[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_.
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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.