Marsh Wren

The Marsh Wren has reddish-brown upperparts boldly streaked with black and white, pale buffy underparts, a whitish line above the eye, mostly unmarked reddish wing coverts, and a short, barred tail that is usually held cocked upright.  Length: 5 inches  Wing span: 6 inches.

Photograph © Greg Lavaty

Sexes similar.

None.

Juveniles are similar to adults.

Marsh Wren inhabit marshes with cattails, rushes, or cordgrass.

Photograph © Greg Lavaty

Marsh Wrens eat insects, but also spiders and snails.

Marsh Wrens forage in dense grasses or sedges.

Marsh Wrens breed from southern Canada south to the upper Midwest. They winter from the southern U.S. south, but are resident on the West Coast. The population may be declining in the east but increasing in the west.

Male Marsh Wrens may build a number of “dummy” nests, from which the female chooses the one to be used.

Male Marsh Wrens may have several mates.

The song consists of a rattling trill.

The Marsh Wren’s nest is an oval mass of sedges or grasses with a side entrance. It is placed low in dense cattails or bulrushes.

Number: Usually lay 4-5 eggs.
Color: Pale brown with darker markings.

Incubation and fledging:
The young hatch at about 13-16 days and fledge at about 12-16 days, though remaining dependent on the adults for another several weeks.

Published by the Smithsonian Institution between the 1920s and the 1950s, the Bent life history series of monographs provide an often colorful description of the birds of North America. Arthur Cleveland Bent was the lead author for the series. The Bent series is a great resource and often includes quotes from early American Ornithologists, including Audubon, Townsend, Wilson, Sutton and many others.

Bent Life History for the Marsh Wren – the common name and sub-species reflect the nomenclature in use at the time the description was written.

The name given by Wilson for this species is now restricted to the long-billed marsh wrens inhabiting a rather limited breeding range on the Atlantic slope from Rhode Island to Virginia. Outram Bangs (1902) has shown that the wrens from this region have the extensively white lower surface, as so clearly depicted in Wilson’s plate; and he has given a new name to the wrens of the interior of New England and the Middle West.

Julian G. Griggs spent a large portion of the summer of 1937 on Jamestown Island in the James River, Va., studying the habits of this wren in some detail. He has kindly sent me his extensive unpublished manuscript, giving the results of his observations, with the privilege of quoting from it. On this island, “about 750 or nearly one-half of the island’s 1,600 acres are marshland, which occupies much of the southern part of the island. Five narrow, parallel ridges of wooded land extend like so many fingers into the marshy area where this study was made. A narrow, branching, brackish, tidal creek extends up into the morass. Dominants in the marsh are Peltandra and marsh grass. The latter dominates creek banks and other slightly raised portions, while the former, because of its greater preference for water, occupies the lower areas of the marsh. Here and there among the grass are groups and individuals of marsh alder, groundsel-tree, knotweed, Kosteletzkya virginica, rosemallow, and swamp milkweed. Cattails and longbills are usually associated together, but this is not the case at Jamestown.”

In other portions of the range of this subspecies, this long-billed marsh wren doe8, occasionally at least, nest in the narrowleaf cattails (Typha augustifolia), notably in coastal Connecticut and coastal Virginia. Chreswell J. Hunt (1904), however, gives a somewhat different impression of the haunts of this wren; his studies were “made along the tidewater creeks which empty into the Delaware River near Philadelphia. These creeks have high and wooded banks on one side, while on the other side for the most part lie low stretches of alder swamps, covered during the summer with a rank growth of spatter-dock, calamus, wild rice, and pickerel-weed, with here and there a clump of rosemallow or a gorgeous cardinal flower. It is here that countless numbers of these little birds find a congenial summer home.”

Aretas A. Saunders writes to me: “Here in Connecticut are many bits of cattail marsh. Some contain both broad and narrow-leaved cattail, and marsh wrens. Others contain only broad-leaved cattail and no marsh wrens. They will nest where there are rushes (Scirpus) or reeds (Phragmites).”

Nesting: The nests studied by Mr. Griggs were mainly in the marsh-grass habitat, and within this area the wren builds its nest in various plants. “The only condition necessary is that there be a place of anchorage 15 or more inches above ground. As a consequence of this, marsh alder and groundsel trees are extensively used for nesting sites. A few nests were found in Hibiscus, and one in some poison ivy, which happened to grow at the right angle. In a situation where alder is common, most nests are made in its branches; likewise, where marsh grass (Spartina) is thick a majority of nests are built among its stems. In the locality examined a majority of wren nests were fairly close to the water, because higher ground there afforded the right habitat. In a slough just north of the island, nests were found over 150 yards from the nearest water.”

The average height above ground of 21 nests measured early in June was 33 inches, one was only 15 inches, and two were 72 and 78 inches up in groundsel-trees (Baccharis). None of the nests in the marsh grass, even later in the season when it averaged 8 feet high, were over 5½ feet above ground; but one was recorded in a groundsel-tree that was 9 feet up. The usual number of dummy nests per female varied from two to four, rather fewer than reported for some other subspecies.

Mr. Hunt (1904) says that—

along the Pensauken Creek each patch of calamus has its pair of Wrens, and each pair build, on an average four nests. In this locality the globular nests are generally built among the calamus stalks or in the crotch of an alder or elder bush. A visit to these swamps, on May 30, 1904, showed each pair of birds to have three nests almost completed, while the foundation for a fourth was in most cases already started. They seem to work on all of them at once. I watched a Wren with a piece of building material in his bill. First he carried it to one nest and started to stick It into that, then he flew away with it to another nest and finally he inserted It into the walls of the third, every little while stopping to sing a snatch of his merry song.

Robert Ridgway (1889) says of the nests that he observed in the marshes of the Potomac River, near Washington, D. C.: “Although usually fastening the nest to upright sedge—or reed-stalks, the writer has found several that were built in small willow trees, at heights varying from 6 to 15 feet above high tide.” Evidently this long-billed marsh wren often differs from other races of the species elsewhere, in its choice of nesting sites.

The long-billed marsh wren must be added to the long list of birds that have been known to use pieces of snakeskin in their nests, for Josiah H. Clark (1899) found a nest near Rutherford, N. J., that was “lined with a cast-off snake skin, which was about a foot long.”

Eggs: This subspecies seems to lay fewer eggs, on the average, than do some other races. Mr. Griggs found that the average for 23 clutches was 4.3. Two nests had 3 eggs, 12 had 4, and 9 had 5 eggs.

I cannot see that there is any difference in the appearance of the eggs from those of the species elsewhere. The measurements of 40 eggs average 16.0 by 12.3 millimeters; the eggs showing the four extremes measure 17.3 by 13.2, 14.2 by 11.7, and 15.2 ‘by 11.2 millimeters. These eggs are all in the United States National Museum.

Having included so much in the life history of the prairie marsh wren, it seems unnecessary to add much more here, as the general habits of these two races are very much alike. Mr. Hunt (1904) has this to say about the midnight song of the long-billed marsh wren: “At all hours of the night the Marsh Wren’s notes may be heard ringing across the marsh. Drifting with the tide, in an open boat, among these swamps I have heard this night song at its best. There is a pleasant surprise in store for the bird-lover who has missed it. This night song is no doubt the same as that sung in the daylight but the night gives to it a certain charm. One must hear it mingled with the quivering call of a Screech Owl and the ‘quawk, quawk’ of Night Herons to fully appreciate it.”

Mr. Griggs adds the following items of interest: “About two-thirds of the nests studied were destroyed between the time of the laying of the first egg and the time at which the young were ready to fly. In a very few cases the nests were damaged somewhat, or a hole was torn in the bottom, but in the great majority eggs and young disappeared without the nest being damaged. About a dozen small rodents were discovered in Jamestown Island nests. One, caught inside a dummy nest, proved to be a rice rat (Oryzomys palustris patustris). Mr. Crook, of Williamsburg, found fragments of eggs and an adult bird, which had been eaten by some small mammal that jumped into the water at his approach. Only a few scattered feathers, the skull, leg, and wing bones of the parent bird remained.”

Only about six watersnakes and about the same number of blacksnakes were seen in the marsh. “At no time was a snake seen suspiciously close to a wren nest. One blacksnake, in a bayberry bush about 15 feet from a nest, was watched for some time. His movements were accompanied by audible rustlings, but the wren never noticed him.”

Wind and rain caused considerable damage to several nests, which the wrens deserted. “Nests in Iva or Bacclaris were often insecurely attached. One such hung by two strands of grass for days. Young wrens were in it at the time and, although every slight breeze caused it to swing from side to side, the parent bird seemed oblivious to the precariousness of the situation.”

He says that territorial boundaries were very flexible. “I have seen males fly back and forth between dummy nests spread out over a distance of a hundred yards or more. Seldom was one wren seen chasing another. At no time was a wren noticed flying after a bird of another species.”

Wayne (1910) says that, in South Carolina, “this form is an abundant autumn, winter, and late spring visitant. My earliest record is September 4, 1895, and the latest May 17, 1897. During the migrations it is most abundant in October and April, when it is commonly found on the salt marshes. In winter, however, the birds prefer the freshwater marshes on the rice plantations, and I have seen more than a hundred individuals in the course of a few hours in such situations.”

DISTRIBUTION

Range: Southern Canada to central Mexico.

