_Egg dates._--Massachusetts: 26 records, May 21 to July 11; 15 records,

May 30 to June 10, indicating the height of the season.

New Brunswick: 13 records, June 13 to 28; 9 records, June 5 to 19.

New York: 19 records, May 30 to July 16; 10 records, June 2 to 11.

Nova Scotia: 13 records, June 7 to 28; 9 records, June 13 to 20

(Harris).

DENDROICA VIRENS WAYNEI (Bangs)

WAYNE'S BLACK-THROATED GREEN WARBLER

CONTRIBUTED BY ALEXANDER SPRUNT, JR.

HABITS

It was a silent world, this great cypress swamp where I sought the nest

of the Wayne's black-throated green warbler in the company of the man

whose name it bears and who first made it known to science. A vast

flooded expanse of trees and water--colorful, eerie, and mysterious--it

was a realm of gray-green gloom. Huge trunks towered on all sides; long

aisles of wine-dark mirror-smooth water stretched illimitably away among

the buttressed columns. The grayness that predominated, from the

furrowed knees and smoother trunks of the great trees to the shrouds of

moss festooned from their branches, was relieved here and there by

contrasting splotches of bright green overhead where occasional shafts

of brilliant sunlight penetrated the canopy of feathery foliage.

Our dugout made no sound as it slid along. Only the slight splash of the

paddle entering and leaving the water gave evidence of any means of

propulsion. Now and again the silence was broken by the calls echoing

down the flooded aisles--the clear whistle of the prothonotary warbler

ringing sweetly, the full-voiced carol of the yellow-throated warbler,

the strident call of the pileated woodpecker answered by the distant cry

of a hunting red-shouldered hawk. Occasionally the deep, resonant

"whoo-aw" of a barred owl reverberated solemnly among the cypresses, and

once a sombre anhinga flapped ahead of the dugout to plunge cleanly into

the still water in full career.

But above these evidences of swamp life, above the swish of breaking

bass, the crashing splash of a disturbed alligator, the clamor of a

startled heron or ibis, sounded one persistent call from the high

branches--a song of seven notes, five on the same tone, one ascending,

the last descending. It was this call that drew us on, the song of the

bird whose nest we sought that morning, Wayne's warbler, the southern

race of the black-throated green warbler.

To find the black-throated green warbler in a cypress swamp might seem

strange indeed to one who knows the species in its spruce and balsam

highlands, in the rhododendron and laurel thickets of the Blue Ridge, in

the evergreens of the Adirondacks and Maine! Yet here it is, one of the

characteristic avian dwellers of the warm swamplands of the South

Carolina Low Country, arriving in the spring to nest amid the green

cypress twigs, the drooping limbs of the magnolias, and the majestic

spread of the live oak.

When Arthur T. Wayne of Mount Pleasant, S. C., discovered the first

nesting of this race he was sure it was not a typical _Dendroica virens

virens_, and on April 25, 1918 he sent a male to Outram Bangs of the

Museum of Comparative Zoology in Cambridge, Mass. Later he sent him six

other specimens. Upon comparing them with specimens of _D. v. virens_,

Bangs (1918) described it as a different race, giving it the name of the

discoverer, _waynei_. Extracts from his published material are

illuminating. He states, for instance, that "this series proves to

represent a form easily distinguishable from true _Dendroica virens_

(Gmelin). I take great pleasure in naming it after the keen

ornithologist and excellent observer and collector who discovered it,

and who noticed its peculiarities even without sufficient material with

which to compare it."

The subspecific differences are mainly a duller coloration, less

yellowish, and of a paler shade, and the throat patch more restricted.

Its principal variation from _virens_ is its much smaller and more

delicate bill. As Bangs points out, "measurements of a bill so small do

not convey the same impression that an actual comparison of specimens

does. The bill of the new form when compared to that of _D. v. virens_,

appears not more than two-thirds as large." Certainly this is true. So

marked is the difference that a specimen of _waynei_ placed amid a score

or more of _virens_ can easily be picked out even at some distance.

