Part 39

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:

The Golden-cheek is not a bird of the forest, being seldom met with in the tall timbered areas in the wilder valleys along the rivers, or in the tall trees which fringe the streams in the cañons; but its favorite haunts are among the smaller growth of trees, on the rough wooded hillsides, and which covers the slopes and "points" leading up from the cañons, and the boulder strewn ridges or "divides" which separate the heads of the creeks. The trees which compose this growth consist chiefly of mountain cedar (juniper), Spanish or mountain oak, black oak, and live oak on the higher ground, and live oak and Spanish oak clumps or thickets on the lower flats among the foothills, interspersed in some localities with dwarf walnut, pecan and hackberry. All these trees grow on an average from 10 to 20 feet high, the cedar often forming almost impenetrable "brakes". Whatever space remains among the oaks and cedars is generally covered with shin oak brush, which is a characteristic feature of the region. The cedar or juniper appears to possess some peculiar attraction for this bird for they are seldom found at any great distance from cedar localities, and they seem to divide the greater part of their time between the cedars and Spanish oaks, searching for Insects, with occasional visits to other oaks, walnuts, etc., but seldom descending as low as the shin oak brush, which averages four to five feet. It is quite probable that future observations will show, that some favorite insect food which comprises a portion of their "bill of fare," is found among the cedar foliage.

_Spring._--The golden-cheeked warblers arrive in central Texas about the middle of March, sometimes a little earlier or later. The adult males precede the young males and females by about 5 days. Mr. Attwater (Chapman, 1907) says: "The song of the male is the first unmistakable notification of its arrival and within a few days it is quite common and the females are also observed. In the localities described the Golden-cheeked Warbler is by no means a rare bird, and it is by far the most abundant of the few Warblers, which breed in the same region."