Breeding range: The long-billed marsh wren breeds north to southern British Columbia (Chilliwack, Lac la Hache, and Springhouse); northern Alberta (Peace River Landing and the Athabaska Delta); southern Saskatchewan (Prince Albert and Indian Head); southern Manitoba (Lake St. Martin, Winnipeg, and probably Chemawawin); southern Ontario (Lake Nipissing and Ottawa); southern Quebec (Blue Sea Lake and Montreal); and New Brunswick (probably Woodstock and Midgic). East to New Brunswick (probably Midgic); and south through all the Atlantic Coast States to southern Florida (Eldred). South to southern Florida (Eldred and Charlotte Harbor); the Gulf coast to. southeastern Texas (Port Arthur and Cove); southern Illinois (Horseshoe Lake); southern Missouri (Marionville); southern Kansas (Wichita); southern Colorado (Alkali Lakes and Saguache); central Utah (Marysvale); central Nevada (Ruby Lake and Carson); southwestern Arizona (Yuma); and southern California (Calipatria and Escondido). From North Carolina to Texas the species is confined almost entirely to the coastal marshes.

Winter range: The long-billed marsh wren occasionally winters almost as far north as it breeds. The regular winter range is discontinuous. In the west it winters north to southwestern British Cohunbia (Vancouver and the Okanagan Valley). East to southern British Columbia (Okanagan Valley); southeastern Washington (Walla Walla); eastern Oregon (Malbeur Lake); central Utah (Bear River Marshes and St. George); southern Nevada (Searchlight); southern California (opposite Yuma); Sonora (Sonoyta); and Sinaloa (Mazatlan). South to Sinaloa (Mazatlan); and Baja California (Santiago and San Jose del Cabo). West to the Pacific Ocean. The eastern winter range is north to Texas (El Paso, Del Rio, Fort Clark, and Dallas); southern Louisiana (Cameron and Mandeville); western Florida (Pensacola); eastern Georgia (Okefenokee and Augusta); and coastal North Carolina (Swanquarter and Cape Hatteras). East to the Atlantic Ocean. South to southern Florida (Royal Palm Hammock); the Gulf coast to southern Texas (Harlingen and Brownsville); and central Mexico (Veracruz, Jalapa; Hidalgo, Miraflores; and Jalisco, Ocotlan). West to Jalisco (Ocotlan), Chihuahua (Chihuahua), and western Texas (El Paso).

The above range includes the entire species, which has been divided into 10 subspecies or geographic races. The long-billed marsh wren (T. p. palustris) breeds along the Atlantic coast from Rhode Island to Virginia; Wayne’s marsh wren (T. p. waynei) breeds on the coast of North Carolina; Worthington’s marsh wren (T. p. griseus) breeds from South Carolina to Florida, east coast; Marian’s marsh wren (T. p. marianae) is found along the Gulf coast from Charlotte Harbor, Fla., to Mississippi; the Louisiana marsh wren (T. p. thryophilus) occurs in the coastal district of Louisiana and Texas; the Alberta marsh wren (T. p. laingi) breeds in Alberta and western Saskatchewan; the prairie marsh wren (T. p. iliacus) breeds from the Great Plains and Prairie district east to Quebec, New Brunswick, and New England; the western marsh wren (T. p. plesius) breeds from central British Columbia, central Washington, central Oregon, and northeastern California, east to central Colorado; the tule wren (T. p. paludicola) breeds in the coastal district from southwestern British Columbia to southern California; and the Suisun marsh wren (T. p. aestuarinus) breeds in the interior of California from Napa and Solano Counties south to Tulare County.

Spring migration: Some late dates of spring departure are: Florida: Daytona Beach, April 30. Georgia: Athens, May 24. North Carolina: Chapel Hill, May 7. Texas: Lytle, May 12. Early dates of spring arrival are: Georgia: Macon, March 27. North Carolina: Raleigh, April 14. District of Columbia: Washington, April 17. Pennsylvania: Harrisburg, April 26. New Jersey: Cape May, April 28. New York: Rochester, April 27. Massachusetts: Cambridge, April 23. Vermont: Burlington, May 12. Kentucky: Bowling Green, May 5. Ohio: Sandusky, April 19. Ontario: Hamilton, April 6. Indiana: Bloomington, April 7. Michigan: Ann Arbor, April 20. Illinois: Chicago, April 25. Wisconsin: Madison, April 11. Missouri: St. Louis, April 13. Iowa: Sioux City, April 17. Minnesota: Minneapolis, April 20. Manitoba: Margaret, May 3. Kansas: Harper, April 4. Nebraska: Lincoln, April 6. South Dakota: Yankton, April 23. North Dakota: Wahpeton, April 17. New Mexico: Albuquerque, April 6. Colorado-Boulder, April 3.

Fall migration: Some late dates of fall departures are: British Columbia: Okanagan Landing, November 18. Wyoming: Laramie, October 15. Colorado-Boulder, October 22. New Mexico: Mesilla, October 16. Manitoba: Aweme, October 16. North Dakota: Argusville, October 21. South Dakota: Sioux Falls, October 26. Nebraska: Nebraska City, November 12. Kansas: Lawrence, October 22. Minnesota: St. Vincent, October 15. Wisconsin: Milwaukee, October 23. Iowa: Keokuk, November 21. Missouri: Forsyth, October 2. Michigan: Vicksburg, November 6. Illinois: La Grange, October 17. Indiana: Indianapolis, November 9. Ontario-Toronto, October 15. Ohio-Oberlin, November 19. Kentucky: Danville, October 27. Massachusetts: Boston, November 30. New York: New York, October 20. New Jersey: Elizabeth, October 20. Pennsylvania: Jeffersonville, October 29. District of Columbia: Washington, November 16. North Carolina: Raleigh, October 19. Georgia: Athens, October 11.

Early dates of fall arrival are: North Carolina: Chapel Hill, September 20. Georgia: Athens, September 24. Florida: Fernandina, September 17. Texas: Lytle, October 10.

Casual record: A specimen was collected at Godthaab, Greenland, in May 1823.

Egg dates: Alberta: 28 records, May 26 to July 8; 18 records, June 16 to 20.

California: 113 records, March 24 to July 22; 63 records, May 4 to 31, indicating the height of the season.

Florida: 39 records, May 2 to August 13; 20 records, May 24 to June 11.

Illinois: 48 records, May 26 to July 27; 29 records, May 30 to June 8.

Massachusetts: 31 records, May 27 to June 28; 17 records, June 6 to 18.

New York: 60 records, May 12 to August 13; 42 records, June 4 to 24.

Virginia: 37 records, April 25 to June 30; 26 records, June 6 to 26.

Washington: 31 records, March 15 to June 27; 16 records, March 15 to April 22; 10 records, June 8 to 27. Wisconsin: 12 records, May 26 to July 14.

WORTHINGTON’S MARSH WREN

TELMATODYTES PALUSTRIS GRISEUS (Brewster)

HABITS

This is the rather small, decidedly gray race of the long-billed marsh wrens that is apparently resident in the Atlantic coast region from South Carolina to northern Florida.

In naming this race, William Brewster (1893) compares it with the northern race as follows: “Black of upper parts much duller and less extended than in palustris, usually confined to the extreme sides of the crown and a short narrow area in the middle of the back, and in extreme specimens almost wholly absent. Brown of sides, flanks, and upper parts pale and grayish. Dark markings of the under tail-coverts, flanks, sides, and breast faint, confused and inconspicuous, sometimes practically wanting.”

The haunts and habits of Worthington’s marsh wren are very similar to those of the more northern coastal race. It is confined almost entirely during the breeding season, and probably for the rest of the year, to the extensive salt marshes along the tidal creeks. Wayne (1910) says that, near Charleston, S. C., when he was a boy, these “birds fairly swarmed throughout the high marshes bordering these creeks and it was not uncommon to find from 25 to 50 nests in a few hours of careful search.” In Florida, according to Arthur H. Howell (1932), “the birds live in the wettest and boggiest parts of the salt marshes, chiefly on the borders of the tidal creeks, where their nests are fastened to the growing stems of the rushes, at a height of 2 or 3 feet above the water.” Ivan B. Tomkins (1932) has found it along the Savannah River to about 2 miles west of Savannah, or about 17 miles inland from the outer islands, including the river ricefields, where the water is either fresh or brackish according to the height of the river.

Nesting: Mr. Howell says that these wrens “breed in loose colonies, often of considerable size.” The nests “are constructed of dead leaves of rushes and marsh grasses woven together and lined with fine grasses and down from the cattails.” What few nests of this wren I have seen are similar to nests of the species I have seen elsewhere. A nest m my collection was taken by Mr. Worthington in Nassau County, Fla.; it was 3 feet above the mud, suspended amongst the grass, near the edge of a creek in a salt marsh; it held six fresh eggs on July 6, 1906.

Eggs: What eggs I have seen are similar to the eggs of other long-billed marsh wrens, showing the usual variations. Mr. Wayne (1910), however, says: “On several occasions, between the years 1877 and 1879, I remember distinctly having found pure white eggs of this form with a speck or two of purplish shell markings at the larger end. The eggs are, however, normally chocolate color, but sometimes of a paler shade and spotted with brownish olive. From four to six eggs are laid. * * * Three broods are certainly raised, for I have taken eggs as late as August 9.”

The measurements of 40 eggs in the United States National Museum average 15.4 by 11.2 millimeters; the eggs showing the four extremes measure 16.5 by 12.2, 15.5 by 12.5, and 14.1 by 11.1 millimeters.

