Contributed by Alfred Otto Gross
[Published in 1958: Smithsonian Institution United States National Museum Bulletin 211: 53-80]
The meadowlark is the outstanding and the most characteristic bird of the American farm. It is revered by the farmer not only because of its charming simplicity and its cheerful, spirited song, but also for its usefulness as a destroyer of harmful insects and the seeds of obnoxious weeds. The coming of the meadowlark in the early spring, while the fields are still brown, is a thrilling event. His arrival is made known by his plaintive but not complaining or melancholy song as he stands mounted atop some tall tree in a grassy meadow, with his bright yellow breast surmounted by a black crescent gleaming in the morning sun.
The meadowlark has the build and the walk, as well as the flight, of the quail; and since it frequents the marshes, especially in its winter quarters, it has sometimes been called the marsh quail. This name has probably lead many a hunter to think of it as a game bird. Fortunately in recent years fewer meadowlarks are killed for food, and this may be at least one factor responsible for the increasing numbers as well as the extension of its nesting range.
When I first came to Maine 35 years ago the meadowlark was a comparatively rare bird in the southern part of the state. Since that time it has steadily increased in numbers, until today almost every suitable meadow and grass field has its quota of meadowlarks. Similar increases in the number of meadowlarks have been reported from other sections of its range. Milton B. Trautman states in a letter that he counted 400 pairs of meadowlarks while walking through suitable fields, during the course of a few days in the Buckeye Lake region, Ohio. He estimated the amazing number of 1,400 pairs as nesting in the area, an average of 1 meadowlark for every 7 acres, or about 91 to the square mile.
In 1906 - 1908 I conducted the fieldwork of a statistical survey of the birds of Illinois for the Illinois Natural History Survey. In making the census counts, I walked many times through fields and woods over the length and breadth of the state. An assistant traveled at 30 yards distant and parallel to my line of march and was responsible for measuring the distance of each field traversed in terms of paces, which later were translated into feet. The species and the numbers of birds flushed in a strip 50 yards in width, including those flying across the strip within a hundred yards to our front, were recorded. Thus we covered all types of crops and vegetation during all conditions of weather and at all seasons of the year to obtain a comparative sample of the birdlife. During the summer months alone an area equivalent to 7,793 acres was covered, on which 85 species of birds were recorded. The meadowlark proved to be the most abundant of the native Illinois birds, being represented by 1,025 individuals, or 13.2 percent, of the total bird population. There was an average of 85 meadowlarks to the square mile for the whole area traversed. As the birds were unequally distributed, never occurring, for example, in woodlands or among shrubbery, their numbers rose to 266 to the square mile in stubble, 205 in meadows, 160 on untilled lands, 143 in pastures, and 131 on wastelands, but fell to 10 per square mile in fields of corn.
The meadowlark population varied in numbers from the northern to the southern part of the state, 100 in northern Illinois being represented by 175 in the central and by 215 in the southern part. The center of density of the summer meadowlark population at that time was in the southern section, and during the winter months the concentration of meadowlarks in southern Illinois reached an average of 373 per square mile. Many of the birds which nest further north winter in that section of the state.
From various reports I have recently received from the Middle West, it is probable that if the census were repeated today the average meadowlark population would exceed the average of 85 to the square mile obtained during the summer months of 40 years ago.
Spring.--The migration of the meadowlark is a comparatively limited movement, and the bird retires completely from only the most northern sections of its breeding range. It is a regular winter resident as far north as Maine, southern Ontario, and Michigan; and the southern summer residents do not go beyond the Carolinas, Alabama, Louisiana, and southeastern Texas. In spring the migrants reach Missouri and southern Illinois by the middle of March, arriving in the north central states during the first weeks of April, and in Minnesota and the Dakotas usually during the latter part of the month. The first arrivals in Manitoba, Saskatchewan, and Alberta appear in the last of April or the first week of May. The vanguard of the migrants reaches southern New England about the middle of March, and a marked movement extends well into April.
According to William Brewster (1886b), the meadowlarks are among the birds which migrate exclusively by night. He states: "Species which migrate exclusively by night habitually feed in or near the shelter of trees, bushes, rank herbage or grass, and when not migrating are birds of limited powers of flight and sedentary habits, restricting their daily excursions to the immediate vicinity of their chosen haunts. As a rule they are timid, or at least retiring disposition, and when alarmed or pursued seek safety in concealment rather than extended flights."
Meadowlarks migrate by night because they are either afraid to venture on long exposed journeys by daylight, or unable to continue these journeys day after day without losing much time in stopping to search for food. By taking the nights for traveling they can devote the days entirely to feeding and resting in their favorite haunts.
Milton B. Trautman (1940) in his observations at Buckeye Lake, Ohio, differs from the conclusions reached by Brewster. He writes:
Few Eastern meadowlarks were seen or heard migrating in very late evenings and early mornings, but many more were observed in the daylight hours. In late March and April individuals and loose flocks of as many as 60 flew northward at a low elevation across the lake. Loose flocks of 5 to 100 birds were often observed flying during spring and fall. The flocks generally flew a short distance and began to feed. Presently, those in the rear rose into the air and, flying over the flock, alighted in front to feed again. This maneuver was many times repeated. When the flock reached an obstruction, such as woods, cattail swamp, or lake, it flew over in a long loose column. The flocks traveled in this leisurely manner 2 to 6 miles an hour. Sometimes the flocks stopped feeding and flew 1 to 3 miles at a low elevation before dropping to the earth to feed again.
At Ithaca, N. Y., G. B. Saunders has found that the first meadowlarks to be seen early in the year are males. As early as January young birds which may be classed as vagrants are reported. Upon the advent of warmer weather more vagrants which have wintered only a short distance to the south wander in, feeding in manured fields and about farm buildings. Stormy weather often covers the fields with snow and sends them into barnyards where they may pick their food along with domesticated animals. Not infrequently many of these early meadowlarks perish in the long blizzards which put an end to their food supply.
Later, usually about the middle of March, the first migrants appear. These are old males, few in number and quiet in manner, which have wintered far to the south. By the end of March the migrant males become abundant. Song is less common among these early flocks of migrants than among the first resident males, which come a week or so later. When these arrive in the latter part of March they are active in the mornings and late afternoon, but during midday they often retire to a common feeding ground which the birds from different territories share without any apparent hostility. Their early song, although sweet and full of spirit, is not of the brilliance which characterizes it in April when the first resident females arrive.
Groups of migrants and resident males continue to arrive until the latter part of April. The last resident males establish themselves in areas left by the first wave of migrants, or carve their territories from the domains of earlier males unable to defend their original holdings. The migrants remaining in flocks resume their journey northward.
The vanguard of migrant females arrives 2 weeks or more after the first resident males and are followed closely by the first resident females. The coming of the resident females stimulates the first song peak of the males, whose songs become longer and more brilliant and animated. They engage in territorial combats over females, in defense displays and sexual flights, as well as posturing and sexual displays for the benefit of the female. The female is at a much lower sexual pitch at this time and responds only by preliminary stages of posturing such as the erection of the body to a vertical position, pointing her bill upward and twitching her wings. Late in April she reaches the necessary sexual level and begins building the nest and laying the eggs.
Other migrant and resident females, which are young birds, continue to come in April and May and even in June and July. These birds, which are late in maturing, become mates of polygamous or late arriving males, or remain unmated.