The southern limit of the breeding range of _virens_ appears to be the

high mountains of Carolina and Georgia and northern Alabama, usually at

elevations of more than 4,000 feet; _waynei_ is confined to a coastal

strip (in some cases less than 5 miles from the ocean) so that the

intervening area between it and _virens_ averages about 300 miles. In

all that distance, no northern black-throated green warbler appears

except in scattered and isolated instances. The migratory route of

_waynei_ is as yet imperfectly known, but since _virens_ is so scarce

along the lower Atlantic coast as to be virtually absent, and since it

has never yet been secured or reported along the Carolina and Georgia or

northern Florida coasts, it would seem that any specimen seen in those

localities would be _waynei_.

_Spring._--Wayne (1910) said of this bird in South Carolina: "This

species arrives with great regularity [Charleston County] as the

following dates will show, viz., March 26, 1890; March 27, 1900; March

27, 1912; March 23, 1916. It is not common until the middle of April and

its passage through the coast region requires so long a time that one

not acquainted with the migrations of birds might readily believe that

it bred here ... that this species should remain on the coast until

June, and not breed is very surprising."

At that time he was, of course, unaware that the species contained two

races, but, as Outram Bangs has pointed out (1918), these March arrival

dates in coastal South Carolina occur when "true _D. virens_ is still in

winter quarters in Mexico and Central America." Thus, it will be seen

that the migration times must vary considerably, and the arrival of the

coastal race is in advance of the true species, indicating a different

and less distant winter home, another phase to consider when comparing

the two.

There is almost a complete dearth of additional information on arrival

dates in other southern states. My records of South Carolina arrival

dates in recent years do not vary much from Wayne's, and he has no

earlier ones. I have but once encountered _waynei_ in spring elsewhere

than in South Carolina, this being a specimen observed in full song in

Rhetta Lagoon, Cumberland Island, Ga., on April 15, 1932. However, that

it was, in fact, a migrant is beyond all question for it is not present

in its United States range in late fall and winter.

In his description of the race in 1918, Bangs stated that "it would seem

not unlikely that the South Carolina form is resident and non-migratory,

and I hope Mr. Wayne will be able to prove whether or not this is so."

This belief of Bangs' was carried into the A. O. U. Check-List (1931)

which gives the range of this form as "resident in the coastal district

of South Carolina." This is not the case; _waynei_ does not remain in

winter, and is therefore not resident but migratory, as I have

previously pointed out (1932).

The migration of this race is as yet imperfectly known. While any

coastal migrant black-throated green warbler would probably be referable

to it, as _virens_ appears to keep to the interior when travelling

north, as a matter of fact there are almost no records of migratory

occurrences. S. A. Grimes (MS.) tells me that he has never observed any

black-throated green warbler in the area about Jacksonville, Fla., where

he lives and where he is much afield, having had years of experience.

Earle K. Greene (MS.) similarly states that his experience of over two

years in the Okefenokee Swamp in southern Georgia failed to produce "a

single individual." Strictly in line with his observations are those of

Francis Harper (MS.) whose experience in the Okefenokee is even more

extensive than Greene's. He writes in response to my request, "I have

never found the slightest trace of the bird there." This is strange, as

the Okefenokee would seem to be typical habitat for the Wayne's Warbler,

but it evidently does not occur there.

_Courtship._--Nothing is known of the courtship behavior of this bird,

owing to the difficulty of observation, the very restricted range of the

bird, and the dearth of local observers.

_Nesting._--Wayne was under the impression that he was the discoverer of

the first known nest of this race, but search of the literature reveals

that he was in error, though the first nests found were not recognized

as those of _waynei_. Wayne secured the first eggs, and these still

appear to be the only ones in existence, as all other breeding records

deal with young birds. Authentic breeding information is exceedingly

scanty, and since this is the case, all of the instances are mentioned

herewith.

The first recorded breeding was in coastal North Carolina, and is

mentioned by Pearson and the Brimleys (1919). They included it under the

black-throated green warbler, as the species at that time had not been

divided into two races. One nest was found at La Grange, Lenoir County,

in June 1905 (Smithwick), and the other at Lake Ellis, Craven County,

June 1910. Adults were seen feeding very young birds.

