BOSTON PUBLIC LIBRARY
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3 9999 06317 641^
NATURAL HISTORY
OF THE
KING RAIL
NUMBER 67
UNITED STATES
DEPARTMENT OF THE INTERIOR
BUREAU OF SPORT FISHERIES AND WILDLIFE
NORTH AMERICAN FAUNA
This publication series includes monographs and other reports of scientific in-
vestigations relating to birds, mammals, reptiles, and amphibians, for professional
readers. It is a continuation by the Bureau of Sport Fisheries and Wildlife of the
series begun in 1889 by the Division of Ornithology and Mammalogy (Department
of Agriculture) and continued by succeeding bureaus — Biological Survey and
Fish and Wildlife Service. The Bureau distributes these reports to official agen-
cies, to libraries, and to researchers in fields related to the Bureau's work;
additional copies may usually be purchased from the Division of Public Docu-
ments, U.S. Government Printing Office.
Reports in North American Fauna since 1950 are as follows (an asterisk indi-
cates that sale stock is exhausted) :
*60. Raccoons of North and Middle America, by Edward A. Goldman. 1950. 153 p.
*61. Fauna of the Aleutian Islands and Alaska Peninsula, by Olaus J. Murie;
Invertebrates and Fishes Collected in the Aleutians, 1936-38, by Victor B.
Scheffer. 1959. 406 p.
*62. Birds of Maryland and the District of Columbia, by Robert E. Stewart and
Chandler S. Bobbins. 1958. 401 p.
*63. The Trumpeter Swan; Its history, habits, and population in the United
States, by Winston E. Banko. 1960. 214 p.
*64. Pelage and Surface Topography of the Northern Fur Seal, by Victor B.
Scheffer. 1961. 206 p.
65. Seven New White-winged Doves From Mexico, Central America, and South-
western United States, by George B. Saunders. 1968. 30 p.
m. Mammals of Maryland, by John L. Paradiso, 1969. 193 p.
67. Natural History of the King Rail, by Brooke Meanley. 1969. 108 p.
NATURAL HISTORY
OF THE
KING RAIL
MAR 3 1 2000
King Rail near Jacksonville, Fla. (Photograph by Samuel A. Grimes)
NATURAL HISTORY
OF THE
KING RAIL
By Brooke Meanley, Wildlife Biologist
Patuxent Wildlife Research Center
Division of Wildlife Research
BUREAU OF SPORT FISHERIES AND WILDLIFE
NUMBER 67
UNITED STATES
DEPARTMENT OF THE INTERIOR
Walter J. Hickel, Secretary
Leslie L. Glasgow, Assistant Secretary for
Fish and Wildlife, Parks, and Marine Resources
FISH AND WILDLIFE SEKVICE
Charles H. Meacham, Commissioner
BUREAU OF SPORT FISHERIES AND WILDLIFE
John S. Gottschalk, Director
North American Fauna, Number 67
Published by
Bureau of /Sport Fisheries and Wildlife
May 1969
U.S. GOVERNMENT PRINTING OFFICE
WASHINGTON : 1969
For sale by the Superintendent of Documents, U.S. Government Printing Office
Washington, D.C. 20402 - Price 60 cents
Contents
Page
Introduction 1
History and Systematic Position 4
History 4
Systematic position 6
Relationship to the Clapper Rail 6
Distribution and Migration 10
Distribution 10
Migration 12
Local movements 14
Ecological Relations 16
Louisiana gulf coast marshes 16
The delta marsh 16
The subdelta marsh 17
The prairie marshes 18
Southern ricefields 20
Florida 25
South Carolina Low Country 27
Savannah National Wildlife Refuge 29
Upper Savannah River Valley 32
Chesapeake Bay Country 32
Tidewater Virginia 32
Virginia Eastern Shore 34
Maryland Eastern Shore 35
Inner Coastal Plain of Maryland 38
Delaware Bay, Del 39
Great Lakes region 41
North-central prairie marshes 42
Northern Great Plains 42
Description 43
Size 43
Adult plumage 43
Legs and feet , 45
Bill 45
Tongue and lining of mouth 45
Eye 45
Notes on sexing and aging 45
Molting 46
Breeding Biology 48
Homing 48
Territories 48
Defense of territories 49
Courtship behavior 50
Mating call and pair formation 50
Other calls 50
Display 51
Courtship feeding 53
(V)
VI CONTENTS
Page
Prenesting activity 53
Calling 53
Symbolic nest building 54
Copulation 54
Nesting period 54
Nest site and materials 57
Egg laying and clutch size 61
Clutch size 61
Description of eggs <- . 61
Weight of eggs 62
Incubation 62
Hatching 64
Nesting success and survival 65
Breeding status of first-year birds 65
Development and Behavior of Captive Rails 66
Development of Young 66
First-day chick 66
One to thirty days 67
Thirty to sixty days 68
First winter plumage 72
Miscellaneous notes on behavior of young 72
Sleeping 72
Competition 73
Bathing 73
Winter behavior of captive rails 73
Foods 75
Arkansas ricefields 77
Texas ricefields 79
Louisiana ricefields 79
Upper St. Johns River, Fla 79
Currituck Sound, N..C 79
Patuxent River, Md 80
Beaver Dam, Wis 80
Chicago, 111 80
Feeding Behavior 81
Pellet casting 81
Feeding young 82
Regional observations 83
Arkansas ricefields 83
Delaware Bay marshes 83
Savannah National Wildlife Refuge 84
Some unusual observations 85
Mortality Factors 86
Manmade objects 86
Predation 86
Hurricanes 88
The King Rail as a Game Bird 90
Methods of hunting 90
Patuxent River, Md 91
Eagle Lake area, Tex 93
Other areas 93
Summary 95
Literature Cited 98
CONTENTS VII
Page
Appendix 1 — Methods of Capturing for Banding 103
Types of capturing devices 103
Long-handled dip or clap net 103
All-purpose or cloverleaf trap 103
Tending traps 105
Age for banding King Rail chicks 106
Need for banding data 106
Appendix 2. — Local Names 107
ILLUSTRATIONS
Frontispiece, King Rail near Jacksonville, Fla ii
Figure
1. A King Rail walking 3
2. Mated King Rail and Clapper Rail collected in Delaware 8
3. Approximate breeding range and principal distribution of the
King Rail in North America 10
4. Louisiana coastal marshes 17
5. Prairie marsh, Grand Chenier, La 18
6. Southern bulrush, fall panicum, and alligatorweed in the
Prairie Marsh type 21
7. King Rail wading through ricefi eld 22
8. Arkansas Grand Prairie near Stuttgart 23
9. Nest of King Rail in wet rice stubble 24
10. Habitat of King Rail, Indian River County, Fla 26
11. King Rail habitat on Seminole Indian Reservation, Glades
County, Fla 27
12. Floodgate and ricefield canal near Savannah, Ga 28
13. South Carolina Low Country: ricefield nesting habitat along
Savannah River, Jasper County 30
14. Alligatorweed in canal at Savannah National Wildlife Refuge.- 31
15. Big cordgrass habitat, Nanticoke River marsh, Md 33
16. Winter abode of King Rail, Rappahannock River near Tappa-
hannock, Va 34
17. Switchgrass habitat of King Rail, Elliott Island, Md 36
18. Nest and eggs of King Rail in Nanticoke River marsh, Vienna,
Md 37
19. Nest and 9 eggs of King Rail in brackish marsh, Long Marsh
Island, Eastern Bay (of Chesapeake Bay), Md 38
20. Taylor's Gut at low tide, Kent County, Del 39
21. Displays of the King Rail 52
22. Canopy on King Rail nest in roadside ditch, Arkansas Grand
Prairie 58
23. King Rail nest in roadside ditch near Stuttgart, Ark 58
24. King Rail incubating in nest of cattails in roadside ditch,
Arkansas Grand Prairie 59
25. King Rail incubating in open nest along roadside ditch,
Mamou, La 59
26. Distraction display of King Rail near nest 64
27. Downy young King Rail, 31 days old, with juvenal plumage
beginning to develop 69
28. Ventral view of 31-day-old King Rail showing development of
white juvenal plumage in sternal and abdominal regions and
crural tract 70
VIII CONTENTS
Page
29. Fifty-day-old King Rail with juvenal plumage nearly com-
plete 71
30. Captive King and Clapper Rails at Patuxent Wildlife Research
Center, Laurel, Md 74
31. Foods of the King Rail in a brackish bay marsh, Kent County,
Del 77
32. Regurgitated King Rail pellets from Dorchester County, Md__ 82
33. Method of hunting railbirds in Patuxent River wildrice marshes,
Maryland r 91
34. Railbird boats tied up after the hunting season 92
35. Method of hunting railbirds in southern ricefields 94
36. All-purpose or cloverleaf trap and drift fence in shrub swamp
at Patuxent Wildlife Research Center 104
37. All-purpose or cloverleaf trap 104
Photographs are by the author unless otherwise credited.
Introduction
The King Rail {R alius elegans Audubon), largest of North Amer-
ican rails, is indeed an elegant bird, as its Latin name implies. Its
striking appearance (fig. 1), secretive nature, and association with a
variety of wetland habitats make it a favorite of bird students and
rail hunters. The King Rail is found in most of the eastern half of
North America, from the Atlantic coast to the Great Plains and from
the Gulf of Mexico to southern Canada. It is most abundant in the
fresh and brackish tidal marshes of the Atlantic and Gulf Coastal
Plain, the domestic ricefields of Arkansas, Louisiana, and Texas, and
the marshes of southern Florida. It is fairly common in parts of the
Midwest Prairie and Great Lakes region.
I began my studies of this interesting bird in 1950 in the Arkansas
ricefields, and have continued them until 1967, both in the field and
in the laboratory.
Many of the field observations, particularly those of courtship
behavior, were made from an automobile which served as an admirable
mobile blind. Such a blind was used to follow courting rails along
roadside ditches in Arkansas and Louisiana, making it possible to
study the detailed nuptial courtship behavior of 20 different pairs
and the prenuptial behavior of four. Under these conditions it was
possible also to distinguish the sexes by their behavior rather than
by their size differences, which are sometimes difficult to ascertain in
the field.
The highly vocal nature of the King Rail and its characteristic
calls, varying with different conditions, enhance the value of field
observations and made the call-count census a practical technique.
Studies of growth and development of the young were made with
captive birds, which are quite tractable if obtained early in life from
nests or hatched from eggs in incubators.
Studies of breeding biology were made mostly on the Arkansas
Grand Prairie in the vicinity of Stuttgart from 1950 through 1955.
Subsequent studies on life history and ecology were made at Mamou,
Evangeline Parish, La. ; Broadway Meadows near Woodland Beach,
Kent County, Del.; the Pee Dee River at Georgetown, S.C. ; the
Savannah National Wildlife Refuge, Jasper County, S.C; and the
Patuxent Wildlife Research Center, Laurel, Md.
l
2 NORTH AMERICAN FAUNA 67
I have supplemented my own observations with a review of the
published studies of others and have attempted to bring all informa-
tion on the King Rail together into a monographic treatment.
My discussions in this paper include the history of the discovery
of the King Rail as a distinct species by Audubon in 1834 and its
systematic position in relation to the Clapper Rail, as taken mostly
from the literature. The discussions of other topics are largely from
my own observations, but supplemented with literature reports. The
principal topics include distribution and migration; ecological rela-
tions; physical characteristics; breeding biology; development and
behavior of captive rails ; foods and feeding ; mortality factors ; and
the King Rail's position as a game bird. Appendixes include methods
of capturing and banding and a list of local names. Aquatic plant
names used in the text are from Hotchkiss (1950) unless otherwise
indicated.
I am indebted to many persons for assistance with this project.
Anna Gilkeson Meanley, my wife, assisted with the field work over a
7-year period in Arkansas and Louisiana. E. R. Kalmbach, former
Director of the Denver Wildlife Research Center, made the sketches
of courtship displays and offered encouragement and many sugges-
tions during the early phases of the study in Arkansas. Other col-
leagues from the Bureau of Sport Fisheries and Wildlife who were
most helpful in various phases of the work include Nancy C. Coon,
John W. Aldrich, Van T. Harris, Lucille F. Stickel, Paul A. Stewart,
Robert E. Stewart, Charles C. Sperry, Neil Hotchkiss, Francis M.
Uhler, Frederick C. Schmid, Glen Smart, Johnson A. Neff, Robert G.
Heath, Luther C. Goldman, and David K. Wetherbee. Anthony J.
Florio of the Delaware Game and Fish Commission was helpful in
Delaware studies. I am grateful to Samuel A. Grimes of Jacksonville,
Fla., for his photograph of a King Rail used as the frontispiece.
NATURAL HISTORY OF THE KING RAIL
Figure 1. — A King Rail walk-
ing. One foot is placed in
front of the other, producing
a single line of tracks.
./"'
History and Systematic Position
HISTORY
John James Audubon published the first description of the King
Eail as a distinct species. The Great Red-bieasted Rail or Fresh-water
Marsh Hen, as he called it, was introduced with, the publication of his
painting in Birds of America (Audubon, 1834, plate 203). A year
later a description of this new rail appeared in his Ornithological
Biography (Audubon, 1835, p. 27-32).
Alexander "Wilson, Audubon's predecessor, encountered this same
species but thought it was the adult form of the Clapper Rail {Rallus
longirostris) , "following the current opinion of gunners that it was
a very old example of that species" (Stone 1908, p. 110). Elaborating
on this point, Audubon (1835, p. 27) stated:
No doubt exists in my mind that Wilson considered this beautiful bird merely
the adult Rallus crepitans [Rallus longirostris crepitans], the manners of which
he described, as studied at Great Egg Harbour, New Jersey, while he gave in
his works the figure and colouring of the present species. My friend, Thomas
Nuttall, has done the same, without, I apprehend, having seen the two together.
Always unwilling to find fault in so ardent a student of nature as Wilson, I felt
almost mortified when, after having in the company of my worthy and learned
friend, the Reverend John Bachman, carefully examined the habits of both
species, which in form and general appearance, are closely allied, I discovered
the error which he had in this instance committed. Independently of the great
difference as to size between the two species, there are circumstances connected
with their habits which mark them as distinct. The Rallus elegans is altogether
a fresh-water bird, while R. crepitans never removes from the salt-water
marshes . . .