The following accounts of territory and courtship are based primarily on an exhaustive treatment of the subject contained in an unpublished thesis, submitted at Cornell University in 1932 by George Bradford Saunders ("A Taxonomic Revision of the Meadowlarks of the Genus Sturnella Vieillot and the Natural History of the Eastern Meadowlark Sturnella Magna Magna Linnaeus").
Territory.--Soon after his arrival during the latter part of March, the resident male leaves his companions and selects a territory, preferably a grassland or meadow, because of the great abundance of food as well as his decided liking for this type of habitat. The size and shape of the territory depend chiefly on the area of suitable land available, the local abundance and strength of competing males, the relative concentration of food supply, and certain barriers and individual range requirements of the male. The size of the territory may be increased as a result of polygamous relations, particularly if the females choose widely separated nesting sites, but the average size of 15 territories at Ithaca, N. Y., was found to be 7 acres. There is a decided difference between the total area of the territory and that which is regularly used. The more concentrated the food supply the less need there is for foraging, and the smaller the area frequented. Of two territories studied by G. B. Saunders throughout the breeding season of 1931, one contained 9 and the other 20 acres; but due to the abundance of food in a meadow separating the two families, one monogamous and the other with three females, this common feeding ground was shared. In each territory, however, the area of land used regularly was the same, 6 acres.
Important to the male are the various commanding perches from which he can survey his territory. Telephone wires or electric power lines with unobstructed views often furnish favorite song and lookout posts. Mounds of earth, farm implements, and fence posts provide perches near the ground. During the first few days of his occupancy he visits them from time to time and selects one for his primary headquarters. Here he sings and watches during the day, usually roosting nearby at night. This territorial center is frequented faithfully during the entire season, unless his routine is changed by polygamy, in which case secondary headquarters are often established nearer his mates. As he may have as many as three females, each having two broods, the chief center of interest in the territory may change as each female reaches the peak of sexual responsiveness.
By intimidating songs and alarms, displays and disputes, the male meadowlark defends his domain against the encroachment of covetous rivals. It is clear that from the beginning of his tenure the male has a definite conception of his territorial acreage and chases all resident males of his species beyond these boundaries. The competitors may come to blows, but it is usually a matter of vehement displays or competitive singing, ending when the vanquished bird takes wing. The loser may be pursued rapidly as far as the boundary line; the victorious male then returns to the sentinel station, singing a spirited flight song as he flies. G. B. Saunders describes a territorial flight he observed on March 25, 1929: "My attention was drawn to two birds fighting savagely in the grass. From a distance I could see the flashing white rectrices and was able to identify them as meadowlarks. One male was on top of the other jabbing him fiercely with his long bill. Then they rolled about for a moment wrestling and stabbing with their feet locked together. Instead of taking wing they hopped at each other, grappled, and again fell on their sides. Wings were held loosely and white tail feathers flashed repeatedly as their tails opened and closed spasmodically. After more than a minute of jabbing, one bird arose and flew, pursued hotly by the victorious contestant who gave a jubilant flight song during the chase."
Courtship.--The arrival of the females on the breeding territory stimulates the resident males, who by this time are well prepared for an animated and lively courtship, to a frenzied rivalry that often becomes furious. Two rival males have been seen tumbling about on the ground on their backs with their feet firmly locked together, striking at each other with their bills in mortal combat.
The courtship is featured by elaborate displays, spectacular flights, and intensive singing. G. B. Saunders describes a performance he observed from a blind on April 16, 1931: "Today instead of witnessing the usual routine I observed the first resident female seen since the preceding fall serve as the center of attraction for three competing males. As I reached the blind all four birds took wing and began a most exciting and spectacular chase in which they zig-zagged back and forth, describing circles 200 yards in diameter and maintaining for the most part a steady flight, but occasionally sailing on set wings or giving pulsating strokes. Throughout most of the exhibition the female was pursued by all the males, but now and then two of the latter would engage in a private chase after each other (rarely striking in midair) only to return quickly to the magnetic female. Finally all four came down to the spot from which they had flown. The female began walking about, feeding in the short grass; occasionally she paused to give a conversational chatter that impressed me as being softer, finer, and more modulated than the alarm chatter. The males vied in following her, first one then another arching his body, pointing his bill up, and flying jerkily toward her at an elevation of from 3 to 6 feet. At times they would walk near her with quick, short steps, their bodies held vertically, bills pointed to the zenith, wings twitching so rapidly that the remiges (particularly the tertiaries) described a blurred arc above their backs, and tails convulsively spreading and flashing the white areas. Then they would spring into the air, fanning their wings powerfully but jerkily for six or eight strokes. This 'jump flight' apparently serves two purposes, that of displaying to the female, and of observing and intimidating other males.
"In this way the four birds proceeded for some 50 yards, the female for the most part apparently uninterested but occasionally pointing her bill, twitching her wings and tail, and revealing her excitement. The males at such times would attempt to intimidate each other with violent displays. The female would chatter her approval. After several minutes all four birds flew out of sight, but very soon one male returned followed by the female. Now and then a new note which sounded like the beert of the nighthawk was given. She resumed her feeding, while he continued to post himself nearby. Instead of making himself tall and slim however, he fluffed out his body feathers until the yellow breast he presented to her gaze was a broad, flat golden shield set with a shining black gorget. He continued to make advances, pointed his bill upward now and then, flirted his wings, etc. She chattered or gave the weet, weet, weet call in answer to almost every song. It is noteworthy that while he alone was displaying, he did not sing. Then, resuming his perch at headquarters, he sang brilliantly for 19 minutes, averaging 11 songs per minute. During the next 4 days he continued to spend much of his time near her, frequently displaying and posturing, but she seldom displayed, usually continuing to feed quietly near him."
The usual courtship display of the male is summarized by Saunders as follows: "Taking a direct stance near the female he raises his body to its full height, stretches his neck to its full length, and points his bill to the zenith. The tail is fanned, showing all the white, and is also jerked up and down; the wings are flirted rapidly over the back, either simultaneously or alternately; and the breast feathers are fluffed out to form a lovely shield of contrasting yellow and black. The beert note may be given. He may spring from the ground as has been described, even flashing his tail in midair.
"The female's reaction to this performance is to raise her body to its full height, stretch her neck, and point her bill; flashing her wings and tail in answer to his song and chattering or giving a beert.
"Throughout this period they spend much of their time together. When he is aloft singing she is usually feeding or perching nearby. As a rule, however, she frequents an elevated perch much less often than the male and seems less sure of herself while doing it. If he is singing, she often answers each song with soft conversational chatter, 'dzert-tet--tet-tet-tet.' They seem to enjoy their companionship very much, remaining together in long flights across the territory to feeding grounds and maintaining this proximity while feeding; and occasionally indulging in a sexual flight, the male singing a beautiful flight song. After such a flight he often repeats his sexual advances in a more or less obvious manner, but she responds either weakly or not at all. While feeding, they pass hours in which little if any show of sexual interest is witnessed."