Continually impressed with the birds' presence in coastal South Carolina

so late in spring, Wayne sought evidences of nesting and, on April 11,

1917, saw a female carrying nesting material in a large cypress swamp in

Charleston County, but could not locate the nest. On the twenty-eighth

of the same month he detected both a male and female in the same

procedure but again failed to find the nest. His (1918) comment on this

follows: "The brief account of this bird written in 'Birds of South

Carolina' is, in the main, correct. Although I had never found it

breeding when the book went to the press I was absolutely certain that

it really bred on the coast." A year later, on April 28, 1918, he saw

another female engaged in nest building, and again was unable to find

the nest. Those who knew Wayne's untiring energy in such work can

readily understand the extreme difficulty experienced in locating this

elusive bird's home. It was on this last date that he secured the type

specimen from which Bangs described the race. The following year finally

brought success. Wayne (1919) states:

"On March 20, 1919, I visited the place where the type specimen was

taken. * * * A few males were heard singing from the topmost branches of

tall, gigantic, deciduous trees, and were also seen to fly into very

tall pines." He again visited this spot on April eighteenth with Henry

Moessner and the latter located a nest. It "was built in a live oak

tree and on the end of a horizontal branch among twigs * * * absolutely

concealed * * * about 38 feet above the ground." Wayne climbed a nearby

tree and with Moessner's help from below, attempted to pull the oak limb

toward him in order to reach the nest, when "sad to relate, without a

moment's warning, the limb snapped off and the four fresh eggs that the

nest contained were dashed in fragments on the ground."

The nest itself was preserved, and Wayne describes it as "small and

compact, measuring 1-3/4 inches in height and 1-1/2 inches in depth. It

is constructed of strips of fine bark and weed stems, over which is

wound externally the black substance that is invariably present in the

lining of the nests of Bachman's Warbler (_Vermivora bachmanii_). The

interior * * * is chiefly composed of a beautiful ochraceous buff

substance, doubtless from the unfolding leaves of some fern, and a few

feathers."

On the twenty-eighth, ten days after this nest was found, Wayne returned

to the swamp with the Misses Louise Ford and Marion Pellew and found "a

very young bird just from the nest and unable to fly more than a few

feet, being fed by the male parent, which shows that the birds breed

irregularly."

The party proceeded to another part of the swamp where a female was seen

to enter a large magnolia. "Miss Ford * * * saw the female go to her

nest * * * built near the extremity of a long drooping magnolia limb,

but on the horizontal portion of it and about 25 feet above the ground."

This nest held four heavily incubated eggs, these being the first ones

actually taken. This nest had a quantity of caterpillar silk binding the

fibres of Spanish and hypnum moss outside, and was "lavishly lined with

the beautiful ochraceous buff substance from young fern leaves, as in

the first nest."

Edward S. Dingle (MS.) writes that "on the morning of April 25, 1923, a

Wayne's warbler was observed building in a cypress tree; the bird

collected material from the ground and also from the trunk of a large

cypress nearby. The male was not seen." On the third of May following, I

accompanied Wayne and Dingle to the site; there Dingle located the nest,

climbed the tree, and secured it, with four eggs. This nest was 62 feet

from the ground and 5 feet out from the trunk. This is the third, and

last nest from South Carolina with eggs, on which data are extant. All,

with the exception of the first, were in Wayne's collection at his

death, and are now in the Charleston (S. C.) Museum. The sites in each

case, were found by Wayne, but the nests were actually located by

Moessner, Dingle, and Miss Ford.

Commenting on these discoveries, Wayne (1919) states: "I have known this

bird ever since May 4, 1885 when I took a male in Caw-caw Swamp,

Colleton County, S. C., while on a collecting trip with my friend, the

late William Brewster. I gave the bird to him in the flesh, and in his

collection it still remains. The nest and eggs have remained unknown

until brought to light by this season's research."

Russell Richardson (1926) reported black-throated green warblers in the

Dismal Swamp, on the North Carolina side, in June. No evidence of

nesting was found by him, and he did not, apparently, realize that the

birds he saw were _waynei_. In 1932 Drs. W. R. McIlwaine and J. J.