J. d'Arcy Northwood (1956, p. 224), commenting on Audubon's
discovery, said:
The king rail was one of Audubon's scoops. Here was a large rail, not particu-
larly rare, that lived unknown and undescribed under the noses of the experts
in Philadelphia. Audubon realized that it was distinct from the clapper rail of
the salt marshes, with which it had been confused, and named it the Great Red-
breasted Rail or Fresh-Water Marsh Hen.
The type locality given in the American Ornithologists' Union
Check-list of North American Birds (1957, p. 152) is Kentucky, South
Carolina, Louisiana, and north to Camden, N.J., and Philadelphia —
Charleston, S.C.
Although Audubon collected and observed the King Rail in several
localities prior to his field studies in company with Bachman in the
4
NATURAL HISTORY OF THE KING RAIL 5
Charleston, S.C., region, it is apparent from his original description
(Audubon, 1835) that he finally decided that it was indeed a new spe-
cies on the basis of their work in that region. It would seem that the
species distinctness may even have been brought to his attention by
Bachman, who in a letter to Audubon sent from Charleston, S.C., and
dated December 27, 1832, posed this question :
May not the Northern Marsh Hen, be the Bird which we here call the Fresh
Water M. Hen & our Ash coloured one that keeps to the Marsh be peculiar to
the South? I should like to have this matter ascertained.
In a letter from Charleston dated March 27, 1833, Bachman, again
referring to the Marsh Hen, said (Deane, 1929, p. 180, 184), "My
opinion first expressed [in the letter of December 27, 1832] is every
day strengthened."
Audubon (1835, p. 27-28) reported that he caught a female at Hen-
derson, Ky., on May 29, 1810, and also collected a female near Camden,
N.J., in July 1832.
Stanley C. Arthur ( 1937, p. 503) , a biographer of Audubon, believed
that Audubon's painting of the Fresh-water Marsh Hen was made at
New Orleans in 1821 when he spent the winter, spring, and fall there.
Audubon obtained birds from the city market and from two hunters
engaged to collect for him, and painted over 100 birds from this area
(including work in the St. Francisville area in West Feliciana Parish) .
An entry in Audubon's journal, dated December 20, 1821 (Corning
1929, p. 224), says that he "Recd a nondescript rail." And an entry
made the next day says, "Drew a streaked Rail." This may have been
a King Rail, but if it was, apparently it was not recognized as a new
species at that time.
An interesting letter from Rodolphe M. deSchauensee, Curator of
Birds at the Academy of Natural Sciences in Philadelphia, dated
February 7, 1962, sheds some light on the possibility of an existing
type specimen or cotype :
I have gone into the question of the type specimen of the King Rail mentioned
in your letter of February 1 with what I think- are interesting results.
Some years ago Fletcher and Phillips B. Street gave to the Academy a collec-
tion of birds which had belonged to Edward Harris who was a friend of Audu-
bon. On looking through our collection I found in this lot an immature specimen
of the King Rail. In vol. 3 (p. 28) of Audubon's Ornithological Biographies he
says "I killed one female in New Jersey, a few miles from Camden, in July, 1832
in company with my friends Edward Harris and Mr. Ogden . . ."
In the Elephant Folio (vol. 3) pi. 203 engraved in 1834 two birds are shown,
an adult and an immature. The bird in Harris's collection agrees very well both
in color and measurements with the bird depicted as the immature specimen. As
the bird was collected in 1832 the plate engraved in 1834 and Audubon's original
description published in 1835, there is every reason to suppose that this is the
bird shown on the plate.
In view of all the above I feel that it is justifiable to regard this specimen
as a cotype. Audubon described an adult male, a female and an immature. If
6 NORTH AMERICAN FAUNA 67
these three birds all existed today they would of course all be cotypes. In the
plate the female is not figured, only the adult male and young.
If this specimen were to be accepted as a cotype, the type locality
would have to include Camden, N.J. However, in his original descrip-
tion of R. elegans, Audubon (1835) said that most of his observations
of this species were in South Carolina.
SYSTEMATIC POSITION
The King Rail belongs to the order Gruiformes, which in North
America includes the cranes, limpkins, rails, gallinules, and coots.
Birds of this group mostly inhabit wetland environments, particularly
marshes.
The suborder Grues includes the families Gruidae (cranes) and
Aramidae (limpkins). Rails, gallinules, and coots belong to the sub-
order Ralli, which contains a single family, Rallidae. In North Amer-
ica this family comprises seven genera and nine species.
The three North American species of the genus Rallus, R. elegans
(the King Rail), R. longirostris (the Clapper Rail), and R. limicola
(the Virginia Rail) , have laterally compressed bodies which facilitate
passage through dense marsh vegetation; rather long, slender, and
slightly curved bills which are as long as or longer than the tarsi, and
longer than the heads; large, strong legs; long, slender, unwebbed
toes; short, rounded wings (with vestigial claws); short, tip-up,
pointed tails less than half as long as the wings ; flanks conspicuously
barred with white ; olive or grayish dorsal regions which are striped
with black or dusky markings; and buffy or rufescent breasts. R. ele-
gans is larger than R. limicola, which it resembles in color, and is
more rufescent than races of R. longirostris but is about the same size
as that species.
Two races of the King Rail are generally recognized : Rallus elegans
elegans of North America, and Rallus elegans ramsdeni, the Cuban
form. Apparently a third form, Rallus elegans tenuirostris, occurs in
the fresh- water marshes of the Valley of Mexico. There is a difference
of opinion concerning the systematic position of tenuirostris, some
authors assigning it to Rallus elegans and others to Rallus longirostris.
The recent work of Warner and Dickerman (1959) seems to indicate
that the plumage and inland distribution of this form are more like
that of Rallus elegans.
RELATIONSHIP TO THE CLAPPER RAIL
Some ornithologists believe that King and Clapper Rails are merely
races of the same species. Structurally and behaviorally they are simi-
lar. The plumages of several Clapper races closely resemble that of
the King Rail. Their breeding ranges overlap in numerous coastal
brackish marshes, in at least one of which there is absolute evidence
of interbreeding resulting in the production of viable eggs.
NATURAL HISTORY OF THE KING RAIL 7
Oberholser (1937, p. 314-315) , in discussing the relationship of these
two species, stated that —
it remains yet to determine the status of the king rail, Rallus elegans, of the
Eastern United States, and its single subspecies, Rallus elegans ramsdeni, of
Cuba. This is an unusually difficult matter to decide, and one concerning which
there may well be difference of opinion. The chief external characters separating
the king rails from the clapper rails consist in the much more reddish bend of
the wing, and in the rich rufescent-olive tinge of the upper parts of the former
birds, this involving both the centers and margins of the feathers. There is little
or no trenchant difference in behavior, voice, nest building, or other habits
between these two species. Neither one of the external characters of plumage
above mentioned, nor any difference in size or proportions, is entirely trenchant
when all the races of Rallus longirostris are included.
The occurrence of King and Clapper Rails in the same breeding
grounds has been observed by several ornithologists. Robert E. Stewart
(personal communication) observed a King and a Clapper Rail
together with brood at Chincoteague Island on the coast of Virginia in
June 1951. He has also on numerous occasions observed King and
Clapper Rails together in the tidal marsh along Ape Hole Creek, a
tributary of Pocomoke Sound, Somerset County, Md. H. M. Stevenson
reported seeing a Clapper Rail walking directly in front of a King
Rail at Alabama Point, Ala., June 6, 1965 (Stewart, J. R., 1965,
p. 553). In April 1956, I collected a King Rail and a Clapper Rail
from the same pond at Grand Chenier, Cameron Parish, La. In this
area, the narrow chenier (stranded rim of the sea or old shoreline)
serves somewhat as a barrier between the fresh and salt marsh, and
these two species merely have to walk a hundred yards or so to be
together. It is difficult to separate the two species in the field in the
gulf coast marshes, although the breast of the resident Clapper race,
R. I. saturatus, is duller brown in contrast to the more rufescent breast
color of the King Rail.
On the South Atlantic coast, Ivan R. Tomkins (1958, p. 11) en-
countered a similar situation near Savannah, Ga. He wrote :
This brackish area, a place of transition from fresh to salt, has some peculiar
situations in respect to bird habitats. In the middle of Elba Island I have seen
both King and Clapper Rails on territory so close together that both birds were
in view at the same time.
In the New York City region, John Bull (1964, p. 169) reported
11 specimens and 19 sight records of King Rails in coastal salt marshes
and a January record of two King Rails feeding with a Clapper Rail
on a mud flat at Lawrence.
On May 18, 1960, John S. Webb and I observed a King Rail and a
Clapper Rail together in a brackish tidal marsh along the Delaware
Bay near Fleming's Landing, Kent County, Del. The mated pair (fig.
2) were observed on their nesting territory on numerous occasions
thereafter and were collected on June 11. The nest was also located
on that date, and the five eggs were removed and placed in an incuba-
tor. Despite the fact that optimal incubation conditions were main-
NORTH AMERICAN FAUNA 67
Figure 2.— Mated King Rail (subadult female) left, and Clapper Rail (adult
male) right, collected at Taylor's Gut, Kent County, Del., June 11, 1960. Eggs
of pair were fertile. (Photograph by Frederick C. Schmid.)
NATURAL HISTORY OF THE KING RAIL 9
tained (64 percent relative humidity and 37.8° C. forced draft)
(Wetherbee, 1959) the embryos died between the 17th and 19th days
of incubation. The embryos appeared to be normal, and the deaths
were believed to have been accidental rather than indicative of genetic
incompatibility.
Subsequent observations revealed that King and Clapper Rails
frequently were found together in the extensive brackish bay marshes
in the Taylor's Gut area known as Broadway Meadows and located
between Fleming's Landing and Woodland Beach, Del. (Meanley
and Wetherbee, 1962, pp. 453^57; Meanley, 1965, pp. 3-7).
During the breeding seasons of 1960-64, a series of specimens was
collected in the Broadway Meadows marsh for plumage analysis.
Specimens were obtained at three stations: (a) the upper reaches of
the brackish marsh at Fleming's Landing, where King Rails only
were observed ; ( b ) the outer brackish marsh at Woodland Beach on
Delaware Bay, where Clapper Rails only were observed; and (<?) the
intermediate area between these two stations at Taylor's Gut, where
both Kings and Clappers occurred. Specimens from the intermediate
area showed a wide variation from typical King plumage to typical
Clapper plumage (table 1).
In addition to the localities mentioned, there are undoubtedly many
other such areas in the brackish marshes of the Atlantic and Gulf
Coastal Plain where mixed King Rail and Clapper Rail populations
occur. In fact, almost any Coastal Plain river that has extensive brack-
ish marshes and a sizable fiddler crab population is a potential King-
Clapper mixing ground.
Table 1. — Specimens randomly collected from Taylor's Gut, Del., an area of mixed
King and Clapper Rail populations
Species and age Sex Collection
date
King Rail:
Adult Male May 13,1961
Do Female - Apr. 15,1963
Do do Aug. 23,1963
Do .do _ May 29,1964
Do do July 30,1964
Subadult do >June 11,1960
Do do June 30,1960
Juvenile do... July 14,1964
Clapper Rail:
Adult.. Male 'June 11,1960
Do do June 25,1960
Do do June 29,1964
Do Female. May 13,1961
Do.... ..do Aug. 30,1963
Hybrid(?):
Adult Male Aug. 23,1963
Do do Do.
Do Female Aug. 30,1963
1 Paired and active nest found.
348-693 O— 69-
Distribution and Migration
DISTRIBUTION
Unlike the Clapper Rail, which in the eastern United States is
mainly restricted to a rather narrow band of salt marshes along the
Atlantic and gulf coasts, the King Rail is found throughout the east-
ern half of North America. In general, its breeding range extends
from the Gulf of Mexico to southern Canada and from the Atlantic
coast to about the 100th meridian in the Great Plains (fig. 3).
Scale in Miles
400 »QO jOO
Figure 3. — Approximate breeding range and principal distribution of the King
Rail in North America (black shaded area along Gulf and Atlantic Coasts and
in Lower Mississippi Valley and peninsular Florida indicates main wintering
range).
The boundaries of the breeding range as given in the American
Ornithologists' Union Check-list (1957, p. 152) and supplemented
by additional records, mostly from the distribution files of the Section
of Migratory Non-Game Bird Studies, Migratory Bird Populations
Station, Laurel, Md., and from Audubon Field Notes, are as follows :
The northern boundary extends from southeastern North Dakota
10
NATURAL HISTORY OF THE KING RAIL 11
(western Dickey County), central Minnesota (Otter Tail and Hen-
nepin Counties), southern Wisconsin (Jamesville, Madison, Racine),
central Michigan (Saginaw Bay), southern Ontario (St. Clair Flats
to Toronto), and New York (Buffalo, Branchport, Ithaca, Long
Island), to Massachusetts. The western boundary extends from south-
eastern North Dakota, eastern Nebraska, western Kansas (Cheyenne
and Meade Counties), and central Oklahoma, to southern Texas
(Corpus Christi). Collections of specimens at Brownsville, Tex., on
September -27, 1911, and April 2 (year not given) (Griscom and
Crosby, 1925, p. 527) suggest the possibility of breeding in that area.
The eastern boundary extends from Massachusetts southward along
the Atlantic coast to the southern Everglades. The southern boundary
includes the gulf coast region, in places virtually to the edge of the
gulf itself.
Records of occurrence near or beyond the limits of the normal
breeding range are as follows: St. John's, Newfoundland (October
20, 1935) ; Wellington, Prince Edward Island, Canada (March 28,
1917) ; Ottawa, Ontario (May 7, 1896) ; Crane Lake, Ontario (July
31, 1931) ; Port Perry, Ontario (April 21, 1923) ; Bucksport, Maine
(November 22, 1909) ; Fargo, N. Dak. (October 15, 1925) ; Key West,
Fla. (November 2, 1895) ; Dry Tortugas, Fla. (May 1961) (W. B.