I can find nothing in the recorded habits of this wren that is peculiar to the race. Mr. Wayne says that—

a mouse (Hesperomys leucopus), which lives in the marsh and builds a nest similar to that of the wren, commonly takes possession of the nest and often eats the eggs as well as the young. * * * At the present time the birds are rare and confined to a few restricted and widely separated localities, the great cyclone of August 27 and 28, 1893, having almost exterminated them. This form is non-migratory, and I understand that it is abundant on the coast of Georgia. If the birds were migratory the places of those that were destroyed by the cyclone of 1893 would be filled by migrants from Florida and Georgia. This, however, has not been the case, showing conclusively that this form is nonmigratory.

MARIAN’S MARSH WREN

TELMATODYTES PALUSTRIS MARIANAE (Scott)

HABITS

Although much of the earlier literature on the status and distribution of Marian’s marsh wren is decidedly confusing, it now seems safe to conclude that it is the resident, breeding form on the west coast of Florida. The 1931 Check-list gives as its range, “Gulf coast from Charlotte Harbor, Florida, to Mississippi.” The latest authority on Florida birds, Arthur H. Howell (1932) calls it “an abundant resident on the Gulf coast from St. Marks south to Old Tampa Bay. * * * Marsh Wrens are not known to breed at Pensacola, and we found no breeding colonies from that point eastward until we reached St. Marks; from there southward they are abundant in suitable marshes as far as Tarpon Springs. Pennock reported a few birds seen in Charlotte Harbor, April 11 and 13, 1921, at which time he took a specimen and observed a nearly completed nest.”

The Rev. H. E. Wheeler (1931) made a survey of the breeding colonies of Marian’s marsh wren on the coast of Alabama, establishing the fact that this wren breeds in all suitable coastal marshes throughout the whole Gulf coast of that State, but he evidently found no breeding records for Mississippi. He, also, made an exhaustive study of the previous literature relating to this wren on the Gulf coast and to its supposed occurrence on the Atlantic coast, which is well worth reading, as an interesting history of the confusion that has existed, and to which the reader is referred, as space will permit only a few quotations from his excellent paper.

A perusal of the literature that appeared prior to 1932 will reveal some confusing statements. Dr. Louis B. Bishop (1904) made the following surprising comment: “Originally described from the west coast of Florida and believed to be resident there and restricted to that locality, it has only recently become evident that the real home of Marian’s Wren is in the salt marshes that fringe the coast of North Carolina. There it is common in spring, breeds, and occasionally remains in winter, as I took one on Pea Island: 30 miles north of Hatteras: on Feb. 8, 1901, and found it tolerably common there in January, 1904.” He found fresh nests there in May, and, after he left, nests with eggs were sent to him from there.

Ridgway (1904) gave the range as south Atlantic coast of United States from North Carolina to South Carolina and western Florida. And the 1910 Check-list gave it as breeding on the coast of North Carolina and wintering south to South Carolina and the west coast of Florida. Arthur T. Wayne (1910) backs up Dr. Bishop by saying that “it breeds, as far as is known, only on the North Carolina coast.” He records it as a migrant in South Carolina, where he says that “they are common until the beginning of November, when the great majority migrate southward, but a few winter regularly among dense reeds which grow in profusion on some of the coast islands.” Strangely enough, the 1931 Check-list does not record ~my marsh wren as breeding on the coast of North Carolina! It has since become evident that all the Atlantic coast records of supposed Marian’s marsh wrens, published prior to 1930, are referable to the new dark-colored race, Telmatodytes palustris waynei, recently described by Dingle and Sprunt (1932). This race is so strikingly similar to marianae in most of its characters that the confusing of the two is not surprising. The status of waynei as the breeding marsh wren of the North Carolina coast has recently been confirmed by Dr. Alexander Wetmore (1941) and his assistants, as one of the results of their field work there during 1939.

In his original description of marianae, W. E. D. Scott (1888b) says: “The great difference between this species and palustris is in the conspicuous barring of the upper and under tail-coverts and the feathers of the flanks, and olive instead of rufous brown coloring throughout, with the much darker coloration of underparts.” The differences between marianae and waynei are not so well marked and are much less conspicuous. Dingle and Sprunt (1932) say: “A satisfactory comparison of Telmatodytes palustrus waynei with marianae is not possible on account of inadequacy of specimens of the latter. In size, waynei seems to be slightly larger than the Florida form; in color it is quite similar, except that there is more white on the under parts.”

In order to satisfy me as to the characters that separate waynei from marianae, James L. Peters and James C. Greenway helped me examine the series of both forms in the Museum of Comparative Zoology in Cambridge, where there are 31 specimens from the west coast of Florida and 18 specimens from North and South Carolina. It seems to me that, in the Florida birds, the sides of the head and neck average darker, the black space on the back is rather more extensive and the flanks are browner than in the Carolina birds; also, the breast, in adults at least, is more inclined to be mottled with dusky, and there is much less white on the under parts. These are only average differences and are rather slight, but the Carolina bird seems to be far enough removed geographically to warrant its recognition as a subspecies.

Ridgway (1904) gives the best description of Marian’s marsh wren as follows:

Similar to T. p. palustris, but smaller, the coloration much darker; pileum usually entirely black or with black largely predominating; white streaks of interscapular region narrower, sometimes almost obliterated; brown of scapulars, rump, etc., darker, the upper tail-coverts (sometimes the whole rump) usually barred with dusky; sides and flanks more extensively, and usually darker, brown than in T. p. palustris, the chest often strongly shaded pale brown or brownish buff; frequently the chest or sides (or both) are speckled with dusky, and sometimes the sides and flanks are barred with darker brown; under tail-coverts distinctly, often broadly, barred with brown or dusky; mandible usually dusky for much the greater part of its length.

As to its haunts in the vicinity of Tarpon Springs, Fla., the type locality, W. E. D. Scott (1890) says: “I have found them most commonly on the salt water marshes at the head of tide water, but have detected them in the saw-grasses of the fresh water lakes and ponds that I have investigated for at least ten miles back from the coast.”

Nesting: Arthur H. Howell (1932) writes: “In the extensive salt marshes at the mouth of Pithlachascotee River, near Port Richey, we found these Wrens breeding commonly in the dense growth of Juncus, standing 4 to 5 feet high. On May 28, 1918, we collected several well-grown young birds. In the marsh at Elvers, June 2, 1919, D. J. Nicholson observed several nests from 5 to 9 feet above the ground in mangrove trees.”

There are two sets of eggs in my collection, taken by C. J. Pennock, near St. Marks, Fla., that came from nests 2 feet above the ground in saw grass.

D. J. Nicholson wrote to Mr. Wheeler (1931) that these wrens near New Port Richey “nest among Juncus (J. roemerianus), a sharp-pointed rush, and principally in mangrove trees from 5 to 14 feet above the mud in salt marshes. The tree-nesting may seem strange to you and it was quite a surprise to me when I found them nesting under such odd circumstances. I think high water and rats had something to do with this nesting custom here, and it may be a comparatively recent habit.” From the same source of information, Mr. Wheeler goes on to say:

Nests of rnarianae found in mangrove trees were fastened to the forks of small limbs, generally at their ends, or in the tops of small mangrove bushes. * * * Occupied nests are lined with soft shredded grasses, and sometimes with feathers, and they are so cleverly woven together that they are a complete protection against rain. None have even been found that were damp inside. Although the marsh wrens nest in colonies, the nests of marianae are seldom less than 40 feet apart. On the east coast of Florida Nicholson counted four to six “dummy nests” to every occupied nest of griseus; but In the colonies of marianae on the west coast near Elfers, he found only one or two bachelor nests to one that was occupied.

Referring to the coast of Alabama, he writes:

It was on the tidal flats, or rather monotypic marshes, of Heron Bayou that we found marsh wrens nesting, enough to satisfy the heart of any ornithologist. This region of vast and almost impenetrable marshes Is known to the fishermen as West Heron Bay. Several narrow bayous penetrate the grass-grown region, one of them widening into a so-called lake. In such a region, in the tall bladed grasses, which grow higher than the rushes, and nearer open water, we found the marsh wrens numerous. They were singing near their neatly built nests, their entrancing songs being much in the tempo of the songs of the Prairie Marsh Wren. * * *

The nest of Marian’s Marsh Wren differs in no essential way from the nest of other closely-related species or subspecies. It is globular in shape, well secured to the taller marsh grasses, and usually about 2 or 3 feet above high tide. Oftentimes the nest can be detected from a moving skiff. The bachelor nests, which are unlined, are in the proportion of four or five to one which is lined and occupied.

Eggs: Marian’s marsh wren seems to lay fewer eggs than the northern races of the species, usually three to five. These are like the eggs of the species elsewhere and show the usual variations, some being quite pale, but most of them being of a deep, rich chocolate-brown; many have a wreath of darker spots about the larger end. The measurements of 40 eggs average 15.4 by 12.2 millimeters; the eggs showing the four extremes measure 16.4 by 12.7, 16.1 ‘by 12.9, 14.2 by 11.9, and 15.0 by 11.0 millimeters.

Behavior: Mr. Wheeler (1931) writes: “Contrary to our expectation, we did not find these wrens particularly shy. The breeding birds were very easy to approach; and although they did not remain long on open perches they seemed quite unmindful of our invasion of their territory, singing joyously all the while, and often within 2 or 3 feet of us. Often and again the males would reappear and perch in plain view on the side of the tallest reed, and that without interruption of their song. If we could have walked through the thick vegetation at low tide with a Graflex camera, we might have gotten pictures of the birds in action.”

WAYNE’S MARSH WREN

TELMATODYTES PALUSTRIS WAYNEI  (Dingle and Sprunt)

Comment on this subspecies was made under Marian’s marsh wren, to which the reader is referred. Since that account was written, the above subspecies has been accepted and will appear in the next Checklist.

