boddaert)

PROTHONOTARY WARBLER

PLATES 4-6


HABITS

I do not like the above name for the golden swamp warbler. The
scientific name _Protonotaria_, and evidently the common name, were
apparently both derived from the Latin _protonotarius_, meaning first
notary or scribe. I sympathize with Bagg and Eliot (1937), who
exclaimed:

What a name to saddle on the Golden Swamp-bird! Wrongly compounded in
the first place, wrongly spelled, wrongly pronounced! We understand that
Protonotarius is the title of papal officials whose robes are bright
yellow, but why say "First Notary" in mixed Greek and Latin, instead of
Primonotarius? Proto is Greek for first, as in prototype. Why and when
did it come to be misspelled Protho? Both Wilson and Audubon wrote
Protonotary Warbler, a name seemingly first given to the bird by
Louisiana Creoles. Both etymology and sense call for stress on the third
syllable, yet one most often hears the stress laid on the second. Here,
certainly, is a bothersome name fit only to be eschewed!

The scientific name cannot be changed under the rules of nomenclature,
but a change in the common name would seem desirable. However, the name
does not make the bird or detract from its charm and beauty. It will
still continue to thrill with delight the wanderer in its swampy haunts.

The center of abundance of the prothonotary warbler as a breeding bird
in this country is in the valleys of the Mississippi River and its
tributaries, notably the Ohio, the Wabash, and the Illinois Rivers. Its
summer range extends eastward into Indiana and Ohio, northward into
southern Ontario, Wisconsin, Michigan, and Minnesota, and westward into
Iowa, Nebraska, Kansas, Oklahoma, and eastern Texas--wherever it can
find suitable breeding grounds.

It also breeds in the Atlantic Coast States from Virginia to Florida.

It is essentially a bird of the damp and swampy river bottoms and
low-lying woods, which are flooded at times and in which woodland pools
have been left by the receding water. Perhaps this warbler abounds more
than anywhere else in the valley of the lower Wabash, where William
Brewster (1878) found it to be--

one of the most abundant and characteristic species. Along the shores of
the rivers and creeks generally, wherever the black willow (_Salix
niger_) grew, a few pairs were sure to be found. Among the button-bushes
(_Cephalanthus occidentalis_) that fringed the margin of the peculiar
long narrow ponds scattered at frequent intervals over the heavily
timbered bottoms of the Wabash and White Rivers, they also occurred more
or less numerously. Potoka Creek, a winding, sluggish stream, thickly
fringed with willows, was also a favorite resort; but the grand
rendezvous of the species seemed to be about the shores of certain
secluded ponds lying in what is known as the Little Cypress Swamp. Here
they congregated in astonishing numbers, and early in May were breeding
almost in colonies. In the region above indicated two things were found
to be essential to their presence, namely, an abundance of willows and
the immediate proximity of water. * * * So marked was this preference,
that the song of the male heard from the woods indicated to us as surely
the proximity of some river, pond, or flooded swamp, as did the croaking
of frogs or the peep of the Hylas.

Dr. Chapman (1907) writes of this bird in its haunts:

The charm of its haunts and the beauty of its plumage combine to render
the Prothonotary Warbler among the most attractive members of the
family. I clearly recall my own first meeting with it in the Suwanee
River region of Florida. Quietly paddling my canoe along one of the many
enchanting, and, I was then quite willing to believe, enchanted streams
which flowed through the forests into the main river, this glowing bit
of bird-life gleamed like a torch in the night. No neck-straining
examination with opera-glass pointed to the tree-tops, was required to
determine his identity, as, flitting from bush to bush along the river's
bank, his golden plumes were displayed as though for my special benefit.

Dr. Lawrence H. Walkinshaw (1938) says that the golden swamp warbler
"nests rather abundantly along southwestern-Michigan rivers. * * *
Winding streams, bordered densely with oak, maple, ash, and elm, shallow
ponds with groups of protruding willows and flooded, heavily shaded
bottom-lands are favorite nesting habitats for the Prothonotary Warbler
(_Protonotaria citrea_). Such habitats occur along the banks of the
Kalamazoo River and its tributary the Battle Creek River in Calhoun
County, Michigan."

_Territory._--The males arrive on the breeding grounds a few days or a
week before the females come and immediately try to establish their
territories, select the nesting sites, and even build nests. Dr.
Walkinshaw (1941) writes:

The Prothonotary Warbler is a very strongly territorial species. When a
male takes possession of a certain area he continually drives off all
opponents if he is able. At certain areas in Michigan I have watched
these birds battle intermittently for two or three days, usually for the
same bird house, one male finally taking possession. In addition I have
observed them to drive off House Wrens (_Troglodytes aedon_),
Black-capped Chickadees (_Penthestes atricapillus_) and Yellow Warblers
(_Dendroica aestiva_). * * * The male Prothonotary Warbler selects the
territory, selecting the nesting site before he becomes mated for the
first nest, but thereafter both birds inspect the new nest sites.

On observations made near Knoxville, Tenn., Henry Meyer and Ruth Reed
Nevius (1943) found that--

three males established territories. Male I arrived April 14. By the
next day he was singing on an area 550 feet long and for the most part
not more than 200 feet wide. It included three kinds of habitats: (a) a
grassy terrace on which several nesting boxes were located, (b) river
banks densely covered with small trees and bushes, and (c) a small open
orchard which constituted the connecting link between the terrace and
the river bank. Male II arrived on April 18 and occupied a narrow
territory along a brook confined by wooded slopes and which contained
two lotus ponds. The area was about 400 feet long and 100 feet wide. A
nesting box was on a stake above one of the ponds. Male III appeared May
5 in the terraced area being claimed by Male I. During the day, the 2
males sang energetically and flew often only a few inches apart. Male I
maintained his territory and Male III disappeared.

There were a number of nesting boxes on the area that the males
investigated, carrying nesting material into some of them while they
were waiting for the females to arrive. The mate of the first male came
on April 20, and--

on this day this pair communicated by their full call-note. Twice the
male was seen pursuing the female rapidly in a small semi-circle and
pausing, called a soft, full note which was later heard only when the
two sexes were together.

The mate of Male II came April 22, four days after the latter's arrival.

Combat with other species found within the territories of these birds
was observed. Combat with the Bluebird was most frequent but one or more
indications of opposition was noticed with the Flicker, Downy
Woodpecker, Acadian Flycatcher, Tufted Titmouse, Robin, and Cardinal.

The males sing persistently and energetically from the time that they
arrive on their territories, hoping to attract their mates, but they are
not always successful, especially in regions where the species is rare
or not very common, and their nest-building brings no occupant. Edward
von S. Dingle writes to me that, at Summerton, S. C., a male
prothonotary warbler built a nest in a low stub, but no female was ever
seen. He sang frequently and remained in the vicinity for several weeks.
And Frederic H. Kennard, in far-away Massachusetts, mentions in his
notes that he saw one and watched it for several days, June 16-20, 1890.
"He sang loudly and clearly and sweetly, and seemed to like a particular
place by the side of the river, for when I returned later in the day, he
was still there, on the other side of the river." On June 19, he watched
him for half an hour. He was always in the same locality. On a later
search, no nest or no mate could be found.

_Courtship._--Brewster (1878a) gives the following full account of this
performance:

Mating began almost immediately after the arrival of the females, and
the "old, old story" was told in many a willow thicket by the little
golden-breasted lovers. The scene enacted upon such occasions was not
strikingly different from that usual among the smaller birds; retiring
and somewhat indifferent coyness on the part of the female; violent
protestations and demonstrations from the male, who swelled his plumage,
spread his wings and tail, and fairly danced around the object of his
affections. Sometimes at this juncture another male appeared, and then a
fierce conflict was sure to ensue. The combatants would struggle
together most furiously until the weaker was forced to give way and take
to flight. On several occasions I have seen two males, after fighting
among the branches for a long time, clinch and come fluttering together
to the water beneath, where for several minutes the contest continued
upon the surface until both were fairly drenched. The males rarely meet
in the mating season without fighting, even though no female may be
near. Sometimes one of them turns tail at the outset; and the other at
once giving chase, the pursuer and pursued, separated by a few inches
only, go darting through the woods, winding, doubling, now careering
away up among the tree-tops, now down over the water, sweeping close to
the surface until the eye becomes weary with following their mad flight.
During all this time the female usually busies herself with feeding,
apparently entirely unconcerned as to the issue. Upon the return of
the conqueror her indifference, real or assumed, vanishes, he receives
a warm welcome, and matters are soon arranged between them.

_Nesting._--The prothonotary warbler and Lucy's warbler are the only two
American warblers that habitually build their nests in cavities, usually
well concealed. The normal, and probably the original, primitive,
nestling sites are in natural cavities in trees and most nests are still
to be found in such situations today. The prothonotary is not at all
particular as to the species of trees, nests having been found in many
kinds of trees, although perhaps a slight preference is shown for dead
willow stumps. Nor is it particular as to the size or condition of the
cavity, or its location, though quite often choosing one over water or
near it. The height above the ground or water varies from 3 feet or less
to as much as 32 feet but there are more nests below 5 feet than there
are above 10, the height of the majority being between 5 and 10 feet.
The size and shape of the cavity are of little concern; if the cavity is
too deep, the industrious little birds fill it with nesting material up
to within a few inches of the top; sometimes a very shallow cavity is
used, so that the bird can be plainly seen from a distance as it sits on
the nest. The old deserted holes of woodpeckers or chickadees are
favorite nesting sites; the entrances to these have often been enlarged
by other agencies, or are badly weathered. In very rotten stumps, the
warblers have been known to excavate partially or to enlarge a cavity.

The nests built by the males in early spring, referred to above, are
probably rarely used as brood nests and might be classed as dummy nests.
The family nest is built almost entirely by the female, with
encouragement and a little help from her mate, who accompanies her to
and from the nest and in the search for material; much of the soft,
green moss used extensively in the nest is often obtainable from fallen
logs and stumps in the vicinity.

Brewster (1878a) mentions a nest taken from a deep cavity that "when
removed presents the appearance of a compact mass of moss five or six
inches in height by three or four in diameter. When the cavity is
shallow, it is often only scantily lined with moss and a few fine roots.
The deeper nests are of course the more elaborate ones. One of the
finest specimens before me is composed of moss, dry leaves, and cypress
twigs. The cavity for the eggs is a neatly rounded, cup-shaped hollow,
two inches in diameter by one and a half in depth, smoothly lined with
fine roots and a few wing-feathers of some small bird."

In Dr. Walkinshaw's (1938) Michigan nests, "moss constituted the bulk of
the nesting material in nearly all cases, completely filling the nest
space whether it was large or small. On top of this the nest proper was
shaped and a rough lining of coarse grape-bark, dead leaves, black
rootlets procured from the river-banks, and poison-ivy tendrils was
added. Above this a lining of much finer rootlets, leaf-stems, and very
fine grasses was used."

In addition to the materials listed above Meyer and Nevius (1943)
mention hackberry leaves, hairs, pine needles, horsehair, and cedar bark
in their Tennessee nests. They say that from 6 to 10 days were required
for nest construction, and that from 3 to 5 days more elapsed before the
first eggs were laid. Their four nests were all in bird-boxes; one was
in an orchard over plowed ground, one over a lotus pond in a wooded
ravine, and two were over lily pools near buildings.

Dr. Walkinshaw (1938) publishes a map showing the location of 21 nesting
boxes along the winding banks of the Battle Creek River, in Calhoun
County, Mich., and writes: "Of the 28 nests found during 1937, 19 were
in bird-houses over running water, 6 were in stubs over water (2 of
which were over running water), and the other 3 were in natural holes
back from the river bank. Of 44 nests found from 1930 through 1937,
excluding the 21 in bird-houses, six were over running water in old
woodpecker holes, one in a bridge-support in a slight depression, and
nine in natural holes over standing water. Seven were in old woodpecker
holes from two to a hundred and sixty feet back from the river-bank."

Many and varied are the odd nesting sites occupied by prothonotary
warblers. Dr. Thomas S. Roberts (1936) writes:

The vagaries of this bird in choosing artificial nesting-places are
shown by the positions of the following nests. On the La Crosse railroad
bridge: in a cigar-box nailed on the engine-house on top of the draw; on
one of the piers; in a metal ventilator-cap four inches in diameter,
that had fallen and lodged just at the point where the draw banged
against the pier, and close under the tracks; in a shallow cavity in a
piece of slab-wood nailed to a trestle-support close under the road-bed
of the railroad; these all far out in the middle of the Mississippi
River. Still others are: in a Bluebird box on a low post by a
switching-house and busy railroad platform; in a cleft in a pile in the
river; in a tin cup in a barn, to reach which the birds entered through
a broken pane of glass; in a pasteboard box on a shelf in a little
summer-house; in an upright glass fruit jar in a house-boat; and other
similar situations. In most cases the birds had to carry the
nesting-material long distances, especially to the places on the bridge.

John W. Moyer (1933) relates an interesting story that was told to him
by people living in a farm house along the Kankakee River. A pair of
these warblers built their nests and raised their broods for three
consecutive seasons in the pocket of an old hunting-coat, hung in a
garage; each year the man cleaned out the nest and used the coat in the
fall, and the next spring the birds used it again. M. G. Vaiden tells me
of a similar case.

Nests have been found in buildings, on beams and other supports. Louis
W. Campbell (1930) reports two on shelves in sheds, one in a small paper
sack partly filled with staples and another in a coffee can similarly
filled. Nests in cans in various situations have been found a number of
times, and others have been reported in a tin pail hung under a porch,
in a mail box, in a box on a moving ferry boat, in a Chinese lantern on
a pavilion, and in an old hornets' nest.

Dr. Walkinshaw writes to me: "At Reelfoot Lake, Tenn., during July,
1940, I found 8 nests of the prothonotary warbler, all built a few feet
above the water in small natural holes in cypress knees. Evidently these
are regular late-summer nesting sites." The knees were farther under
water earlier in the season. Most of his 76 Michigan nests were over
water, or less than 100 feet from it; but 10 were 300 or more feet away
from it and 2 were over 400 feet away. M. G. Vaiden tells me of a pair
that nested in the tool box of a log-loading machine that was in daily
operation, hauling logs.

_Eggs._--From 3 to 8 eggs have been found in nests of the prothonotary
warbler, from 4 to 6 seem to be the commonest numbers, 7 is a fairly
common number, and at least 3 sets of 8 have been reported; in the
J. P. Norris series of 70 sets are 34 sets of 6, 15 sets of 7, and
2 sets of 8.

The eggs vary in shape from ovate to short ovate, and they are more or
less glossy. The eggs are undoubtedly the most striking of the warblers'
eggs, with their rich creamy, or rose-tinted cream, ground color, boldly
and liberally spotted and blotched with "burnt umber," "bay," "chestnut
brown," and "auburn," intermingled with spots and undertones of "light
Payne's gray," "Rood's lavender," "violet-gray," and "purplish gray."
There is quite a variation in the amount of markings, which are
generally more or less evenly scattered over the entire egg; some are
sparingly spotted and blotched, while others are so profusely marked as
almost to obscure the ground color (Harris).