Robertson, personal communication) ; and Tlacotalpan, Veracruz,
Mexico (January 18, 1901).
The King Rail's principal wintering range coincides with that part
of the breeding range where the species is most abundant, in the tide-
water country from the Delaware Valley to southeastern Georgia, and
southward through interior Florida into the Everglades, westward
through the gulf coast marshes and the rice belts of Louisiana and
Texas, and north into the Arkansas rice belt.
The King Rail is a regular winter resident along the Atlantic
coast as far north as New York City and in the Mississippi Valley
to southeastern Missouri. Southernmost winter records in the United
States are from the Lower Rio Grande Valley, in the vicinity of
Brownsville, Tex. (December 28, 1911, and January 10, 1923) (Gris-
com and Crosby, 1925, p. 527) .
Some of the numerous winter records (mainly from the distribution
files of the Section of Migratory Non-Game Bird Studies, Migra-
tory Bird Populations Station, Laurel, Md.) north of the princi-
pal winter range are as follows: LaSalle, Ontario (December 15,
1930) ; Lome Park, Toronto, Ontario (December 26, 1960) ; Fal-
mouth, Maine (December 17, 1899) ; Cambridge, Mass. (December 30,
1896) ; Cape Cod, Mass. (December 30, 1951) ; Hillsdale, Mich.
(December 11, 1896) ; Detroit, Mich. (February 6, 1907) ; Port
Huron, Mich. (December 6, 1902) ; Vicksburg, Mich. (February 6,
1909) ; Prudensville, Mich. (December 7, 1938) ; Monroe County,
12 NORTH AMERICAN FAUNA 67
Mich. (December 30, 1934, and February 8, 1934) ; Bayside, Long
Island, N.Y. (December 24, 1924) ; Miami and Meade Counties, Kans.
(late December) ; and Montauk Point, Long Island, N.Y. (Decem-
ber 27, 1951).
In the New York City region, Bull (1964, p. 169) reported 15
winter specimens, 5 in December, 7 in January, 1 in February, and
2 the first week in March. Eleven of the fifteen were taken in salt
marshes, and there were 19 sight records in salt marshes, nearly all
in December and January.
On the basis of extensive field observations by several ornitholo-
gists, including Robert E. Stewart, Milton B. Trautman, D. J. Nich-
olson, T. D. Burleigh, Oliver H. Hewitt, and myself, and as a result
of an intensive literature review, the most important areas of con-
centration probably have been determined.
The King Rail occurs in greatest numbers in the vast coastal marsh
and ricefield area of southern Louisiana. Other areas supporting
high populations include the coastal marsh-rice belt of Texas; the
Arkansas rice belt ; the fresh and brackish tidal marshes of the Caro-
linas and Georgia; the Everglades, the Kissimmee Prairie, and the
St. Johns River marshes of Florida; and the tidal marshes of the
Delaware Valley and Chesapeake Bay. The Lake Erie marshes of
northern Ohio and the St. Clair Flats opposite Detroit, Mich., are
two important concentration areas in the North Central States.
MIGRATION
Throughout most of its range the King Rail is migratory. Evi-
dence of movements between wintering and breeding grounds is based
on recoveries of banded birds, and birds heard calling overhead at
night, striking beacons, and appearing in odd places such as city
streets during periods of migration.
The Atlantic Coastal Plain, particularly its outer section, and the
Mississippi Valley are important fly ways of the King Rail. The
occurrence of King Rails near the Atlantic coast during migration is
due to movements to and from breeding grounds in that area. King
Rails commonly breed at many places less than 50 miles from the
coast. Several known localities include Butler Island near Darien,
Ga. ; Savannah National Wildlife Refuge, Jasper County, S.C. ;
Georgetown, S.C. ; Currituck Sound, N.C. ; Norfolk, Va. ; the coastal
sea islands and Delaware Bay marshes.
A King Rail collected by I. N. Gabrielson in the Atlantic coast salt
marshes at Wachapreague, Va., August 25, and several taken by
hunters in September at Chincoteague, Va., indicate the probable
route of migration of at least some northeast Atlantic coast breeding
birds.
During 7 years' residence in the lower Mississippi Valley, I heard
migrating King Rails regularly every spring at Alexandria, La., and
NATURAL HISTORY OF THE KING RAIL 13
Stuttgart, Ark. On the night of March 11, 1956, single King Rails
were heard calling as they migrated northward over the city of Alex-
andria at 8:30, 9:30, and 11 p.m. They appear to be less vociferous
while migrating in the fall. The most commonly uttered call of mi-
grating King Rails is a chur-r-r-r-r (the r like the German "R").
Another call occasionally given is chac-chac-chac-.
Probably most fall migration takes place after molting, which is
completed about the first of September. In Delaware, I have collected
flightless birds in the last week in August that would still have been
flightless through the first week in September. However, some rails
collected in late August had nearly or completely renewed their flight
feathers.
The fall departure schedule for three species of rails at the Pa-
tuxent Wildlife Research Center, Laurel, Md., was determined by a
trapping and banding program extending from midsummer to early
winter. King Rails were the first to leave the area (the last by late
September) ; they were followed by Soras (the last by early Novem-
ber) , and lastly by Virginias (the last of December) .
David C. Hulse (personal communication) wrote that a definite
influx of King Rails is noticed annually at Decatur, northern Ala-
bama, in late September : "Local birds are still here and at this time
must be augmented by migrants. Departure is gradual and by late
October rails become gradually scarce."
King Rails breeding at the southern limit of their range in the
gulf coast region are probably permanent residents or may perform
short coastwise migrations.
Winter records for the Middle Atlantic and North Central States
suggest the possibility of permanent residency by some individuals.
In the Chesapeake Bay region of Maryland there are two records of
King Rails banded in August and recovered in the same marsh the
following January. Also, a 6-week-old chick banded July 12, 1968,
at the Patuxent Wildlife Research Center, Laurel, Md., was recovered
December 12, 1968, at the same place.
As of January 1966 there have been only two recoveries of King
Rails that migrated an appreciable distance from the point of band-
ing. Only one of these was a direct recovery (bird recovered within
12 months of banding date) . A 2-week-old chick was banded at Stutt-
gart, Arkansas County, Ark., on June 2, 1952, and recovered at Cut
Off, Lafourche Parish, La., December 1, 1952, having traveled a dis-
tance of about 350 miles. The other recovery concerned a King Rail
banded at Lassie, Wharton County, Tex., June 9, 1949, and recovered
at Brookville, Montgomery County, Ohio, 1,000 miles away, on May
2, 1951. A King Rail banded at Ruthven, Palo Alto County, Iowa,
August 25, 1951, and recovered at Lake View, Sac County, Iowa,
14 NORTH AMERICAN FAUNA 67
September 10, 1951, had traveled some 60 miles and was probably
migrating.
Spring arrival and fall departure dates are given in table 2. Some
of these dates are questionable; those for the interior northern part
of the range may be more reliable than those along the Atlantic coast
where so many King Eails winter.
LOCAL MOVEMENTS
Information on local movements was obtained at two marshy im-
poundments at Patuxent Wildlife Kesearch Center, Laurel, Md., dur-
ing the summers of 1965 and 1966. In 1965, four pairs of King Kails
nested in an impoundment known as Knowles Unit 2 ; in 1966, no King
Rails nested there because of deep water. Knowles 2 has an area of 20
acres, of which about 10 acres are shrub swamp or marsh or a mixture
of the two. Knowles Unit 1, the other impoundment, is larger, but
contained only about 6 acres of marsh and shrub swamp at the time.
One pair of King Rails nested there in 1965 and 1966.
Table 2. — Spring arrival and fall departure dates for migrating King Rails
Location Arrival Departure Source
Mississippi Flyway:
Stuttgart, Ark Feb. 15 to Mar. 30 Mid- October to mid- Meanley, unpublished.
November.
Missouri... Mar. 21 to 31... Late October Widmann, 1907, p. 59.
Buckeye Lake, Ohio Apr. 11 to 20. Late September Trautman, 1940, p. 229.
Chicago, 111.. Mid-April Late October Ford, 1956, p. 33.
Knox County, Ind. Apr. 15 to 30. Section of Migratory Non-
game Bird Studies, Migra-
tory Bird Populations
Station, unpublished.
Kansas Apr. 18 (median date) Johnston, 1964, p. 611.
Vicksburg, Mich Apr. 19 Oct. 10 (latest) Rapp, 1931, p. 7.
Southern Michigan Apr. 21 to May 10 Late August through Wood, 1951, p. 146-147.
September.
Southern Minnesota Late April Late September Roberts, 1936, p. 440.
Clay County, Iowa May 1 to 7 Early September Tanner and Hendrickson,
1956, p. 54.
Decatur, Ala Late October Hulse, personal communica-
tion.
Atlantic Flyway:
Raleigh, N.C Late March. Brimley, 1917, p. 299.
Western Pennsylvania Mid to late April Todd, 1940, p. 183-184.
New York, N.Y Late April.. Cruickshank, 1942, p. 156.
Connecticut River Valley September, early Bagg and Eliot, 1937, p. 178-
Massachusetts and October. 180. Sage and Bishop,
Connecticut. 1913, p. 48.
Broadway Meadows September Meanley, unpublished.
Kent County, Del.
Laurel, Md September Meanley, unpublished.
One of a pair of nesting King Rails trapped and banded on June 12,
1965, in Knowles 2, was first recaptured on August 2, approximately
200 yards from where it was banded. This bird was recaptured a second
time on August 7, approximately 360 yards from the second site and 500
yards from the original site.
The young of the 1965 nest in Knowles 1 were hatched on July 2 and
3. Earlier, one of the adults was banded and color-marked while incu-
bating. Both adults and the brood remained within 100 feet of the
nest most of the time until at least July 24, a period of 3 weeks.
NATURAL HISTORY OF THE KING RAIL 15
On July 8, 1966, the same color-marked adult, its mate, and their
brood of eight 2-week-old chicks were trapped in Knowles 1, 50 feet
from where the color-marked adult had nested the previous year. The
unhanded adult and the chicks were banded when trapped.
On July 16, two of the eight chicks were captured and the rest of
the family was observed on an island in Knowles 2, 0.4 mile from the
site of their original capture in Knowles 1 on July 8. To reach the
island in Knowles 2, the 3 -week-old flightless chicks had not only
walked nearly half a mile but had swum across 50 feet of open water
that had a depth of 3 feet.
Ecological Relations
The King Rail probably occurs in a wider variety of. habitats than
any other rail. The species ranges from coastal salt and brackish
marshes to shrub swamps and occasionally is found even in upland
fields near marshes where it forages for grasshoppers and grain, and
where it sometimes nests.
The distribution of the King Rail's habitat coincides rather closely
with that of the muskrat {Ondatra zibethicus) . Muskrats create opti-
mum habitat for rails by opening up marshes and producing networks
of pathways leading to plunge holes. When the tide goes out, water is
trapped in the holes, and rails use them as drinking places. Muskrat
trails are also favorite places for crayfish burrows. The crayfish are a
prime food of the rails and are usually carried to the tops of muskrat
houses for eating.
Because of the geographic as well as the local variation in habitats
of the King Rail, the ecological relations will be discussed on a regional
basis.
This chapter will cover both my own observations on the ecology of
the King Rail in Louisiana, Arkansas, South Carolina, Delaware, and
Maryland, and those of other authors in different States or areas.
LOUISIANA GULF COAST MARSHES
The King Rail and the Louisiana Clapper Rail, a brownish form
virtually indistinguishable from the King Rail in the field, occur to-
gether in some sections of the Louisiana coastal marsh. This vast marsh
area of more than 4 million acres is divided into three major divisions,
each of which is a distinct habitat type and will be discussed sepa-
rately : the delta marsh, subdelta marsh, and prairie marsh (St. Amant
1959, p. 97-101) (fig. 4).
The delta marsh
The delta marsh, near the mouth of the Mississippi River, comprises
only 7 percent of the total coastal marsh area of Louisiana. Important
plant species of the delta marsh are cattails (Typha spp.) , roseau cane
or reed (Phragmites communis) , common three-square (Scirpus ameri-
canus) , dog-tooth grass (Panicum repens) , giant cutgrass (Zizaniopsis
miliacea), saltmarsh cordgrass (Spartina alterniflora) , delta duck
potato (Sagittaria platyphylla) , alligatorweed (Altemanthera phil-
oxeroides), and water hyacinth {Eichhomia crassipes).
16
NATURAL HISTORY OF THE KING RAIL
17
LOUISIANA MARSHES
■ DELTA MARSH
X) SUB-DELTA MARSH
= PRAIRIE MARSH
| KING RAIL CENSUS AREA
• PLACE NAMES
g J/^CUT OFF
Figure 4. — Louisiana gulf coast marshes.
Oberholser (1938, p. 109, 201) reported the collection of a King
and six Clapper Rails in the delta marsh at the mouth of the Mississippi
River, indicating their presence in this habitat type.
The subdelta marsh
The subdelta marsh, comprising 74 percent of the Louisiana coastal
marsh, extends westward from the Mississippi River Delta to Cow
Island and Chenier au Tigre in Vermilion Parish. Both its fresh-water
and brackish marshes are of two types, floating or with a firm floor of
clay. The predominant plant species in both types is "paille fine" or
maidencane (Panicum hemitomon). Associated with it are cattail,
southern bulrush (Scirpus calif ornicus) , sawgrass (Cladium jamai-
cense), wapato (Sagittaria latifolia), alligatorweed, and water hya-
cinth. In brackish areas either saltmeadow cordgrass (Spartina patens)
or Olney's three-square (Scirpus olneyi) is dominant; the latter is
dominant if there is management (burning) for muskrat production.
Salt marshes in this area are dominated by needlerush (Juncus roe-
marianus), saltmeadow cordgrass, and saltmarsh cordgrass.