Dingle and Sprunt (1932) describe it as “similar to Telmatodytes palustris palustris, but smaller; bill shorter and more slender; wing, tail and tarsus average shorter; upper parts darker, inclining more to olive brown; head and nape sooty black, the majority of specimens showing a short, faint median streak; black dorsal area of greater extent; tail and under tail coverts more heavily barred; flanks richer brown; these, and sides of breast with more or less barring.”

“A satisfactory comparison of Telmatodytes palustris waynei with marianae,” these authors say, “is not possible on account of inadequacy of specimens of the latter. In size, waynei seems to be slightly larger than the Florida form; in color it is quite similar, except that there is more white on the under parts.”

The breeding range of this form seems to be on the coast of North Carolina, and it is found in South Carolina and Georgia in winter. See remarks under marianae. Its habits are apparently no different from those of the other coastal races of the species.

LOUISIANA MARSH WREN

TELMATODYTES PALUSTRIS THRYOPHILUS Oberholser

HABITS

The long-billed marsh wren of the coast region of southern Louisiana is very much like Marian’s marsh wren and evidently closely related to it. Ridgway (1904) makes this comparison: “Similar to T. p. marianae, but paler and still smaller; pileum always extensively brown medially, often mostly brown; brown of scapulars, rump, etc., lighter, sometimes approaching broccoli brown or drab; upper tail-coverts unbarred, or with bars very indistinct; under parts never (?) speckled, but chest more or less strongly tinged with brownish buff, and sides and flanks extensively brown. Differing from T. p. palustris in decidedly smaller size, duller brown of upper parts, and more extensively brown under parts.”

Dr. Oberholser (1938) says that “it lives in the marshes and in the high grass of the coast meadows, among the reeds, rushes, grasses, and similar kinds of vegetation,” which indicates that it is similar in all its habits to the other southern coastal races.

The measurements of only five eggs of the Louisiana marsh wren are available. The eggs are in the United States National Museum. The eggs showing the four extremes measure 16.0 by 12.4 and 15.3 by 11.5 millimeters.

ALBERTA MARSH WREN

TELMATODYTES PALUSTRIS LAINGI Harper

HABITS

Dr. Francis Harper (1926) describes this wren as “nearest to T. p. iliacus, but paler on scapulars, rump, upper tail-coverts, and flanks; median area on forehead and crown more distinct. (T. p. plesius is a much browner and duller bird than laingi.)”

The Athabaska Delta, where the type specimen of this race was taken, is probably the northernmost point at which any long-billed marsh wrens breed. Dr. Harper gives, as the range of this subspecies in summer, Alberta and western Saskatchewan, and says that it seems to intergrade with the prairie marsh wren in south-central Saskatchewan, and that the area of intergradation “may coincide with the approximate boundary between the prairies on the east and the plains on the west.”

The habits of the Alberta marsh wren are apparently similar in all respects to those of the prairie marsh wren.

There is a very pretty nest of this wren in the Thayer collection in Cambridge, taken near Fort Saskatchewan, Alberta, on June 2,1900. It was “attached to tules in a muskeg” and was constructed chiefly of a downy substance that looks like cattail down, very compactly felted and reinforced with interwoven strips of the tules, or other marsh plants, which bound the whole structure very firmly together; its walls are so thick and solid that it must have been practically rainproof; it is about 7 inches high and about 4 inches in diameter. There is another nest, from the Little Red Deer River, Alberta, that is more normal for the species, having been attached to the stems of bulrushes and made of the usual materials.

A. D. Henderson, of Belvedere, Alberta, writes to me of an experience that was new to him: “On July 8 I pushed my canoe into a large bed of tules where marsh wrens were singing, leaving it close to an empty nest. To enable me to find the canoe again without difficulty after wading the tule bed, I attached a bunch of white cotton to the tops of tall tules. On July 12 I returned to the same spot and found that the wren had profusely bedecked the nest with the cotton I had left nearby. It was put on quite loosely and not woven into the structure.”

The eggs of the Alberta marsh wren apparently show the usual variations common to the species. The measurements of 23 eggs average 16.3 by 12.3 millimeters; the eggs showing the four extremes measure 16.8 by 12.2, 16.4 by 13.1, 15.2 by 12.7, and 15.9 by 11.9 millimeters.

PRAIRIE MARSH WREN

TELMATODYTES PALUSTRIS ILIACUS Ridgway

HABITS

The old familiar type name, long-billed marsh wren (T. p. palustris), has been restricted to the wrens of this species living on the Atlantic slope from Rhode Island to Virginia, and the birds of interior New England and the middle west are now known by the above names. Such are the vagaries of name-shifting that our old friend of the Massachusetts marshes is now called the “prairie” marsh wren, though hundreds of miles away from the nearest prairies! In describing this race, Mr. Bangs (1902) writes:

At present there are confused under the name Cistothorus palustris (Wilson) two quite distinct birds; one, true C. palustris, breeding in the salt and brackish marshes of the Atlantic coast from Connecticut southward; the other inhabiting the inland fresh-water marshes and extending north to Massachusetts, Ontario and southern Manitoba. The former, a small bird, has the chin, throat and belly pure white and the breast is usually white also, though sometimes faintly clouded with pale brownish, with the rump, upper tail-coverts and scapulars dusky brown. The latter is a decidedly larger form, in which the chin, throat and belly are buffy or brownish white, the breast much more distinctly clouded with brownish and the rump, upper tail-coverts and, scapulars reddish brown.

Wilson’s plate shows a decidedly white-breasted bird, to which he gave the name palustris; there can be no doubt, therefore, that the Atlantic coast bird should carry the type name.

The prairie marsh wren is naturally not evenly distributed throughout its wide range. Marshes of the type it requires are often widely scattered, or entirely lacking over large areas. Small, isolated marshes of less than an acre in extent are usually avoided, but where the larger marshes contain suitable vegetation the wrens may be very numerous and their nests more so.

The favorite haunts of the prairie marsh wren are the large freshwater marshes of the interior, where there is a dense growth of cattails (Typha angustifotia and T. latifolia) , bulrushes (Scirpus lacustris), sedges (Carex), or wildrice (Zizania aquatica), which are often mixed with tall marsh grasses of various kinds, or with a scattering growth of buttonbush (Cephalanthus) and other small bushes. In eastern Massachusetts we sometimes find them along the banks of tidal rivers, where the water is brackish and where there is a thick growth of tall reeds and salt-marsh grasses. I have found them, also, in pure stands of wildrice bordering a sluggish inland river.

Dr. Charles W. Townsend (1905) tells of a large marsh in Essex County, Mass., in which “the growth of rushes and grasses is rank and tall, and among these a multitude of Long-billed Marsh Wrens live and build their nests. The rush-like plants in which they breed are chiefly as follows, belonging to several widely separated families: great bulrush (Scirpus lacustris), horse-tail (Equisetum timonsum), sweet flag (Acorus calamus), blue joint-grass (Calamagrostis canadensis), reed canary-grass (Phalaris arundinacea).”

Spring: Very little seems to be known about the migrations of the marsh wrens. Elon H. Eaton (1914) says: “Evidently they migrate at night, and high in the air, so as to see their way and escape their enemies more successfully.” They arrive in central New York from May 4 to 16.

Dr. Wilfred A. Welter (1935) has given us such a fine life history of the prairie marsh wren, based on extensive observations at Ithaca, N. Y., and at Staples, Mimi., that I cannot do better than to quote from the results of his work. At both places he found that the average date for the arrival of the males was May 10 and that the females came between May 20 and 28. Males begin to select and defend their breeding territories soon after their arrival. He says:

The preferred habitat is not, as one might suppose, a dense tangled mass of dried and broken cat-tails, remnants of the preceding season, but a comparatively open area with a few tattered stalks and an abundance of some species of Carex.

* * * Fighting over territorial rights between males Is, to a large extent, a matter of outbluffing the opponent. A male approaching too closely to the boundary of another’s area is challenged by the song of the rightful owner. This is usually sufficient for the intruder, but sometimes the challenge Is accepted by the visitor giving voice to his emotions and continuing to transgress upon the area In question. The first male in this case fluffs out his feathers to impress the other and, if necessary, flies at his opponent. The usurper usually reciprocates by flying at his neighbor a time or two and then, at least in all instances observed, becomes the vanquished and departs from the scene of battle.

* * * In an area 400 by 650 feet in the Renwick Marsh at the head of Lake Cayuga eight males took up residence In the spring of 1931. * * * The cattail-sedge association was greatly preferred to the grass association by the male birds in selecting territories. * * * Typha augustifolia is much preferred to T. latifolia as a nesting site. * * * The male territories in the favored area were noticeably smaller than in the grassy area. A single monogamous male occupied a territory of from 13,000 to 15,000 square feet, while in the grass association this was extended to approximately 30,000 square feet. The territory of a polygamous male, on the other hand, was considerably larger than that of a monogamous male nesting in the same sort of vegetation. * * * This difference in size can readily be accounted for by the fact that the female birds do not tolerate each other during the nesting season. As a result those males intent upon leading dual lives must separate the objects of their affection as widely as possible.

Courtship: The courtship of the marsh wren is expressed in song and in display. According to Dr. Welter (1935), “song does not seem to be as important in attracting the female as display. Of course the song originally attracts the prospective mate into the territory and then display becomes first in importance. When the females begin to arrive from the south the males sing almost constantly.” The songs at this tune often average about 25 per minute, but during nest building the songs are less numerous and the intervals between singing periods become longer.