Nesting.--The meadowlark is primarily a bird of the grasslands, meadows, and pastures; and it is in such places that we usually find its nest. I have also found them in corn, alfalfa, and clover fields and weedy orchards, as well as in grassed islands among plowed fields. The nest is made of dried grasses lined with finer materials. In Illinois I have found nests lined with small amounts of horsehair; in Maine, wiry grasses and even pine needles are sometimes employed for this purpose. Most of the nests have a dome-shaped roof constructed of grass more or less interwoven with the attached and growing parts of the clump of grass or weeds against which it is built. The interior of the nest is open to view from only one side, and this opening may be more or less obscured by overhanging grasses. Sometimes there is a covered passageway to the nest especially to those built in a field where the tall grass was not cut during the previous season. Some of the nests are so well hidden that they are difficult to find, and are discovered only when the bird is flushed by the accidental encroachment of someone walking through the field. The colors and markings of the plumage blend so perfectly with the surroundings that if a nesting bird could restrain its fear, a person might pass within inches of a nest and never be aware of its presence. Most nests that I have found were those I nearly trampled under my feet. Most nests are built in a small depression of the ground, the depth of which may be augmented by some excavation by the bird until it is about 1 to 2 1/2 inches deep. Skutch writes of a nest he found in an alfalfa field near Ithaca, N. Y., on May 26, 1931; it was a sparse structure of grasses only half covered over and set in a depression of the ground so that the upper side of the five eggs were about level with the surface of the field.
The nest varies in size and bulk according to the situation in which it is found. Of five nests measured, the average total height was 7 inches, the outside diameter 6 1/2 inches and the inside measurements of the nesting cavity approximately 4 by 5 inches. The average opening is 3 1/4 inches wide and 4 inches high.
A. C. Bent found an unusual nest containing three eggs in an unusual site at Sea Isle City, N. J., on June 23, 1928. Located in short grass on Black Rail Marsh, it was made of dried coarse grasses, completely arched over with a thick, dense canopy of coarse dry grasses and weeds, and was much like the nest of the black rail in appearance. He found another nest containing four eggs among the beach grass at Chatham, Mass., on May 28, 1904, sunk into the sand just back of the crest of the beach.
F. W. Rapp reports a meadowlark's nest, containing four eggs, which was located within 9 feet of the track of the Grand Trunk Western Railroad. Trains running at a high rate of speed, making much noise and jarring the ground, apparently did not disturb the birds. The nest, in short grass, was completely covered over with dried grasses and the entrance was away from the tracks.
Robert L. Denig (1913) reports unusual conditions under which a meadowlark nested at Wakefield, Mass., where the U. S. Marine Corps conducted rifle practice during the summer of 1909. Mounds of earth about 3 feet high were built to elevate the firing points at 100-yard intervals. The meadowlark built its nest on the far side of the 400-yard mound directly in line with the target, so that the muzzle of the rifle of the man lying on the mound was directly over the nest and not more than 2 feet above it. At first when the firing skirmish line was about 400 yards distant, the birds would fly away; but as the practice continued they became more and more accustomed to the noise; they would allow the men to approach nearer and nearer before leaving the nest and would return at once when the firing ceased at that point. As the time came for the eggs to hatch, one of the birds would remain on the nest throughout the firing, even when the gun was being discharged directly over its head, not more than 2 feet away. Finally, the eggs hatched, and the young birds were brought up, so to speak, "under fire."
G. B. Saunders provides the following notes on nest building: At Ithaca, N. Y., the meadowlark begins nesting in late April or early May. The time required for building the nest varies from 3 to 18 days. One pair began carrying nesting material on April 20 and completed the nest on May 8, another began building on April 21 and finished on May 8. Still another began carrying grass on June 3 and finished her nest and laid her first egg on June 6. In every case except the last one mentioned, several nests were started and worked upon. Usually the first beakfulls of nesting materials are deposited at different places and at first there seems to be no concentration on any particular location. During these first days copulation with the male takes place. When the female is responsive to the advances of the male she crouches close to the ground, shortens her neck, and points her bill upward at an angle. She flutters and flirts her wings and lifts and spreads her tail. The male displays a few feet distant, and the copulatory act is finally achieved with no sounds being uttered by either bird.
Trips with nesting material are most frequent early and late in the day, but may sometimes continue during midday as well. The details of nest building are presented in the following typical case. On May 12, 1931, a nest which Saunders had under observation was scarcely begun, but both cup and roof had been started. A natural depression 2 inches deep had been slightly modified by the female, who used her bill for the retouching. Into this cavity a thin layer of last year's grass blades had been laid. Much of the tuft of grass surrounding the cup had been arched over it and woven together, being secured by a few long dry grass stems woven among the growing blades. In half a morning's work she had both the cup and roof well started. The male gave no assistance in the enterprise and offered none later.
On May 13 as well as on the two following.days, the female made regular trips with material every 5 to 10 minutes. She usually remained at the nest from 35 seconds to 2 minutes placing the material that she had brought. On this day the nest was half completed; the lining of the bowl was much deeper but was still flimsy. On May 15 the nest was complete except for occasional additions of material. The first egg was laid on May 17. Occasionally the building continues for several days after egg laying is begun.
Saunders' observation of the nest-building process is thus summarized:
Following the choice of a nesting site, the customary first step is to prepare the earthen foundation for its cup of withered grass. There may be a natural depression, a hoofprint or similar hollow already present, in which case the female remodels it by using her bill as a combination pick and forceps tool. In some soil the marks of her heak remain after the young have departed. The use of the bill for digging the soil is not surprising, for the habit is often shown in feeding, when the meadowlark employs it to probe for insects and grain and to dislodge clods of earth.
Once the hollow is satisfactory, the adjacent grasses or other growing plants are pulled over the pit and interlaced, or secured by the addition of long stems or blades of dead grass until they form a more or less complete dome which later conceals the eggs from view and protects them from the sun and rain. Other nests are not below the general surface level but are built entirely above it, there being a front step as a result of this variation in architecture.
The cup and nest lining are usually fashioned while the dome is in the process of construction, first one part and then another receiving the attention of the female. Many more than a hundred loads of dried grass going to the making of the finished home. Although many authors credit both sexes of the eastern meadowlark with the job of building, I have never observed a male sharing in this activity. Perhaps he does, but such a male would be an exceptional individual, and a far more helpful mate than any of the dozen males which had their intimate lives scrutinized daily during my study at Ithaca, N. Y.
G. B. Saunders, the first to discover the common practice of polygamy among meadowlarks, reports that the secretive nature of the females and the inconspicuousness of their nests are two of the principal reasons why the eastern meadowlarks have been able to keep their polygamous habits a secret for so long. Although meadowlarks breed in every one of the 48 states and are abundant in most of them, no mention has appeared in the voluminous literature on Sturnella regarding the frequent bigamy of the males of this subspecies. His intensive field work in more than 20 territories at Ithaca, N. Y., in 1931 revealed that about 50 percent of the males were polygamous. One of them was found to have three females, all of which were nesting at the same time. He adds that among the several reasons why polygamy is common among meadowlarks is the fact that the females are not hostile to one another as they are in many other species; they feed together, associate with the male together, and often nest within 50 feet of each other. Another is that the males are repeatedly attracted by desirable females.
Eggs.--The number of eggs of a set of the meadowlark varies from three to seven, but sets of five eggs are most common. Sets of four are more usual in the second brood nests of the season. Birds breeding in the southern part of the nesting range on the average lay smaller sets.