J. P. Norris (1890b), in his description of his 70 sets, describes 2
eggs in each of 2 sets as "unmarked, save for four or five indistinct
specks of cinnamon." These were in sets of 6 eggs each. Pure white,
unmarked eggs were once taken by R. M. Barnes (1889). Dr. Walkinshaw
(1938) gives the measurements of 78 eggs as averaging 18.47 by 14.55
millimeters; the eggs showing the four extremes measured =20= by 15, 19
by =16=, and =17= by =13= millimeters.

_Incubation._--The eggs are laid, usually one each day, very early in
the morning; Dr. Walkinshaw (1941) says between 5:00 and 7:00 a. m. in
Michigan; Meyer and Nevius (1943), in Tennessee, saw the female enter
the nest to lay as early as 5:00 a. m. on May 2, and as early as 4:44 on
May 23, remaining in the nest from 28 to 36 minutes on different
occasions. The period of incubation is recorded as 12, 13-1/2, and 14
days by different observers; about 13 seems to be the average, according
to Dr. Walkinshaw (MS.), probably depending on conditions and the method
of reckoning. Incubation seems to be performed entirely by the female,
but the male feeds her to some extent while she is on the nest.
Incubation starts the day before the last egg is laid.

_Young._--Meyer and Nevius (1943) write:

The adults shared feeding duties, and both removed fecal sacks. During
the first three days the female steadily brooded the young. One female,
observed from 4:55 to 8:10 a. m., when the young were one day old, spent
a total of 70 minutes off and 155 minutes on the nest. Trips from the
nest lasted an average of 8.6 minutes, while periods on the nest
averaged 19.4 minutes. * * * At one nest when the young were eight days
old, activities were noted during the eight and one-half hours from 8:30
a. m. to 5:30 p. m. The young were fed an average of 16 times an hour.
* * * The adults were seen carrying spiders and insects, small green
caterpillars frequently being used. Mr. H. P. Ijams saw a male offer a
10-day old nestling a may-fly. An incubator-hatched bird accepted
egg-yolk, ants, ant larvae, crickets, earthworms, and spiders.

They say of the development of the young: "The young on the day of
hatching had orange-red skin. The mouth lining was red. Down was
distributed over the frontal and occipital areas of the capital tract,
spinal tract, femoral, altar, and humeral tracts. Feather sheaths of
the alar tracts penetrated the skin the first day after hatching. On the
second day after hatching the eye-slits began to open. Feather sheaths
of the humeral, femoral, and crural tracts emerged on the third day;
those on the dorsal and ventral tracts emerged on the fourth day, and
those of the capital and caudal tracts on the fifth day. On the fifth
day the sheaths began breaking." During the next five days the young
developed rapidly and became more and more active, and on the tenth day
began to leave the nest.

Young observed by Dr. Walkinshaw (1914) at Reelfoot Lake "averaged 11
days of age when leaving the nest in 1939, while 21 young in Michigan
during 1939 and 1940 remained in the nest for a period of 10-3/4 days."
Of the comparative nesting success in the two localities, he says:

In Michigan from 1930 through 1940, 121 nests of the Prothonotary
Warbler were observed. Only 28, or 23.14 per cent, were successful. Out
of 413 eggs, 159 (38.47 per cent) hatched and 100 young were fledged
(.87 per total nest; 3.78 per successful nest). The fledgling success
was 25.66 per cent of eggs laid. More failures in Michigan resulted in
more nestings by individual birds.

In Tennessee during 1939, 30 nests were observed until terminated or
successful; 19 were successful (63.33 per cent) while out of 139 eggs,
78 hatched and all the young lived to leave the nest or 56.11 per cent
fledging success of eggs laid; 2.6 young were fledged per total nest;
4.1 per successful nest.

He also notes that in Michigan the species is typically single-brooded
if the first nesting is successful, but that in Tennessee it is
typically double-brooded.

_Plumages._--According to Dr. Dwight (1900) the natal down, located as
indicated above, is brownish mouse-gray. Ridgway (1902) gives rather the
best description of the juvenal plumage as follows: "Pileum, hindneck,
back, and scapulars dull olive-greenish; wing-coverts, tertials, rump,
and upper tail-coverts slate-gray, tinged with olive, the middle and
greater wing-coverts narrowly tipped with light olive-greenish,
producing two very indistinct bands; secondaries, primaries, and
rectrices as in adults; sides of head pale yellowish olive; chin,
throat, and chest dull light grayish olive, darkest on chest rest of
under parts dull white, passing on sides and flanks into olive-grayish."

In very young birds, according to Dr. Dwight, there is a variable amount
of brownish wash on the back, which fades out to gray. And Dr. Chapman
(1907) says that the white on the inner webs of the tail feathers is
more restricted than in adults and more or less mottled with blackish.
This first plumage is followed in June and July by a partial postjuvenal
molt involving all the contour plumage and the wing coverts but not the
rest of the wings or the tail. The young bird now becomes a golden swamp
warbler, the young being nearly like the adults, the females being
considerably duller in color than the males and having less white in the
tail.

The crown and hind neck in both sexes, both old and young, are washed
with dusky or olive in the fall. Spring plumages are produced by very
slight wear without molt. There is one complete, annual molt in late
summer.

_Food._--Very little seems to have been recorded on the food of the
prothonotary warbler. It is evidently highly insectivorous, obtaining
most of its food from the trunks and branches of trees and shrubs and
from fallen logs. Brewster (1878) says: "This Warbler usually seeks its
food low down among thickets, moss-grown logs, or floating débris, and
always about water. Sometimes it ascends tree-trunks for a little way
like the Black-and-White Creeper, winding about with the same peculiar
motion."

Dr. Roberts (1936) lists "ants, and other insects and their larvae," as
its food. Some of the food of the young is mentioned above, most of
which is doubtless included in the food of the adults. Spiders, beetles,
may-flies, and other insects should be included, as well as many
caterpillars and the larvae of water insects. Audubon (1841) says: "It
often perches upon the rank grasses and water plants, in quest of minute
molluscous animals which creep upon them, and which, together with small
land snails, I have found in its stomach."

_Behavior._--Brewster (1878a) observes:

When seen among the upper branches, where it often goes to plume its
feathers and sing in the warm sunshine, it almost invariably sits nearly
motionless. Its flight is much like that of the Water-Thrush (either
species), and is remarkably swift, firm, and decided. When crossing a
broad stream it is slightly undulating, though always direct. * * *

In general activity and restlessness few birds equal the species under
consideration. Not a nook or corner of his domain but is repeatedly
visited through the day. Now he sings a few times from the top of some
tall willow that leans out over the stream, sitting motionless among the
yellowish foliage, fully aware, perhaps, of the protection afforded by
its harmonizing tints. The next moment he descends to the cool shades
beneath, where dark, coffee-colored water, the overflow of the pond or
river, stretches back among the trees. Here he loves to hop about on
floating drift-wood, wet by the lapping of pulsating wavelets; now
following up some long, inclining, half-submerged log, peeping into
every crevice and occasionally dragging forth from its concealment a
spider or small beetle, turning alternately his bright yellow breast and
olive back towards the light; now jetting his beautiful tail or
quivering his wings tremulously, he darts off into some thicket in
response to a call from his mate; or, flying to a neighboring
tree-trunk, clings for a moment against the mossy bole to pipe his
little strain or look up the exact whereabouts of some suspected insect
prize.

_Voice._--The same gifted writer and careful observer (Brewster, 1878)
gives the following good account of the distinctive song of this
warbler:

The usual song of the Prothonotary Warbler sounds at a distance like the
call of the Solitary Sandpiper, with a syllable or two added,--a simple
_peet, tweet, tweet, tweet_, given on the same key throughout. Often
when the notes came from the farther shore of a river or pond we were
completely deceived. On more than one occasion, when a good opportunity
for comparison was offered by the actual presence of both birds at the
same time, we found that at the distance of several hundred yards their
notes were absolutely undistinguishable; nearer at hand, however, the
resemblance is lost, and a ringing, penetrating quality becomes apparent
in the Warbler's song. It now sounds like _peet, tsweet, tsweet,
tsweet_, or sometimes _tweet, tr-sweet, tr-sweet, tr-sweet_. When the
bird sings within a few yards the sound is almost startling in its
intensity, and the listener feels inclined to stop his ears. The male is
a fitful singer, and is quite as apt to be heard in the hot noontide or
on cloudy days, when other birds are silent, as during the cool morning
and evening hours. The ordinary note of alarm or distress is a sharp
one, so nearly like that of the Large-billed Water Thrush (_Siurus
motacilla_) that the slight difference can only be detected by a
critical ear. When the sexes meet a soft _tchip_ of recognition common
to nearly all the Warblers is used. In addition to the song above
described the male has a different and far sweeter one, which is
reserved for select occasions,--an outpouring of the bird's most tender
feelings, intended for the ears of his mate alone, like the rare evening
warble of the Oven-Bird (_Siurus auricapillus_). It is apparently
uttered only while on the wing.

Although so low and feeble as to be inaudible many rods away, it is very
sweet, resembling somewhat the song of the Canary, given in an
undertone, with trills or "water-notes" interspersed. The flight during
its delivery is very different from that at all other times. The bird
progresses slowly, with a trembling, fluttering motion, its head raised
and tail expanded. This song was heard most frequently after incubation
had begun.

Dr. Roberts (1936) refers to this flight song, as delivered "after the
manner of the Maryland Yellow-throat, * * * consisting first of the
usual rapid monotone of five or six notes and ending with a pleasing,
varied warble, full and strong in some of its notes and far sweeter than
the usual utterance."

Dr. Walkinshaw (1938) says of the usual song: "Uttered at the rate of
five or six times per minute, the song lasts slightly over one second.
It is given all day long from the time of arrival until the young have
left the nest and has been heard as late as the 16th of August (1931).
The frequency is much greater during the early nesting season and during
the earlier hours. During midday on warmer days the number of times per
hour seems much less. Later, from four until near sundown, it again
increases. During late nesting, when the young are about to leave the
nest, the rate again decreases, but it is heard several days after the
young leave the nest." Aretas A. Saunders tells me that the songs are
pitched at C´´´´ or B´´´, and the call note, _tseek,_ at A´´´.

_Field marks._--The golden swamp warbler could hardly be mistaken for
anything else. The rich, brilliant yellow of the head and breast,
sometimes almost orange on the head, only slightly paler in the female,
the absence of wing bars, and the large amount of white in the tail will
distinguish it.

_Enemies._--Dr. Walkinshaw (1941) says that the house wren is a serious
competitor with the prothonotary warbler in Michigan, contending with it
for nesting sites in the bird-boxes.

The cowbird is a persistent enemy of this warbler in spite of its
hole-nesting habits; perhaps if the warbler nested in deeper holes it
might find some relief from this pest. Among 70 sets of eggs of this
warbler in the J. P. Norris collection, 18 contain cowbirds' eggs. Dr.
Friedmann (1929) found no less than 36 records of such parasitism in the
literature, and says: "As many as four eggs of the Cowbird have been
found in a single nest together with four of the Warbler's. There are
several cases on record of doubled-storied nests of this bird, with a
Cowbird's egg buried in the lower story. Such cases are, however, not
common, and usually the Warbler seems to make no attempt to get rid of
the strange eggs." E. M. S. Dale wrote to me of a nest, found near
Toronto in 1933, that contained seven eggs of the cowbird and none at
all of the warbler!

Snakes sometimes destroy the eggs or young.

_Fall._--Dr. Walkinshaw (1938) says that "the majority of the
Prothonotaries leave our rivers [Michigan] by the second or third of
July. One may canoe some years a good many miles during the latter part
of July or the early part of August without finding a single
Prothonotary, whereas in other years many groups can be found. The
majority evidently are early migrants. Very few remain until late August
or early September, the latest date being September 9, 1934, at Battle
Creek."

The 1931 A. O. U. Check-List states that this warbler apparently crosses
the Gulf of Mexico in migration "and is not found in Mexico north of
Campeche," but probably some migration is along the coast of Texas and
Mexico, as suggested by George G. Williams (1945).

Dr. Chapman (1907) says: "The route of the Prothonotary Warbler in its
fall migration is interesting; the breeding birds of the Middle Atlantic
States apparently pass southwest to northwestern Florida and then take a
seven-hundred-mile flight directly across the Gulf of Mexico to southern
Yucatan, instead of crossing to Cuba and thence to Yucatan."

Alexander F. Skutch writes to me: "Unrecorded from Guatemala, the
prothonotary warbler is a rare bird of passage and very rare winter
resident in the more southerly portions of Central America. When
Carriker published his list of Costa Rican birds in 1910, he had a few
records from the highlands--apparently of migrating birds--and from the
Pacific lowlands, but none from Caribbean lowlands. But on March 4,
1934, I found it not uncommon at Puerto Limón, where I saw one among the
royal palms in Vargas Park, and several among the shrubbery about the
outlying cottages, all within a hundred yards of the Caribbean Sea. It
has been recorded a number of times from the Canal Zone, but it is not
common there. It is almost always seen in the vicinity of water."

_Winter._--Apparently the main winter range is in Colombia and perhaps
Venezuela. Referring to Magdalena, Colombia, P. J. Darlington, Jr.
(1931), writes: "The Prothonotary Warbler swarms during the winter in
the mangroves at Sevillano and in the fresh swamps at Cienaga. It was
seen also in bushes on the sea beach at Donjaca September 15, and along
the Rio Frio River in the edge of the foothills, where it was especially
common in February. The birds usually occur near water, but numbers were
noted again and again in yellow-flowering, acacia-like trees on the
border of stump land and dry forest, far from water."


DISTRIBUTION

_Range._--Eastern United States to northwestern South America.

_Breeding range._--The prothonotary warbler breeds =north= to
southeastern Minnesota (Cambridge, Lake Pekin, and La Crescent); central
Wisconsin (New London and Shiocton); southern Michigan (Hesperia,
Lansing, and Ann Arbor); northern Ohio (Toledo and Cleveland); extreme
southern Ontario (Rondeau); western New York (Buffalo and Oak Orchard);
northern West Virginia (Parkersburg); central Maryland (Seneca and
Bowie); and southern Delaware (Gumboro). =East= to southern Delaware
(Gumboro); eastern Virginia (Dyke, near Alexandria, and Dismal Swamp);
and the Atlantic coast to central Florida (Lake Gentry and Padgett
Creek). =South= to central Florida (Padgett Creek and possibly
Puntarossa); the Gulf coast to southeastern Texas (Cove, Houston, and
Bloomington). =West= to central Texas (Bloomington, Fort Worth, and
Gainesville); central Oklahoma (Norman and Oklahoma City); eastern
Kansas (Emporia and Manhattan); northwestern Iowa (Lake Okoboji); and
southeastern Minnesota (Rochester, Red Wing, and Cambridge).