“The display of the male is quite simple but interesting. When the female is near he will take up his station a foot or two above her, fluff out his breast feathers and under tail coverts, and jauntily cock his tail over his back so that it almost touches. He now resembles a tiny ball of feathers perched among the reeds. As he becomes more animated he beats his partially folded wings up and down rapidly and sways his head dizzily from side to side. The female probably will fail to notice him, or at least she will not indicate any interest, and, after pursuing her and displaying for several minutes, he will burst into song and fly to another portion of the territory.”

The sexual organs of the male are well developed when he arrives, but those of the female are not, so that she has to avoid him until she is ready. Dr. Welter continues:

During the period of nest construction she reaches the height of her development and is ready for the mating act. When the male approaches her at this dine, singing, she climbs up a cat-tail stalk and gives the trili which has already been described. Then she beats her wings rapidly, points her bill toward the zenith, and places her tail well over her back. The male goes through the courtship display previously described. At the proper time he climbs upon the back of his mate, heats his wings rapidly as the cloacae come in contact and copulation is completed. The whole procedure takes but a few seconds. Both remain in the immediate vicinity for a short time, the male with feathers fluffed out and tail up, the female quiet and demure.

It is usually the male who tries to induce the female into copulation but on one occasion the female was observed going through the behavior leading to the mating act to entice the male. In this instance the act had been completed 25 minutes previously. The male, not giving the proper response, was chased by the female among the eat-tails and it Is not known whether she was successful or not.

Dr. Welter believes that the male is “essentially polygamous while the female is not.” Several of the territories were inhabited by one male and two females, and in one doubtful case it was thought that a male had three mates. There was another doubtful case of polyandry, where a female had no regular mate, and her nest was placed between the territories of two mated males.

Nesting: The prairie marsh wren nests in wet marshes, where the water is from a few inches to 2 or 3 feet deep, along the banks of tidal rivers where the water is brackish (in Massachusetts), along sluggish inland streams, around the shores of ponds, and in inland marshes or sloughs. It seems to prefer to build its nest in the narrowleaf cattail (Typha angustifolia), seldom using the broadleaf species (T. latifolia). Early in the season, before the green flags have grown to sufficient height, I have found the nest in some thick bunch of the dead flags of the previous season, but the new green flags are much preferred. The nests are usually placed 1 to 3 feet above water, seldom higher, and are securely fastened to two or more stems of the cattails.

Nests are less often placed in bulrushes (Scirpus), sedges (Carex), wildrice (Zizania), tall marsh grasses, or even small bushes. In North Dakota we found these wrens nesting around the edges of the sloughs in either dead or green cattails, or in the bulrushes. Near Lake Winnipegosis, Manitoba, we found a nest firmly attached to the canes of bulrushes, 4 feet above the water; it was within 4 feet of a canvasback’s nest and was lined with down from the nest of the duck. The nest is said to be shaped like a coconut, or globular, but some that I have seen have been egg-shaped with the pointed end at the bottom. The entrance is a small round hole, usually near the top.

Dr. Welter (1935) gives an elaborate account of the building of the brood nest, which is done almost entirely by the female, and which requires 5 to 8 days, beginning 6 to 15 days after her arrival. He writes:

The initial effort in building consists of lashing the supporting plants together and In this way form a cup-like foundation upon which the remainder of the nest rests. Carex and Calamgrostis are the chief materials used in this part of the structure. The outer walls which are composed for the most part of long strips of cattail leaves and stems and leaves of sedges and grasses is the roughest part of the structure. Water-soaked materials, often more than a foot long, are used here as they are more pliable and can more easily be woven together. The first strands are woven around the long axis and others, as the nest assumes shape, are put in at various angles. Some of these strands are fastened to the supporting structure by actually weaving these stems into the nest. Some of the growing leaves are also woven into the outer walls. If the support is a sedge or a grass, leaves may form a good share of the periphery. An opening is left on one side about two-thirds of the distance from the bottom of the nest. At this stage a dummy would be complete. The walls average at least a half inch in thickness and the external measurements of the entire structure approximate seven and five inches for the vertical and horizontal diameters, respectively. Inner diameters average five and three inches.

The outer shell is a small part of the completed structure, and only 2 days are required to build it. The remainder of the work is done from the inside and one must take a nest to pieces to get an idea of its arrangement. Grass and sedge leaves and small stems are used to form the second layer. This gives the walls firmness and tends to fill in the large air spaces which are necessarily present among the coarse materials of the outer walls.

The next layer to be added seems to function as an insulating region. Cat-tail down, feathers, small unidentified rootlets, entire plants of Lemna, and decayed fragments of Typha and Carex are the materials most often used. These are also placed into the structure in a wet condition so that, when dry, they form a compact and tight-fitting region which serves as a non-conductor of heat, cold, and moisture.

The innermost region is composed of finely shredded pieces of the vascular materials of the plants forming the outer layers. A large proportion of it is very fine strips of sedges and grasses of the preceding year. Feathers of almost any available sort are used here. Those from the following birds have been identified: Red-winged Blackbird, Virginia Rail, American Bittern, Pheasant, Ruffed Grouse, and domestic chicken. The projection at the opening is a part of this inner lining. This “door-step” or sill is always present in the female nest but is lacking in the nests of the male. It is possible, therefore, to determine the sex which built a given nest l)y checking for the presence of this sill. This projection forms the floor of the opening and extends farther into the nest than any other part of the lining. * * *

One wonders what the function of this door-step might he. Perhaps it serves as a protection to the eggs and young as the nest, owing to the uneven growth of the supporting plants, often assumes a distorted position which would allow the contents to roll out were it not for this structure. In like manner when the nests are placed in sedges or grasses winds alter the nests to such an extent that the young or eggs would be endangered if no sill were there to prevent the catastrophe.

Several observers have reported mud in the lining of the nests, but Dr. Welter and others have failed to note it; perhaps some mud may be brought in accidentally with material secured from the muddy floor of the marsh; it seems doubtful if the wrens every carry in mud intentionally.

The long-billed marsh wrens are notorious for building extra or dummy nests, which are almost never occupied as brood nests. These are built by the males, mainly during the 10 days or so intervening between the arrival of the males and the coming of the females. Anywhere from 1 to 10, usually not more than half a dozen, are more or less incompletely constructed by a single male within the limits of his territory. We do not fully understand the reason for these extra nests; several theories have been advanced to account for the habit, which is not wholly confined to this species, but none of the theories appears wholly satisfactory. The most plausible theory seems to be that it gives the birds an outlet for surplus energy during the period of sexual activity, for it almost always ends soon after the females arrive and mating takes place. These male nests are never as fully completed as are the brood nests; they usually do not go beyond the first stage mentioned above, and are often abandoned before they reach even that stage of completion. There is little evidence that they are ever used as brood nests, or as sleeping places for the males, or as territorial land marks.

A. D. DuBois mentions in his notes a nest that was of the “usual construction except that the top of the nest was covered by green leaves bent over and woven together over the top. All the previous nests observed here, having the green leaves woven over (nearly a dozen so noted) were empty nests.”

Milton B. Trautman (1940) noted that, out of 208 nests, observed at Buckeye Lake, Ohio, 161 had their openings facing toward the south or west. There was one colony, “which was an exception, for 11 of 19 nests opened toward the northeast.”

Eggs: The marsh wren’s set may consist of 3 to 10 eggs; the larger numbers are rare; 5 or 6 seem to be the commonest numbers. They are generally ovate, sometimes more rounded and rarely more pointed; they are not glossy unless heavily incubated. Marsh wrens’ eggs are unique in color, the general effect being dull brownish, “Verona brown” to “snuff brown,” or the color of dry, powdered baking chocolate. The ground color varies from “snuff brown” to pale “pinkish cinnamon”; it is generally evenly sprinkled with minute dots, or very small spots of darker shades of brown, often partially, or wholly obscuring the ground color; these markings are sometimes concentrated into a ring or a cap at the large end. F. W. Braund tells me that “light or stony gray” eggs are often found in Ohio. I have seen eggs with a pinkish ground color and reddish brown spots that resembled the eggs of the house wren, but these are rare. Very rarely an egg, or a whole set of eggs, is pure white and unmarked.

The measurements of 40 eggs average 16.5 by 12.4 millimeters; the eggs showing the four extremes measure 17.8 by 12.1, 17.6 by 13.3, 15.0 by 13.0, and 17.6 by 11.2 millimeters.

Young: The incubation period, as noted by several observers, is about 13 days, and the young remain in the nest for about the same length of time, or a day longer, if not disturbed; Dr. Welter (1935) says 14 days. Incubation seems to be performed wholly by the female, and she feeds the young while they are in the nest; the male assists in this afterward. Following are some of Dr. Welter’s observations on the young:

The type of food delivered to the young by the female is determined to a certain extent by the age of the nestlings. At first this consists of very small juicy morsels such as mosquitoes and their larvae, larval Tipulids, midges, and ether delicate forms. The mother brings a whole beakful of food to the nest at one time and parcels it out to the hungry occupants. * * * During the morning and evening approximately 10 trips are made per hour with food, but during midday this number is somewhat reduced.