According to Bendire (1895):
The eggs of the Meadowlark vary considerably both in shape and size; the majority are ovate, while others are short, elliptical, and elongate ovate. The shell is strong, closely granulated, and moderately glossy. The ground color is usually pure white; this is occasionally covered with a pale pinkish suffusion, and it is very rarely pale greenish white. The eggs are more or less profusely spotted, blotched, and speckled over the entire surface with different shades of brown, ferruginous, pale heliotrope purple, and lavender; these markings generally predominate about the larger end of the egg, and are rarely heavy enough to hide the ground color.
In some sets the markings consist mainly of a profusion of fine dots; in others the spots are well rounded and fewer in number; and again they occur in the shape of irregular and coarse blotches, mixed and finer specks and dots; in fact, there is an endless variation in the style of markings.
The average measurement of a series of two hundred and one specimens in the United States National Museum collection is 27.75 by 20.35 millimeters, or 1.09 by 0.80 inches. The largest egg measures 30.78 by 22.61 millimeters, or 1.21 by 0.89 inches; the smallest, 21.59 by 18.29 millimeters, or 0.85 by 0.72 inch.
Ninety-five eggs weighed by G. B. Saunders had an average weight of 6.6 grams, a minimum weight of 5.4 grams, and a maximum weight of 7.7 grams. Eggs of any one clutch are usually similar in size, coloration, and weight. He found that of 85 eggs in 20 nests found at Ithaca, N. Y., 14 eggs, or 15.5 percent, were sterile.
Incubation.--When an average clutch of five eggs is laid, incubation may begin with the deposition of the third, the fourth, or the fifth egg, but more frequently begins with the laying of the fourth egg. The incubation period is usually 14 days, but under certain unusual conditions it may be only 13 or as many as 15 days.
Although the male has been credited with a share in incubation, Saunders has never witnessed any such cooperation in the many nests that he has closely observed. Once incubation has begun the female remains on the eggs most of the day, leaving only long enough to feed. The nest is never left at night unless she is frightened by an intruder. During 12 hours and 40 minutes of daylight, one female spent 9 hours and 40 minutes, or 76 percent of the time, on the nest. The longest absences from the eggs were during the middle of the day when the temperature was highest. On cool or rainy days less time is spent away from the nest.
While incubating, the female is continually active. These activities include listening to songs and sounds of approaching danger, "humming" when the male sings, turning the eggs, feeding on insects which come within reach of the nest, probing in the nest, rearranging nest materials, preening, etc. The female on the nest often responds to the flight song of the male by voicing low, sweet chuckling notes that are unlike any others uttered by the meadowlarks. The softness of these sounds precludes their detection by man at distances greater than 15 or 20 feet. The eggs are turned many times during the day, and in the course of 1 hour in the late afternoon a female was observed to turn them five times. The eggs do not hatch in the order that they are laid; for example, the fourth egg may hatch several hours in advance of the third. Individual variation in development causes differences of several hours or even a day in the hatching time.
Young.--The young at the time of hatching, according to G. B. Saunders have a smooth orange-red skin; the bill and nails are flesh color; and the natal down is pearl gray. The down is longest on the capital and spinal tracts and shorter on the humeral and femoral tracts. When the down is dry it fluffs out and appears quite abundant, particularly on the spinal tract. On the head the down is localized chiefly above the eyes and on the occiput. The dark sheaths of juvenal feathers are visible on the dorsal surface of the head and on the spinal and dorsal regions, and less easily discernible in the humeral, alar, femoral, and crural tracts.
Shortly after hatching, the nestling reacts to the food call of the female and holds up its mouth in a wobbly and uncertain manner, at the same time uttering weak notes, see see or seep seep. The female during the first few days spends long hours in the nest brooding the nearly naked young. The young are fed by both the male and female but the male feeds them much less often. During the first 2 days the young evacuate in the nest, being too weak and lacking the instinct to void their droppings outside the doorway. Later, by the third or fourth day, each youngster may be observed to turn about and expel the mucous-covered sac beyond the rim of the nest. These sacs are removed by the adults but in the early stages of the development of the young they may be eaten. Egg shells are removed to a considerable distance by the female immediately after the young are hatched. Infertile eggs are usually left in the nest during the entire period of occupation.
By the third or fourth day a slitlike opening appears in the eyelids, so that the youngster can see whenever it is fed or disturbed. At other times the eyes remained closed. The position of the young birds is now more upright and alert, and the wings have grown enough to be useful as props for maintaining balance. The legs are still almost useless; there is little muscular coordination and they are well sprawled out at the sides.
By the fifth day the eyes are fully opened and the voice is stronger. The nestlings now face the opening of the nest, expectantly waiting for food. Growth is rapid, and the juvenile plumage is rapidly acquired. When disturbed, the young now exhibit signs of fear. Wing exercises and stretching of the legs and neck are indulged in frequently.
By the eighth day the young are very alert and receptive to sounds coming from outside the nest. They may be seen frequently preening their feathers, apparently to facilitate the unsheathing process. During the remaining days in the nest the young become so active the nest is wrecked and the roof worn away, exposing the nestlings to view and to the hot, direct rays of the sun. When thus exposed to the sun they pant violently in order to control their temperature. Sometimes they leave the nest, but return after being fed.
By the eleventh or twelfth day the birds normally take their final leave of the nest, although if molested they may desert it as early as the eighth day. Feather growth of the wing tracts has proceeded sufficiently by the eleventh day to allow the nestling to fly, in case flight is necessary. However, the newly departed young meadowlark seldom takes to its wings during the first few days except to make short jumps in the grass. The young are fed by the adults for a period of at least 2 weeks or longer after they leave the nest. Their food call is a loud bisyllabic tseup, tseup, and it is by these notes that they are located and fed by the adults. The second nest may be started within 2 or 3 days of the desertion of the first one. While the female is building and laying she continues to feed the first brood, but when the second incubation is begun the male assumes the major part of the work of caring for the young of the first brood, which are about 3 weeks old at this time.
Gradually they learn to catch insects for themselves and become more and more independent. When they are able to shift for themselves, they are apparently chased out of the territory by the male. They probably do not travel far before September, when they acquire their first winter plumage.
Four birds taken from a nest when 8 days old were raised in captivity by G. B. Saunders. Since they were given long hours of freedom in their native fields, their development and habits were similar to those of wild juveniles. On the fifteenth day they all took dust baths, fluffing and shaking their plumage as adults would, Following this exertion they drank heartily from a basin of water. On the sixteenth day they began prying into the soil with their bills, which marked the inception of the boring habit which is so typical of the adults. On the seventeenth day they began to stand high on their legs and to hold bodies erect whenever they heard a sound which startled them. On the twentieth day one was observed to take a thorough bath in the water, after which he spent several minutes in a systematic dressing of his plumage, during which he apparently used his oil gland frequently. On the twenty-second day, two of the four began feeding for themselves; before that Saunders had been feeding each bird about 175 grasshopper nymphs daily. On the same day one of the two that had begun feeding themselves gave a rolling chatter very similar to that uttered by the adults.