The prothonotary warbler has been recorded as casual or accidental
=west= to southeastern Nebraska (Powell and Lincoln); southeastern South
Dakota (Yankton and Sioux Falls); and central Minnesota (Brainerd).
=North= to southern Ontario (London and Hamilton); central New York
(Ithaca); Massachusetts (Northampton, Amherst, and Concord); New
Hampshire (Concord); and Maine (Matinicus Island and Calais).

_Winter range._--The winter home of the prothonotary warbler is in
Central America and northwestern South America where it has been found
=north= to northwestern Costa Rica (Bolson); Nicaragua (Escondido
River). =East= to northwestern Venezuela (Mérida and Encontrados); and
western Colombia (San José de Cucuta and Villavieja). =South= to
southwestern Colombia (Villavieja); and northwestern Ecuador
(Esmeraldas). =West= to northwestern Ecuador (Esmeraldas); western
Colombia (Antioquia); western Panamá (Paracote and David); and Costa
Rica (Puntarenas and Bolson). It has been reported to occur in winter in
Campeche and on Cozumel Island, Mexico, and casually or accidentally in
Cuba (Habana), Jamaica, and St. Croix, Virgin Islands.

_Migration._--The probable route of the prothonotary warbler between its
summer and winter homes is across the Gulf of Mexico, from the Yucatan
peninsula where it occurs in both spring and fall migration. The casual
or accidental occurrences of this warbler in Cuba (Habana); Jamaica; and
St. Croix, Virgin Islands, are in migration.

Late dates of spring departure are: Colombia--Villavieja, February 5.
Panamá; Canal Zone--Barro Colorado, March 10. Nicaragua--Edén, March 23.
Quintana Roo--Cozumel, April 6. Cuba--Habana, April 4.

Early dates of spring arrival are: Yucatán--Mérida, March 28.
Jamaica--Black River, February 28. Cuba--Habana, March 31.
Florida--Pensacola, March 18. Alabama--Booth, April 4.
Georgia--Fitzgerald, March 21. South Carolina--Yemassee, March 27.
North Carolina--Greenville, April 6. Virginia--Suffolk, April 10.
Mississippi--Gulfport, March 18. Louisiana--Morgan City, March 10.
Texas--Cove, March 28. Arkansas--Huttig, March 31. Missouri--St. Louis,
April 17. Kentucky--Bowling Green, April 5. Illinois--Murphysboro, April
17. Ohio--Berlin Center, April 18. Michigan--Grand Rapids, May 3.
Iowa--Iowa City, April 26. Wisconsin--Madison, May 2. Minnesota--Red
Wing, May 7. Oklahoma--Tulsa, April 2. Kansas--Manhattan, April 26.
Nebraska--Blue Springs, April 30.

Late dates of fall departure are: Nebraska--Watson, September 1.
Oklahoma--Oklahoma City, September 14. Texas--Kemah, September 11.
Wisconsin--Racine, September 22. Iowa--Sioux City, August 31.
Michigan--Three Rivers, September 13. Ohio--Columbus, October 5.
Illinois--Oak Park, October 17. Kentucky--Lexington, October 6.
Tennessee--Elizabethton, October 19. Louisiana--Monroe, October 8.
Mississippi--Deer Island, September 27. North Carolina--Raleigh, August
26. South Carolina--Charleston, September 17. Georgia--Atlanta, October
8. Yucatán--Chichén-Itzá, October 18.

Early dates of fall arrival are: Florida--Fort Myers, August 8.
Yucatán--Chichén-Itzá, October 7. Honduras--Tela, September 8.
Nicaragua--Río Escondido, September 2. Costa Rica--Bonilla, August 28.
Panamá--Obaldia, September 15. Colombia--Gaira, September 11.

_Banding records._--Banding provides a hint as to the life-span of the
prothonotary warbler. One banded as an immature on June 16, 1940, in
Convis township, Calhoun County, Mich., was color banded when it
returned to the same place in 1942. Subsequently it was identified by
the colored band on May 14, 1944, and May 10, 1945.

_Casual records._--The prothonotary warbler was reported at Nassau,
Bahamas, on August 29, 1898. It has been twice reported at Bermuda: one
shot from a flock in the fall of 1874, and another specimen collected in
November 1903. A single bird was observed at Mammoth Hot Springs,
Yellowstone Park, Wyo., on September 10, 1931. There are two records for
Arizona. On May 1, 1884, a specimen was taken near Tucson at an altitude
of 2,300 feet, the highest record of the species in the United States.
Another specimen was taken September 8, 1924, at Cave Creek, 4 miles
northeast of Paradise in the Chiricahua Mountains.

_Egg dates._ Florida: 8 records, April 18 to May 9; 5 records, April 28
to 30.

Illinois: 79 records, May 6 to June 21; 46 records, May 20 to June 4,
indicating the height of the season.

Iowa: 56 records, May 15 to June 26; 36 records, May 27 to June 6
(Harris).




LYMNOTHLYPIS SWAINSONII (Audubon)

SWAINSON'S WARBLER


CONTRIBUTED BY EDWARD VON SIEBOLD DINGLE

PLATES 7-9


HABITS

"The history of our knowledge of Swainson's Warbler," write Brooks and
Legg (1942), "is a curious one, falling into definite periods." This
bird was discovered in the spring of 1832 by the Rev. John Bachman "near
the banks of the Edisto River, South Carolina." His discovery of the
bird is described as follows: "I was first attracted by the novelty of
its notes, four or five in number, repeated at intervals of five or six
minutes apart. These notes were loud, clear, and more like a whistle
than a song. They resembled the sounds of some extraordinary
ventriloquist in such a degree, that I supposed the bird much farther
from me than it really was; for after some trouble caused by these
fictitious notes, I perceived it near to me and soon shot it" (Audubon,
1841). Dr. Bachman took five specimens; then, up to the spring of 1884,
Swainson's warbler remained almost a lost species, for according to
Brewster (1885a) there is no record of more than eight or nine birds
being collected. Wayne, through collections and field work near
Charleston, opened a productive 25-year period in the history of
_swainsonii_, in which many valuable contributions were made by various
observers. From 1910 to 1930 the name _swainsonii_ was practically
absent from the pages of current ornithological literature.

Brewster (1885a) has given us the best description of the bird's haunts
in the low country:

The particular kind of swamp to which he is most partial is known in
local parlance as a "pine-land gall." It is usually a depression in the
otherwise level surface, down which winds a brook, in places flowing
swiftly between well-defined banks, in others divided into several
sluggish channels or spreading about in stagnant pools, margined by a
dense growth of cane, and covered with lily leaves or other aquatic
vegetation. Its course through the open pine-lands is sharply marked by
a belt of hardwood trees nourished to grand proportions by the rich soil
and abundant moisture. Beneath, crumbling logs cumber the ground, while
an undergrowth of dogwood (_Cornus florida_), sassafras, viburnum, etc.,
is interlaced and made well-nigh impenetrable by a network of grapevines
and greenbriar. These belts--river bottoms they are in miniature--rarely
exceed a few rods in width; they may extend miles in a nearly straight
line.

The writer has had a long acquaintance with Swainson's warbler in the
low country of Carolina. Except during September (fall migration) the
birds were almost never seen out of sight of substantial growths of
cane, even when the nests were built in bushes, low trees, or vines.
This has been the experience of practically all observers and, as Brooks
and Legg (1942) remark, "an idée fixe among ornithologists" existed; the
familiar description of habitat by Brewster (1885a) became a dictum:
"Briefly, four things seem indispensible to his existence, viz., water,
tangled thickets, patches of cane, and a rank growth of semi-aquatic
plants."

Hence, the ornithological world received a surprise to learn that
_swainsonii_ was a summer resident and breeder in different localities
of high altitude in the Appalachian Chain. Although several observers
have found the bird nesting beyond the limits of the Coastal Plain, even
in Piedmont territory, as La Prade (1922) did at 1,050 feet above sea
level, it was E. A. Williams (1935) who first detected it in a truly
mountainous terrain. During two successive summers he found birds near
Tryon, N. C., "in open woods."

Loomis (1887) was quite prophetic when, in recording a Swainson's
warbler from Chester, S. C., "in the heart of the Piedmont Region, one
hundred and fifty miles from the coast," he wrote: "It awakens the mind
to the possibility of an Up-Country habitat, yet awaiting discovery,
where the true centre of abundance will finally be located."

The efforts of Brooks and Legg (1942) have shown Swainson's warbler to
be a locally common summer resident in south-central West Virginia up to
an elevation of 2,000 feet above sea level; no positive evidence of
breeding has been found, but it undoubtedly does breed. In Tennessee,
Wetmore (1939) has found the bird in mountainous country at 3,000 feet.

The question naturally arises, Did Swainson's warbler always inhabit
higher altitudes, or is this a recent extension of range and partial
change of habitat? The answer will probably never be found; but study of
changing conditions in its low country habitat for the past several
decades may throw light on this interesting problem. Within the writer's
experience the canebrake areas have long been exposed to forest fires,
timber cutting, overgrazing, drainage, and the construction of a
hydroelectric project, as a result of which thousands of acres of
timbered swampland are now under water.

_Spring._--The birds that winter in Jamaica enter the United States
through Florida, but it is probable that those from Yucatán make a
direct flight across the Gulf to the delta of the Mississippi. The
earliest recorded spring arrival in the United States was on March 22,
1890, on the lower Suwanee River. The same year the species was taken at
the Tortugas, March 25 to April 5 (Chapman, 1907). The earliest arrival
near New Orleans, was March 30, 1905 (Kopman, 1915). Meanley (MS.)
records it from central Georgia on March 31, 1944. Swainson's warbler
reaches the vicinity of Charleston, S. C, during the first week of
April, the earliest being the fifth of that month.

_Nesting._--Nests are built in bushes, canes, masses of vines, and
briers; 10 feet seems to be the maximum height from the ground, while
some nests have been found as low as 2 feet. The average elevation would
be around 3 feet. As many nests are built over dry ground as over water.
The nest is quite bulky and loosely constructed; a typical one in situ
looks like a bunch of leaves lodged in a bush or cane, as the stems
point upward. The outer walls of the nest are composed of various leaves
such as oak, gum, maple, tupelo, and cane; the inner walls are usually
of cane, while the lining is of pine needles, black fiber of moss
_Tillandsia_, cypress leaves, rootlets, or grass stems. Sometimes
horsehair is also present.

[AUTHOR'S NOTE: A few more notes on the nesting of Swainson's warbler
may well be added to the above general statements. Brewster's (1885b)
nests, taken by Wayne in the low country of South Carolina, are
evidently typical for that region. All four of these nests were in
canes. Wayne (1886) says that the nests "are generally built in canes,"
but he has also found them "in small bushes, and in one instance in a
climbing vine, by the side of a large public road." Brewster (1885b)
gives the measurements of two of his nests; the smallest of the four
measures--

externally 3.50 in diameter by 3.00 in depth; internally 1.50 in
diameter by 1.50 in depth; the greatest thickness of the rim or outer
wall being 1.00. * * * The nest June 27 is very much larger, in fact
quite the largest specimen that I have seen, measuring externally 5.00
in diameter by 6.00 in depth; internally 1.50 in diameter by 1.25 in
depth; with the rim in places 1.75 thick. It is shaped like an inverted
cone, the apex extending down nearly to the point of junction of the
numerous fascicled stems which surround and support its sides. Its total
bulk fully equals the average nest of our Crow Blackbird, while it is
not nearly as finished a specimen of bird architecture. Indeed it would
be difficult to imagine anything ruder than its outer walls,--composed
of mud-soaked leaves of the sweet gum, water oak, holly, and cane,
thrown together into a loose mass, bristling with rough stems, and
wholly devoid of symmetry or regularity of outline. The interior,
however, lined with pine needles, moss fibre, black rootlets, and a
little horse-hair, is not less smooth and rounded than in the other
specimens.

Troup D. Perry (1887), with his friend George Noble, found no less than
24 nests near Savannah, Ga., in 1887; some of these were in gall or
myrtle bushes and one was in a saw palmetto 2-1/2 feet high. S. A.
Grimes has sent us a photograph of a nest on the broad leaf of a saw
palmetto (pl. 7). Albert J. Kirn (1918) says of the nesting sites of
Swainson's warbler in Oklahoma: "A well shaded clump of trees in the
woods, such a place as would suggest itself for a Wood Thrush, yet not
exactly so, with considerable 'buck brush' undergrowth, but no grass or
weeds is selected for a nesting site. In the top of this 'buck brush'
usually about two feet high the nest is built; about half of the nests
found were close to the river bank--the Little Caney River. All but two
were built in the brushy undergrowth. These two were fastened to briers
and slender brush and were higher up, 3.5 and 4 feet."

F. M. Jones wrote to Brooks and Legg (1942) of a nest found in
southwestern Virginia: "This nest was in a very dense growth of
rhododendron bushes close to a stream of water where the sunlight never
penetrated. It was 5 ft. 6 in. up, built in the forks of a slender beech
limb which grew across the top of a rhododendron bush (_R. maximum_) and
partly supported by the top of the rhododendron. * * * The outside of
the nest measured 7 in. wide by 5 in. deep and the inside 2 in. wide by
1-13/16 in. deep."

It is evident, from the above and other similar accounts that, at higher
elevations northward and westward, Swainson's warbler nests in bushes
and vines where there are no canes to be found.]

_Eggs._--Swainson's warbler usually lays three eggs; sets of four are
rare and of five very rare. Although there are records of nests
containing two incubated eggs or two young birds, these probably
represent incomplete sets or cases where an egg or a nestling has been
destroyed.

Eggs are quite globular, the two ends sometimes scarcely
distinguishable; the shell is thick and has a distinct polish; the
ground color is white with a bluish tinge; however, a set of three eggs
in the writer's collection had a faint greenish tinge, while several
observers describe sets of pale pink or buffy white.

Rarely, spotted eggs are found. Wayne (1910) says: "Spotted eggs are,
however, very rare and I have found only four or five nests containing
them." The only spotted egg the writer has found is in the set referred
to above; of these, two are immaculate, while the third is "faintly
though distinctly speckled around the larger end with reddish brown"
(Dingle, 1926).

Brewster (1885b) describes a set collected by the late Arthur T. Wayne:
"One is perfectly plain; another * * * has two or three minute specks
which may be genuine shell markings; while the third is unmistakably
spotted and blotched with pale lilac. Over most of the surface these
markings are fine, faint, and sparsely distributed, but about the larger
end they become coarser, thicker, and deeper colored, forming a
well-defined ring or wreath."

Burleigh (1923) writes: "Unlike all the descriptions I had read, and the
few eggs I had seen, these were light pink in ground color and dotted
distinctly over the entire surface with light brown spots, this almost
forming a wreath at the larger end of one egg." These eggs were found
near Augusta, Ga., and the parent was secured.

Wayne (1910) was of the opinion that two broods are raised in a season.