As the nestlings grow the insects brought to the nest become appreciably larger in size. Ground, diving, and long-horned beetles, caterpillars of various assortments, sawflies and other hymenoptera, and other accessible forms now constitute the diet of the ever-hungry young. Sometimes the insect is so large that the young bird experiences difficulties in swallowing it. In such Instances the female takes the hexapod to the side of the nest, chops and tears it into several smaller morsels, and then brings It back for a second trial which is usually a success. * * *

Even when the nestlings are very young, little time during the day is given to brooding. Usually after a feeding or two the young are brooded for a few minutes and then feeding is resumed. My records show a total brooding of 18 minutes per hour when the young are 2 days old. As the nestlings increase in size the brooding periods become shorter and the intervals between such periods become longer, so that, after the first week, they are discontinued during the hours of daylight. * * *

The excreta, enclosed in their envelopes, are removed by the female after feeding. These droppings are usually carried some distance from the nest and deposited, but occasionally the female has been observed eating them. * * *

When the young are small the faecal material is deposited in the bottom of the nest. As the nestlings increase in size, however, they maneuver about until they assume a position facing away from the entrance, and the dropping is ejected on the periphery or the nest. During the later period of nest lire the young succeed in ejecting the excrement with such force that it is carried over the side of the nest and drops to the ground.

Other waste materials, such as eggshells, infertile eggs, or any young that might die in the nest are carried away. The young increase in weight very rapidly, from about 0.87 gram at hatching to about 11.08 grams at the end of the twelfth day. Meantime the nest has become enlarged and worn as the young increase in size. The young may leave the nest on the twelfth day, if disturbed, but normally not until the fourteenth day. Occasionally one will return to the nest for shelter, but they usually spend the nights perched in the dense flags. The parents care for them for at least 2 weeks, though after the first 10 days they are able to secure some of their own food. The family group remains together through the summer and wanders about at some distance from the nesting place.

It seems to be the consensus that two broods are raised in a season, but not a third. Dr. Welter (1935) found no evidence of a third brood. “The female begins her second nest about 2 weeks after the young of the first have left the nest. The majority of the nests, then, in the regions studied would be started between July 15 and August 1, with the last week in July the most active period.” Probably while the female is building the second nest the male is busy with the first brood and is not very active in building dummy nests.

Plumages: Dr. Welter (1936) has published another excellent paper on the development of the plumage in the young marsh wren and on subsequent molts, to which the reader is referred for details; it is fully illustrated with drawings and photographic halftones. It is evident from the photographs that the young bird is practically fully feathered in the juvenal plumage before it leaves the nest, though the wings are Lot fully developed and the tail is still rudimentary. Dr. Dwight (1900) says that the natal down is white. In the juvenal plumage the young wren is much like the adult, but the crown is uniformly dull black, without the dividing brown area; the white streaks on the back are very faint or lacking; and the white superciliary stripe is indistinct. Dr. Dwight says that the first winter plumage is “acquired by a partial postjuvenal molt beginning about the middle of August which involves the body plumage, the wing coverts, probably the tertiaries, but not the rest of the wings nor the tail,” young and old becoming practically indistinguishable. Dr. Welter (1936) differs from Dr. Dwight, as to the extent of this molt, saying: “Juvenals collected during the fall of 1931 which are now in the Cornell Collection show a molt of both rectrices and remiges.” These two authorities also differ as to the prenuptial molt. Dr. Dwight says that the nuptial plumage, in both adults and young birds, “is acquired by a complete prenuptial moult as indicated by the relatively unworn condition of the feathers when the birds arrive in May.” He had no positive evidence of the molt, however. Dr. Welter could “find no evidence of a prenuptial molt in the series of specimens examined.” The plumage of birds living in such dense vegetation must be subjected to rather severe abrasion, which might require a renewal of plumage oftener than once a year; and it may be that the prenuptial molt takes place during the late winter or very early spring, before the birds arrive on their breeding grounds. Dr. Witmer Stone (1896) agrees with Dr. Dwight’s view, and I have seen some half a dozen specimens, taken in North and South Carolina, Florida, New Mexico, and Mexico, between February 23 and March 28, that show various stages of a complete prenuptial molt. Whether these are adults or young birds I do not know.

Food: The marsh wren feeds almost entirely on insects and their larvae, which it obtains on the marsh vegetation or on the floor of the marsh. Dr. Welter (1985) says that “much of the food is obtained near or from the surface of the water. * * * It is not unusual to observe the bird as he sights a juicy morsel fly into the air and capture it in the manner of a flycatcher. Insects as large as dragonflies are taken in this way. * * * Coleoptera and Diptera assume the highest rank while various other orders are represented to a lesser degree. Carabidae and Dytiscidae occur more frequently among the beetles than any other forms while a large percentage of the Diptera belong to the Tipulidae.”

F. H. King (1883) reports from Wisconsin that “of 14 stomachs examined one ate 1 ant; one, a caterpillar; one, 3 beetles; three, 3 moths; one a small grasshopper; one, 5 grasshopper eggs; one, 1 dragon-fly; and one a small snail.” Mosquito larvae are probably prominent in the food, as are larvae of other flying insects, diminutive mollusks, and aquatic insects. Forbush (1929), referring to Massachusetts, says that “in the salt marsh at high tide, it feeds on insects which crawl up on the grass and reeds, and at low tide it feeds largely on minute marine animals which it finds on or near the ground.”

Behavior: The marsh wren is much more often heard than seen. As we drift along some quiet stream bordered by extensive cattail marshes, we hear all about us the gurgling, bubbling songs, or the chattering, scolding notes of the birds, but not one is in sight in the dense jungle of flags. Perhaps one may explode into the air, rising a few feet above the cattails with an outburst of enthusiastic song and drift down again into cover; or we may see one make a longer flight from one part of the marsh to another, buzzing along on slow, direct, steady flight with rapid wing beats. If we watch quietly, curiosity may prompt one to come peering at us with furtive glances from the shelter of his retreat, clinging with feet wide apart to two swaying stems like a little acrobat doing the “splits”; his tail is held erect or pointed saucily forward and his head is lifted so high that head, body, and tail seem to form a feathered circle. He climbs nimbly up and down the reeds like a feathered gymnast, flow gliding down to the base to pick some food from the water, now gleaning along the stems, and again swinging jauntily from a swaying toll. He is the embodiment of active energy, always in motion, never still for a moment, and always chattering, scolding, or singing. He is a shy and elusive little mite; if we make the slightest motion while watching his antics, he vanishes instantly into the depths of his reedy jungle.

Although most of the marsh wrens probably live in harmony with their neighbors in the marsh, some, perhaps many, have formed the bad habit of sucking the eggs of least bitterns and red-winged blackbirds, as reported by several observers. For example, Dr. Chapman (1000) saw one of these wrens puncture all the eggs in two nests of least bitterns, and he attempted to photograph the bird in the act; the wren did not eat the contents of the eggs, though it may have returned to do so later; it looked like a case of pure viciousness. And Dr. A. A. Allen (1914) says that “of 51 nests of the Redwing observed in a limited area, the eggs of 14 were destroyed” by marsh wrens, and it is not at all uncommon to find one or more of the eggs of a nest with neat, circular holes in one side, such as would be made by the small, sharp beak of a wren.” One that he watched “began to drink the contents much as a bird drinks water. After a few sips, it grasped the eggshell in its beak and flew off into the marsh, where it continued its feast.” Dr. Welter (1935) evidently thinks that such behavior is exceptional for he says: “Many nests of other species of birds were under observation in the marsh and at no time were punctured eggs found or other indications of egg eating by the Marsh Wren observed.”

Voice: Wilson (Wilson and Bonaparte, 1832) evidently did not admire the vocal powers of the marsh wren, saying that “it would be mere burlesque to call them by the name of song,” for “you hear a low, crackling sound, something similar to that produced by air bubbles forcing their way through mud or boggy ground when trod upon”; this is a fair description of some of the notes; hut he apparently was not referring to the full song, parts of which are quite musical. F. Schuyler Mathews (1921) says that the song “ripples and bubbles along in a fashion similar to that of the Winter or House Wren, but with a glassy tinkle in tone not characteristic of the songs of the other species and a tempo perceptibly more rapid than that of the House Wren’s music.” Dr. Charles W. Townsend (1905) writes: “The song begins with a scrape like the tuning of a violin followed by a trill with bubbles, gurgles, or rattles, depending no doubt on the skill or mood of the performer, at times liquid and musical, at other times rattling and harsh, but always vigorous. It ends abruptly but is generally followed by a short musical whistle or trill, as if the Wren were drawing in its breath after its efforts. I have heard one sing fifteen times in a minute.”

Dr. Welter’s (1935) description is only slightly different; he dissects the song into three parts; first a grinding sound consisting of two to five notes with somewhat the quality of the aac notes of the white-breasted nuthatch; then comes the more musical “warble-like” part, which reminds him “of a sewing machine of the older sort being run rapidly, but of course it is less metallic and more musical. It has much of the spontaneity of the House Wren’s song but is otherwise quite distinct. This middle section begins at a low pitch, climbs upward, and then descends again.” The third section he calls a trill, which is again “quite low but lacks the harshness of the beginning of the song. * * *

“This entire song is given during May and most of June. Toward the end of the month, however, the last part is often omitted and often neither the beginning nor the end is heard.” The song period seems to cease entirely in August, but the full song has been heard in October, which may mean that a second song period occurs in fall.

The marsh wren is a persistent singer, chiefly during the early morning and the evening hours, but during the height of the season it sings all day and often at night. Only the male sings. He sings while clinging to the reeds or while moving among them; he indulges in his most delightful flight song while flying above the vegetation from one part of his territory to the other; or, rising in the air to a height of several feet, he flutters down to cover again in full song.