Juvenal plumage acquired by a complete moult. Above, clove-brown, the feathers broadly edged with buff palest on the nape, those of the back having double subapical spots of russet. Median crown stripe, and superciliary line cream-buff. Wings sepia-brown, the primaries and secondaries obscurely barred on the outer web with darker brown and edged with pale vinaceous cinnamon shading to white on the first primary, the tertials clove-brown broadly edged with buff and having a row of partly confluent vinaceous cinnamon spots on either side of their shafts producing a barred effect, . . . the rest of the wing coverts obscurely mottled with light and dark browns and edged with buff, the alulae with white. The three outer pairs of rectrices are white with a faint dusky subapical shaft-streak, the next pair largely white and the others hair-brown confluently barred with clove-brown, and whitish edged. Below, including "edge of wing" pale canary-yellow, nearly white on the chin, the sides of the throat, breast, flanks, crissum and tibiae washed with pinkish buff, streaked and spotted with brownish black which forms a pectoral band. Bill and feet pinkish buff, the former becoming slaty, the latter dull clay color.
First Winter Plumage acquired by a complete post-juvenal moult beginning about September first after the juvenal dress has been worn a long time, young birds and old becoming practically indistinguishable.
Above, similar to the previous plumage, but all the browns even to the wing and tail quills much darker, often black, and distinct barring rather than mottling, the rule. The feathers of the back have large single subapical spots of rich Mar's-brown crossed by two faint dusky bars, and the primary edgings are usually grayer. Below, a rich lemon-yellow (including the chin and supraorbital dash) veiled with buff edgings and a black pectoral crescent is acquired completely veiled with deep buff and ashy edgings. The streakings below are heavier and darker, many of the feathers with subapical russet spots and the wash on the sides is deeper and pinker.
First Nuptial Plumage acquired by wear which is excessive by the end of the breeding season producing a dingy brown and white appearance above with yellow and black below. The subapical spots of the feathers of the back are almost entirely lost by abrasion and the same force scallops out the light portions of the tertiaries, wing coverts, and tail. Neither the yellow nor the black below fades very appreciably, but the shining denuded shafts of the feathers project far beyond the abraided barbs. The yellow seems even to be intensified by the loss of paler barbules.
Adult Winter Plumage acquired by a complete postnuptial moult in September. Usually indistinguishable from first winter dress.
Adult Nuptial Plumage acquired by wear as in the young bird.
Female: In natal down and juvenal plumage the sexes are indistinguishable. Later the female differs only in slightly duller colors and a more restricted black area on the throat. The moults are exactly the same as in the male.
Abnormal plumages involving albinism and melanism are known to occur in the meadowlark. The majority of the cases of albinism which have been reported are actually only partially albinistic; in most the brown of the upperparts is white or whitish, whereas the yellow of the underparts seems to be retained in varying degrees of intensity .
James Savage (1895) collected an albino meadowlark near Buffalo, N. Y., in which "The usual brown of the upper parts was of a pale buff color with the pattern of the feather markings indistinctly discernible, while the yellow on the breast was as pure as in an ordinary Lark."
Louis S. Kohler (1915b) gives an account of a partial albino meadowlark he observed near Bloomfield, N. J.: "On October 7th during the afternoon while strolling over the fields I came upon a partly albino bird. This bird was of normal plumage except the tail and wings in which parts, more than half the feathers were devoid of color. This bird during its association with others of its kind was continually being attacked and presented a very bedraggled appearance from their frequent onslaughts and was forced into solitude by them at close intervals. But in spite of their pugnacity it always returned to the vicinity of its tormentors and was immediately set upon and driven off."
G. B. Saunders states: "There is an albino eastern meadowlark in the Cornell University museum which has upperparts and wings whitish, the bill pale brown, the jugular crescent buffy brown, but the yellow underparts nearly normal." There are many other similar cases of partial albinism in the meadowlark but I have discovered no report of a pure albino eastern meadowlark.
Chas. H. Townsend (1883) describes a melanistic specimen collected in New Jersey as follows: "The upper plumage is of the normal color, while the whole head, neck and under parts are perfectly black. There is the faintest possible trace of yellow along the sides, and no white feathers in the tail, which is very dark above and below."
Food.--Few birds of the agricultural areas can claim a higher rank in its economic relations to man than does the meadowlark. During the summer months most of its food consists of insects and closely allied forms. It eats practically all of the principal pests of the fields and is particularly destructive to the dreaded cutworms, caterpillars, beetles, and grasshoppers. In the autumn, and especially in winter, when insect life is scarce, it resorts in a large measure to seeds. It does feed on certain grains useful to man, such as corn, wheat, rye, and oats; but most of these are waste left behind at harvesttime. It seldom disturbs these cereals when growing or before being harvested. I have seen flocks of them in weedy cornfields where apparently they were feeding exclusively on seeds of smartweeds and ragweeds. Meadowlarks have been known to eat certain fruits such as wild cherries, very small part of their subsistence.
An account of the food habits of the meadowlark among the sand hills of North Carolina in winter is given by M. P. Skinner (1928):
During the winter the number of Meadowlarks remained quite constant, although there were temporary variations each day. But in February it became noticeable that some of the winter birds were leaving. They seem to stand the cold weather, but snows cover their usual food and then these birds may be found in very unusual places, on any little patch of bare ground they can find, and about barns and stockyards.
During the winter in the Sandhills the Meadowlarks depend largely on seeds and waste oats for food, but also catch caterpillars, cutworms, earthworms, and as many kinds of insects as they can. These foods are secured on the ground and in the short stubble and grasses. At times these birds seem to give preference to seeds and at other times to feed almost entirely on insects even during the depth of winter when insects might be supposed to be scarce. For securing the two different kinds of food, the Meadowlarks use quite different methods. When after seeds they hunt through the grass and weeds, stopping occasionally to gather seeds from the standing or fallen stalks. When they find places where the seeds are numerous on the ground, they both scratch with their feet and dig with their bills. If there is a wind blowing, they usually fly to the lee side of the field and then advance on foot across it and against the wind. This is apt to scatter the flock especially as one individual often has better luck than another, and the unsuccessful ones usually hunt up new places for themselves rather than share the first ones' success. Even when scattered over a large field the flock retains its organization, and when one bird leaves, the others usually follow one by one at short intervals until all have left. When they are feeding on insects the Meadowlarks move more rapidly, and perhaps separate more. Then, they do not search the ground or dig with their bills, but they look very closely at the bases of the bunches of grass as they pass by. At times they appear to find insect-catching very profitable at the stockyards and near barns.
Occasionally a Meadowlark takes both insects and seeds indiscriminately. Such a bird came walking through the rough at the edge of a golf links; like a Flicker, it thrust its bill into the soil experimentally every step or two. At the foot of a tuft of grass it dug out two white grubs and ate them, then it walked over to a spray of dried everlasting, pulled it down and ate several seeds while holding the stalk down under one foot.
In Florida and sections of southern United States more of the food during the winter months consists of insects, chiefly beetles but also cutworms, caterpillars, and grasshoppers. Howell (1924) has found the meadowlark to be an important enemy of the cotton-boll weevil in the south. Since it feeds regularly upon this insect during the winter months, it very materially reduces the number which might otherwise descend on the cotton crop the following season.
Investigations in South Carolina and other southern states as far west as Texas, according to Beal, McAtee, and Kalmbach (1927), have substantiated accusations that the meadowlark is guilty of destroying sprouting corn.