Murray visited Dismal Swamp on May 23-26, and "found Wayne's warblers

rather common." From Murray (1932) we find that they "heard two singing

males on May 23rd as we came down the Washington Ditch to the Lake; two

males singing on the 24th near the entrance to the Feeder Ditch * * *

and six males on the 26th." They also found two family parties of adults

feeding small birds on the 24th. One of these parties was near the mouth

of the Feeder Ditch; the other a half mile up the Jericho Ditch from the

Lake (Drummond). * * * The young birds were out of the nest and could

fly well. They looked like big bumble-bees buzzing across from one tree

to another; staying rather high up. The adults ranged low in gathering

food, both male and female feeding the young birds.

_Eggs._--The eggs of _waynei_ are similar to those of _virens_. Wayne

has described them (1919) as "of a white or whitish color speckled and

spotted in the form of a wreath around the larger end with brownish red

and lilac." The sets previously described are the only ones of which the

writer is aware, and may be the only ones in collections. Whether any

have ever been secured outside of South Carolina is doubtful.

Measurements of Wayne's two sets average 16.79 by 12.25 and 15.12 by

12.03 mm., a trifle under the average for eggs of _virens_. The breeding

records for the Dismal Swamp (Virginia) and two localities in North

Carolina, concern young birds only.

_Plumages._--Data available are not sufficient for a detailed

description of the plumages but they are probably the same as those of

_virens_.

_Food._--No positive information on the food of _waynei_ exists, as far

as I can ascertain, except that in July 1939, G. H. Jensen examined the

stomach contents of a single specimen secured by Howell and Burleigh at

Murrells Inlet, S. C., June 5, 1932. It was full and contained 100

percent animal matter, consisting of 3 Lepidoptera larvae, 98 percent; 1

_Formica_ sp., 2 percent. That the race is insectivorous goes without

saying, but more than that remains to be worked out. Howell (1932) cites

Barrows as saying that _virens_ consumes plant lice, span-worms, and

leaf-rollers together with berries of poison ivy. Probably _waynei_

indulges similar tastes.

_Behavior._--Wayne's warbler is essentially a high-ranging bird. It

spends much of its time amid the topmost branches of cypress, magnolia,

gum, and other swamp trees, rarely descending to even mid-sections of

this characteristic growth while feeding. Highly restless and

exceedingly active in movements, it is constantly on the go and, as a

consequence, is rather difficult to see and study satisfactorily, the

oft-repeated song being the best indication of its whereabouts. As might

be supposed, the female is even more elusive, and flits about like some

swamp wraith, silent and mysterious. The failure of as keen an observer

as Wayne to locate the nests of building females gives an idea of its

secretiveness.

In these respects it differs materially from _virens_, at least in my

experience with that race, which is frequency found at rather low

elevations. Doubtless the type of growth is responsible, for _virens_ is

a spruce-balsam-hemlock dweller, and these evergreens are dense trees

with branches often beginning only a few feet from the ground, so that

it can be seen and watched rather easily.

While several authors have referred to _virens_ as a tame bird, the same

cannot be said for _waynei_. In years of experience with the latter, I

have always found it shy and retiring. Singing freely enough if unaware

of observation, it often ceases when it detects an intruder, and since

the song is one of the surest means of locating it, great care has to be

taken in moving about, particularly near the nest.

The nest is impossible to find without watching the female, for it is

more often than not completely invisible from the ground. _D. v. waynei_

is found in the same habitat with yellow-throated and parula warblers,

but, unlike them, never utilizes the hanging clumps of Spanish moss

(_Tillandsia_) in which they invariably nest. I have climbed a tall

cypress and collected a nest and eggs of _D. d. dominica_ while _waynei_

was singing in the near vicinity. The preference of _waynei_ for heavy,

old-growth swamp forests is so marked that if this timber is cut out,

the bird disappears from the area completely, even though other growth

is left standing. In the South Carolina Low Country, this characteristic

is shared by both Bachman's and Swainson's warblers, both of which nest

in heavy swamps.

_Voice._--Though it was the cuckoo which Bryant characterized as "a

wandering voice," he might well have written the words with respect to

this tiny warbler for the bird is heard far more readily than it is

seen. As a songster it is all but indefatigable. Perhaps this is because

the depths of the cypress swamps and the old "backwaters" are cooler

than the surrounding highlands, but no matter how warm the day, or close

the atmosphere, the constantly reiterated, seven-note song resounds

through the air most of the day. The ornithologists I have guided to the

haunts of _waynei_ all agree that the song is very close to that of

_virens_. Perhaps it is a shade more deliberate and studied, as might be

expected of a southerner! However, to all intents and purposes, it is

the same song. I am inclined to describe it as slower and more

pronounced, but after all the difference is minor.