[AUTHOR'S NOTE: The measurements of 50 eggs average 19.5 by 15.0
millimeters; the eggs showing the four extremes measure =21.6= by 14.2,
20.8 by =16.0=, =18.0= by 14.1, and 19.5 by =13.5= millimeters
(Harris).]

_Plumages._--[AUTHOR'S NOTE: Ridgway (1902) describes the juvenal
plumage of Swainson's warbler as follows: "Head, neck, back, rump, upper
tail-coverts, chest, sides, and flanks plain brown (varying from
broccoli to bister); rest of under parts whitish or dull pale yellowish,
more or less clouded with brown; middle and greater wing-coverts
indistinctly tipped with cinnamon-brown; otherwise like adults, but no
trace of lighter superciliary nor darker postocular stripes." Specimens
that I have seen in this plumage are more nearly "cinnamon-brown" than
the colors named above on the back and wing coverts, and the latter show
very little evidence of cinnamon tips.]

The postjuvenal molt, which evidently includes only the contour plumage
and the wing coverts, occurs early in the summer; I have seen young
birds beginning to acquire the first winter plumage as early as June 12,
and others that had nearly completed the molt on July 20; these birds
were not yet fully grown. Wayne (1910) writes: "I have taken young birds
which were as large as the adults and which were acquiring their
autumnal plumage as early as June 2, but it must be borne in mind that
the season in which these young were taken (1906) was exceptionally
advanced."

Brewster (1885a) describes the young bird in its fall plumage as
follows: "Entire upper parts rich olive strongly tinged with
reddish-brown, the crown scarcely deeper-colored than the back, the
wings a trifle redder; loral stripe blackish; superciliary stripe tinged
with yellow; under parts strongly yellowish, otherwise like the adult."

The nuptial plumage is apparently assumed by wear and fading, the
reddish-brown and yellowish colors becoming much duller. There are no
specimens available of either young or adult birds that indicate a
prenuptial molt.

The postnuptial molt seems to occur mainly in August, but perhaps
earlier, and is evidently complete; I have seen birds in full, fresh
autumn plumage as early as August 28. This fresh plumage is similar to
the spring plumage, but the crown and back are nearly uniform brown, the
crown is darker than in spring, the back is browner than in spring, and
the breast and flanks are more or less clouded with grayish.

_Food._--Howell (1924) says that "four stomachs of this bird from
Alabama contained remains of caterpillars, spiders, and Hymenoptera
(ants, bees, etc.)."

Brewster (1885a) considered the principal food to be small coleopterous
insects, "as well as some small green worms that are found on water
plants, such as the pond lily (_Nymphaea odorata_) and the Nelumbium
(_Cyamus flavicomus_).

_Behavior._--Swainson's warbler is an unsuspicious bird and can be
easily observed in its haunts where the vegetation is not too dense and
tangled and the tree canopy overhead partially open. The neutral color
of the bird is often apt to conceal him in the shadowy undergrowth.
Singing males usually remain on the same perch during their periods of
song, apparently disinclined to move. He often sings from the ground
during insect hunting; Meanley (MS.) says: "It was so wrapped up in its
song as to be absolutely unconcerned; it sang at my very feet with its
head thrown back, its beak pointing perpendicularly toward the sky,
pouring forth its resounding melody in the best of warbler fashion."

The female is a close sitter, and the observer has usually to touch her
before she leaves the nest. Grimes (1936) writes: "This bird would not
leave her eggs until _pushed_ off, and when I held my hand over the nest
she straddled my fingers in trying to get back onto it. * * * When I did
drive her away from the nest she fluttered along on the ground in the
manner of a crippled bird, her actions manifestly intended to induce me
to follow. This bird certainly was not badly frightened, for within a
few minutes she was back on her nest, accepting deerflies from my
fingers and swallowing them with apparent relish."

Brewster (1885a) gives an admirable portrayal:

His gait is distinctly a walk, his motions gliding and graceful. Upon
alighting in the branches, after being flushed from the ground, he
assumes a statuesque attitude, like that of a startled Thrush. While
singing he takes an easier posture, but rarely moves on his perch. If
desirous of changing his position he flies from branch to branch instead
of hopping through the twigs in the manner of most Warblers. Under the
influence of excitement or jealousy he sometimes jets his tail, droops
his wings, and raises the feathers of the crown in a loose crest, but
the tail is never jerked like that of a _Geothlypis_, or wagged like
that of a _Siurus_. On the contrary, his movements are all deliberate
and composed, his disposition sedentary and phlegmatic.

_Voice._--The bird student who hears the song of Swainson's warbler as
he sings in his wooded retreat is fortunate, for it is one of the
outstanding warbler songs and, once heard, leaves a lasting impression
upon the listener. At a distance it bears much resemblance to the songs
of the hooded warbler and the Louisiana water-thrush. Close up, however,
the appealing quality, lacking in the other two, impresses the listener
strongly. The song has, in the majority of individuals, a highly
ventriloquial effect, but the writer has listened to birds whose notes
did not in the slightest degree possess this quality.

The song varies in length and number of notes but can be separated into
two distinct parts; the first few notes are uttered rather slowly, the
last ones more rapidly and on a descending scale. The second part
closely follows the first, with no apparent separation. Brooks and
Legg (1942) write: "It might be translated as _whee, whee, whee,
whip-poor-will_, the first two (or three) introductory notes on an even
pitch, the last _whee_ a half-tone lower, and the slurred phrase with
_will_ separated into two syllables, and accented on the _whip_ and on
the _wi_-part of the _will_. The last phrase sounded at times remarkably
like one of the songs of the White-eyed Vireo."

When the singer begins his performance, the bill is pointed directly up,
and he seems entirely unconscious of anything but his own musical
efforts. "During his intervals of silence," says N. C. Brown (1878), "he
remains motionless, with plumage ruffled, as if completely lost in
musical reverie." Brewster (1885a) adds:

It is very loud, very rich, very beautiful, while it has an
indescribably tender quality that thrills the senses after the sound has
ceased. * * * Although a rarely fervent and ecstatic songster, our
little friend is also a fitful and uncertain one. You may wait for hours
near his retreat, even in early morning, or late afternoon, without
hearing a note. But when the inspiration comes he floods the woods with
music, one song often following another so quickly that there is scarce
a pause for breath between. In this manner I have known him to sing for
fully twenty minutes, although ordinarily the entire performance
occupies less than half that time. Such outbursts may occur at almost
any hour, even at noontide, and I have heard them in the gloomiest
weather, when the woods were shrouded in mist and rain.

Several times the writer has seen males when the inspiration had not
quite come to them; the bird would throw back its head but utter only
one or two opening notes of his song.

The call note is a chip, which Brewster calls "a soft _tchip_
indistinguishable from that of _Parula americana_." But Murray (1935)
writes that it is "more throaty and full-bodied than that of most
Warblers." Brooks and Legg (1942) describe it as "clear, penetrating
chirps, having (to our ears) much the same quality as do the chirps of
the Mourning Warbler. They are not quite so loud, but have a more
ringing quality than those of the Hooded Warbler."

_Field marks._--[AUTHOR'S NOTE: Swainson's warbler is a plainly colored
bird, with no conspicuous field marks. It is brownish olive above and
whitish below, with no white in either wings or tail; there is a whitish
line over the eye and a dusky streak through it; but the bill is long
and sharply pointed.]


DISTRIBUTION

_Range._--Southeastern United States to southern Mexico.

_Breeding range._--Until about 1935 Swainson's warbler was considered to
be confined in summer to the southern canebrakes and coastal marshes. It
is now known to breed =north= to extreme southern Illinois, probably
(seen in breeding season to Olive Branch, Duquoin, and Mount Carmel);
southeastern Kentucky (Big Black Mountain); central to northern West
Virginia (Charleston, Mount Lookout, Sutton, and Buzzard Rocks,
Monongalia County); and southeastern Maryland (Pocomoke River Swamp).
=East= to eastern Maryland (Pocomoke River Swamp); eastern Virginia
(Warwick County and Dismal Swamp); eastern North Carolina (New Bern,
Lake Ellis, and Red Springs); eastern South Carolina (Summerton,
Charleston, and Yemassee); eastern Georgia (Savannah and Okefinokee
Swamp); and northeastern Florida (Jacksonville). =South= to northern
Florida (Jacksonville, Oldtown, Whitfield, and Pensacola) and southern
Louisiana (Mandeville, New Orleans, and Baton Rouge). =West= to eastern
Louisiana (Baton Rouge, Bayou Sara, and Jena); central Arkansas (Camden
and Conway); extreme northeastern Oklahoma (Copan); and central Missouri
(Concordia).

Within this large breeding area are two almost discontinuous breeding
ranges: the coastal and swamp range long considered the only home of the
species; and the more recently discovered mountain home along the slopes
of the Allegheny Mountains from northern West Virginia nearly to the
Georgia line where it has been found to an altitude of nearly 3,000
feet.

_Winter range._--The winter home of the Swainson's warbler is very
imperfectly known from a dozen or more specimens, most of which are from
Jamaica where it has been listed as a rare winter resident. There are
records also from the Swan Islands (March 1); Santa Lucia, Quintana Roo;
Pacaytain, Campeche; and the city of Veracruz. Two specimens have been
taken near Habana, Cuba; one on September 25, the other in April; and
one near Guantánamo on January 18, 1914.

_Migration._--Dates of spring departure are: Jamaica, April 8.
Cuba--Habana, April 14.

Early dates of spring arrival are: Florida--St. Petersburg, March 25.
Alabama--Autaugaville, April 3. Georgia--Savannah, March 25.
South Carolina, April 1. Louisiana--New Orleans, March 30.
Mississippi--Biloxi, March 31. Tennessee--Memphis, April 20.
Texas--Point Bolivar, April 17.

Late dates of fall departure are: Texas--Kemah, September 27.
Tennessee--Sulphur Springs, September 9. Mississippi--Gulfport, October
6. South Carolina--Charleston, October 10. Georgia--Savannah, October
18. Alabama--Greensboro, September 6. Florida--Pensacola, October 2;
Sombrero Key (4 struck lighthouse November 10).

Dates of fall arrival are: Tamaulipas--Matamoros, August 29. Jamaica,
October 1.

_Casual records._--A specimen was recorded near Corsicana, Tex., on
August 24, 1880; another was collected at Kearney, Nebr., on April 9,
1905; and one near Holly, Prowers County, Colo., on May 12, 1913.

_Egg dates._--Florida: 3 records, May 7. Georgia: 35 records, May 4 to
July 13; 19 records, May 29 to June 17, indicating the height of the
season. South Carolina: 28 records, May 2 to June 30; 14 records. May 12
to June 12 (Harris).




HELMITHEROS VERMIVOROS (Gmelin)

WORM-EATING WARBLER


PLATE 10


HABITS

The breeding range of the worm-eating warbler covers much of the central
portion of the United States east of the prairie regions. Its center of
abundance seems to be in the vicinity of Pennsylvania, but it breeds
less abundantly northward to southern Iowa, New York, and New England
and southward to Missouri and to northern Alabama and Georgia, as well
as in much of the intervening wooded region, where it is essentially a
woodland bird.

The distribution, migration, and habits of this warbler were but poorly
understood by the early writers on American birds, and neither Wilson
nor Audubon ever saw its nest; the latter's description of the nest,
probably from hearsay, is entirely wrong. Frank L. Burns writes to me:
"Bartram neglected to list this species, although he had furnished the
type to Edwards 35 years earlier, and from the information furnished by
the youthful Bartram it doubtless received its name, which is a misnomer
perpetuated by Gmelin in his _Motacilla vermivora."_ Mr. Burns says
further on in his notes: "I searched for 10 seasons before I found my
first nest, and oddly enough it was through the parent bird carrying a
'worm' to its young; nevertheless I have since thought that a more
fitting name for the species would have been hillside or laurel
warbler."

Hillside warbler would not be a bad name for this bird, which shows a
decided preference for wooded hillsides covered with medium-sized
deciduous trees and an undergrowth of saplings and small shrubbery.
Often a running stream with numerous swampy places, overgrown with brier
tangles and alders, bounds the base of the hill as an additional
attraction. It is seldom seen outside of its favorite woods and returns
year after year to the same chosen haunts.

W. E. Clyde Todd (1940) says that in western Pennsylvania "wooded slopes
are its chosen abodes, the shadier and cooler the better. * * * Deep
ravines, down which trickle little streams, and the slopes of which
support good stands of deciduous trees, with plenty of shrubbery and
bushes for cover, are favorite resorts." In Ritchie County, W. Va.,
William Brewster (1875) found it "most partial to the retired thickets
in the woods along water courses, and seldom or never found in the high
open groves."

_Spring._--The northward movement of the worm-eating warbler evidently
begins in March, as the earliest arrivals from the Bahamas, the West
Indies, and Cuba reach southern Florida during the first week in April.
From its main winter resorts in Central America the flight seems to be
partially across the Gulf of Mexico. Professor Cooke (1904) says in
part: "The time of arrival on the coasts of Louisiana and Texas is about
the same as in southern Florida. * * * Houston is the southernmost point
in Texas from which it has been recorded to date, and Alta Mira is the
northernmost point of record in Mexico. Since the species is apparently
not common west of Louisiana or north of Vera Cruz, it is probable that
the principal line of migration is from Yucatan and the coast
immediately west of Yucatan directly north to the northern coast of the
Gulf of Mexico." According to Williams (1945) the species is common on
the coast of Texas in spring, and it probably migrates along the coast.
Thence the migration proceeds northward through the Mississippi Valley
and through the Atlantic Coast States east of the Alleghenies, the
warblers reaching the more northern breeding grounds by the middle of
May, where nesting activities begin as soon as mates have been selected.

_Nesting._--Evidently Thomas H. Jackson, of West Chester, Pa., was the
first to report the discovery of the nest of the worm-eating warbler; he
published an account of it in the American Naturalist for December 1869,
from which Baird, Brewer, and Ridgway (1874) quote as follows: "On the
6th of June, 1869, I found a nest of this species containing five eggs.
It was placed in a hollow on the ground, much like the nests of the
Oven-Bird (_Seiurus aurocapillus_), and was well hidden from sight by
the dry leaves that lay thickly around. The nest was composed externally
of dead leaves, mostly those of the beech, while the interior was
prettily lined with the fine, thread-like stalks of the hair-moss
(_Polytrichium_). * * * So close did the female sit that I captured her
without difficulty by placing my hat over the nest."

This nest was quite characteristic of the species. Mr. Burns writes to
me: "The nest, well hidden under a drift of dead forest leaves, never
varied in composition in over a hundred examples examined by me, in
partly skeletonized leaves and the characteristic reddish-brown lining
of the flower stem of the hair moss." Every one of 50 nests found by Mr.
Jackson was lined with these flower stems, and out of 34 nests reported
by Dr. Samuel S. Dickey (1934) only one failed to contain this material,
being lined with "black and gray horsehair." Samuel B. Ladd (1887) says
that "sometimes fine grass and horse-hair are used as part of the
lining." Dr. Chapman (1907) writes: "Nests taken by J. N. Clark at
Saybrook, Connecticut (C. W. C.) are composed of decayed leaves and
lined with stems of maple seeds." And there are probably a few other
exceptions to the rule.