Both banding returns and collections of birds substantiate the oc-
currence of King Rails in the subdelta marsh. A King Rail banded
at Stuttgart, Ark., in June 1952, was recovered at Cut Off, Lafourche
18 NORTH AMERICAN FAUNA 67
Parish, in the subdelta marsh, in December 1952. Cut Off is approxi-
mately 25 miles south of New Orleans. King Rails were collected at
Chenier au Tigre, January 1, 3, and 5, 1934, by A. M. Bailey (Ober-
holser 1938, p. 199) ; at the same location on March 31 and April 1,
1947, by I. N. Gabrielson; and at Avery Island, May 7, 11, 13, and 15,
1930, by E. G. Wright (Oberholser, 1938, p. 199).
I made a census, based on calls, in Terrebonne Parish, 1.2 miles south
of Dulac, on January 3, 1963, to determine the abundance of King
Rails in the area (table 3) . The birds were heard calling from what
appeared to be an abandoned silted-in canal where shallow ponds were
interspersed with dense patches of vegetation dominated by clump
grass (Spartina spartinae) . In a 1-mile strip, 50 feet wide, 19 King
Rails were counted. Short-billed Marsh Wrens (Cistothorus platensis) ,
Soras, Virginia Rails, and Common Gallinules {Gallinula chloropus)
were also common in this same census strip.
The prairie marshes
The prairie marshes in the southwestern part of the Louisiana gulf
coast (Vermilion and Cameron Parishes) comprise 19 percent of the
total area. Near the gulf coast much of the prairie marsh is bisected
by ridges known as cheniers (stranded rims of the sea or old shore-
line) that parallel the coast (fig. 5). Cheniers extend in straight
lines for many miles and in most places are wide enough only for a
Figure 5. — Prairie marsh, Grand Chenier, La., March 1956. Mixed King and
Clapper Rail populations sometimes occur in the same marsh type in this area.
Both species were collected from the same half-acre pool near here, April 1956.
NATURAL HISTORY OF THE KING RAIL 19
road bordered on each side by a line of live oaks. In some places there
are a few houses. In some sections, cheniers separate fresh and brackish
marshes.
Table 3. — Abundance of King Rails in certain areas, as indicated by censusing
Location Number of King Rails Date Cover type
Black and Pedee Rivers, 25 in 100 acres » 2 Apr. 10-12, 1961 Giant cutgrass, cattail,
Georgetown County, S.C. big cordgrass, arrow-
arum.
Savannah River, Jasper 14 in 13 acres 13.. Apr. 20, 1961 Softstem bulrush.
County, S.C.
10 miles south Fellsmere, 30 in 100 acres 3 May 8, 1964 .-. Maidencane.
Indian River County, Fla.
Savannah National Wildlife 46 along 7-mile route < . Apr. 12, 1960 Giant cutgrass, cattail,
Refuge, Jasper County, sawgrass.
S.C.
Dulac, Terrebonne Parish, 19 along 1-mile route 4. Jan. 3, 1963 Clump grass, or needle
La. cordgrass.
4 miles north Creole, 24 along 1-mile route *. Jan. 5, 1963 Fallpanicum.
Cameron Parish, La.
3 miles north Pecan Island, 20 along 1-mile route 4. Jan. 4, 1963 Fallpanicum.
Vermilion Parish, La.
Stuttgart, Arkansas County, 22 along 6-mile route *. Apr. 1955 -- Rice stubble, broom-
Ark, sedge, cattail, softrush.
1 Males only.
2 Two-stage sampling.
• Strip census.
* Roadside count.
In this area salt marshes near the coast are dominated by a salt-
grass-saltmeadow cordgrass-saltmarsh cordgrass association. Land-
ward from this association, brackish marshes extend north to the
Creole and Grand Chenier ridges. Principal plants of the brackish
marsh are saltmeadow cordgrass, Olney's three-square, and saltmarsh
bulrush (Scirpus robustus). In the transition areas between brackish
and fresh water, such plants as giant cutgrass, bull tongue, pickerel-
weed (Pontederia cordata), and wild millet (Echinochloa crusgalli)
grow. The fresh-water marshes lie mainly north of the Grand Chenier
and Creole ridges. In the higher parts of those marshes the following
plants are found : bull grass (Paspalum boscianum) , lake grass (Pas-
palum distichum), dotted smartweed (Polygonum punctatum),
squarestem spikerush (Eleocharis quadrangulata) , and delta duck
potato. Sawgrass is the climax type in the lower parts of the fresh-
water marsh.
In the vicinity of the cheniers, King and Clapper Rails occur close
to one another or together. Referring to this situation, Lowery (1955,
p. 227) made the following statement :
The King and Clapper Rails are extremely similar in appearance and are,
for the most part, simply ecological representatives of each other. The former
generally inhabits fresh-water marshes and is widespread in the interior of the
United States; the latter is always found on or near the seacoast in brackish-
or salt-water marshes. . . . There are brackish marshes in which both breed
side by side without intermingling ; . . .
I find it difficult to believe that the two species do not interbreed in
the prairie marshes. Several King-Clapper pairs (and their nests and
20 NORTH AMERICAN FAUNA 67
eggs) have been collected recently in Delaware brackish marshes, in-
dicating that they do sometimes interbreed when they occur together
(Meanley and Wetherbee, 1962, p. 453-457) .
Near the village of Grand Chenier, I collected both Kings and Loui-
siana Clapper Eails from the same small pond on the south side of the
chenier. The dominant vegetation in the immediate area was clump-
grass. On the south side of this narrow chenier, in the brackish
marshes, the gulf coast form of the Clapper Rail is the dominant
species, but the marsh on the landward side is the King Rail's domain.
Rivers, such as the Mermenteau, and canals crossing the chenier ex-
tend the brackish water landward, and occasionally storm tides also
affect large areas of the marsh, extending salt water into the fresh-
water zone and changing the habitat. This area may well be described
as a mixing ground of plants and animals. A common avian associate
of the rails breeding in the clumpgrass and saltgrass marsh was the
Mottled Duck (Anas fulvigula).
I also encountered four King Rail broods, still in downy black
plumage, and three single adults 4 miles north of Grand Chenier on
July 23, 1955. At this station the marsh was composed of a mixture
of southern bulrush, cattail, a Sagittaria* probably land folia, and
water-hyacinth.
A census of King Rails, based on calls, was made in a marsh border-
ing the Pecan Island road, 2 miles south of the old Intracoastal Canal,
Vermilion Parish, on January 4, 1963. Twenty birds were counted in
20 minutes along a 1-mile strip approximately 200 yards wide, at 6
p.m. (table 3). The dominant vegetation in the census area was fall
panicum (Panicum dichotomiflorum) . A similar census was made 5
miles south of the Intracoastal Canal on the east side of the road to
Creole, Cameron Parish, on January 5, 1963. Between 5 :30 p.m. and
6 p.m. 24 birds were counted along a 1-mile strip approximately 200
yards wide (table 3) . The dominant vegetation types in the area were
southern bulrush and fall panicum (fig. 6). Soras were also abundant
in the same habitat.
SOUTHERN RIGEFIELDS
The gradual shift in the domestic rice (Oryza sativa) growing in-
dustry from the South Atlantic coast to the South Central States of
Louisiana, Texas, Arkansas, and Mississippi after the Civil War
opened up a new marsh habitat for King Rails and other water birds.
Much of the land where rice is grown today was once a vast natural
tall-grass prairie in which the Greater Prairie Chicken (Tympanu-
chus cupido) was abundant. Harmon, Thomas, and Glasgow (1960, p.
153) reported that approximately 3 million acres in this area were
devoted to rice growing by 1958, and that this acreage wintered 4
million ducks and geese. Many aquatic plants grow in ricefields, and
NATURAL HISTORY OF THE KING RAIL
21
Figure 6. — Southern bulrush (Scirpus californicus) , fall panicum {Panicum
dichotomiflorum) , and alligatorweed (Alternanthera philoxeroides) in prairie
marsh type near Creole, Cameron Parish, La., January 5, 1965. Between 5 :30
and 6 p.m., January 5, 1963, 24 King Rails were counted along a 1-mile transect
through this marsh.
virtually all produce seeds utilized by a variety of water birds. Rice-
fields furnish an optimum all-purpose habitat for King Rails for
nearly 6 months during the summer half of the year, and a source of
food for them in winter (fig. 7) .
On the gulf coast prairie of Louisiana and Texas, rice planting be-
gins in March. Some early varieties are harvested by late July, but
most fields are harvested from early August to early October. The
planting season in Arkansas is about 2 weeks later, and harvest is
from late August to early November. The fields are irrigated by wells
or by canal systems fed from reservoirs or bayous. Water remains on
the fields for 3 or 4 months and is maintained at a constant level of
from 6 to 10 inches.
On the Arkansas Grand Prairie I found the nesting density of
King Rails in one ricefield to be at least one nest per 15 acres, a figure
based on the location of five nests in a 75-acre field in July. These nests
were located by a team of men walking abreast and systematically
covering the field to remove a pest plant, the coffeebean {Sesbania
exaltata) . The height of the nesting season was several months past,
and these nests probably represented a renesting effort or a second
22
NORTH AMERICAN FAUNA 67
Figure 7. — King Rail wading through ricefield toward nest on dike, Stuttgart,
Ark., July 13, 1952.
nesting after an earlier first successful nesting. It is not to be con-
strued that this is an average nesting density for Grand Prairie
ricefields.
Since much of the rail nesting in rice country is completed before
the rice is high enough to provide nesting cover, a better idea of nest-
ing density could be obtained from nest counts or mating call counts
in the spring when most of the rails are found in roadside ditches
and canals and occasionally in rice stubble. As an example, in April
1955 I located 22 occupied nesting territories along 6 miles of con-
tinuous roadside ditch beginning 2 miles north of Stuttgart, Ark.
(table 3 and fig. 8).
In Evangeline and Jefferson Davis Parishes in Louisiana I found
many nests in roadside ditches where the dominant vegetation was
paille fine (maidencane) and softrush (Juncus effusus). In Arkansas
Grand Prairie ditches in 1952 and 1953, nests were found mainly in
stands of softrush, cattail, common spikerush {Eleocharis palustris),
and lake sedge ( Carex hyalinolepis and C. lacustris) , a plant which
grows to a height of 3 feet or more and forms very dense stands that
persist intact through the winter. Lake sedge was available for nest-
ing cover earlier than any plant in the roadside ditches. Ten years
NATURAL HISTORY OF THE KING RAIL
23
Figure 8. — Arkansas Grand Prairie near Stuttgart, Ark., May 1952. Rails nest
in roadside ditches in April and May. Vegetation in ditches is mostly softrush
(Juncus effusus) and two sedges (Carex stipata and C. hyalinolepis) . In June,
rails move into ricefields (left of ditch) for late nesting or renesting as rice
begins to form nesting cover.
later (1962), in those same ditches, awl-fruited sedge (Carex stipata)
was the dominant plant, and four of six nests located during May
of that year were constructed of this plant.
Old rice stubble is sometimes used for nesting. On the southwestern
Louisiana rice prairie where farming is less diversified than on the
Arkansas Grand Prairie, many farmers let the stubble fields lie out
through the winter and spring for cattle grazing. In one such wet
stubble field at Mamou, Evangeline Parish, I located two rice-straw
nests on May 5, 1957 (fig. 9).
During the summer, when the rice is growing and the fields appear
as a vast green marshland, virtually all King Rails in the rice belt
frequent the fields. Some are renesting, and others are wandering
about with their broods in search of crayfish, minnows, and aquatic
insects which abound here.
Nesting associates of the King Rail in Louisiana ricefields are the
Fulvous Tree Duck (Dendrocygna bicolor), the Purple Gallinule
(PovphyrvZa martinica), the Least Bittern (Ixohryclms exilis), and
along the southern border of the rice belt the Mottled Duck. The most
common bird in the area is the Red- winged Blackbird (Agelavus
phoenicew) . In the northern part of the principal Louisiana rice
belt, at Mamou, Evangeline Parish, I found the Long-billed Marsh
Wren (Telmatodytes palustris) nesting in rice.
24 NORTH AMERICAN FAUNA 67
Figure 9. — Nest of King Rail in wet rice stubble, Mamou, La., May 5, 1957.
Photographed as found.
In 1955 a rice farmer at Mamou located six Fulvous Tree Duck
nests in a 400-acre block of rice, and in the same locality I found 22
active Purple Gallinule nests in a 10-acre section of a 25-acre ricefield.
In the Arkansas ricefields the Purple Gallinule is an uncommon
breeding bird. There are no breeding records for the Fulvous Tree
Duck in Arkansas ricefields, but there are several early fall occurrence
records. The Short-billed Marsh Wren has been found nesting in
Arkansas ricefields in August and September.
King Rails are more secretive in winter than at other seasons and
often are present in good numbers in some localities although seldom
or never seen. For 5 years in the Arkansas rice belt, I was in the field
daily without ever seeirg one in the dead of winter. Yet they were
present, as a mink trapper brought me several each January.
On the Arkansas Grand Prairie, I have often come across trails
forming tunnels which often continue for many feet beneath the
matted vegetation of a ditch bank. These trails appear to have been
made by some mammal, yet many tell-tale signs, particularly the
characteristic regurgitated pellets and roundish droppings about
the size of a silver dollar, are proof that King Rails use them. Regard-
less of whether these trails are made by rails, mink (Mustela vison),
rabbits (SyUilagus floridanm), or rats (unidentified) , trapping indi-
cates that they are used by all four species. King Rails also spend
the winter in small marshy tracts along the bayous that dissect the
Grand Prairie.
At Stuttgart, Ark., a King Rail used a long water pipe about IV2
feet in diameter and running from a pumphouse to a small reservoir
NATURAL HISTORY OF THE KING RAIL 25
as its winter retreat. From late November to mid-December, I stopped
by almost daily to see its fresh tracks leading from the pipe to the
mudflat of the reservoir.
A rather surprising wintering habitat is the cutover longleaf pine
(Pinus pakbstrls) land of central Louisiana. Bobwhite and Wood-
cock hunters flush King Rails from little damp spots or seepage
areas in the bluestem {Andropogon tener and A. diver gens) range.
Crayfish, a prime food of the rail, also are found there.