Arthur H. Howell (1932) describes the song of _virens_ as "a drowsy,

drawled ditty of four or five notes, _wee-wee-wee-su-see_, the next to

last note on a lower pitch and the final one distinctly higher." This

portrays quite well the song of _waynei_, except for the number of

notes, which are much oftener seven than less, the first five being

exactly alike, the sixth descending, and the seventh ascending.

Frank M. Chapman (1907), quoting Gerald Thayer, says of _virens_ that

"most of the individuals in a region sing nearly alike ... but about one

in forty does queer tricks with its voice. Among the commonest of these

tricks is the introduction into all parts of the song of a pronounced

quaver or tremulo.... The song is sometimes disguised almost past

recognition." He states further that the "deliberate song of five

(sometimes six or eight) notes, is the one usually described in books."

I have never noted any "quaver or tremulo" in the song. It may occur,

but in the scores of times I have heard the song it has not taken place.

Nor can I recall any song of eight notes. Occasionally, _waynei_ will

utter only five notes, but this is the marked exception and not the

rule. Certainly, individuals in a given region sing exactly alike, and

indeed, all the specimens I ever heard sounded alike, except for the

occasional slight variation in number of notes.

_Fall._--The length of stay of _waynei_ in its summer range has not yet

been determined with certainty. Few departure dates have been recorded,

but in all probability the bird is a rather early migrant. Occurrence of

the song decreases markedly after the nesting season, making the birds'

movements much more difficult to trace. It will be recalled that young

were noted flying on May twenty-fourth, in the Dismal Swamp of Virginia.

South Carolina birds were seen to fly "a few feet" on April

twenty-eighth, almost a month earlier. The North Carolina records show

that young were noted "in June", probably early in the month. That

multiple broods are raised is also something of an open question, though

it seems that in South Carolina two are raised. Henry H. Kopman (1904)

states that on July 30, 1897, he took one at Beauvoir, Miss., on the

Gulf coast, and later comments (1905) that "Professor Cooke [W. W.] is

inclined to think" that the Beauvoir bird was a stray. Probably it was a

stray, and in view of what we know today, the chances are that the bird

was a specimen of _waynei_. Many of the birds of course linger much

later than that; on September 29, 1935, Earle R. Greene [MS.] noted one

at Lake Mattamuskeet, N.C. This is doubtless a rather late date and may

be taken as about the limit of its stay along the Atlantic coast.

Enough remains to be learned about this most interesting race to keep

students busy. The highly attractive type of habitat, the marked

isolation of nesting pairs even in a restricted range, the active

character and handsome appearance of the bird itself, all these combine

to render Wayne's warbler distinctive and appealing.

DENDROICA CHRYSOPARIA Sclater and Salvin

GOLDEN-CHEEKED WARBLER

HABITS

This elegant warbler is confined in the breeding season to a very narrow

range in south-central Texas, the timbered parts of the "Edwards

Plateau" region. It has been reported as breeding in Bandera, Bexar,

Comal, Concho, Kendall, Kerr, and Tom Green Counties, and rarely north

to Bosque and McLennan Counties. It winters in the highlands of southern

Mexico and Guatemala.

The golden-cheeked warbler was entirely unknown to early American

ornithologists. William Brewster (1879) gives the following brief

account of its early history: "The original specimens were procured by

Mr. Salvin in Vera Paz, Guatemala. Since that time, with the exception

of a male obtained by Mr. Dresser, near San Antonio, Texas, about 1864,

no additional ones have apparently been taken. The specimen mentioned by

Mr. Purdie was taken by George H. Ragsdale in Bosque County, Texas,

April, 1878." The bird is now well known in the limited region outlined

above, and many specimens of the birds, their nests, and their eggs have

found their way into collections.

The first comprehensive account of its habits was given to Dr. Chapman

(1907) by H. P. Attwater, of San Antonio, Tex. He says of its summer

haunts in the counties named above:

Life histories of North American wood warblers, Part 1 (of 2) · 第 37 / 46 段