Most observers agree that the worm-eating warbler prefers to nest on
hillsides, either sloping or steep, but a number of nests have been
found on the sides of deep, shady ravines, or on steep banks. Mr. Ladd
(1887), however, states: "I have observed that these birds are not
confined necessarily to hill-sides, as was heretofore supposed, as I
have taken three sets on level ground and in rather open places, with
little shade. The experience of Mr. Thomas H. Jackson of this place,
who has taken ten nests this year, corroborates this fact."

The nests are generally well concealed under a canopy of dead leaves,
drifted by the wind and lodged against a maple, beech, dogwood, or ash
sapling, or under hydrangea, laurel, or rhododendron bushes, or under
some bunch of weeds or other obstruction. They are sometimes concealed
under the roots of a tree or in a cavity in a bank where they are
protected somewhat by fallen leaves.

_Eggs._--The number of eggs laid by the worm-eating warbler varies from
3 to 6, but the set usually consists of 4 or 5. The eggs are ovate or
short ovate, sometimes rather pointed, and only slightly glossy. The
white ground color is speckled and spotted with shades of "russet,"
"vinaceous russet," and "auburn," intermingled with "light brownish
drab" and "light vinaceous-drab." The markings, usually more thickly
grouped at the large end, vary considerably, some eggs being boldly
marked, while others are almost immaculate, or have just a few pale
freckles of "light brownish drab" and "fawn." The measurements of 50
eggs average 17.4 by 13.6 millimeters; the eggs showing the four
extremes measure =20.8= by =14.5=, and =15.5= by =12.7= millimeters
(Harris).

_Incubation._--Frank L. Burns (1905) writes:

Incubation does not always commence immediately after completion of set,
particularly if the season be young. It is probable that the second
night witnesses the beginning of that period and, as far as my
experience goes, I believe it is performed by the female alone. The male
feeds her when covering newly hatched young.

The home-coming of a brooding bird, after a brief airing and feeding, is
heralded several hundred yards distant by frequent _chips_ and short
flights from branch to branch near the ground, in leisurely fashion and
circuitous route, until at length, arriving above the nest, she runs
down a sapling and is silent. The bird is a close sitter and if
approached from the open front will often allow a few minutes' silent
inspection, eye to eye, at arm's length, sometimes not vacating until
touched, then she runs off in a sinuous trail, not always feigning
lameness before the young are out. When disturbed with young in the nest
she will flutter off with open wings and tail, and, failing to lead one
off, will return with her mate, who is seldom far off at this period,
circling about the nest or intruder, and, if the young are well
feathered, she will dash at them, forcing them from the nest and to
shelter. Once this brave little bird dashed at me and ran up to my
knee, scratching with her sharp little claws at every step. On the
return the birds always make the vicinity ring with their protests--a
quickly repeated _chip_. The period of incubation in one instance was
thirteen days.

_Young._--Mr. Burns continues:

Young fear man soon after their eyes are open, and a menacing finger
will cause them to scamper out and away, repeated replacing in the nest
proving of no avail after they became panic-stricken. At three days of
age they made no outcry but opened their mouths for food, which
consisted of a species of white moth, or "miller," and soft white grubs,
supplied by either of the parent birds. At that period they were naked
except a fluff on head and wing quills, just showing feathers at tips.
In the presence of an intruder and absence of the parents, they will sit
motionless if not threatened, and, but for the blinking, beady eyes, one
might mistake them when well fledged, at very close range, for dead
leaves. The head stripes became visible under the nestling down on the
seventh day, and they left the nest ten days after leaving the shell, in
the one case I have kept record of. The parents keep the young together
for several days at least, just how long is impossible to say. One brood
is all that is reared in a season, I think.

_Plumages._--Dr. Dwight (1900) calls the natal down "brownish
mouse-gray," and describes the juvenal plumage as follows: "Whole body
plumage and the wing coverts cinnamon, palest on the abdomen. Wings and
tail olive-brown edged with olive-green. Two indistinct lateral crown
stripes brownish mouse-gray. A transocular streak dusky." Ridgway's
(1902) description is somewhat different: "Head, neck, and under parts
buff, the pileum with two broad, but strongly contrasted, lateral
stripes of wood brown or isabella color; a postocular streak of the same
color; back, scapulars, rump, and upper tail-coverts wood brown or
isabella color; wing-coverts light buffy olive, the middle and greater
broadly but not sharply tipped with cinnamon-buff; remiges and rectrices
grayish olive-green, as in adults." Young birds seem to vary
considerably in the color of the upper parts.

A partial postjuvenal molt occurring in late June or early July involves
all the contour plumage and the wing coverts but not the rest of the
wings or the tail. The young bird in its first winter plumage is
practically indistinguishable from the adult at that season, except for
the juvenal wings, in which the tertials are lightly tipped with rusty
brown.

There is apparently no spring molt, but a complete postnuptial molt
occurs in July. Spring birds are slightly paler, grayer and less buffy
than in the fall. The sexes are practically alike in all plumages.

_Food._--As I have said, the name worm-eating warbler seems to be
somewhat of a misnomer for this bird. Edward H. Forbush (1929) writes:
"I find no records of any consumption of earthworms by this species,
which although a typical ground warbler spends some of its time hunting
among the branches of trees, where it finds span-worms. It also hunts on
the ground in damp places frequented by army-worms. Nevertheless these
are not worms but caterpillars. Probably, however, in its perambulations
and peregrinations upon the surface of the earth the bird now and then
does pick up a small earthworm, for earthworms form a staple food for
many birds when the ground is moist."

Arthur H. Howell (1924) says: "Little is known of the food of this
species, but it seems doubtful whether it lives up to its name of
worm-eater.' Two stomachs of this bird from Alabama contained remains of
weevils, beetles, bugs, caterpillars, and Hymenoptera." Howell (1932)
further reports: "The stomachs of three individuals taken in Florida in
April contained small grasshoppers, caterpillars, sawfly larvae,
beetles, and spiders. One dragon-fly, one bumblebee, and one 'walking
stick' were also included in the contents." Professor Aughey (1878)
included the worm-eating warbler among the birds seen catching locusts
in Nebraska.

_Behavior._--Brewster (1875) gives the best account of the activities of
the worm-eating warbler as follows:

They keep much on the ground, where they _walk_ about rather slowly,
searching for their food among the dried leaves. In general appearance
they are quite unique, and I rarely failed to identify one with an
instant's glance, so very peculiar are all their attitudes and motions.
The tail is habitually carried at an elevation considerably above the
line of the back, which gives them a smart, jaunty air, and if the
dorsal aspect be exposed, in a clear light, the peculiar marking of the
crown is quite conspicuous. Seen as they usually are, however, dimly
flitting ahead through the gloom and shadow of the thickets, the
impression received is that of a dark little bird which vanishes
unaccountably before your very eyes, leaving you quite uncertain where
to look for it next; indeed, I hardly know a more difficult bird to
procure, for the slightest noise sends it darting off through the woods
at once. Occasionally you will come upon one winding around the trunk of
some small tree exactly in the manner of _Mniotilta varia_, moving out
along the branches with nimble motion, peering alternately under the
bark on either side, and anon returning to the main stem, perhaps in the
next instant to hop back to the ground again. On such occasions they
rarely ascend to the height of more than eight or ten feet. The males
are very quarrelsome, chasing one another through the woods with loud,
sharp chirpings, careering with almost inconceivable velocity up among
the tops of the highest oaks, or darting among the thickets with
interminable doublings until the pursuer, growing tired of the chase,
alights on some low twig or old mossy log, and in token of his victory,
utters a warble so feeble that you must be very near to catch it at all,
a sound like that produced by striking two pebbles very quickly and
gently together, or the song of _Spizella socialis_ heard at a distance,
and altogether a very indifferent performance.

_Voice._--Aretas A. Saunders has contributed the following study of the
song of this warbler:

The song of the worm-eating warbler is a simple trill, varying from
1-2/5 to 2-1/5 seconds in length. It is usually all on the same pitch,
but a few songs rise or fall a half tone, and one record I have rises a
full tone and then drops a half tone at the end. The quality is not
musical, but rather closely resembles some forms of the chipping
sparrow's song. The pitch varies from G sharp´´´ to F sharp´´´´, one
tone less than an octave.

The majority of songs are a continuous trill, that is, the notes are too
fast to be separated and counted by ear. I have three examples that are
broken into short, very rapid notes. Two of these were of 18 notes and
one was of 28. Most of the songs vary in loudness, becoming loudest in
the middle, or beginning loud and fading away toward the end. One record
becomes louder toward the end and ends abruptly.

Francis H. Allen describes in his notes a song "remarkably like that of
the chipping sparrow, but more rapid than is usual with that species, I
think, and perhaps shorter, though not so short as the chippy's
early-morning song. The bill quivers with the song, but does not close
between the _chips_. The bird sang constantly as it flitted about,
usually 10 or 20 feet from the ground, seeming to prefer dead branches
and twigs."

Almost everyone emphasizes the resemblance of the song to that of the
chipping sparrow. Burns (1905) says: "I can distinguish no difference
between the notes of this species and the Chipping Sparrow; the first
may be a trifle weaker perhaps." But, in some notes recently sent to me,
he writes: "The song has often been described as easily mistaken for
that of either the chipping sparrow or slate-colored junco, but by no
means by an expert. The notes of the worm-eater have a buzzing or
bubbling quality not easily described, but are quite distinct from the
flat notes of the species named above." And Eugene P. Bicknell (1884)
writes: "The songs of no other three birds known to me are more alike
than those of the Worm-eating Warbler, the Chipping Sparrow, and the
Slate-colored Snowbird." He is in agreement with Saunders and Burns that
this bird sings from the time of its arrival until the last of June or
early July, but he also says: "On July 10, 1881, several of these birds
were silently inhabiting a small tract of woodland, their first season
of song having passed; here, on August 14, and again on the 21st, they
were found in fine plumage and in full song." Evidently there is a
cessation of singing during the molting period.

Burns (1905) says of the song: "The series of notes may be uttered while
perched, or creeping about the lower branches of the trees, sapling
tops, bushes or fallen brush, or while on the ground. With slightly
drooping tail and wings, puffing out of body plumage, throwing its head
back until the beak is perpendicular, it trills with swelling throat an
unvarying _Che-e-e-e-e-e-e_, which does not sound half so monotonous in
the woods as does the Chippy's lay in the open."

Dr. Chapman (1907) adds: "Mr. W. DeW. Miller of Plainfield, New Jersey,
tells me that he has on two occasions heard a flight song from this
species. It is described by him as much more varied and musical than the
ordinary song, though lacking in strength. It was given as the bird flew
through the woods at an even level, not rising above the tree-tops, as
does the Oven-bird and other flight singers."

_Field marks._--When seen walking around on the ground the worm-eating
warbler might be mistaken for an ovenbird, but the conspicuous black
stripes on the head of the former are quite distinctive, very different
from the head markings of the latter. Moreover, the ovenbird is
distinctly spotted on the breast, whereas the warbler has a plain,
unmarked breast and no conspicuous wing bars. Except for the bold
stripes on the head it is just a plain olive and buffy warbler in all
plumages.

_Enemies._--Says Burns (1905): "This Warbler's enemies are wood-mice,
red squirrels and hunting dogs; the latter will sometimes push up and
overturn the nest; an occasional weasel or blacksnake may destroy a few
young. The percentage of loss while in the nest cannot be high."

Friedmann (1929) regards the worm-eating warbler as a "rather uncommonly
imposed upon species" by the eastern cowbird. "Twenty-one definite
records, and as many more indefinite ones have come to my notice."

_Winter._--Dr. Alexander F. Skutch contributes the following: "Widely
distributed as a winter resident in Central America, the worm-eating
warbler appears to be everywhere very rare. It occurs from Guatemala to
Panamá on both coasts, and upward in the mountains to at least 5,000
feet. On February 26, 1935, I found one in the forest on Barro Colorado
Island, Canal Zone, which appears to represent a slight southward
extension of the known range. I have recorded this rare visitant from
every part of Central America below 6,000 feet in which I have made an
extended sojourn during the months of the northern winter, yet only one
or two in each locality, except on the Finca Mocá on the Pacific slope
of Guatemala at 3,000 feet above sea-level, where in one day--January
21, 1935--I saw three. The worm-eating warbler is found in the Tropics
beneath dense thickets or in the undergrowth of the forest, usually near
the ground; but at times one will rise to the lower branches of the
trees to investigate curled dead leaves caught up among them. It is
solitary rather than social in its habits.

"The records of the occurrence of this warbler in Central America are
too few to indicate clearly the dates of its arrival and departure. I
found one at Tela, Honduras, on August 19, 1930; but the next early
record is for October 14, at the same locality. Griscom quotes a record
by Dearborn for the occurrence of this warbler at Patulul, Guatemala, on
April 2; but except for this, the latest record I have seen is from El
General, Costa Rica, March 11, 1939."


DISTRIBUTION

_Range._--Eastern United States to Panamá.

_Breeding range._--The worm-eating warbler breeds =north= to
northeastern Kansas (Lawrence); possibly central southern Nebraska (Red
Cloud); probably south-central Iowa (Des Moines); probably southern
Wisconsin (Wyalusing, Madison, and Milwaukee); northeastern Illinois
(Hinsdale); southern Indiana (Terre Haute, Bloomington, and
Indianapolis); central Ohio (Columbus, East Liverpool, and possibly
Cleveland); southern New York (Penn Yan and Albany), and southern
Connecticut (New Haven and Saybrook). It has been found in summer north
to London, Ontario; Northampton, Ipswich, and North Eastham,
Massachusetts. =East= to Connecticut (Saybrook); Long Island (Newtown);
northern New Jersey (Elizabeth and Morristown); eastern Pennsylvania
(Norristown and Philadelphia); northern Delaware (Wilmington); central
Maryland (Baltimore; rarely east of Chesapeake Bay); eastern Virginia
(Cobham and Dismal Swamp); central North Carolina (Chapel Hill and
Statesville); northwestern South Carolina (Caesars Head, Mount Pinnacle,
and Sassafras Mountain); and northern Georgia (Brasstown Bald and
Atlanta). =South= to northern Georgia (Atlanta); central Tennessee
(Nashville and Wildersville); northern Arkansas (Newport and Winslow);
and, occasionally, extreme northern Texas (Bowie County and
Gainesville). =West= to northern Texas (Gainesville); northeastern
Oklahoma (Jay); and eastern Kansas (Lawrence). It has been recorded in
summer, but with no evidence of breeding, at Red Cloud, Nebr., and at
London and Vineland Station, Ontario.