This habit seems to be confined to the migrating or wintering flocks before they have broken up for the breeding season and is probably occasioned by a scarcity of other available food. North Carolina seems to be the most northerly state in which this objectionable trait of the meadowlark manifests itself. Corn planted in March is most susceptible to attack and cases may be frequently encountered where whole fields must be replanted, resulting in a delayed and less profitable crop. In attacking the sprouts the birds usually drill a small conical hole down to the germinated kernel which they eat, leaving the tender sprout exposed to the withering effect of sun and air.
F. E. L. Beal (1926), in a report of a detailed analysis of the contents of 1,514 stomachs of meadowlarks, found that 74 percent consisted of animal food and 26 percent of vegetable matter. The animal food consisted of practically all insects, chiefly "ground" species such as beetles, bugs, grasshoppers, and caterpillars, with a few flies, wasps, and spiders. Of the various insects eaten, crickets and grasshoppers are the most important, constituting 26 percent of the food of the year and 72 percent of the food in August. Of the 1,514 stomachs collected at all seasons of the year, 778, or more than half, contained remains of grasshoppers, and one was filled with fragments of 37 of these insects. Next to grasshoppers, beetles are the most important food item of the meadowlark, food amounting to about 25 percent. Forty-two adult May beetles and numerous white grubs of this beetle, a most destructive insect, notably to grasses and grain, were found. Among the weevils the cotton-boll and alfalfa weevil were the most important economically. Caterpillars, including many cutworms, form a constant element of the food and in May constitute over 24 percent of the entire food. Adult moths and butterflies are seldom eaten. The remainder of the insect food is made up of ants, wasps, and spiders, with some bugs, including chinch bugs, and a few scales.
The vegetable food, according to Beal (1926), consists of grain and weed and other hard seeds. Grain was found chiefly in stomachs collected in winter and early spring; hence it represented waste material. Clover seed was found in only six stomachs and but little in each. Seeds of ragweed, barnyard grass, and smartweed are eaten from November to April, inclusive, but during the rest of the year are replaced by insects.
As for the food and behavior of meadowlark young, G. B. Saunders says that within an hour after the meadowlark is hatched it receives its first meal of cutworms, other small insects, and spiders. Young grasshopper nymphs which the female has mashed between her mandibles may be included in these early meals. When the adult arrives at the nest, insects can be seen projecting from her bill, and these she feeds to the young by squatting on her tarsi in front of the entrance and putting morsels well down the throat of each youngster. When the young are satisfied she resumes her brooding. During brooding, a nestling may get hungry, in which case the female raises her breast and reaching down into the nestling's open mouth, gives it either some insects which were left over, or a meal of regurgitated food. That she regurgitates is clearly shown by the pumping action of her neck and head. During the first few days there is no pronounced change in the routine of the female, for she continues to spend long hours in the nest brooding the nearly naked young.
When the young later become stronger, hardier, and somewhat insulated by feathers, the female spends much less time at the nest and feeds the young no regurgitated food. By the time the young are about 6 days old they are receiving the usual fare of grasshoppers and larvae, plus ground beetles, crickets, and other heavily chitinized insects. During later nest life their hunger must be appeased about every 5 to 10 minutes early and late in the day, and at intervals of about every 15 minutes during the hotter hours. Most of these trips are made by the female, whereas the male makes few visits and is much less solicitous in his attentions to the young. The female averages nearly a hundred trips a day to the nest during the 12 days the young are in the nest. The food daily given each nestling weighs 8 to 20 grams, a weight equivalent to that of about 100 to 300 small grasshopper nymphs. Saunders estimates, on the basis of various methods of determination, that a 10-days' supply for 10 nestlings, when the chief food is grasshoppers, would be 5,000 to 7,000 grasshoppers. These figures again emphasize the great economic importance of the meadowlark.
Voice.--The plaintive and very pleasing whistled notes of the meadowlark, heard on its arrival, stand out among my most delightful memories of early spring on an Illinois farm. There, where a tall Osage orange tree stood at the edge of a rolling meadow, a meadowlark came each year to announce his arrival. This song may be rendered by the words Ah-tick-seel-yah or Heetar-see-e-oo, but others have translated it variously such as Spring-o'-the-yeear; Peek-you can't see me; Toodle-te, to-on, etc. There is an infinite number of variations of the territory song, but all have much the same quality. This song is not only the first heard from the meadowlark in spring, but is the one repeated from the singing posts throughout the season.
The meadowlark is known to alternate the versions of its song. Frances H. Allen (1922) writes of a bird he observed on an April morning:
He had four or more songs in his repertoire. The first, which was repeated a number of times in succession, resembled the opening notes of the white-crowned sparrow's song, but had three high notes on the same, pitch, instead of two, before the lower one: 'ee-ee-ee-hew.' It was a beautiful song and so different from anything we commonly hear from the meadowlark that I did not suspect its author at first. . . . then the bird began to alternate this song with another which seemed a good musical complement to it. This second song began low and ended high. It was something like 'hew-hew-he-hee,' the third note shorter than the others. After a few alternations of these two songs the bird dropped the first and sang only the second a number of times, but dropped that in turn and finally took up two or three simpler and more normal songs, of which one, at least, was sweeter than most meadowlark songs.
The peak of singing activity, when the most beautiful songs may be heard, occurs during the first part of the breeding season, prior to incubation. During incubation there is a distinct lull in singing which lasts until the return of sexual activities in preparation for the second brood. Another lull occurs during the rearing of the second brood and lasts until fall, when singing is again renewed. In sections of the country where the meadowlark is represented by individuals during all seasons, its characteristic territory song may be heard throughout the year, even during the winter months.
The versatile meadowlark has also a flight song, a truly ecstatic performance. Prefacing the flight song with a few notes from a perch, it flies swiftly upward, sometimes spirally into the air. It vibrates its wings rapidly and utters penetrating and chattering notes in rapid concert not unlike that of the bobolink. After flying more or less in a circle, it slowly descends to the ground. This song too is variable but is very different and not at all suggestive of the ordinary song.
The songs of the eastern and western meadowlark have frequently been compared. Albert Brand (1938) who has made a study of vibration frequencies of passerine bird song, found that for the eastern meadowlark the highest note had 6,025, the lowest 3,150, and the approximate mean 4,400 vibrations per second. Those of the western meadowlark are much lower in pitch--3,475 for the highest, 1,475 for the lowest, and 3,475 for the approximate mean.
When the meadowlark is alarmed or excited it nervously flits and twitches its tail, exposing the white tail feathers. This behavior is accompanied by a sharp nasal call note, which changes to a rolling chatter followed by a plaintive but pleasing whistle. G. B. Saunders describes the call notes of the meadowlark in detail, as follows:
The day-old nestling first voices his calls for food with a faint 'tseep, tssep, seep, seep' or 'tsp, tsp.' As he gains strength this utternace is a lisping 'sweet, sweet, sweet.' By the seventh or eighth day the note becomes a bysyllabic 'tscheep, tscheep,' 'tscheep' or 'tschip', tschip'.' All of these notes are of the same general type. When out of the nest, the juvenile's call is a loud peeping 'tseup', tseup' ' or 'sweet, sweet,' similar to the 'weet, weet' notes of the adults.