FLORIDA
A. H. Howell (1932, p. 202), in commenting on the status of the
King Rail in Florida during the early 1930's, said it was probably
most numerous in the Everglades and big marshes of the upper St.
Johns River. D. J. Nicholson of Orlando (personal communication,
1962), who has made an intensive study of Florida birds since 1900,
told me the King Rail is still a common to abundant breeding bird in
many parts of central and southern Florida, although extensive drain-
age projects in the area have destroyed thousands of acres of marsh
habitat. In addition to the two areas mentioned by Howell, Nicholson
included the open wetlands of the Kissimmee Prairie as an important
King Rail area. He added that the King Rail is common in the
St. Johns River marshes in Seminole, Orange, Volusia, Brevard, and
Indian River Counties; and is found in good numbers nesting on
Merritt Island, Brevard County, "both in the salt marshes near
Wayne's Clapper Rail, as well as in numerous fresh water ponds on
that island."
S. A. Grimes (personal communication) reports that, in northern
Florida, the King Rail occurs in most of the fresh-water marshes
of Duvall County. Two of the several nests he found were in open
cypress bayheads.
A. D. Cruickshank (personal communication), in writing from
Brevard County, said that the King Rail is decidedly more common
there in winter than during the breeding season, with peak numbers
usually coming in late December and January. Apparently the local
population is augmented by migratory populations from north of
Florida. Evidence of local abundance in this area is based on the
annual Audubon Society Christmas bird count conducted within a
15-mile radius of Cocoa, Fla. (Cruickshank et al., 1953-66). The
numbers of King Rails reported has ranged between 11 and 93 over
the past 14 years and averaged 40 per year. Cruickshank reports that
the best localities are (a) fresh-water marshes around Lake Poinsett,
a large lake in the St. Johns River, and (b) fresh- water marshes on
Merritt Island.
During the period May 4-8, 1964, I examined some marshes in
Indian River, Osceola, and Glades Counties. Approximately 10 miles
south of Fellsmere, Indian River County, at the junction of State
348-693 O — 69—3
26 NORTH AMERICAN FAUNA 67
Highways 60 and 512, I found King Kails common where maiden-
cane and pickerelweed formed a high percentage of the vegetation of
the wetter marshes (fig. 10). Apple snails (Pomacea palmdosa), the,
Figuee 10. — Habitat of King Rail, 10 miles south of Fellsmere, Indian River
County, Fla., April 1967. Marsh vegetation in foreground is mostly maideneane
(Panicum hemitomon) and pickerelweed (Pontederia cordata), in background
sawgrass (Cladium jamaicense). White waterlily (Nymphaea odorata) in
pond left of center. Forest community is pond cypress (Taxodium ascendens).
The density of King Rails in this area was estimated at 30 birds per hundred
acres.
major food of the Limpkin (Aramus guarauna), and the eggs of
these snails, were scattered abundantly throughout the wetter marshes,
but were absent from the drier ones. Houses of the round-tailed
muskrat (Neofiber alleni) were abundant in both wetter and drier
marshes. Limpkins and rails use the tops of these houses as their
"dinner tables." King Kails were most commonly in the drier marshes.
I estimated a density of approximately 30 birds per hundred acres
in a tract on the east side of Highway 512 (table 3).
On the Brighton Seminole Indian Reservation, Glades County, I
heard and saw King Rails in marshes composed largely of pickerel-
weed, bull tongue (Sagittaria laneifolia) , and dotted smartweed (fig.
11). While oh a trip through the reservation in January 1958, I saw
two very dark-plumaged King Rails in a small pickerelweed marsh.
NATURAL HISTORY OF THE KING RAIL 21
Figure 11. — King Rail habitat on Seminole Indian Reservation, Glades County,
Fla., May 5, 1964. Predominant plants in marsh are bull tongue (Sagittaria
lancifolia), pickerelweed (Pontederia cordata), and dotted smartweed (Poly-
gonum punctatum) .
SOUTH CAROLINA LOW COUNTRY
King Kails are common to abundant in many fresh-water and
brackish tidal-river marshes of the South Carolina Low Country.
These marshes are along the famous rice rivers of colonial times : the
Ashepoo, Black, Combahee, Edisto, Pee Dee, Santee, Savannah, Wac-
camaw, Wando, and others. It was in such marshes that domestic rice
was grown until about 1915. Remnants of the old ricefield dikes and
canals built by slaves are still evident in the marshes (fig. 12).
The dominant vegetation type of most sections of the marshes today
is giant cutgrass. Because of the blanched appearance of the giant
cutgrass in winter, these marshes were referred to by the early
explorer-naturalists as the "white marsh." Giant cutgrass provides
excellent escape and nesting cover for rails but apparently is of no
food value to them, although Purple Gallinules, Red-winged Black-
birds, and Bobolinks (Dolichonyx oryzworus) feed on its flowers and
seeds.
A survey of the marshland in the Low Country from the lower
Cape Fear River at Wilmington, N.C., to the Altamaha River at
Darien, Ga., seems to indicate that the King Rail is largely associated
with the "white marsh" zone of these coastal rivers. It should be
emphasized, however, that many secondary plant communities occur
within this zone with varying King Rail population densities depend-
ing upon local ecological conditions.
NORTH AMERICAN FAUNA 67
Figure 12. — Floodgate and ricefield canal near Savannah, Ga., April 1960. The
land is irrigated as the tide rises from the Savannah River. King Rails nest
in giant cutgrass (Zizaniopsis miHacea) seen along right-hand edge of canal.
The high population density in this area is probably due to good nesting
cover and the abundance of the red-jointed fiddler crab (Uca minax), a
favorite food of the rail.
In the Low Country, March and April appear to be the best months
for censusing, as King Rails are more vociferous at this time than dur-
ing any other period of the year. A narrow strip of marshland border-
ing a river is the most suitable place for censusing.
The marshland on the west side of the Pee Dee River and one of its
tributaries, the Black River, for a distance of 8 miles north of George-
town, S.C., totalling 3,000 acres, was selected as a sampling area.
The sampling design was suggested by Dr. Don W. Hayne of the
Institute of Statistics, North Carolina State College, Raleigh, N.C.
Georgetown is approximately 8 miles inland from the coast on
Winyah Bay at the confluence of the Pee Dee and Waccamaw Rivers.
The Black River flows into the Pee Dee about 2y2 miles north of
Georgetown. The mean tidal range at Georgetown is Sy2 feet. At
Georgetown the river is slightly brackish, and big cordgrass (Spartina
cynosuroides) is an important plant component of the marsh, espe-
cially along old ricefield canals. Six 10-acre plots were composed
mainly of the following plants : big cordgrass, 35 percent ; Olney's
three-square, 20 percent; cattail, 14 percent; giant cutgrass, 13 per-
cent; arrow-arum (Peltandra virginica), 11 percent; softstem bul-
rush, 5 percent; and river bulrush (Seirpvs fluviatilis) , 2 percent.
Vegetation analyses were made also of three 10-acre plots 8
miles north of Georgetown on the Black River. On the Black River,
NATURAL HISTORY OF THE KING RAIL 29
giant cutgrass formed 52 percent of the marsh vegetation, arrow-
arum 36 percent, and cattail 12 percent.
For the actual census, the entire marsh area, as shown on U.S.
Geological Survey maps, was blocked off into 64 primary sampling
units, each a 630-foot-wide transect extending from the edge of the
river to the land. Each transect was divided into 10-acre plots, the
number in a transect depending upon the width of the marsh at that
location along the river.
Ten of the 64 transects were randomly selected as primary sampling
units, and one 10-acre plot from each was chosen for censusing. At
least 1 hour between 5 and 8 a.m. or 5 and 7 p.m. during the period
April 10 through 12, 1961, was spent in each plot counting calls.
The number of male King Rails in a transect was estimated by
multiplying the number of 10-acre plots in that transect by the num-
ber of birds heard in the census plot. The estimated total of 755 males
in the 3,000-acre marsh was derived by multiplying the average num-
ber of birds per sample transect by the total number of transects. The
density estimate of 25.2 male birds per 100 acres was calculated by
dividing the total population by 30 (the number of 100-acre units in
the 3,000-acre marsh). Since some of the calling King Rails were
undoubtedly already mated, and most of the others would be eventu-
ally, the average number of breeding rails per 100 acres could be
inferred to be 25 pairs (table 3).
Sampling indicated that the density of the King Rail population
was higher at Georgetown in the Pee Dee River marshes than several
miles up river along its tributary, the Black River. One 10-acre plot
at Georgetown had six calling rails, and four other plots had four
each. None of the Black River plots had more than two rails. The
higher density at Georgetown could be attributed to the higher pro-
portion of red-jointed fiddler crabs (Vca mlnax).
Savannah National Wildlife Refuge
The Savannah National Wildlife Refuge in South Carolina and
Georgia, near Savannah, is about 25 miles -upriver from the ocean on
what was formerly a rice plantation (fig. 13) and is divided by the
Savannah River. The larger acreage is on the South Carolina side.
Its marsh is the fresh tidal type, with a tide which rises about 1 foot.
Giant cutgrass is the dominant vegetation of much of the marsh
on the refuge. The old growth of cutgrass forms a nearly pure stand,
and has an average height of about 5 feet, but will average higher
when the new growth matures. Arrow-arum, dotted smartweed, and
swamp smartweed {Polygonum hydropiperoides) are scattered about
the marsh, particularly along the edges and on high spots. There are
numerous small holes made by the red-jointed fiddler crab, an impor-
tant food of the King Rail, along the tidal creeks and edges of the
marsh. While making a survey of King Rail populations in April
30
NORTH AMERICAN FAUNA 67
, ? '•• ..
Figure 13. — South Carolina Low Country : ricefield nesting habitat in Jasper
County along the Savannah River, fall 1958.
NATURAL HISTORY OF THE KING RAIL 31
1960, 1 estimated a density of one pair per acre in the cutgrass marsh
bordering the river. A year later I also heard many King, Virginia,
and Sora Eails calling at night in the same marsh.
Much of the refuge canal system and some of the ponds are choked
with alligatorweed, a plant that forms extensive mats upon which
rails, gallinules, coots, herons, and several species of ducks do much
of their foraging for aquatic insects (fig. 14). Small patches of giant
S&
0*4 M
*****
< ■■"■ ..
Figuke 14. — Alligatorweed (Alternanthera philoxeroides) in canal at Savannah
National Wildlife Refuge, Jasper County, S.C., April 1960. This plant forms
spongy, extensive mats upon which rails, gallinules, coots, herons, and ducks
forage for aquatic insects, fish, amphibians, and crustaceans. Such mats have
an obvious value to birds that utilize its growth form to facilitate their quest
for food. Note alligator on far side of canal. Alligators feed on many forms
of animal life including various water-birds such as rails.
cutgrass in which Purple Gallinules nest are sparsely distributed
along the canals. I located many King Kail pairs with feeding terri-
tories along sections of the canals. Some of these territories were not
more than 20 feet square, indicating the high food productivity of these
aquatic mats. All of the King Rails that I observed feeding in the
choked-up canals, however, nested on the other side of the dike in a
deep-water impounded marsh containing a mixture of giant cutgrass,
sawgrass, cattail, royal fern (Osmunda regalis), buttonbush (Ceph-
alanthus occidentalis) , and myrtle {Myrica cerifera) .
I made a roadside count of calling males on April 12, 1960, along
a 7-mile route beginning at the north entrance to the refuge on U.S.
32 NORTH AMERICAN FAUNA 67
Highway 17, continuing along the east dike, through the string of
island hammocks, and ending at the Savannah Eiver just inside the
south entrance on U.S. Highway 17. On this census, made between
6 and 7 p.m., 46 males were tallied (table 3) .
On April 20, 1961, I estimated the breeding King Rail population
in a nearly pure softstem bulrush (Scirpus validus) marsh along
U.S. Highway 17A, about 2 miles north of Savannah, Ga. The bulrush
averaged about 5 feet in height, and the marsh had a firm bottom
covered with 1 to 2 inches of water. A 13-acre section was marked
off into transects, and King Rail territories were then spot-mapped
on the basis of three calls from any one area. This mating-call count
indicated a breeding population of 14 males in the 13-acre tract
(table 3).
Upper Savannah River Valley
King Rails also nest further upriver in the Savannah River Valley
section of the Upper Coastal Plain in South Carolina. Norris (1963,
p. 2, 19) described the typical nesting habitat as a "Carolina bay" —
an oval-shaped water-filled depression with rank growths of maiden-
cane and other aquatic plants.
CHESAPEAKE BAY COUNTRY
Tidewater Virginia
Tidewater Virginia is the section of the Middle Atlantic Coastal
Plain that extends from the fall line (the line separating the Piedmont
Plateau from the Coastal Plain) to the Chesapeake Bay. It is dissected
by numerous rivers, the largest of which are the Potomac, the Rappa-
hannock, the York, and the James.
The King Rail is common throughout the year in much of Tidewater
Virginia and usually occurs in greatest numbers in marshes where
big cordgrass is dominant. Big cordgrass is one of the best cover plants
for King Rails in Tidewater because of its height and occurrence in
fairly dense stands, and because it retains its life form throughout
most of the year (fig. 15). In the early 1960's, I found King Rails
common in the big cordgrass marshes at Norfolk, West Point, and
Tappahannock.
Other marsh types, especially Olney's three-square, wild rice
(Zizcmia aquatica), and cattail, are important for the King Rail,
but there is less acreage of these types, and wild rice does not provide
cover in the winter. During the winters of 1958 and 1961, 1 encountered
several muskrat trappers who were inadvertently catching King Rails
in the extensive Rappahannock River brackish marsh flats across the
river from Tappahannock. These flats are dominated by Olney's
three-square (fig. 16). Several King Rails were removed from muskrat
NATURAL HISTORY OF THE KING RAIL
33
Figure 15. — Big cordgrass (Spartina cynosuroides) (tall plant) and arrow-
arum (Peltandra virginica) (broad-leaved plant next to water) along tidal
creek, Nanticoke River marsh, Wicomico County, Md., August 1967. Big
cordgrass usually grows along the margins of tidal guts in brackish bay
marshes, but may form extensive, nearly pure stands in brackish tidal-river
mashes. It is one of the most important cover types for King Rails in the
Chesapeake Bay region. (Photograph by Luther Goldman.)