_Winter range._--In winter the worm-eating warbler is found =north= to
southern Tamaulipas (Altamira); northern Florida, casually (Blue Springs
and Amelia Island), and the Bahamas (Abaco, Nassau, and Great Inago).
=East= to the Bahamas (Great Inago); Jamaica and central Panamá (Río
Chepo). =South= to Panamá (Río Chepo, Barro Colorado, and Chiriquí).
=West= to western Panamá (Chiriquí); Costa Rica (Escasú and Volcán
Tonorio); El Salvador (Mount Cacaguatique); Guatemala (Dueñas, Patulul,
and Naranjo); southern Chiapas (Huehuetán); western Veracruz (Jalapa);
Hidalgo (Pachuca); and southern Tamaulipas (Altamira).

_Migration._--Late dates of spring departure are: Panamá--Darién March
16. Costa Rica--El General, March 19. El Salvador--Barra de Santiago,
April 8. Guatemala--Patulul, April 2. Yucatán--Mérida, April 9.
Cuba--Habana, May 1. Bahamas--Abaco, April 29. Florida--Seven Oaks,
May 14. Georgia--Cumberland, May 7. Alabama--Barachias, May 1.
Mississippi--Biloxi, April 27. Louisiana--Avery Island, April 23.

Early dates of spring arrival are: Florida--Pensacola, March 26.
Georgia--Savannah, April 4. South Carolina--Mount Pleasant, April 7.
North Carolina--Bat Cave, April 16. Virginia--Richmond, April 19. West
Virginia--Morgantown, April 4. District of Columbia--Washington, April
21. Pennsylvania--Beaver, April 29. New York--Jones Beach, April 20.
Louisiana--Grand Isle, April 3. Mississippi--Bay St. Louis, April 5.
Tennessee--Chattanooga, April 15. Kentucky--Bowling Green, April 3.
Indiana--Brookville, April 17. Ohio--Columbus, April 18.
Texas--Brownsville, March 29. Missouri--St. Louis, April 15.
Iowa--Keokuk, April 21.

Late dates of fall departure are: Missouri--St. Louis, September 20.
Ohio--Austinburg, September 23. Kentucky--Middlesboro, September 27.
Tennessee--Athens, October 5. Mississippi--Biloxi, October 11.
Louisiana--Monroe, September 30. New York--Balston, September 23.
Pennsylvania--Atglen, October 10. District of Columbia--Washington,
September 13. West Virginia--Bluefield, September 19. Virginia--Salem,
October 24. North Carolina--Andrews, October 11; Raleigh, November 3.
South Carolina--Charleston, October 11. Georgia--Atlanta, October 10.
Florida--Fernandina, October 3.

_Casual records._--A specimen was collected in Bermuda on October 4,
1899. An individual was present at Wood Pond near Jackson, Somerset
County, Maine, September 1 to 12, 1935; and one was reported seen at
Mayagüez, Puerto Rico, on October 15, 1943, following a small hurricane.

_Egg dates._--Connecticut: 7 records, May 27 to June 29.

New Jersey: 4 records, May 21 to 30.

Pennsylvania: 75 records, May 15 to June 30: 45 records, May 24 to June
5, indicating the height of the season (Harris).




VERMIVORA CHRYSOPTERA (Linnaeus)

GOLDEN-WINGED WARBLER


CONTRIBUTED BY WINSOR MARRETT TYLER

PLATES 10, 11


HABITS

The golden-winged warbler is one of the daintiest among this group of
gay-colored little birds. Its plumage is immaculate white below and
delicate pearl-gray on the upper parts, the crown and wings sparkle with
golden yellow, and on the throat and cheeks is a broad splash of jet
black.

It is only within comparatively recent years that we have become well
acquainted with the goldenwing: the older ornithologists, Wilson,
Audubon, and Nuttall, knew it only as a rather uncommon migrant,
drifting through from the south, and they had no idea where it bred. At
a much later date J. A. Allen (1870) says of it: "This beautiful warbler
has been taken, so far as I can learn, but few times in the western part
of the State; it seems to be more common in the eastern, where it
breeds." He cites the first record of the finding of a nest in the State
in 1869. There is, however, an earlier record of its nesting. Dr. Brewer
(1874) states: "Dr. Samuel Cabot was the first naturalist to meet with
the nest and eggs of this bird. This was in May, 1837, in Greenbrier
County, Va."

William Brewster (1906), speaking of the bird in 1874, when he first
found it in eastern Massachusetts, says: "If the species inhabited any
part of the Cambridge Region before the year just mentioned, it was
overlooked by several keen and diligent collectors, among whom may be
mentioned Mr. H. W. Henshaw and Mr. Ruthven Deane." Since that time the
bird has increased in numbers here until at present it is common in
suitable localities.

_Spring._--The goldenwing appears in eastern Massachusetts about the
middle of May, or sometimes a little earlier, at the time when many of
the resident warblers are arriving on their breeding-grounds. At this
season the bright green leaves are beginning to open in the thickets and
trees on the borders of woodlands where the goldenwing finds its food;
and under the trees in the wooded swamps where the bird will build its
nest, fresh new growth--skunkcabbage, ferns, and a host of spring
plants--is pushing through the dead leaves, spreading a green carpet on
the forest floor. But even thus early in the year, when the trees are
nearly bare, it is not easy to see as it feeds high up in the trees, far
out near the tips of the branches. Indeed, but for its queer little
song, we should rarely suspect that it had come back to its summer home.

_Nesting._--The golden-winged warbler builds its nest on the ground,
generally raised somewhat by a substratum of dead leaves. The nest is
supported by stalks of herbs--often goldenrod or meadow rue--or by fern
fronds, or it may be hidden deep in a clump of grass, or it may lean
against the base of a small shrub or tree with grass all about it. The
leaves above the nest develop as the season advances and soon completely
conceal it, and the plants, by their growth, may raise the nest a little
above the ground. The cup of the nest is made chiefly of long strands of
dry grass and narrow strips of grapevine bark, with a few hairs in the
lining. This fine, flexible material is pressed down on the inside by
the weight of the incubating bird and the nestlings, becoming smooth and
firm like a mat, whereas on the outside wall the long grass blades and
fibrous vegetable shreds are left free and, protruding loosely in all
directions for some distance from the cup, produce a disorderly, unkempt
appearance, like a little loose handful of fine hay.

Edward H. Forbush (1929) quotes an account of the goldenwing by Horace
O. Green who has had an extensive experience with the species and who
gives the following interesting details of the construction of the nest:

The nest of the Golden-wing usually has a bottom layer of coarse dead
leaves on which is placed a ring of large dry leaves, arranged with the
points of the leaves downward, so that the leaf stems stick up
noticeably around the edges of the nest proper, which is built within
and upon this circular mass of leaves, and is made of rather wide strips
of coarse grass or rushes, and usually has considerable grape vine bark
interwoven in it. The nest lining is coarse and rough, sometimes the
eggs being laid on the rough grape vine bark, and in some nests other
coarse fibers are used. A very characteristic feature of the nest lining
is fine shreds of light reddish-brown vegetable fiber, which at first
glance might easily be mistaken for dry needles from the pitch pine--but
careful examination shows it to be the inner layers of the bark from the
grape vines. The nest is very bulky for the size of the bird and Is
rather loosely put together by crossing the materials diagonally, so
that it slightly resembles a rather coarse basket-work. I never saw a
nest of this species which had a soft lining, such as many other
warblers use--the eggs are apparently always deposited on rough
material.

The general color of the nest is very dark, especially just after a
rain, when the materials of which it is composed look almost black--this
being one thing which helps to distinguish these nests from those of the
Maryland Yellow-throat, which generally builds a much lighter colored
nest, lined with fine grass, and sometimes with horse hair. Another
small point of difference which is noticeable on close examination is
that the lining in the Yellow-throat's nest is usually of a much finer
and lighter colored material, and appears to be woven in horizontally,
or at least to show some traces of such a design, especially around the
upper edge--while the Golden-wing closely adheres to the diagonal
criss-cross pattern with the loose ends of the nesting materials
sticking up at an angle above the rim of the nest cavity.

Mr. Green describes the surroundings of the nest thus:

For their summer home these birds prefer the border of deciduous woods,
where tall trees give plenty of shade, to an adjacent clearing with a
growth of briers, bushes and grass, and the nest is usually placed just
outside the line of the forest proper, but within the shade of the
trees. A meadow wholly surrounded by woods is frequently selected. The
ideal place to search for a nest of the species is in one of those
woodland meadows, which has a clear brook flowing through it, with
briers, tussocks of grass and a fresh growth of goldenrod scattered
around in profusion, with birch trees and wild grape vines growing near
the edges where the meadow meets higher ground--and all this bordered by
tall oak, chestnut and maple trees which furnish an abundance of shade
to the vegetation of the meadow itself.

J. Warren Jacobs (1904) describes the nest much as above and adds: "The
opening is not straight down, but slightly tilted, the jaggy leaf-stems
and bark sometimes reaching two or three inches above the rim of the
nest proper. As incubation advances, the rough rim on the lower edge of
the nest becomes broken down, and by the time the young birds are ready
to leave, this part of their home is worn smooth by the attendant
parents."

He gives the measurements of 17 nests as follows: "Outside 3.6 to 5.0
inches in diameter, and 3.0 to 5.0 inches in depth; and on the inside,
from 1.7 to 2.5 inches in diameter by 1.3 to 2.5 inches deep." These
measurements agree very closely with the records of several other
observers. Jacobs continues: "Seemingly before the birds have had time
to complete their nest, the female begins the deposition of the eggs.
Generally, where I had opportunity to watch the nests daily, or at
intervals between the beginning and completion of the set, the eggs were
laid on consecutive days, but in two or three instances it was noticed
that the laying missed a day."

_Eggs._--The set for the golden-winged warbler may consist of anywhere
from 4 to 7 eggs; 5 is perhaps the commonest number, but 4 is a common
number, and the larger numbers are increasingly rare. The eggs are ovate
or short ovate, and have only a slight luster. They are white or creamy
white, with a wide variety of markings in "auburn," "argus brown," "Mars
brown," "hazel," "Hay's brown," "liver brown," and "burnt umber," with
underlying speckles or spots of "light brownish drab" and "light
vinaceous drab." There is, also, much variation in the amount of
markings, some being very sparingly speckled and others are quite
heavily marked, with some of the spots assuming the proportions of
blotches. Occasionally small hairline scrawls, or scattered spots, of
brown so dark as to appear almost black, are found. The markings are
usually denser toward the large end. The measurements of 50 eggs average
16.7 by 13.0 millimeters; the eggs showing the four extremes measure
=18.6= by 13.0, 16.8 by =13.7=, =15.5= by 12.5, and 15.9 by =12.3=
millimeters (Harris).

_Young._--Jacobs (1904) states that the incubation period is 10 days and
that the young birds are able to leave the nest 10 days after hatching.
In a nest which Maunsell S. Crosby (1912) watched closely, the eggs
hatched on June 1 and the young flew on June 10.

The fledglings are delicate little birds, brownish olive on the back,
washed with yellow below, and have two widely separated yellow wing
bars. They have astonishingly long legs and soon become very active,
fluttering about in the shrubbery and clinging to the branches. Walter
Faxon (1911) in speaking of them gives this lively picture which could
well be applied to them soon after leaving the nest: "In appearance and
habit they were grotesque little fellows, clinging with their
disproportionately long legs to the low herbage, like peeping Hylas in
the springtime clinging to the grasses and weeds above the surface of
the water. The little thread-like natal plumes still waving from the
tips of their crown feathers enhanced the oddity of their appearance."
Mr. Faxon, to be exact, is speaking here of some young birds of mixed
parentage, but his words apply equally well to the behavior and
appearance of the young of _chrysoptera_ which he and I watched year
after year together. Both parents are very attentive to their young
brood, bringing to them food which they find both on low plant growth
and high in the overshadowing branches.

The fledglings call to their parents with a very characteristic note, a
little quavering, high, fine chirp which I find written in my journal
_crrr_ and _tzzz_. It suggests somewhat a note of young chipping
sparrows, but is less sharp and crisp. In form it also resembles the
call of the young cowbird, but again it is gentler and weaker in tone.
Mr. Faxon (1911) refers to it as the "cricket note." The young birds
acquire their first winter plumage about a month after they leave the
nest, and hence to the eye are indistinguishable from their parents, but
as they still continue to use the call of their babyhood, they may be
recognized as immature birds even when they are feeding high up in the
trees.

_Plumages._--[AUTHOR'S NOTE: I can find no description of the natal
down. Dr. Dwight (1900) describes the juvenal plumage, in which the
sexes are practically alike, as "above, grayish or brownish olive-green.
Wings and tail slate-black edged chiefly with bluish plumbeous gray, the
coverts and tertiaries with olive-green. Below, pale olive-yellow, the
throat dusky. Transocular streak dusky. * * *

"First winter plumage acquired by a partial postjuvenal moult, beginning
early in July, which involves the body plumage and wing coverts, but not
the rest of the wings nor the tail, young and old becoming practically
indistinguishable." He describes the young male in this plumage as--

above, plumbeous gray veiled with olive-green edgings; the crown bright
lemon-yellow veiled posteriorly only. Below, grayish white, with yellow
edgings here and there, the chin, jugulum, lores and auriculars
jet-black veiled slightly with pale buff. Broad submalar stripes joining
at angle of the chin, and superciliary lines white. Outer half of median
and greater coverts bright lemon yellow forming an almost continuous
wing patch, lesser coverts plumbeous gray, edged with olive-green.

First nuptial plumage acquired by wear, through which the buff edgings
of the black areas, the olive edgings of the back and the yellow edgings
below are almost completely lost, the plumage becoming clear gray,
white, yellow and black.

Of the female, he says: "In first winter and other plumages olive-gray,
dusky on the lores and auriculars, replaces the black areas of the male,
and olive-yellow marks the crown. Above, the plumage is greenish; the
submalar stripes are grayish." Subsequent plumages are acquired by a
complete postnuptial molt in late June and July and by wear in early
spring.]

_Food._--Little exact information has been gathered regarding the food
of the goldenwing. The insects it feeds on are mainly so small that it
is generally impossible to identify them. Jacobs (1904) states: "Once I
saw a female carry a small brownish butterfly to her young; and several
times I have discovered the birds taking small smooth green worms--such
as strip the leaves of their green coat, leaving the ribbed skeleton--to
their nestlings. The legs of a spider protruded from a bird's bill as
she approached her nest."

The little pale green larva which Jacobs mentions impresses us as the
chief article of food, as we watch the birds. It is 1/2 to 3/4 inch long
and appears to have a smooth, hairless skin. These larvae are obtained,
I believe, chiefly in the large trees.