This wren also has several alarm, call, or chattering notes. According to Dr. Welter (1935) —

the kek kek or tschuk Is given by the female. The male’s note sometimes resembles this also but can usually he distinguished by its more grating nature and may be described as rrek. A series of notes is usually given together so the rrek’s do not sound very distinct as they roll into each other producing a chattering. The kek notes, however, while also given together, maintain their identity. The female has a hissing sound that she gives If too closely pressed by the male. Preceding copulation the female has been heard to give a trill like that at the end of the male’s song.

The call notes of the young are quite similar to those of the adult. The nestling, when the female arrives with food, gives a beady peep or peet. At first these notes are scarcely audible but us the young become older and stronger the peet is clearly heard. As the young leave the nest the peet gradually develops into a queck. It is much more squeaky than the adult kek and also lacks the woody quality. The notes of the juvenal become more and more like those of the adult until they are indistinguishable.

He says that the songs of the young males begin late in August and are entirely different from those of the adult. They reminded him at first of “the efforts of a not altogether successful Catbird,” but they were “given in a more rasping manner. The grating notes of the beginning and the trill at the end are usually omitted by young birds.”

Mr. Trautman (1940) “timed an isolated singing male whose territory was in a small stand of cattail and found that between 10 p. m. and 3 a. m. his average was 9 songs a minute.” Another, in a similar situation, sang at the rate of 11 songs a minute between 1:40 a. m. and 2:50 a. m. on a moonlight night. The singing slowed down during the middle of the day, between 10 a. m. and 2 p. m., to 4 songs a minute. “The amount of singing done by these birds declined sharply after mid-August, and by September 5, only an occasional, half-hearted song could be heard.”

Aretas A. Saunders writes to me: “The song of this bird is rather low-pitched and guttural, or sometimes squeaky. It consists of a series of rapid notes, so rapid as to call the result a trill, but more frequently slow enough to count the number. In 26 of my records, without trills, the number of notes varies from 8 to 16 and the average number is 12. In a majority of the songs the notes are all equal in time, but some have portions where the notes are more rapid in part of the song. These portions are sometimes the beginning and sometimes the end, or occasionally in the middle of the song.

“The pitch of the notes varies from C” to C”‘. One record is all on one pitch (B”‘). A number of others are all on one pitch except the first or the final note, but others vary in numerous ways. The greatest variation in pitch in any one song is 2½ tones, and the average 1½ tones. I have occasionally seen a bird sing a flight song, when the song is somewhat more prolonged than I have described, but I have never succeeded in getting a record of this song.

“In spite of the simplicity of this song the individuals vary it considerably. I have recorded five different songs from one individual. The quality sometimes changes from guttural to squeaky in the same song. The time of songs varies from 11/5 to 2 seconds, though flight songs are probably longer.”

Field marks: One hardly needs field marks to recognize a long-billed marsh wren, for it is wrenlike in appearance and behavior, and no other wren lives in such wet marshes. If perchance it is seen in the drier part of a marsh or meadow, it can be distinguished from the short-billed marsh wren by the blackish, unstreaked crown, the white line over the eye, and the black upper back streaked with white.

Enemies: Hawks and owls would have difficulty in capturing these active little birds as they dive into their dense retreats. Red-winged blackbirds are often seen chasing wrens for reasons stated above. Dr. Welter (1935) mentioned three small mammals, meadow mice, jumping mice, and Bonaparte’s weasels, as probably guilty of destroying some eggs and young. He says that Dr. A. A. Allen has seen bronzed grackles eating the young and has found bumble bees occupying the nests. Fleas, lice, and hippoboscid flies sometimes damage the young.

Fall: Dr. Welter writes: “There is no marked exodus of birds from the marsh at a given time in the fall. At first the young of the year remain in family groups but, as the time of departure approaches, there is an apparent flocking together of young birds, usually near the water’s edge. At this time 25 or 30 birds may be observed together feeding near the surface of the water. * * * The first birds to leave are the adults and some of the young of the first brood.” No adults were found after September 10; the birds that remain after that date are young birds, mostly those of the second brood, either in juvenal plumage or molting out of it. “As these birds complete the molt they, too, depart for their winter homes so that, by October 20, only a few scattered individuals remain. By the first of November these, also, have departed.”

Elon H. Eaten (1914) describes the departure thus:

On one occasion while I was concealed in a blind watching for ducks to enter the marsh, I saw the last representative of this species leave the marshes at the foot of Canandaigua Lake. It was a cool night late in October when the moon was at the full. The little fellow uttered a feeble warble which attracted my attention and then rose from near my station, fluttering higher and higher into the air until lost at an elevation of about 300 feet, where I caught my last glimpse of him against the full moon. The following morning when I visited the marsh no more wrens were left. Evidently they migrate at night, and high in the air, so as to see their way and escape their enemies more successfully.

Winter: Most of the prairie marsh wrens migrate in fall and spend the winter in Mexico or along the Gulf coast to western Florida. But some few individuals remain in their summer haunts all winter in the shelter of the dense cattail marshes. There are winter records for Massachusetts, Connecticut, New York, and Ohio. It may be that they are more common in winter than we realize, for they are silent and remain well hidden in the marshes, where they are hard to find.

WESTERN MARSH WREN

?TELMATODYTES PALUSTRIS PLESIUS Oberholser

HABITS

The western marsh wren breeds in the Great Basin regions of the western United States, from central British Columbia, Washington and Oregon, and northeastern California eastward to the Rocky Mountains in central Colorado and southward into New Mexico. Its winter range extends into Mexico.

Ridgway (1904) describes it as “very similar in coloration of upper parts to T. p. iliacus, but the brown averaging paler and decidedly less rufescent; upper tail-coverts usually more or less distinctly barred with dusky, and middle rectrices more distinctly barred; color of flanks, etc., conspicuously different, being pale wood brown, pale isabella color, or pale broccoli brown instead of bright buffy cinnamon or tawny-buff, the under tail-coverts usually more or less distinctly barred; wing and tail averaging decidedly longer (especially the tail), culmen averaging slightly shorter.”

The haunts of the western marsh wren are evidently similar to those of the other races that breed in the fresh-water marshes and sloughs in the interior.

Nesting: Dawson and Bowles (1909) give a very good description of the nest of this wren as follows:

The Marsh Wren’s nest Is a compact ball of vegetable materials, lashed midway of cat-tails or bulrushes, living or dead, and having a neat entrance hole in one side. A considerable variety of materials is used in construction, but In any given nest only one textile substance will preponderate. Dead cat-tail leaves may be employed, in which case the numerous loopholes will be filled with matted down from the same plant. Fine dry grasses may be utilized, and these so closely woven as practically to exclude the rain. On Moses Lake, where rankly growing bulrushes predominate in the nesting areas, spirogyra is the material most largely used. This, the familiar, scum-like plant which masses under water in quiet places, is plucked out by the venturesome birds in great wet banks and plastered about the nest until the required thickness is attained. While wet, the substance matches its surroundings admirably, but as it dries out it shrinks considerably and fades to a sickly light green, or greenish gray, which advertises itself among the obstinately green bulrushes. Where this fashion prevails, one finds it possible to pick out immediately the oldest member of the group, and it is more than likely to prove the occupied nest.

The nest-linings are of the softest cat-tail down, feathers of wild fowl, or dried spirogyra teased to a point of enduring fluffiness. It appears, also, that the Wrens of ten cover their eggs upon leaving the nest. Thus, in one we found on the 17th of May, which contained seven eggs, the eggs were completely buried under a loose blanket of soft vegetable fibers. The nest was by no means deserted, for the eggs were warm and the mother bird very solicitous, insomuch that she repeatedly ventured within a foot of my hand while I was engaged with the nest.

A nest in the Thayer collection, taken in Lassen County, Calif., on May 10, 1910, was built in tules 2½ feet above the water. It is a large well-made nest constructed mainly of the fruiting or dry flower clusters of some marsh plant, firmly reinforced and compactly interwoven with narrow strips of tules or other marsh plants, forming a very solid and durable structure; it measures about ‘1 inches in height by about 4 inches in diameter.

Eggs: The eggs of the western marsh wren are indistinguishable from those of the other races of the species. The measurements of 40 eggs average 16.1 by 12.6 millimeters; the eggs showing the four extremes measure 17.3 by 13.2,16.6 by 13.4, and 14.5 by 11.0 millimeters.

Winter: Harry S. Swarth (1917) makes the following interesting observation on the winter distribution of this wren in California:

The known breeding range of the western marsh wren in California is very limited, being merely the restricted northeastern corner of the State, a region which shows strongly Great Basin faunal affinities. In winter, however, plesius is perhaps the most abundant of any form of the species, occurring in numbers over a large part of the State. It is an especially numerous winter visitant in the San Diegan district of southern California. In this region summer is the dry season, a period of such excessive aridity that birds with the needs and proclivities of the marsh wrens are closely limited as to habitat, being restricted to extremely circumscribed areas about the few suitable permanent streams and sloughs. In winter this is all changed. Abundant rains often transform what were dry fields and pastures Into ponds and marshes, while every roadside ditch is running full, and bordered with dense vegetation. In consequence, the visiting marsh wrens are enabled to scatter widely over the country.

He cites a number of records from various points along the coasts of California and Oregon, which indicate that “individuals of this form may occasionally be found in winter at any point, along the coast.” But he shows clearly that the center of abundance in winter is in southern California, “both on the deserts and in the San Diegan region.