34
NORTH AMERICAN FAUNA 67
Figure 16. — Winter abode of King Rail. Rappahannock River near Tappahan-
nock, Va., January 1961. Vegetation is mainly Olney's three-square (Scirpus
olneyi) and saltmarsh cordgrass (Spartina alternifiora).
traps as I watched a trapper run his line. In these and other tide-
marsh habitats, tidal action along the creeks and over adjacent marsh-
lands keeps the water open throughout much of the winter.
At Hog Island, Surry County, in the James River opposite historic
Jamestown, C. C. Steirly found both King and Clapper Rails breeding.
Steirly (1959, p. 47-48) made the following comments about the rail
habitat on the island :
Apparently there is a salinity gradient between the east side of the refuge
and the west side of Cobham Bay. Hog Point might be the dividing line. There
seems to be a slight difference in the tidal vegetation between the two sides of
the refuge although the cord grass marsh seems to be the dominant feature
along the east or down river side. The King Rail is most often seen on the west
side ; however, there is as yet no proof that it does not breed on the east side.
In one of the particular haunts of the King Rail, pickerel weed (Pontederia
cordata) and bulrush (Scripus robustus) occur in some abundance where there
is less tidal fluctuation.
Virginia Eastern Shore
The Eastern Shore peninsula of Virginia lies between Chesapeake
Bay and the Atlantic Ocean. While King Rails would be expected
to occur on the bay side of the peninsula, their presence on the off-
shore barrier islands on the ocean side would seem rather surprising ;
nevertheless, on these salty coastal islands King Rails are found in
NATURAL HISTORY OF THE KING RAIL 35
land-locked fresh-water marshes. Montagna and Wimsatt (1942, p.
434-436) collected a female on Rogue Island 11 miles off the coast be-
tween Hog and Cobb Islands. The specimen had a fully developed
swollen brood patch, and its oviduct contained an egg with shell.
There are several records from Chincoteague Island. Robert E.
Stewart encountered paired adult King and Clapper Rails with a
brood in a salt meadow cordgrass marsh on this coastal island. The
Chincoteague salt marshes are one of the important Clapper Rail
hunting grounds along the Atlantic Coast, and King Rails occasion-
ally turn up in hunters' bags.
One of the best King Rail areas on the bay side of the eastern shore
peninsula is Bullbegger Creek, a tributary of the Pocomoke River.
Big cordgrass is the dominant plant in this creek marsh.
At Knott's Island, at the head of Currituck Sound, partly in Vir-
ginia and partly in North Carolina, A. J. Duvall (1937, p. 462) and
party collected a female King Rail and five chicks along a roadway
in a salt marsh on June 1, 1936.
Maryland Eastern Shore
In Maryland the King Rail is mainly associated with tidal marshes
of the Chesapeake Bay system, and is found in greatest numbers in the
extensive brackish tidal-river marshes of the Eastern Shore, especially
in the vast area of fresh and brackish bay marshes of Dorchester
County (see R. E. Stewart, 1962, for a description of Maryland Chesa-
peake Bay marsh communities). In this area, the following plants are
usually present as pure stands or are found in some combination in
areas where King Rails occur : big cordgrass, broad-leaf and narrow-
leaf cattail (Typha latlfolia and T. angusti folia), Olney's three-
square, switchgrass (Panicum virgatum), softrush, and rosemallow
(Hibiscus moscheutos) .
The importance of big cordgrass in the Maryland section of the
Chesapeake Bay is comparable to that in Tidewater Virginia.
In the brackish tidal-river marsh community of the Choptank River
at Dover Bridge between Talbot and Caroline Counties, a muskrat
trapper caught 50 King Rails in a single season (January 1 to March
15) . Most of the birds were caught where big cordgrass was dominant
but usually mixed with Olney's three-square and switchgrass. Because
of the sparseness of winter marsh cover, King Rails often seek means
of escape and places for hiding different from those used during the
rest of the year. A muskrat trapper on the Choptank River in Mary-
land reports that whenever he surprises a rail along a tidal gut in
the marsh it almost invariably darts into a muskrat hole along an
embankment.
Robert E. Stewart has observed both King and Clapper Rails in the
same big cordgrass marsh along Ape Hole Creek in Somerset County.
36
NORTH AMERICAN FAUNA 67
Boith species are common there, probably because of the abundance of
such prime rail foods as blue crabs (Callinectes saj?idus), mud crabs
(Sesarma reticulatum) , red-jointed fiddler crabs, periwinkle snails
(Littorina in'orata), and salt-marsh snails (Malampus lineatus).
In a brackish bay marsh community at Elliott Island, Dorchester
County, on May 28, 1959, I heard King Rails calling between 11 p.m.
and midnight. Most were calling from the narrow band of big cord-
grass that characteristically borders the sides of Pokata Creek. Soras,
Virginias, and Black Rails (Lateralhts jamaicensis) were heard at
the same time and in the same general area, but mostly in a salt-
meadow marsh type community.
In a fresh bay marsh community north of Savannah Lake, Elliott
Island, King Rails occur where the switchgrass marsh extends inland
for a mile or so forming an understory beneath a loblolly pine (Pinus
taeda) forest (fig. 17). It seems rather strange to flush a King Rail
Figuee 17. — Habitat of King Rail in loblolly pine (Pinus taeda) and switchgrass
(Panicum virgatum) association, Elliott Island, Dorchester County, Md.,
August 1967. (Photograph by Luther Goldman.)
from beneath a stand of loblolly pine. The Short-billed Marsh Wren
was found nesting and wintering in this same pine-switchgrass asso-
ciation. Switchgrass, which attains a height of 5 feet, retains its life
form throughout the year, thus affording excellent cover, especially
in winter when several other marsh plants have deteriorated.
NATURAL HISTORY OF THE KING RAIL
37
On June 10, 1965, I made a King Rail survey of a section of the
Nanticoke River marshes, Wicomico County, across the river from the
town of Vienna. This was a typical muskrat marsh in which Olney's
three-square was dominant, with rosemallow scattered throughout.
Four King Rail nests, located at the bases of rosemallow plants, were
found in a 20-acre section of the marsh. The life form of this plant,
with its cradle-like base and broad leaves forming a protective cover
above, makes it well suited for nest cover (fig. 18) .
Figxtre 18. — Nest and eggs of King Rail in Nanticoke River marsh, Vienna,
Wicomico County, Md., June 10, 1965. Nest found in section of marsh domi-
nated by Olney's three-square (Soirpus obieyi) interspersed with rose mallow
(Hibiscus moschcutos). Four nests found in this marsh were all in rose
mallow. Because of the life form of this plant, the rail does not have to build
a canopy over its nest as it does when using other plants.
38
NORTH AMERICAN FAUNA 6TI
In May 1959, while censusing Red-winged Blackbirds on a number
of small islands in Chesapeake Bay, I was surprised to find King Rails
on almost all of them. The islands provide a brackish environment.
On Long Marsh Island, May 26, 1959, I observed a nest constructed
mostly of saltmeadow cordgrass and containing nine eggs (fig. 19) .
Wi
Figure 19. — Nest and nine eggs of King Rail in brackish marsh, Long Marsh
Island, Eastern Bay (of Chesapeake Bay), Queen Annes County, Md., May 26,
1959. Nest constructed of saltmeadow cordgrass (Spartina patens) and Olney's
three-square (Scirpus olneyi).
Small patches of saltmeadow cordgrass were scattered throughout the
dense growth of hightide-bush (Iva frutescens) on the island. The
King Rail nest was only 15 feet from a Black Duck {Anas rubripes)
nest. On another Chesapeake Bay island (Miller's Island) a King
nest was found in a pure stand of saltmarsh cordgrass.
Inner Coastal Plain of Maryland
Four pairs of King Rails nested in 10 acres of shrub swamp-marsh
mixture at the Patuxent Wildlife Research Center, near Laurel, Md.,
NATURAL HISTORY OF THE KING RAIL
39
during May and June 1965. Softrush, tussock sedge (Oarex stricta),
and arrowhead (Sagittaria sp.) were the common emergent herbaceous
plants. Woody marsh plants included swamp viburnum (Viburnum
nudum), arrow-wood (V. dentatum) , buttonbush, alder (Alnus serru-
lata), winterberry (Ilex verticillata) , red maple (Acer rubrum), and
willow (Salix nigra). In late summer Woodcock (Philohela minor)
were common in this same area.
DELAWARE BAY, DEL.
King and Clapper Kails inhabit the extensive brackish bay marshes
known as Broadway Meadows between Fleming's Landing and Wood-
land Beach, Kent County, Del. Two King-Clapper pairs and their
nests were found in June 1960 at Taylor's Gut approximately halfway
between Fleming's Landing and Woodland Beach (fig. 20).
Figure 20.— Taylor's Gut at low tide, Kent County, Del., September 30, 1963.
Vegetation in this breeding habitat of mixed King and Clapper Rail popula-
tions is mainly saltmarsh cordgrass (Spartina alterniflora) , big cordgrass
(Spartina cynosuroides) , saltmarsh bulrush (Scirpus robustus), and
hightide-bush (Iva frutesccns) . (Photograph by Frederick C. Schmid.)
The section of marsh at Taylor's Gut where mixed populations occur
is more typical of Clapper Rail than of King Rail habitat. The domi-
40 NORTH AMERICAN FAUNA 67
nant vegetation types are saltmarsh cordgrass and saltmarsh bulrush
(table 4) . Hightide-bush borders the tidal guts.
Table 4. — Plant composition at three stations in Broadway Meadows, Del., in 1960
[In percent, based on estimates for five 10-foot-square quadrats at each station; tr.=trace]
Fleming's Taylor's Gut Woodland
Landing (Intermediate Beach
(King Rails area; King Causeway
only) and Clapper (Clapper
Rails) Rails only)
Saltmeadow cordgrass - 50 tr.
Saltmarsh cordgrass. 15 50 70
Saltmarsh bulrush 30 20
Bigcordgrass 5 10 10
Olney's three-square 25
Hightide-bush 5 10
Saltgrass - tr. tr.
Groundsel-bush r tr.
Two miles inland at Fleming's Landing only King Kails were ob-
served. The vegetation at Fleming's Landing is composed mostly of
saltmeadow cordgrass and saltgrass (Distichlis spicata). Occasional
patches of Olney's three-square, big cordgrass, and saltmarsh cord-
grass were distributed through the saltmeadow marsh, and as in the
intermediate area at Taylor's Gut, hightide-bush bordered some of
the tidal guts.
Two miles further toward Delaware Bay along the Woodland Beach
Causeway, Clapper Rails were abundant, but King Rails were not
observed. Saltmarsh cordgrass and saltmarsh bulrush were the domi-
nant plants at this station. Hightide-bush was not present.
It is interesting to note that in the Taylor's Gut area salinity read-
ings are intermediate between those at the other stations (table 5).
Table 5. — Salinity determinations at three stations in Broadway Meadows, Del.
in 1960
[In parts per million. Water samples were analyzed in the chemistry laboratory, Patuxent Wildlife Research
Center, Laurel, Md.; sea strength is 32,000 to 35,000 p.p.m.]
Fleming's Taylor's Woodland
Landing Gut Beach
Causeway
Lowtide 4,380 7,190 7,600
High tide 3,700 5,670 7,480
The red-jointed fiddler crab was abundant at Taylor's Gut during
the period 1959 through 1963, and formed the main food of the rails.
There was a marked diminution in the fiddler crab population in 1964,
and a corresponding decrease in the rail population.
Other breeding birds at Taylor's Gut in order of relative abundance
are the Long-billed Marsh Wren, Red- winged Blackbird, Song Spar-
row (Melospiza melodia) , Seaside Sparrow {Ammospiza maritima) ,
NATURAL HISTORY OF THE KING RAIL 41
Swamp Sparrow {Melospiza georgiana) , Black Duck, and Least Bit-
tern. The muskrat, raccoon (Procyon lotor), and rice rat (Oryzomys
palustris) are common mammals in the area.
GREAT LAKES REGION
In Lake County, 111., Beecher (1942, p. 13-14) found the Carex
Jacustrls consocies or lake sedge-marsh wren community to be the op-
timum breeding habitat of the King Rail. Three nests were located in
5.39 acres (Beecher, 1942, p. 29). Beecher characterizes this marsh
type as follows:
Although the Typha, consocies is so distinctive in its characteristics, there is
considerable overflow of the species presumably finding their optimum within its
bounds into the lake sedge which usually adjoins it in shallower water. Carex
lacustris tends to exist as a closed community ; it is more completely dominant
in its own zone and its boundaries more sharply marked out than those of any
other plant in the hydrosere. To state that it has the same lifeform as Typha
means nothing, since, though much coarser than the grass-like sedges which
follow it, the stalk offers little support. Nests of bittern, gallinule and blackbird
are decidedly less frequent than in cattails, those of the redwing being constructed
on a stool, generally. But the King and Sora Rails and the Prairie Marsh Wren
are much more abundant in this sedge than in cattails, suggesting that it has
qualities of its own. Primarily, it offers the tussock or stool type of substrate so
attractive to rails, and anyone viewing this community for the first time would
appreciate its fitness for the wrens. The growth is denser, less erect and,
doubtless, easier to work.
The King Rail also formerly occurred commonly in the extensive
cattail marshes of the southwestern shore of Lake Erie. On May 30,
1931, Milton Trautman (personal communication) recorded 18 King
Rails in 1 hour in these marshes. Trautman further stated that in
the Sandusky Bay region, on many June and July evenings during the
years between 1925 and 1934, he saw from one to eight broods on
roads adjacent to marshes. At Buckeye Lake in east-central Ohio,
Trautman (1940, p. 229-230) reported more than 50 pairs nesting
annually between 1922 and 1930. Trautman told me that by 1959
only two or three pairs nested there. Surveys in 1961 by Trautman
and others to determine the status of the King Rail in Ohio revealed
that it was disappearing at an alarming rate.