In the following note A. L. Nelson (1933) furnishes an interesting
detail of the bird's diet:

The following observation on the food habits of a Golden-winged Warbler
(_Vermivora chrysoptera_), made in the vicinity of Port Tobacco (Charles
Co.), Maryland on May 6, 1933, seems worthy of mention, inasmuch as
little specific information on the dietary habits of this species has
been recorded. About 1:30 we observed a single individual of this
species actively feeding in a low shrubbery growth of pawpaw (_Asimina
triloba_), which was in full bloom at this date. Closer observation
revealed that the bird was probing about inside the flowers, and
apparently was getting some kind of larvae. Examination of the
flowers revealed that they were infested with a small, brown-headed
lepidopterous larva. Dissection of a large number of flowers indicated
that the infestation was high, the majority of flowers having one larva,
although in many cases two were present. Several infested flowers were
collected for the purpose of rearing the insects to the adult stage
under laboratory conditions. The cycle was completed without difficulty,
the adults emerging within twelve days. These were examined by Dr. Carl
Heinrich of the U. S. National Museum and found to be _Talponia
plummeriana_ Busck, a small brightly colored Tortricid, the only known
food plant of which is the pawpaw.

_Behavior._--A favorite locality for the golden-winged warbler to spend
the summer in eastern Massachusetts may be the border of a wooded swamp
where tall elm and maple trees shade a dense undergrowth of ferns and
other moisture-loving plants, a swamp which runs out toward drier ground
where abounds a growth of gray birches or a tangle of raspberry canes,
wild grapevines, and goldenrod. Such a spot furnishes countless
situations for hiding the nest in the thick vegetation growing in the
half-wet half-dry ground, and also a source of food near at hand in the
high branches of the trees. Much the same conditions exist along the
course of a brook winding through second growth, or near orchards or old
neglected weedy pastures.

Sometimes, as William Brewster (1906) points out, the bird may frequent
"dry hillsides covered with a young sprout growth of oak, hickory or
maple."

In a more southern latitude the habitat may be quite different. Maurice
Brooks (1940), speaking of the bird in the central Allegheny Mountain
region, says: "Shunning the swamps which it frequents in other portions
of its range, it is highly characteristic of the 'chestnut sprout'
association, where the males choose dead chestnuts for perches from
which to sing. It is also fairly common in the pitch and scrub pine
regions on the hills just back of the Ohio river, but becomes less
common toward the eastern portion of the territory with which this
paper deals. It ascends to at least 4,000 feet in Giles Co., Va."

We can watch the little golden-winged warblers best, and often at very
short range, when they are feeding their fledglings recently from the
nest. The little birds sit quietly in the shrubbery near the ground,
waiting for their parents. We can find them easily, for they frequently
utter their characteristic "cricket note," and we can approach them
closely, for they scarcely heed us. The parents, too, when they are
feeding the young birds, pay little attention to us and come fearlessly
to them even when we stand near. At such times they work in a seeming
panic of hurry, flying about in the low growth searching for food, or
visiting the smallest branches high up in the trees, where they cling to
the terminal twigs, hanging like chickadees as they probe among the
curled up leaves (insect nests) for food hidden there, then back to the
waiting young, seemingly in continuous motion and without the slightest
pause in their nervous activity. At this season when the parents are
busy with the young birds, about the third week in June in eastern
Massachusetts, they are so occupied in searching for food that the male
rarely sings.

In two particulars--their tameness, or indifference to our presence, and
the almost complete cessation of singing thus early in the season--the
goldenwing differs from the other common birds which breed in much the
same regions, the chestnut-sided warbler, redstart, northern
yellow-throat, ovenbird, and veery.

Jacobs (1904) speaks of the anxiety of the parent birds if the nest is
disturbed when the nestlings are nearly ready to fly. He says: "If the
hand is placed near the nest at this period of their growth, they will
scramble out and flutter away, all giving vent to their chipping note,
which brings down upon the intruder the wrath of both old birds, who fly
close to his face, snapping their beaks and chipping loudly; then down
upon the ground they fall and feign the broken wing act as long as one
of the young continues to chirp."

_Voice._--The song of the golden-winged warbler is an inconspicuous
little buzzing sound which one might pass by unnoticed, or hearing it
for the first time, might ascribe it to a mechanical sound made by some
insect, not suspecting it to be the song of a bird. Only after we have
become thoroughly familiar with the song do we grasp its definite
character, so that we can pick it out even when we hear it in the
distance among a medley of other voices. In this particular it resembles
the songs of Henslow's and the grasshopper sparrows, which are scarcely
audible, and pass unregarded until well known.

The male goldenwing sings generally from a high perch, often from a
branch bare of leaves; hence, once we find him, we can see him plainly.
When he sings he throws his head back so far that his bill points almost
to the zenith, and sings with it widely open, as if he were pouring out
a great volume of sound. The bird sings freely from his arrival in
spring until mid-June, about a month, often devoting himself to long
periods of singing from the same perch. Later in the season, after the
young have hatched, he sings only fitfully.

The song most often heard is composed of four notes, the first
prolonged, and followed, after an almost imperceptible pause, by three
shorter notes on a lower pitch. All four notes are delivered in a
leisurely manner, drawling in tempo, and might be written _zeee,
zer-zer-zer_. The first note takes up about half the time of the song.
The quality of the voice is buzzing, and when heard near at hand,
slightly rasping, with a lisping suggestion throughout. The song carries
well; curiously it seems little louder when heard at close range, but
from a distance it sounds smoother and, losing much of the buzzing
quality, suggests a long drawn out _thth_, _th-th-th_, like a whispering
wind. Occasionally there may be four short notes, and sometimes only two
following the long initial note.

Like some of the other warblers, notably the black-and-white,
chestnut-sided, and black-throated green, the goldenwing sings two
distinct songs. In the second form the buzzing tone is nearly or wholly
absent. It begins with about half a dozen short notes given in a quick
series on the same pitch, and ends with one long note on a higher key,
_th-th-th-th-th-th-theee_.

I have heard two males singing antiphonally, the responses repeated with
perfect regularity for several minutes.

Of the minor notes the commonest is a short, slightly roughened _dz_.
When much excited both adults use a chattering _tchu-tchu-tchu_,
suggesting in manner of delivery the song of the short-billed marsh
wren, although it is higher pitched and not so loud.

Francis H. Allen (MS.) mentions two other songs, only slightly different
from the above. One goes something like "_tick tick chick chick chick
chick shree_. The _shree_ is a beady note resembling one of the
cedar waxwing's familiar notes." Another song he writes as
"_see-see-see-see-see-see-see-see-see-dz'-dsee_."

_Field marks._--The golden-winged warbler is easy to recognize; it is
the only warbler that combines a blue-gray back and yellow in the wing.
In the two other common warblers with a black throat, the black-throated
blue and the black-throated green, the black runs down the sides a
little way so that the white of the breast comes up in a peak in the
middle of the breast, whereas in the goldenwing the line of division
between the black and white runs straight across. From directly below,
the goldenwing appears wholly black and white, and from this angle is
marked like a chickadee, but a glance at its long, needle-sharp bill
proclaims it a warbler of the genus _Vermivora_.

_Enemies._--Prowling mammals, the enemies of ground-nesting birds, and
predatory hawks are a danger to the bird. In its relation to the
cowbird, Friedmann (1929) reports the bird as "a very uncommon victim."
He says: "I have only six definite records, but the species is listed
as a molothrine victim by Bendire and by Short. As many as four eggs
of the Cowbird have been found in a single nest of this Warbler."

_Fall_ and _winter_.--We lose sight of the goldenwing early in the
season. Silent amid the dense foliage of July and August, the bird is
rarely seen. During the years between 1907 and 1920, when I kept a daily
record of birds seen, I met it only four times in August and only twice
in September, the latest September 12.

Dr. Alexander F. Skutch sends to A. C. Bent the following account of the
bird in its winter quarters: "I am familiar with the golden-winged
warbler in its winter home only in Costa Rica. In this country it
winters on the Caribbean slope from the lowlands up to about 6,000 feet
above sea-level, and on the Pacific slope at least in the region between
2,000 and 4,000 feet. While it appears to be nowhere abundant, I found
it most numerous at Vara Blanca, on the northern slope of the Cordillera
Central at an elevation of about 5,500 feet. Here on one day--November
2, 1937--I saw three individuals, the greatest number I have ever
recorded. This is a region of dense vegetation, subject to much
cloudiness and long-continued, often violent rainstorms--one of the
wettest districts of all Central America. Most of the published records
are from this generally wet side of the country. Yet the bird winters
sparingly in the Basin of El General on the Pacific slope, which during
the first 3 months of the year may be nearly rainless. While in the
Tropics, it appears never to associate with others of its own kind, but
at times may roam about with mixed flocks of other small birds. It may
forage among low, fairly dense, second-growth thickets, or among the
tangled vegetation at the forest's edge, or at times in the forest
itself, or in groves of tall trees, high above the ground. It
investigates the curled dead leaves caught up among the branches, and
devours such small creatures as it finds lurking in their folds. I have
not heard it sing while in its winter home.

"In Costa Rica, it appears to arrive late and to depart early, not
having been recorded before September 15, nor later than April 9. Early
dates of fall arrival are: Costa Rica--San José (Cherrie), September 15
and October 2; La Hondura (Carriker), September 21; Basin of El General,
October 18, 1936; Vara Blanca, October 5, 1937.

"Late dates of spring departure are: Costa Rica--Basin of El General,
April 8, 1936, April 7, 1937, March 30, 1939, and April 9, 1943; Vara
Blanca, April 9, 1938; Guápiles (Carriker), March 30."


DISTRIBUTION

_Range._--Eastern United States to northwestern South America.

_Breeding range._--The golden-winged warbler breeds =north= to central
Minnesota (Detroit Lakes, Onamia, and Cambridge); central Wisconsin (St.
Croix Falls, New London, and Shiocton); northern peninsula or Michigan
(McMillan and Mackinac Island); southern Michigan (Kalamazoo, Locke, and
Detroit); southern Ontario (London and Port Rowan, has occurred north to
Collingwood and Bowmanville); central New York (Medina, Rochester, and
Waterford); central Vermont (Rutland), and northern Massachusetts
(Winchendon, Newton, and Lynn). It has been found in summer and may
possibly breed in southern New Hampshire (Concord and Durham); and
southwestern Maine (Emery Mills and Sandford). =East= to eastern
Massachusetts (Lynn, Boston, and Rehoboth); southern Connecticut (New
Haven and Bridgeport); northern New Jersey (Morristown); central
Pennsylvania (near State College); and south through the mountains to
western North Carolina (Weaverville, Waynesville, and Highlands);
northwestern South Carolina (Caesars Head and Highlow Gap); and northern
Georgia (Young Harris, Margret, and Oglethorpe Mountain). =South= to
northern Georgia (Oglethorpe Mountain and Rising Faun); central
Tennessee (Maryland); northern Ohio (Steuben, Port Clinton, and
Wauseon); northern Indiana (Waterloo); and northern Illinois
(Riverside). =West= to northern Illinois (Riverside); central and
western Wisconsin (Baraboo Bluffs and Durand); and central Minnesota
(Minneapolis, Elk River, and Detroit Lakes). It has been noted in
summer, or in migration, west to St. Louis, Mo.; Lake Quivira and
Lawrence, Kans.; and Omaha, Nebr.

_Winter range._--In winter the golden-winged warbler is found =north= to
central Guatemala (Cobán); and northern Honduras (Lancetilla); casually
or in migration to the Yucatán Peninsula (Campeche and Mérida). =East=
to Honduras (Lancetilla); eastern Nicaragua (Escondido River); Costa
Rica (Guápiles and Guayabo); central Panamá (Lion Hill, Canal Zone); and
central Colombia (Santa Marta region, Bogotá, and Villavicencio); rare
or accidental in western Venezuela (Mérida). =South= to central Colombia
(Villavicencio and El Eden). =West= to northwestern Colombia (El Eden,
Medellín, and Antioquia); western Panamá (Chiriquí); Costa Rica (El
General and Nicoya); and central Guatemala (Cobán).

_Migration._--Late dates of spring departure are: Colombia--Fusagasugá,
March 24. Panamá--Volcán de Chiriquí, April 16. Costa Rica--Vara Blanca,
April 9. Florida--Pensacola, April 22. Alabama--Hollins, May 7.
Georgia--Athens, May 13. South Carolina--Clemson College, May 3. North
Carolina--Raleigh, May 7. District of Columbia--Washington, May 20.
Mississippi--Gulfport, April 18. Missouri--St. Louis, May 25.

Early dates of spring arrival are: Florida--Pensacola, April 5.
Alabama--Barachias, April 22. Georgia--Milledgeville, April 12. South
Carolina--Clemson College, April 21. North Carolina--Asheville, April
23. Virginia--Lynchburg, April 19. West Virginia--Bluefield, April 19.
District of Columbia--Washington, April 24. Pennsylvania--Beaver, April
24. New York--Rochester, April 29. Massachusetts--Belmont, April 28.
Louisiana--Grand Isle, April 6. Mississippi--Gulfport, April 10.
Tennessee--Memphis, April 12. Illinois--Olney, April 17. Indiana--Sedan,
April 27. Michigan--Plymouth, April 30. Ohio--Youngstown, April 27.
Ontario--London, April 30. Missouri--St. Louis, April 18. Iowa--Keokuk,
April 27. Wisconsin--Sheboygan, April 30. Minnesota--Minneapolis, April
30. The golden-winged warbler ranges west to central Iowa in migration,
and in the lower Mississippi Valley is much less abundant in spring than
in fall.

Late dates of fall departure are: Minnesota--Minneapolis, September 30.
Wisconsin--Madison, October 11. Ontario--Point Pelee, September 2.
Ohio--Ellsworth Station, September 23. Michigan--Ann Arbor, October 6.
Indiana--Lyons, September 27. Illinois--Chicago, October 7.
Missouri--La Grange, September 30. Kentucky--Versailles, September 25.
Tennessee--Athens, September 29. Louisiana--New Orleans, September 25.
Mississippi--Gulfport, October 8. Massachusetts--Danvers, September 7.
New York--Brooklyn, October 2. Pennsylvania--Jeffersonville, October 2.
District of Columbia--Washington, September 14. West Virginia--French
Creek, September 15. North Carolina--Piney Creek, October 3. South
Carolina--Chester, September 22. Georgia--Atlanta, October 9.
Alabama--Greensboro, October 4.

Early dates of fall arrival are: Mississippi--Bay St. Louis, July 23.
District of Columbia--Washington, August 8. Virginia--Naruna, August 23.
North Carolina--Highlands, August 15. South Carolina--Charleston, August
20. Georgia--Athens, August 14. Alabama--Greensboro, August 11.
Florida--Pensacola, August 14. Costa Rica--San José, September 15.
Colombia--Bonda, September 6.

_Casual record._--One reported seen at Fort Thorn, N. Mex., in April
1854 by Dr. Joseph Henry. Since no specimen was taken this remains on
the hypothetical list for the State.

_Egg dates._--Massachusetts: 14 records, May 27 to June 24; 9 records,
May 30 to June 7, indicating the height of the season.

Michigan: 33 records, May 13 to June 10; 18 records, May 17 to 30.

New York: 6 records, June 3 to 24.

New Jersey: 7 records, May 25 to June 5 (Harris).