The adult call notes may be expressed phonetically as 'weet, weet, weet.' Those of the female are usually softer and more modulated than those of the male. There is an infinite variation in the expression of these notes. Other conversational calls of the adults are the low pitched and modified alarm notes, 'dzert, dzert' and the 'tet-tet-tet-tet' notes of the chatter. The female often joins them, i. e., 'dzert, tet-tet-tet-tet-tet-tet' in answering the male's song.
The common alarm chatter, 'dzert-tet-tet-tet-tet,' seems to be a modification of the call notes just mentioned. The speeding up due to excitement gives the notes a much harsher quality. The notes 'dzert-dzert' are usually given when a preliminary alarm is uttered. Another note fairly common during the breeding season, but one not heard except at that time, is the queer 'beert' or "nighthawk" note. It may be given as an alarm when the birds are greatly excited, or it may be given during sexual displays and competitions. It is uttered by both sexes.
Aretas A. Saunders has written a very excellent analysis of the song and notes of the meadowlark: "The song of the Eastern meadowlark is a short series of sweet, clear, very high pitched whistled notes. It is loud, carries a long distance, and, when one is near the bird, is rather shrill. The notes are few, compared to those of other birds, and downward slurs from a high to a lower note are frequent. In spite of the few notes, it is exceedingly variable.
"In pitch and time the song is remarkably like human music. The notes are usually on the same eight notes of the octave as in the simpler kinds of human music. The shorter notes are commonly half or a third the length of the longer notes, so that the songs could be recorded on the musical scale, as human music is written, with considerable accuracy. The different songs are easily and quickly recorded by the graphic method. My earliest experiments in recording bird songs were with the meadowlark, and although many of the records have proved to be duplicates, I have at the present time more than a thousand different songs of this species on record. The following data are based on a study of 962 of these records that I have filed and catalogued, the remaining records being still only in my field note books.
"These records show that the songs vary from 2 to 8 notes each, the great majority 3 to 6 notes. There are 4 songs of 2 notes; 65 of 3 notes; 352 of 4 notes; 391 of 5 notes; 132 of 6 notes; 15 of 7 notes and 3 of 8 notes. In spite of the great variation, many records prove to be duplicates, and it is a common experience to hear two or three birds singing the same song, one after the other, and also common to record songs from widely separated localities that are exact duplicates. While the majority of my records are from southwestern Connecticut, I have a good many from various localities in New York, and scattering records from other states. Songs that are common in Connecticut are often equally common in southwestern New York, approximately 400 miles distant. I have also recorded duplicates of Connecticut songs from the vicinity of Dover, Del .
"The pitch of songs varies from C''' to D#'''', a range of l 1/2 tones more than an octave, the highest notes being a little higher than the highest on the piano. The range of individual songs varies from 1 tone to an octave; 12 songs have a range of only 1 tone, and only one has a range of an octave. Nearly half of the records, 446, have a range of 2 1/2 tones, and the average of pitch of all of them is 2.7 tones.
"The duration of meadowlark songs varies from about 2/5 second to nearly 3 seconds, averaging about 1 4/5 seconds. It is difficult to measure short songs accurately with a stopwatch. The time factor of greater interest is the perfect rhythm of the notes and the great number of variations in time arrangement that, with the variations in pitch, go to make the great number of different songs that this species possesses.
"Not only does the meadowlark, as a species, sing a great variety of songs, but each individual has many variations. I once recorded 53 different songs from one individual in less than an hour, and recorded altogether 96 different songs of birds singing in that location in that season.
"Consonant sounds are not prominent in meadowlark songs. In some songs notes are linked together with a liquid consonant sound, like the letter 1 that occurs in about 10 percent of the songs I have recorded. Another consonant sound, which occurs at the beginning of certain notes, most commonly at the beginning of downward slurs, is sibilant and sornds like the letters ts, making a slur sound like 'tseeyah', or something similar. The sound is rather faint however. I have recorded it in less than 5 percent of the songs, but it may be commoner than this indicates, for it is not easily audible from more distant singers.
"In early spring, usually late March and early April, the meadowlark frequently sings two different songs in alternation, usually with a pause of about one second between them. I have eight records of these alternated songs, all different. In most of them one song ends on a high-pitched note and the other on a low pitch, so that they sound something like a question and an answer, and form a pleasing musical combination. All my records but one, recorded at Cross Lake, N. Y., in July, are dated between March 7 and April 11.
"In addition to this form of song, the meadowlark has a flight song, very different in character, that is rather rarely heard. In a good many years I have not heard it at all, whereas in others I have heard it several times, most commonly in late April. The performance begins from a perch, the bird calling at intervals on a rather harsh, nasal, downward slurred note. After several of these notes the bird rises into the air and flies across the meadow singing a song made up of groups of 4 or 5 notes, separated by short pauses. These notes are fricative and not especially musical, nor are they so loud as the common song. Such a song takes 10 to 12 seconds from the beginning notes on the perch until the bird is silent .
"I have heard songs of the meadowlark in every month of the year. The regular period of singing, however, begins in March and lasts until late August. Songs in January are rare, and in 32 years of records I have heard the song in that month only 4 times. In February there is often quite a bit of singing, and in 16 of these years the first song of the year was heard in February, the average date of the first song being February 19. Regular singing, however, does not begin until March, and in 6 years it did not begin until April. The average date of its beginning is March 26.
"I have less full data on the cessation of song, as I have frequently been in places where I could not hear it at the proper season. Five years in Cattaraugus County, N. Y., give an average of August 11 for the last song, whereas 5 years in Connecticut average August 18.
"The song is revived in September or October, and is to be heard quite frequently through the fall until November. In Connecticut 20 years of observations give an average of September 30 for the beginning and November 13 for the end of the fall singing, but such singing is much more erratic than spring singing. Songs in December are rare, though more frequent than in January."
Enemies.--In most sections of its range the eastern meadowlark is not commonly imposed upon by the cowbird. I have never found a nest in New England that contained an egg of the cowbird, and G. B. Saunders states that of over 50 nests studied in Oklahoma and New York, none contained other than meadowlark eggs. However, during the course of a statistical survey of the birds of Illinois in 1906 - 1908 I found four cases of cowbird parasitism: One nest in northern Illinois near Rockford contained three eggs of the meadowlark and one cowbird's egg; of two nests in Champaign County, central Illinois, one contained two meadowlark and three cowbird eggs and the other, three meadowlark and two cowbird eggs, with a broken meadowlark's egg outside of the nest; and a nest near Benton, Franklin County, in southern Illinois, contained two eggs of the meadowlark and two young, one of which, judging from its size and appearance, was a freshly hatched cowbird.
G. Eifrig (1915, 1919) writing on the birds of the Chicago area states that he has repeatedly found nests of the meadowlark with one or more eggs of the cowbird. He also states that one or more or all the eggs of the rightful owner were apparently rolled out. It would seem that the meadowlark is a common victim of the cowbird in the State of Illinois. Milton B. Trautman (1940) found two nests of the meadowlark containing cowbirds eggs at Buckeye Lake, Ohio. Bendire (1895) reports an instance where a second nest was built over one containing the parasitic egg. This is a common habit of certain birds such as the warblers but presumably it is rare in the case of the meadowlark. Herbert Friedmann (1929) has obtained records of cowbird parasitism of the eastern meadowlark from New England, New York, Pennsylvania, Michigan, Illinois, and Iowa, but states that the meadowlark is not a common host.