In Ontario, Baillie (1940, p. 109) reported five breeding localities
(based on the presence of nests or broods) along the southern edge
of Ontario from Lake St. Clair to Toronto : St. Anne's Island, Lake
St. Clair, Lambton County, May 1882 (nest of 13 eggs) ; eastern end
of the north shore of Lake Erie, at Point Abino, Welland County,
May 30, 1894 (nest of 10 eggs) ; north shore of Lake Erie, at Long
Point, Norfolk County, summer of 1921 and 1926 (young) ; western
end of the north shore of Lake Ontario at Toronto, August 22, 1938
(young) ; and at Hamilton, August 6, 1939 (young) .
348-693 O— 69 4
42 NORTH AMERICAN FAUNA 67
NORTH-CENTRAL PRAIRIE MARSHES
Tanner and Hendrickson (1956, p. 54-56) studied the King Rail
in the marshes of Dewey's Pasture Public Shooting Ground, Clay
County, Iowa, from April 1951 to April 1953. Their description of
the habitat in this area is as follows :
• The 402racre research area included 28 marshes lying in the hollows between
gently sloping prairie knolls. These marshes ranged in depth from several inches
to 4 feet and in area from 0.2 acres to 18.0 acres. Of the total 96.4 acres of
marsh, 81.4 acres supported emergent vegetation habitable by rails. The remain-
ing 15.0 acres consisted of open water. The predominant species of emergent
vegetation in the shallowest water along the shores were blue-joint grass
[Calamagrostis canadensis], prairie cordgrass [Spartina pectinata], tussock
sedge and fox sedge [Carex vulpinoidea]. In waters of intermediate depth the
most abundant species of emergent plants were river grass [Fluminea festuca-
cea], lake sedge, sweet flag [Acorus calamus] and water smartweed [Polygonum
coccineum]. In deeper waters the predominant species were broad-leaved cat-
tail, narrow-leaved cat-tail, river bulrush [Scirpus fluviatilis], hard-stemmed
bulrush [Scirpus acutus], pale great bulrush [Scirpus heterochuetus] and large
bur-reed [Sparganium eurycarpum]. The plant names follow Hayden (1943), . . .
... Four of the six nests were found in areas of marsh in which lake sedge
was the predominant vegetation, while the others were found in pure stands of
river bulrush and were attached to plants of that species. Of the four nests
located in the lake sedge cover-type, only one was actually attached to plants
of that species. The others were supported by tussocks of blue-joint grass or
cordgrass, or clumps of hard-stemmed bulrush, which occurred here and there
among the lake sedge.
The estimated number of breeding adults in the 81.4 acres of marsh
was 12. In the same area there was an estimated adult breeding popula-
tion of 54 Virginia Rails and 52 Soras.
NORTHERN GREAT PLAINS
The King Rail rarely breeds in the northern Great Plains, but
R. E. Stewart (personal communication) located a breeding pair in a
prairie pothole in the Missouri Coteau of western Dickey County,
N. Dak., in June 1961. This pothole was a fresh- water type and was
composed chiefly of whitetop grass (Fluminea festucacea) and slough
sedge (Carex atherodes), with an outer border of river bulrush. Sev-
eral pairs of Virginia Rails and Soras were also observed in this same
pothole. Stewart also recorded single King Rails on June 5 and 24,
1963, about 12 miles west of Buchanan, Stutsman County, N. Dak.,
where common cattail and common spikerush were the dominant
plants.
Description
SIZE
The male King Rail is generally larger and heavier than the female.
Males in my study weighed about 100 grams more than females. Six
of nine adult males weighed over 400 grams each, and the average of
all nine was 415.4 grams, whereas the average of nine females was
306.0 grams. These weights do not differ greatly from those of Clapper
Rails (table 6).
Measurements of body length and wing length also reflect the differ-
ence in size of the sexes. These are compared in table 7 along with
measurements of the Clapper Rail, which is somewhat smaller in
these dimensions.
ADULT PLUMAGE
There are no apparent differences between the plumages of the male
and the female King Rail. Ridgway and Friedmann (1941, p. 83)
described the plumage as follows :
Forehead, crown, occiput, and nape deep, rich mummy brown, the feathers of
the forehead and crown with shiny black shafts ; scapulars, interscapulars,
upper and lower back, rump, upper tail coverts, and rectrices deep fuscous to
fuscous-black, the feathers broadly edged with tawny-olive to buckthorn brown,
the edges becoming broader on the more posterior parts, often occupying (be-
tween the two margins) more than half the width of the feather on the long
scapulars and the feathers of the rump and the upper tail coverts, narrow on the
anterior interscapulars ; upper wing coverts deep hazel to bright russet, some
of the outer median and .greater coverts with narrow whitish tips and a con-
cealed narrow subterminal whitish band ; remiges sepia, the outer web of the
outermost primary often slightly paler — Saccardo's umber ; a light strip from the
base of the maxilla over and behind the eye light pinkish cinnamon ; rest of lores,
circumocular area, cheeks and auriculars grayish mummy brown ; lower cheeks
and sides of throat cinnamon ; chin and middle of upper throat white ; lower
throat, breast, and upper abdomen cinnamon becoming paler in the mid-ventral
part of the upper abdomen, the feathers faintly tipped with white on the upper
abdomen, without pale tips on the breast feathers ; middle of abdomen light buff ;
thighs similiar but transversely barred with deep drab to hair brown ; flanks
sepia barred with white, the feathers tipped with white and crossed by two or
three white bars each ; vent similar to flanks ; under tail coverts white, not
buffy, and with sepia areas reduced making the white bars wider; the outer
webs of the lateral ones wholly white ; axillars and under wing coverts deep
rich sepia tipped and crossed by narrow bars of white ; . . .
Ridgeway and Friedmann described the dark and light phase adult
plumages of the King Rail, and suggested that the light phase rarely
43
44
NORTH AMERICAN FAUNA 67
occurs. My recent investigations indicate that the so-called light phase
plumage is probably the result of hybridization or intergradation in
areas of mixed King and Clapper Rail populations, or it may be due
simply to individual variation.
In my collection I have a series of 16 King and Clapper Rail speci-
mens taken from a 1 -square-mile area of brackish marsh in Delaware
(table 1). In this series there are Kings and Clappers with typical
plumages and also gradations from one type to the other. Some of
the specimens appear to be light phase King Rails. Ridgeway and
Friedmann (1941) made no mention of locality, habitat, or the possi-
bility of mixed populations where so-called light-phased birds were
collected.
Table 6. — Weights of King and Clapper Rails
[In grams. All specimens were adults. Bottom line shows mean weights.]
King Rail
King Rail
Clapper Rail
Clapper Rail
(i?. e.
elegans) >
CR. e.
tenuirostris)2
(R.l. crepitans)3
Males Females
(R. l.)«
Males
Females
Males
Females
Males
Females
339.9
253.0
271
220
332.2
300
275
366.0
272.0
306
255
335.0
325
275
367.9
305.0
317
268
300
275
421.0
313.3
331
325
275
427.0
319.0
300
275
436.0
320.0
350
275
438.3
322.0
300
250
453.0
323.0
300
490.0
325.0
325
350
350
350
325
415.5
305.9
306.3
247.7
333.6
320.8
271.4
1 From Arkansas, Delaware, and Louisiana (author's data).
2 All from Mexico (Warner and Dickerman, 1959, p. 50).
3 Both from Delaware (author's data).
4 All from South Carolina, each to nearest 25 grams; race (R. I. crepitans or R. I. waynei) not specified
(Blandin, 1963, p. 33).
Table 7. — Measurements of King and Clapper Rails
[From Ridgway and Friedmann, 1941. All specimens were adults. All measurements are given in mm.
Wing measurements are for the chord, from bend of wing to tip of longest primary]
King Rail (R. e. elegans)
Clapper Rail {R. 1. ciepitans)
Males '
Females 2
Males 3
Females 4
Range Average
Range Average
Range Average
Range Average
Wing
Tail
Exposed
culmen
Tarsus
Middle toe
without
claw
159. 0-177. 0 163. 4
. . 56. 0- 72. 5 65. 9
147. 0-162. 0 154. 3
60. 0- 70. 0 64. 4
142. 5-159. 5 151. 1
55. 0- 69. 0 64. 6
135. 5-160. 0 146. 8
55. 0- 69. 5 61. 9
58. 0- 65. 5 62. 5
.. 52.0-64.0 58.4
50. 0- 63. 0 61. 9
49. 5- 58. 0 54. 0
55. 0- 69. 5 63. 3
48. 0- 56. 0 51. 7
53. 5- 67. 0 59. 6
41. 0- 56. 0 48. 1
.. 50.5-60.5 55.1
46. 0- 56. 0 50. 8
45. 5- 53. 5 48. 8
40. 0- 52. 0 45. 9
1 18 specimens from Illinois, Missouri, District of Columbia, Virginia, Alabama, Louisiana, South Carolina,
and Florida.
2 14 specimens from Illinois, District of Columbia, Maryland, Virginia, Louisiana, and Florida.
3 21 specimens from Massachusetts, New York, New Jersey, Virginia, and North Carolina.
4 17 specimens from New Jersey, Virginia, and North Carolina.
Note.— An incubating female (adult ?) King Rail collected at Stuttgart, Ark., May 1962, had a left wing
measurement (chord) of only 141.0 mm.; a paired female (adult ?) collected at Taylor's Gut, Kent County,
Del., Apr. 15, 1963, had a left wing measurement (chord) of 145.0 mm.
NATURAL HISTORY OF THE KING RAIL 45
The most unusual plumage that I have seen was that of a very dark
brown, almost blackish bird near Lake Okeechobee, Fla., January 1958.
William B. Robertson (personal communication) told me of seeing
several birds with similar dark plumage in the Everglades, and Luther
C. Goldman collected such a specimen near Cape Sable, Fla. Dr. Harry
C. Oberholser examined the specimen and remarked that it had a most
unusual plumage. Apparently he did not make a critical study of it,
and it has since been lost.
LEGS AND FEET
Legs and feet are pale brownish gray. An adult male collected at
Welch, La., January 12, 1963, and two adult males collected in August
1963, in Delaware, had a pinkish-brown color on the inside and out-
side heel areas and immediately above. This heel color is apparently
typical of birds in their second year or older.
BILL
In most adult birds the bill is orange-yellow from the base to at
least the nares in the upper mandible, and usually slightly beyond in
the lower mandible. The outer part of the bill is brownish. However,
one marked wild bird known to be at least 2 years old had a lightish-
brown bill more typical of immatures. A captive immature did not
attain the color at the base of the bill until it was 10 months old. The
color was then yollowish rather than orange-yellow. Young wild birds
2 to 3 months of age had lightish brown bills. The upper mandibles
of these birds were darker.
TONGUE AND LINING OF MOUTH
Tongues and mouth linings of birds 1 year or older, examined
immediately after collection, were a bright orange-red. Young birds
in juvenal plumage, collected during the summer, had yellow tongues
and mouth linings.
EYE
Irides of adult King Rails are reddish-orange while pupils are gray-
ish-black. Eyes of newly hatched chicks are grayish-brown, and 1- and
2-month-old birds have dull-brown irides.
NOTES ON SEXING AND AGING
In a 50-bird sample from Louisiana, examined 3 months after col-
lection in late fall, I was able to sex 47 of 50 birds by weight, and age
36 of 45 by color of the bill and heel. As an aging criterion, the color
of the bill is used most accurately with live or freshly killed birds,
because with time it fades. Wing measurements can also be used as
an aid in sexing birds, since the average for males is nearly 10 milli-
46 NORTH AMERICAN FAUNA 67
meters greated than that for females. From late summer to at least
early winter, the presence of a bursa in young birds will distinguish
them from adults.
MOLTING
Adult King Rails have a complete molt and are flightless for
nearly a month. Young of the year undergo a partial molt which does
not include the tail and flight feathers.
In the Middle Atlantic States the molting season for King Rails
extends from the beginning of the breeding season in late May until
the beginning of fall migration in early October.
In this study, breeding birds found in molt during May and June
were replacing only body feathers. Molt of the remiges and rectrices
was not observed until the first week in July. I have not been able to
ascertain the relation between the partial molt during the breeding
season and the complete molt in the summer. Birds that are renewing
their body feathers while nesting in May and June may be undergoing
prenuptial molts or early postnuptial molts during which the wing and
tail feathers are not dropped.
Bent (1926, p. 262) stated that adult King Rails undergo a partial
molt of the contour plumage during early spring. Eight specimens
collected in the Middle Atlantic States in March and April showed no
signs of molt. An adult female King Rail in the U.S. National Mu-
seum, taken at Alligator Bluff, Kissimmee River, Fla., April 9, 1901,
was molting body feathers when collected. This could well have been
a breeding bird because in Florida this species begins nesting in late
winter. In my records the earliest recorded dates of molting by King
Rails are May 28, 1960, and May 29, 1964, when an adult or subadult
male and female, respectively, were found in breeding condition at
Woodland Beach, Del. The male had pinfeathers on the underside of
the neck, the sternal region, and the crural tract. Feather renewal on
the female appeared to be about three-fourths complete and was pro-
ceeding simultaneously in most areas of the body. Three King Rails in
breeding condition examined at Laurel, Md., June 12 and 18 and July
3, 1965, also were molting body feathers.
Molting rails in breeding condition have previously been reported.
Watson (1962, p. 350) collected molting Spotted Rails {Pardirallus
maculatus) in breeding condition in Cuba; Warner and Dickerman
(1959, p. 50), working near Mexico City, reported two female King
Rails {Rallus elegcms tenuirostris) molting in May during the nesting
season.
My earliest example of a King Rail molting its wing and tail feathers
was at Laurel, Md., July 7, 1965. This bird was flightless. Two other
birds trapped at Laurel, one on July 14, 1967, the other on July 24,
1967, also were flightless. A female collected near Woodland Beach,
NATURAL HISTORY OF THE KING RAIL 47
Del., July 30, 1964, had nearly completed molting. Its new remiges
and rectrices were about half grown, and there was evidence of a late
stage of feather replacement in all tracts except the head and upper
neck regions.