VERMIVORA PINUS (Linnaeus)

BLUE-WINGED WARBLER


PLATES 12, 13


HABITS

Bagg and Eliot (1937) write: "According to Wilson, this species was
discovered by William Bartram, who gave it the descriptive name _Parus
aureus alis caeruleis_ (Blue-winged Golden Tit), and sent a specimen to
'Mr. Edwards' by whom it was drawn and etched. Edwards suspected its
identity with the Pine Creeper of Catesby: hence its present
inappropriate name, _pinus_." As there are other warblers whose wings
are more distinctly blue, those of this warbler being only bluish gray,
the old familiar name, blue-winged yellow warbler, which stood for
many years, seems more appropriate and more truly descriptive.

The blue-winged warbler is a bird of the so-called Carolinian Life Zone,
with a rather restricted breeding range in the Central States and not
quite reaching our northern borders. Its center of abundance in the
breeding season seems to be in southern Ohio, Indiana, Illinois,
northern Kentucky, northern Missouri, and southern Iowa. Its range
extends northeastward to New Jersey, southeastern New York, and southern
Connecticut. It is fairly common in the latter State, and I know of one
small colony in eastern Rhode Island within a mile or two of the
Massachusetts line. North of these points in New England it occurs only
as a straggler or casual breeder. In southern New England I have found
it in rather open situations, in neglected pastures where there is low
shrubbery, brier patches, and bushy thickets around the edges; or in
similar growth along the borders of woods, usually on dry uplands; and
sometimes in the rank growth of tall grasses and weeds near the borders
of swamps or streams.

Frank L. Burns wrote to Dr. Chapman (1907) of its haunts in
Pennsylvania: "This species is here an inhabitant of the rather open
swampy thickets, upland clearings, neglected pastures and fence rows,
where the grass and weeds have not been choked out by a too thick growth
of briers, bushes, saplings and vines." Dr. Lawrence H. Wilkinshaw tells
me that, in southern Michigan, "this species loves deep swampy woods,
where the golden-winged warbler and cerulean warbler are found." This is
quite different from the haunts in which we find it in the east, though
Dr. Chapman (1907) says: "It is not, as a rule, a deep woods warbler,
though I have found it nesting in heavy forest, but prefers rather,
bordering second growths, with weedy openings, from which it may follow
lines or patches of trees to haunts some distance from the woods."

_Spring._--From its winter home in Central America the blue-winged
warbler seems to migrate from Yucatán straight across the Gulf of Mexico
to the Gulf States and along the eastern coast of Texas to Louisiana. It
is apparently very rare anywhere in Florida or the Keys, and along the
Atlantic coast, where it is comparatively rare, it is found at low
elevations. It migrates northward mainly west of the Alleghenies,
seeming to avoid the mountains; the main body of the species seems to
travel through the Mississippi Valley to the centers of abundance in the
central States. Perhaps the birds that settle in southern New York and
New England travel up the Ohio River, drifting through Pennsylvania and
New Jersey to their destination. According to Milton B. Trautman (1940)
this warbler seems to be a rare or uncommon spring migrant in central
Ohio, and "in some migrations only 2 individuals were noted" at Buckeye
Lake; this adds support to the theory that the birds follow the river
along the southern border of the State.

_Nesting._--Although Wilson (1831) gave a very good description of the
nest of the blue-winged yellow warbler, very little was known of its
nesting in southern New England prior to about 1880, when nests were
found in southern Connecticut, where it is now known to be a fairly
common breeder. I found two nests near West Haven, Conn., on June 8,
1910; both were close to the ground but not quite on it; one was in a
clump of blackberry vines, weeds, and grasses, in a swampy corner of a
scrubby lot; the other was in a bunch of grass and rank weeds on some
sprout land among some mixed bushes. Again, on June 1, 1934, I
photographed (pl. 12) a nest near Hadlyme, Conn., on the edge of an
open, neglected field and close to the border of some young woods. It
was built among and attached to the upright stems of a clump of tall
goldenrod. These were all typical of the nests described below.

Massachusetts nests are very rare; Forbush (1929) gives but two nesting
records for this State, and only one for Rhode Island, though I am
confident that its breeds regularly in the latter State. Horace W.
Wright (1909b) gives a very full account of a nest found near Sudbury,
Mass., in some mixed woods, placed between the exposed roots of a
decayed stump and partially concealed by a growth of ferns.

T. E. McMullen has sent me the data for several Pennsylvania nests,
three in old fields, one under a cherry sprout, one under a small bush,
and one 6 inches up in a tussock of goldenrod; another was under a birch
sprout along the edge of an old woods road.

The nest of the blue-winged warbler is unique and quite distinctive,
often shaped like an inverted cone, usually very narrow and very deep
and supported by a firm cup of strong, dead leaves. I cannot improve on
the excellent description given to Dr. Chapman (1907) by Frank L. Burns
as follows:

Outwardly composed of the broad blades of a coarse grass, the dead
leaves of the maple, beech, chestnut, cherry and oak trees; the leaf
points curving upward and inward forming a deep cuplike nest in which
the bird's head and tail seem almost to meet over her back. Occasionally
grass stems, coarse strips or wild grapevine bark, shreds of corn
fodder, and fragments of beech and wild cherry bark appear in the
make-up. Lined most frequently with wild grapevine bark laid across,
instead of bent around in a circle, shredded finest on top, to which is
added an occasional long black horse-hair or split grass stem, with now
and then a final lining of split grass stems in place of fine bark. The
shape varies in accordance to situation, outwardly a short cornucopia, a
round basket, and once a wall-pocket affair, would best describe the
shapes I have noticed.

_Eggs._--From 4 to 7 eggs may be found in the nest of the blue-winged
warbler; 5 seems to be the commonest number, and sets of 6 are not very
rare. The eggs are ovate, with a tendency to short ovate, and they have
only a slight gloss. The white ground color is finely speckled or
sparingly spotted with "chestnut brown," "mummy brown," and "sayal
brown," with under markings in shades of "drab-gray." Some sets have
three or four eggs that are almost immaculate, with one egg sparingly
spotted; other sets occasionally are prominently spotted with
"drab-gray," "light Quaker drab," and "dark vinaceous-drab," or, less
often, with spots of dark "mummy brown." Usually the majority of the
markings are confined to the large end. The measurements of 50 eggs
average 15.7 by 12.5 millimeters; the eggs showing the four extremes
measure =16.8= by =13.0= and =14.2= by =11.6= millimeters (Harris).

_Young._--An egg is laid each day until the set is complete, and
incubation generally begins when the last egg is laid. The period of
incubation is 10 or 11 days, and the young remain in the nest from 8 to
10 days. Mr. Burns gave Dr. Chapman (1907) the following full account of
the nest life:

The task of incubation falls on the female alone. It appears that an
airing is taken in the early morning or a little before midday, and
again in the early evening, though perhaps not regularly every day. I
have not seen the male about the nest with food at this period. The
female will allow a close approach, looking into one's eyes with that
hunted look so common in wild animals, and often flushing without a
protesting note. The period of incubation in the one instance was
exactly ten days.

On June 13, at 6.30 p. m., five young just hatched were blind, naked and
prostrate from chin to sternum. The shells were disposed of immediately,
in what manner I am unable to state; the female was reluctant to vacate.

On June 15, at 2.45 p. m., the young were able to raise their heads
slightly and a fluffy bit of down had appeared about the head, also a
dark stripe along the back bone. The female appeared, accompanied by the
male, and fed the young with small green larvae--such as may be found
on the under-side of oak and chestnut leaves--and then shielded the
callow young from the hot rays of the sun.

On June 16, at 6.30 p. m., when the young were three days old, a downy
puff appeared between the shoulders, wing quills being dark. The
strongest bird had the eyes partly open and the mouth wide open for
food.

On June 18, at 7 p. m., the heads and bodies were no longer
flesh-colored but were well enough covered to appear dark. The eyes were
open. At a _cluck_ from me their mouths flew open. Both parents fed them
with green-colored larvae. When the male rested a moment on a brier
above the nest, the female flew down and drove him away, fed the young,
re-appearing with excrement in her beak, which was carried in an
opposite direction from the regular approach via maple bough and poplar
sapling. The male fed the young from a mouthful of very minute larvae or
eggs, which were gathered from the silken nests in the unfolding leaves
of a nearby poplar; after this (7.30 p. m.) the female covered the young
for the night.

On June 20, at from 6.50 to 7.35 p. m., the young had been seven days in
the nest. They were well feathered and of a yellowish-green cast, the
short tails being tipped with yellow. The parents were more suspicious.
The female came to the maple bough with something in her beak and flew
down to the briers and back again several times before she dropped to
the edge of the nest and fed her young. The male appeared immediately
but swallowed a green grub himself upon discovery of me twenty-five feet
away. The female came again in five minutes with a brownish object in
her bill, but appeared more timid and refused to drop to the nest until
the male set her an example of courage.

On June 21, at 6.12 p. m., the young were fully fledged in green plumage
above and dirty yellow beneath. They showed fear of me for the first
time, eyeing me in the same manner as the parent bird when on the nest.
They were evidently ready to vacate at a moment's notice or hasty
movement on my part. The parents appeared, scolding rapidly. The female
fed the young as soon as I retired to my old stand under a bush, with a
rather large green grub (6.20 p. m.) and flew out to the top of a
blackberry bush, followed immediately by the topmost fledgling. It could
do little more than run. The adults flew to within a yard of my head,
making a great outcry, and in the midst of the excitement the remainder
of the young vacated the nest with feeble _chips_. The male gave his
attention to them, while the female followed me as I beat a hasty
retreat to enable them to collect their little family before dark. Eight
days had elapsed since incubation was completed, and it is not at all
unusual for the young of this species to leave the nest while so tiny
and ragged.

_Plumages._--Dr. Dwight (1900) calls the natal down "mouse-gray," and
describes the juvenal plumage, in which the sexes are alike, as, "entire
body plumage olive-yellow darkest on the back and throat. Wings and tail
slate-gray largely edged with plumbeous gray, the tertiaries and coverts
with olive-yellow; the greater and median coverts tipped with white,
yellow tinged. Rectrices largely white. Lores dusky."

A partial postjuvenal molt begins early in July, involving the contour
plumage and the wing coverts, but not the rest of the wings or the tail.
This molt produces the first winter plumage in which the sexes are very
much alike, the female being duller in color, especially the streak
through the eye, and having less yellow on the crown. Dr. Dwight (1900)
describes the first winter male as "above, bright olive-green,
lemon-yellow on the crown veiled by greenish tips. Below, bright
lemon-yellow, the crissum white or merely tinged with yellow.
Transocular streak black. Wing coverts plumbeous gray, edged with
olive-green, the greater and median tipped with white, yellow tinged,
forming two broad wing-bands."

The birds are now practically adult in plumage. The first and subsequent
nuptial plumages are acquired by wear, which produces little change
beyond removal of the greenish tips. Subsequent winter plumages are
acquired by a complete postnuptial molt each July.

The interesting hybrids between this and the golden-winged warbler are
discussed on pages 3 and 4. Kumlien and Hollister (1903) mention a
probable mating of this with the Nashville warbler.

_Food._--Nothing seems to have been published on the food of the
blue-winged warbler beyond that mentioned above as food given to the
young, which is doubtless eaten by the adults as well. It is apparently
wholly insectivorous, seeking its food near the ground in the weed
patches and underbrush where it lives and among the lower branches of
the trees in its haunts. Probably any small insects that it can find in
such places, as well as their larvae and eggs, including many small
caterpillars, are eaten. Small grasshoppers and spiders are probably
included. Prof. Aughey (1878) observed it catching small locusts in
Nebraska. It is evidently a harmless and a very useful bird in
destroying insects that are injurious to foliage.

_Behavior._--Dr. Chapman (1907) writes: "It is rather deliberate in
movements for a Warbler, and is less of a flutterer than the average
member of the genus _Dendroica_. Some of its motions suggest those of
the tree-inhabiting Vireos, while at times, as the bird hangs downward
from some cocoon it is investigating, one is reminded of a Chickadee."
And he quotes Burns as follows:

Perched inconspicuously near the top and well out on the branchlets of a
tree or sapling, preferably facing an opening, if in a thicket; it is in
itself so minute an object as to be passed unseen by many, more
especially as it is much less active than most of our Warblers. With
body feathers puffed out to a delightful plumpness, except for the
backward sweep of the head while in the act of singing, it remains
motionless for quite a while. When it moves it is with a combination of
nervous haste and deliberation, and its song may be heard from quite
another part of the landscape with no apparent reason for the change.
While it has its favorite song perches, it is quite a wanderer and not
infrequently sings beyond possible hearing of its brooding mate, but
oftener within fifty to two hundred feet of the nest.

_Voice._--Aretas A. Saunders contributes the following study of the
songs of this warbler: "The territory song of the blue-winged warbler
consists of two long, buzz-like notes, the second usually lower in pitch
than the first and rougher in sound, _bzzzzzzz-brrrrrrrr_. The pitch
interval between the two notes varies from one tone to four and a half
tones, but the smaller intervals, one tone and one and a half tones, are
much commoner. The second note is lower in pitch than the first in about
75 percent of my records, and higher in most of the others. In a few
songs the second buzz is a double note, and one may hear both lower and
higher notes from a medium distance, only the lower from a greater
distance, and only the higher when very near the bird.

"The pitch is not high as compared to other warblers, ranging from C´´´
to D´´´´, one tone more than an octave. The territory song commonly
begins on some note from A´´´ to C´´´´. It varies in time from 1-1/5
to 1-4/5 seconds, the first note being either equal to or shorter than
the second. The second note is often twice as long as the first. In some
songs the second note is broken into two notes, and in one record it is
in four short notes, so that the song is essentially like that of the
golden-winged warbler.

"After the birds have been on the breeding grounds for a week or two,
singing of the nesting song begins. This song has the same buzz-like
quality as the other, but it is exceedingly variable, considerably
longer, and hardly ever twice alike. The song often begins with a series
of short notes, like _tsit tsit tsit_, or contains such notes somewhere
in the middle. There are usually long buzzes that change pitch by
slurring upward or downward. On one occasion, I found a bird that sang a
territory song and four different nesting songs. Often the nesting song
is sung in flight. By June this song is heard about as frequently as the
territory song, and in late summer, after the molt, it is the one most
commonly heard.

"The song of this bird is heard from its arrival in spring until early
July, when it ceases for a time. It is usually revived in late July or
early August, and from then on may be heard fairly frequently until the
birds depart about the last of August."

In his notes sent to Dr. Chapman, Burns describes the song as, "a
drowsy, locust-like, _swe-e-e-e-e ze-e-e-e-e_, the first apparently
inhaled and the last exhaled. * * * Another song heard on the first day
of arrival, on one occasion, uttered by several males in company,
possibly transients here, and may be the mating song, suggests the
Chickadee's _che-de-de-e_, _che-dee-e_, and _che-de-de-dee_

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