It is of passing interest to note that eggs of the bob-white quail and bobolink have been found in the nests of meadowlarks, although these instances are not to be classed as parasitism but merely unusual accidents. J. B. Lackey (1913) reports finding eggs of the bob-white in two meadowlark's nests near Clinton, Miss., and Edward R. Ford of Chicago found a meadowlark's nest with four eggs of the meadowlark, one of the cowbird, and one egg of the bobolink .
G. B. Saunders in an examination of 45 adult meadowlarks found 8 contained internal parasites. The tapeworm Anonchotaenia sp. was found in 3 and the parasite Mediorhynchus grandis in 6 birds. The roundworm Diplotriaenoides sp. was found in both Oklahoma and New York birds. Of 5 young in a nest at Ithaca, N.Y., 3 were found to have dipterous larvae, probably of the genus Chrysomyia, in their nasal passages. The meadowlark like most other birds is host to a number of external parasites including lice, ticks, and mites, among which Harold S. Peters (1936) has found the three lice Degeeriella picturata (Osborn), Menacanthus chrysophaeum (Kellogg) and Philopterus subftavescens (Geof.), the three ticks Haemaphysalis leporis-palustris Packard, Ixodes sp., and Amblyomma tuberculatum Marx; and the mite Trombicula hominis Ewing. Occasionally nests of the meadowlark are heavily infested with mites, and G. B. Saunders cites one case where a nest was deserted because of an unusually heavy infestation.
Because the eastern meadowlark has two broods of four or five young during each season, we need not be alarmed at the large number of enemies and of its great mortality. Man, directly or indirectly, is responsible for the loss of a great many meadowlarks and probably he is the most important factor in the control of the species and thus preventing overpopulation. Perhaps the most disastrous but unwitting acts of man is the mowing of alfalfa, clover, and timothy fields in which the meadowlarks nest. In Illinois, while traversing the various sections of the state on foot for hundreds of miles in connection with the statistical bird survey in 1906 - 1908, the loss I noted from this source was appalling. In June and July I saw nest after nest that had been destroyed by mowing machines and it is probably safe to state that more meadowlarks are destroyed by this means, which is repeated year after year, than by any other.
In autumn, when meadowlarks congregated in large flocks in southern Illinois, it was a common experience to see groups of a dozen or more gunners out killing meadowlarks in large numbers, to be carried home for food for themselves and their neighbors. Such practices have been common in some of the southern states in the past, but I am convinced that in recent years there has been less of this kind of destruction because of the more rigid enforcement of protective laws and the general education of the public to the economic value of this bird.
Automobiles, which constitute a menace to certain of our birds, are not such a menace to the meadowlark; however, when the birds frequent dirt roads in autumn to dust their plumage and possibly to pick up stray bits of food, such as grasshoppers, a considerable number have been reported killed.
Since the meadowlark nests on the ground, predatory mammals and birds and probably snakes are responsible for a number of deaths. The domestic cat ranks high as a destroyer of meadowlarks, especially those that nest in fields adjacent to farm homes. Farm dogs, which also roam the fields and which are able to locate the nesting birds through the sense of smell, probably destroy a number of nests. Saunders states that he has seen Bonaparte's weasel attack juvenile meadowlarks.
The examination of the stomach contents of owls and hawks has revealed that the horned and snowy owls, the goshawk, duck hawk, sparrow hawk, red-tailed hawk, red-shouldered hawk, and Cooper's hawk have taken meadowlarks, chiefly during the winter months.
Since the meadowlark is one of the earliest spring migrants, snowstorms frequently cut off its food supply and, the accompanying cold, cause the death of many of the birds. Frederick C. Lincoln (1939) states that during the early part of June 1927 a hailstorm of exceptional violence in and around Denver, Col., killed a large number of meadowlarks and other birds. The ground was strewn with dead birds and many lay dead in their nests where they were incubating eggs or brooding young when the storm broke.
Fall and Winter.--In fall the meadowlarks leave their nesting grounds in Quebec and Ontario during September and October, and by the middle of October the bulk of them have departed. A few individuals may linger on until well into November. Since the meadowlark normally winters in northern United States, the time of departure of migrants is difficult to ascertain.
In southern Illinois during the month of October I have seen immense flocks made up of hundreds of individuals concentrated in the lowlands above Cairo, at the junction of the Mississippi and Ohio Rivers. These flocks were made up largely of birds that had migrated from points farther north. Also, in going through cornfields and stubblelands of this part of the state, I frequently saw smaller companies of them waddling about the clustered stalks. As they paused to inspect me they would hold their bodies in a vertical position with their bills pointed skyward. At the same time they would flick their tails displaying the conspicuous white markings as they opened and closed the fan of feathers .
On the New Jersey coast the meadowlarks start flocking about the middle of August, when it is common to see parties of 20 to 25 individuals. In October the birds band together in large flocks of 200 to 300. Many of these birds pass on farther south, but flocks of 50 to 75 are to be seen throughout the winter. They become much tamer in winter, especially when food is scarce and it is then that they frequent the habitations of man and even enter the towns, where they may be seen in vacant lots feeding in company with English sparrows and starlings. They have also been reported as seen feeding on garbage in alley ways during times of severe blizzards. During the winter months meadowlarks have been flushed from the tall grass of marshes, where the great accumulation of droppings indicated that they had roosted during the night. Meadowlarks have also been known to accompany grackles to their roosts in trees, but this is not common practice.
In recent years meadowlarks have been wintering in increasing numbers in the salt marshes of southeastern Maine in the region of Scarboro, Pine Point, and southward. During October and November as many as 100 to 200 meadowlarks may be started from a single marsh. These birds are probably individuals which had nested in the interior of the state and concentrated on the coast in the autumn .
Fred S. Walker (1910) reports that he has seen meadowlarks at Pine Point throughout the winter. A flock of 30 to 40 were frequently seen in the adjacent marshes.
In very cold weather, when the grasses and weeds of the marsh were buried beneath the snow, they would venture up to the railway station and pick up grain which had fallen from freight cars. . . . In February, when the marsh was deeply covered with snow, I frequently walked out near the river, scraped off snow from small patches of grass and fed the larks with grain--cracked corn, oats, and barley. They evidently relished this, for it was eagerly devoured. On warm days in January and February they often alighted on the telegraph wires and sang.
In South Carolina the meadowlarks arrive in large numbers in October to take up their winter residence in stubble, corn and cotton fields, and in old fields grown up in weeds and brown sedge. These birds, like those that winter along the Maine and New Jersey coasts, spend the nights in the salt marshes. In various parts of the state they swarm about the rice plantations, where they are often killed by hunters who know the meadowlark as the "marsh quail."
At Buckeye Lake, Ohio, M. B. Trautman (1940) writes of
wintering meadowlarks as follows: "In an average winter 10 to
30 birds could be found during a day's field trip, but when the
species was most numerous as many as 210 were seen in a day. The
wintering birds were found in fields and meadows whenever these
were largely free of snow. When there was deep snow the birds
congregated about manure piles, straw stacks, and in barnyards and
adjacent fields where stock was fed."
Eastern Meadowlark* Sturnella magna
*Original Source: Bent, Arthur Cleveland. 1958. Smithsonian Institution United States National Museum Bulletin 211: 53-80. United States Government Printing Office