TULE WREN

TELMATODYTES PALUSTRIS PALUDICOLA (Baird)

HABITS

Professor Baird (1864) was the first to name and describe a western race of this species, based on one specimen from Washington and three from California, and his name still applies to the long-billed marsh wrens of the Pacific coast district from British Columbia to southern California. His brief description gives as its characters: “Bill shorter than tarsus. Tail coverts distinctly banded all across. Bands on tail quite distinct; appreciable on the central feathers.”

Ridgway (1904) gives a fuller description, based on 17 specimens, as follows: “Most like T. p. palustris, but tail-coverts usually barred (especially the upper), middle rectrices more distinctly barred, flanks, etc., deeper brown, bill smaller, and tail decidedly longer; agreeing with T. p. plesius in barred tail-coverts, more distinctly barred tail, and relatively longer tail, but decidedly smaller and with coloration decidedly darker.”

The name tide wren must not be understood to imply that this wren is wholly or even mainly, confined to the tules (Scirpus lacustris occidentalis); although it breeds abundantly in this type of vegetation in the extensive marshes or where it grows around the shores of lakes, it also breeds commonly in the cattail marshes along the intersecting channels or the banks of quiet streams, and in the salicornia of coastal marshes where the supply of cattails is inadequate.

Dr. Gordon D. Alcorn writes to me that the tule wren is abundant in suitable localities in western Washington but says that “the wren population diminishes sharply within a few miles of the Pacific Ocean; this might be due to the scarcity of suitable swamps for nesting purposes. The swamps within this region are mostly of low-growing sedges (Carex), barely reaching 2 feet in height. In most of the swamps the cattails (Typha), of which the wrens are so fond, are scarce or entirely lacking. I regularly visit two wren areas in Grays Harbor County. The first is located on the shores of the harbor and comprises an area of approximately 5 acres. Few cattails are present, the swamp possessing mostly sedges with a few scattered willows. The area almost touches salt water, as the limits of the grounds are bounded on the south and west by the highwater line. I visit also a rather extensive swamp paralleling the ocean beach in the vicinity of Oyhut, Wash. This swamp covers a distance of about 4 miles and is composed of low sedges with a few spiraea bushes.”

Spring: Samuel F. Rathbun tells me that, near Seattle, this little wren is more or less resident in the region, although found more commonly from early spring until late in the fall. “There appears to be a movement of the species during late March and early April; for at this period birds will be heard and seen in and about small marshes or similar localities, in which they are absent during the nesting season.”

Nesting: Dr. Alcorn says in his notes that, in the localities mentioned above, the nests are usually placed within a foot or two of the water; the birds use sedge leaves for building and line the nests with willow cotton; the nests are always fragile structures; and there are not so many decoy nests as are found farther inland.

In the dense cattail swamps farther back from the coast, the nests are more substantial and more typical of the species. In the Thayer collection in Cambridge there are two distinct types of well-built nests, both collected by J. H. Bowles in Pierce County, Wash. One, built 4 feet above the water in dense cattails, where the water was 3 feet deep, is a compact, oval ball, made largely’ of cattail down interwoven with and firmly bound together with strips of the flags; it would have furnished a warm, dry shelter for the young. The other, “woven among coarse marsh grass” in a fresh-water marsh, is a firmly woven and well-made ball of interlaced strips of marsh grasses and reeds, a common type for the species.

Walter E. Bryant (1887) published the following note on a nest found by A. M. Ingersoll in an unusual situation: “A conspicuous nest, containing eggs, was woven among the almost leafless branches of a young willow, five feet above a fresh-water marsh.”

The tule wren seems to be a prolific builder of decoy or male nests. Irene G. Wheelock (1904) says that she has examined “30 in 1 day and found but 1 occupied, and that was the oldest, most tumble down of the lot.” And Dawson and Bowles (1909) say that “in a day Mr. Bowles found 53 nests, only 3 of which held eggs or young. At least 2 broods are raised in a season.” His brother, Charles W. Bowles (1898), throws some light on the use that may be made of these dummy nests: “In the spring of 1896 I found an empty marsh wren’s nest, and on passing by later in the day, saw three nearly fledged young in it. There were also other nests near by, with one or more young in each. It seems to me probable that these duplicate nests are built, if the birds have a large family, for the young to roost in, at least at night, when they are too large to be all contained in one nest, but not yet able to take care of themselves.”

Eggs: The eggs of the tule wren are apparently no different from those of other long-billed marsh wrens. The measurements of 40 eggs average 16.3 by 12.8 millimeters; the eggs showing the four extremes measure 17.6 by 13.0, 16.8 by 13.5, 15.5 by 12.8, and 16.0 by 12.0 millimeters. These eggs are all in the United States National Museum.

Young: Mrs. Wheelock (1904) writes:

Twelve days are required for incubation, and even during this short period the mother is not a close sitter. I have known her to leave the nest for 2 hours in the middle of the day, trusting to the intense heat of the sun to perform her task for her; and but for the thick, moist walls of the cradle, this same sun would have been fatal to the bird life within the shells.

As soon as the eggs hatched in the nest I was watching, I cut a slit in the top of it to look at the young. They were naked, light pink in color, with tiny heads, mere knobs for eyes and buds for wings; each nestling measured 1 inch in length. After this examination I tied up the slit, and before I was a yard away the mother entered the nest again. Four days later the eyes of the young wrens had begun to open, and looked like tiny slits, while a thin buffy down covered the top of their heads and was scattered sparsely over their bodies. As in the young of the long-billed marsh wrens, the ear openings were conspicuously large. Bill and legs had changed from pink to light burnt-orange in color. They were fed by regurgitation for the first 4 days and doubled in weight every 24 hours. When a week old they were commencing to feather, and in 3 days more were nearly ready to leave the nest. They were now fed on larvae of water insects, slugs, and dragonflies, besides other insects, and meals were served four times an hour during most of the day.

These young wrens left the nest, when examined, at an age of 12 days; they were able to glean some of their food but were fed by their parents for 2 weeks longer.

Food: Only 53 stomachs of the California races of this species were examined by Professor Beal (1907),in which animal matter amounted to 98 percent, and vegetable matter, consisting of a few seeds of sedges and one of amaranth, amounted to 2 percent. “Beetles, wasps, ants, bugs, caterpillars, and a few miscellaneous insects, with some spiders and snails, make up the bill of fare.” Bugs—assassin bugs, damsel bugs, stink bugs, leafhoppers, and treehoppers—constituted the largest item, 29 percent. Scales were found in one stomach. Caterpillars and chrysalids amounted to 17 percent, beetles (mainly harmful Species) 16 percent, ants and wasps 8 percent, flies, grasshoppers, dragonflies, and a few other insects 11 percent. Spiders were eaten regularly and made up over 5 percent of the food. One stomach contained 11 small snails.

Not much more need be said about the habits of tule wren after all that has been written about the prairie marsh wren, for their habits are practically identical. Although some suspicion exists, there is no positive evidence that snakes destroy the eggs or young of these wrens, so far as I know. Dr. Gordon D. Alcorn (1931), however, published the following observation, which is at least suggestive; a pair of wrens were much disturbed by the presence of a garter snake near their nests.

The snake was slowly crawling some 2 to 5 feet above the water over the dead cattail leaves and stems in which were located a number of occupied and unoccupied wren nests. The birds remained perfectly silent, but with outspread wings and ruffled feathers darted again and again at the head of the reptile. The snake paid no attention to the birds but continued to ‘explore”, finally approaching a wren nest and entering it. About two-thirds of the snake’s body remained outside the nest while it stayed at the nest for about 30 seconds. The snake was allowed to enter several nests in a similar manner. It was then killed and the stomach examined and found to be empty but for a well-digested slug. Also each nest entered was examined and each found to be empty (undoubtedly “decoys,” as each was unlined and in a conspicuous position.) * * * The snake was without doubt looking for food, either eggs or young birds, in these nests, and was not able to discriminate between occupied and unoccupied nests.

Dawson (1923) writes: “In autumn the Tule Wrens leave the sheltered precincts of the ponds, and go roaming about through dry weed patches and adjacent chaparral. Here they are as noisy and as elusive as ever, and are in nowise awed by their less usual surroundings. There is, doubtless, some invasion from the north and consequent crowding in winter.”

Mrs. Amelia S. Allen tells me that “on September 24, 1941, they were abundant in a damp cow pasture inland in sedge grass.”

SUISUN MARSH WREN

TELMATODYTES PALUSTRIS AESTUARINUS Swarth

HABITS

In naming this race Mr. Swarth (1917) says of its characters: “In coloration aestuarinus is darker than the average of paludicola, especially as compared with southern Californian examples of the latter. Occasional specimens of paludicola, however, from all parts of its range, are quite as dark colored. In dimensions, T. p. aestuarius differs from T. p. paludicola in its greater size throughout, being of about the same dimensions as T. p. plesius. From plesius it differs in its much darker coloration.” This subspecies seems to be an intermediate between the two adjacent races, resembling one in color and the other in size. It also occupies a rather limited breeding range between the other two, mainly in Solano and Sonoma Counties, Calif. The 1931 Check-list gives its range as “west-central California, breeding at the confluence of the Sacramento and San Joaquin rivers, in Napa and Solano Counties, and thence south to Tulare County. In winter spreads beyond its breeding range to Oregon and southern California.” Living as it does in a similar type of country, we should hardly expect to find anything in its habits that is different from those of the neighboring races.

The eggs of the Suisun marsh wren do not differ materially from those of the species elsewhere. The measurements of 27 eggs average 16.0 by 12.7 millimeters; the eggs showing the four extremes measure 17.1 ‘by 13.2, 16.6 by 13.4, 14.4 by 12.4, and 15.0 by 12.1 millimeters.