A captive subadult female began molting wing and tail feathers by
the latter half of July. Two of three adults collected in Delaware on
August 23, 1963, had nearly completed their molts of wing and tail
feathers ; the third had no wing or tail feathers.
On August 3, 1967, 1 took two birds from the Nanticoke River marsh,
Vienna, Md., that had not yet begun to molt.
The molting period for the Clapper Rail in the Middle Atlantic
States is apparently the same as that for the King Rail. A pair of molt-
ing Clapper Rails in breeding condition was collected at Woodland
Beach, June 29, 1964. Only the body feathers were being molted.
Robert E. Stewart (1952, p. 57) trapped and banded many Clapper
Rails at Chincoteague, Va., and made the following notes on their
molt:
During the trapping period [July 16-August 31] most of the adults were under-
going their post-nuptial molt . . . The individual molting period lasts about one
month. The first adult observed in full molt was trapped on July 21. During the
period August 24 to August 31 (period just before hunting season) a total of 11
adults were trapped. Of these only 5 had completed their molt and were capable
of flight, while 4 were in heavy molt, and were completely flightless. Surprisingly
enough the other two adults had not even started to molt and were in very worn
plumage.
In a group of young captive King Rails, the postjuvenal molt was
underway when they were 50 days old. Molting of the body feathers
began before the young could fly, when the flight feathers were about
one-half to three- fourths unsheathed. Another group of young King
Rails, raised in captivity after being hatched on June 7, completed
their postjuvenal molt by the end of the first week of September.
Breeding Biology
Studies of the breeding biology of the King Eail were made mostly
on the Grand Prairie in Arkansas and Prairie Counties, Ark., during
the period 1951-55.
In late winter when rails return to the prairie from more southern
latitudes or simply become conspicuous in areas where they have been
present all winter, the most suitable habitat for the establishment of
nesting territories is the narrow strip of marsh found in roadside
ditches. At this season there is little suitable cover elsewhere. Old rice
stubbles are sometimes used for nesting, but many of these are dried
up or whipped down by winter winds and rains or are plowed under
in the early spring.
HOMING
Some males or females return to the same territory in consecutive
years. An incubating bird of undetermined sex banded on a nest at
Stuttgart, Ark., May 6, 1952, was recaptured the following year on
May 1, on a nest 30 feet from the previous year's nest site. An incubat-
ing bird of imdetermined sex was banded on its nest at the Patuxent
Wildlife Research Center, Laurel, Md., July 3, 1965, and recaptured
in a trap with a mate and brood of eight young on July 8, 1966, 50
feet from the 1965 banding site.
TERRITORIES
Territories occupied by King Eails in roadside ditches consist of
small strips of fresh-water marsh. The dominant plants in most of
these small marsh strips (in order of relative abundance and conse-
quently of relative importance as nesting cover for King Eails in 1952)
were soft- rush, awl-fruited sedge, bottlebrush sedge (Carex comosa),
lake sedge, common spikerush, beakrush (Ehynchospora sp.) , an unde-
termined Graminae, broad-leaved cattail, and smartweed {Polygonwih
sp.).
The schedule of arrival of males in the area and the stage of court-
ship determine size and choice of territory. It is conceivable that the
earlier arrivals manage to claim larger and more suitable territories
than those which arrive later when competition is keener. However,
territorial boundaries are rather fluid during the earlier part of the
courtship period. As additional males move into an area of suitable
48
NATURAL HISTORY OF THE KING RAIL 49
roadside ditch habitat, the large courtship-feeding territories of the
first contingent tend to shrink.
Initial occupation of territories is indicated by the mating call.
During the last week in February and the first week in March 1955, one
male King Kail gave the mating call at various points along 975 linear
feet of roadside ditch. By the second week in March its mating call
was heard from about 500 feet of roadside ditch; its territory then
was about half its original size. The diminishing of the territory was
caused by: (a) Pressure from another courting male, (b) burning of
cover along part of the ditch bank within the original calling territory,
and (e) relatively modest territorial requirements for nesting, particu-
larly if there is plenty of water and ample aquatic animal life for food
in the area about the nest.
Approximate sizes of nesting territories were determined by meas-
uring the distances between three active nests in the same ditch ; from
the center nest it was 298 feet to the nest on one side and 166 feet to
the nest on the other side. The ditch was about 30 feet wide at all three
points.
Defense of territories
King Rails defend their territories both inter- and intra-specifically.
When another King Rail invades a territory, the possessor often pre-
pares to charge by coming to a "freeze," assuming a partial crouch,
drawing in its neck, and slowly ruffling its feathers. It then chases the
intruder on foot and on the wing.
As additional King Rails move into suitable nesting habitat, there
is much fighting, particularly near boundaries of the more desirable
territories. I observed a typical skirmish in a narrow ditch bordering
a secondary road on April 21, 1955. At this unstable territorial bound-
ary, two males attacked eacli other with bill and claws, sparring like
fighting cocks for about 20 seconds. Then the battle suddenly ended,
and the birds moved in opposite directions.
On April 22, 1955, in a rice stubble which appeared to be a common
feeding ground for the occupants of the adjacent section of roadside
ditch, two males (each already paired) "squared off" in a bitter
encounter lasting 3 minutes. There was much chasing both on foot
and on the wing and clashing "fighting cock style." W. E. D. Scott
(Bent, 1926, p. 287-288) reported similar fighting by Clapper Rails
(Rallus longirostris scotti) during the courtship period :
"During the mating season the male birds are very pugnacious and resent
any intrusions from others of the species. At such time I hare seen them have
pitched battles, and finally, one giving in and taking to flight, the victor would
pursue the vanquished on the wing for several hundred feet . . .
Sora Rails, migrating through the Arkansas rice country in spring,
frequent roadside ditches occupied by King Rails on established nest-
50 NORTH AMERICAN, FAUNA 67
ing territories. One King Rail made four passes in running flight at a
Sora in order to evict it from his territory. Virginia Rails received the
same treatment from King Rails.
A call, presumably agonistic, heard when two male King Rails were
in the same territory, could be described as kik-kik-kur-r-r-r-.
COURTSHIP BEHAVIOR
Mating call and pair formation
With the first warm days of late February, the mating calls of King
Rails are heard for the first time in the roadside ditches adjacent to
rice stubbles or other fields. Rails feed in the shallow water of the ditch
and use the broomsedge (Andropogon spp.) on ditchbanks or outside
levees of old ricefields as places of retreat or hiding. Moreover, the
rails use little lanes or pathways, such as those made by cottontails
{Sylvilagm sp.), for traveling in concealment along the ditchbanks.
In late February the only vegetation that offers much concealment to
calling King Rails is the perennial ditchbank sedge which is also the
winter abode of the Short-billed Marsh Wren on the Grand Prairie;
consequently, much of the early season calling emanates from behind
or among clumps of this grass. However, where there happens to be
an old growth of cattails in the ditch, rails may call from this cover.
The male King Rail calls its mate from a concealed, partly con-
cealed, or completely exposed position. The purpose of this call is first
to attract a mate and later, after pair formation, to rally her.
The mating call is one of the least difficult calls to describe. It is
most commonly given as a harsh kik-kik-kik-kik-kik-, but occasionally
varies from a series of kiks to a series of kuks or hups. This variation
may be a matter of interpretation, possibly depending upon the observ-
er's distance from a calling bird. The pitch of the call is steady, but the
tempo increases from time to time. One bird was heard and seen to
give this call continuously for 18 minutes. In the Arkansas ricefields
this call was heard at almost any time during daylight, but less fre-
quently at night. At Elliott Island, Md., in the Chesapeake Bay coun-
try, I often heard the mating call after 10 p.m. D. J. Nicholson (per-
sonal communication) heard dozens of these rails calling all through
the night on the Kissimmee Prairie, Fla., in January and February
1962.
I have never heard a female give the mating call.
Other calls
The most characteristic call of the King Rail, the primary adver-
tising call, is the one that is heard throughout the breeding season. It
may be written as jupe-jupe-jupe-jupe-jupe- or cheup-cheup-cheup-
cheup-cheup- or sometimes as gelp-gelp-gelp-gelp-gelp-. The first
several notes in a series are louder than succeeding ones, and the tempo
NATURAL HISTORY OF THE KING RAIL 51
increases rapidly toward the end of the call when the notes run togeth-
er. One rail gave 25 distinct jupes in a single series, not including those
in the rapid ending which could not be counted. This call carries a
greater distance than the mating call and is somethimes answered by
a number of other King Rails. It is sometimes used when a bird is
startled and occasionally serves as an "all is well" call when a pair of
separated birds are reunited. In addition, I have often observed an
incubating bird using this call when it wishes to be relieved at the nest.
The primary advertising call of the King Rail is slower and more delib-
erate than that of the Clapper, which is usually more of a rapid
chac-chac-chac-chac-chac-.
A call uttered during prenuptial courtship by both the male and the
female, but more frequently a f ter pairing, is a soft and rapid tuk-tuk-
tuk-tuk-tvik-. This sound reminds me somewhat of the clapping to-
gether of the mandibles of the Barred Owl {Strias varia) , and is sel-
dom audible to the human ear beyond 20 or 30 feet. The King Rail
uses the "tuk" call as a rallying call or gives it to indicate its position
to its mate.
Display
The display of the male during prenuptial courtship is relatively
simple and consists mostly in walking about with tail uplifted and
white undertail coverts extended (fig. 21-1) . In this position the white
undertail coverts can be seen from a considerable distance. While
flashing its white undertail coverts, the rail usually flicks its tail up
and down slightly. Females that I observed during the period of pre-
nuptial courtship made no attempt to display.
There were other forms of posturing during the period of courtship
and mating, but apparently the cocked tail and well-exposed white
undertail coverts, accompanied by the mating call, are the principal
means of attracting a mate.
On two occasions I observed what appeared to be another form of
display, the "pursuit display." The circumstances and the behavior
of the male were essentially the same both times. In each case the male
apparently had not succeeded in attracting a mate to his territory. On
March 1, 1955, at 8 :30 a.m., I saw a small and very rufescent King Rail,
later established to be a female, moving along a rice levee bordering a
roadside ditch and approaching a calling male. The female continued
along the water's edge at a slow but steady gait and passed beyond
the male that was standing in the ditch. As soon as she was ahead of
him, the male followed her at a fast walk with head and neck out-
stretched, bill open (but emitting no sound audible at 40 feet), tail
cocked, and white undertail coverts extended (fig. 21-2) .
Following pair formation much of the posturing and calling that
characterized the period of prenuptial courtship continues, at least
52
NORTH AMERICAN FAUNA 67
^.-f>f.
Figure 21. — Displays of the King Rail: (1) In the Advertising Display the tail
is cocked and the white undertail coverts extended. (2) The Pursuit Display
is given when the male pursues the female during prenuptial courtship; the
male walks fast or runs with tail slightly cocked, white undertail coverts
extended, and bill wide open. (3) The Invitational Display is assumed by
the mated male when the female approaches ; the bill points downward and
slowly swings from side to side, and the tail is displayed. (4) A variation of
the Invitational Display consists in wings arched, head turned to one side,
bill open, and tail displayed.
in the earlier phases of nuptial courtship. While pair formation is in
progress, but infrequently during the nuptial courtship period, the
female utters a purr or churr sound, like the purr of a cat, especially
after the male has given the mating call.
The male uses the mating call (kik-kik-) infrequently and with less
vigor when rallying a newly won mate which often strays when
foraging. I observed a good example of the use of this call shortly
after pairing on the evening of March 2, 1955. During an 18-minute
period beginning at 5 :30 p.m., a paired male, while standing partially
concealed on a ricefield border levee, uttered the complete mating
call seven times. Six of the seven times his mate ran to him from a
distance of 100 feet or less where she had been feeding. When the
female came up beside him, the male spread his white undertail coverts
and bent his head and neck so that his bill was perpendicular to and
NATURAL HISTORY OF THE KING RAIL 53
nearly touching the ground (fig. 21-3). From this position, he often
turned his bent head with bill open toward the female. At one of these
meetings the male appeared to be about to mount the female and
begin to rise up with his bill still wide open, but the female evidently
was not ready for copulation and walked away.
I observed the same posture many other times, but the birds were
usually standing in water. On these occasions the bill usually touched
or slightly dipped into the water. Males assumed the pose after pair-
ing, when the feeding female that had been at some distance away
came within 3 or 4 feet of her mate. On one occasion a male under
such circumstances arched his partly opened wings (fig. 21-A) .
Courtship feeding
Courtship feeding, a type of symbolic display that aids in main-
taining the pair bond, was observed during the courtship, egg laying,
and incubation periods of the King Rail. In the Arkansas ricefields,
the crayfish was the only food that I ever saw presented to a female.
In Delaware Bay marshes, the fiddler crab was used for this purpose.
The male usually brings the food item to the female, but sometimes
he may stand where he catches the crustacean, holding it in his bill,
until the female approaches and takes possession.
A mated pair of rails that I observed for a number of days on their
Delaware breeding territory would descend at low tide from the marsh
to a pool in the bed of a tidal creek. The female would usually stand
in the pool while the male hunted food for her. He would frequently
run up the winding creek bed for 25 yards or so, catch a fiddler crab,
and run back to present it to the female. Why he often traveled such
distances when there were plenty of fiddlers nearby is not known.
During a 2-hour period of observation in an Arkansas ricefield, I
saw the male of a pair catch seven crayfish, five of which he presented
to his mate.
PRENESTING ACTIVITY
Calling
As the nesting season approached, the mating call and undertail
covert flashing by the male all but ceased, and the addition of a num-
ber of calls, mostly soft or subdued, increased the repertoire of the
mated pair. Paired rails used such calls as rallying devices when sep-
arated or as reassuring answers to one another's calls when together.
A call frequently given by both birds, particularly as nesting
approached, was a very soft poyeek-poyeek-poyeek-poyeek-poyeek-,
or wyeek-wyeek-wyeek-ivyeek-ivyeek-, which seemed to act as an
inquiry of the whereabouts of the mate.
Several males gave one of the more unusual calls, a deep booming
sound requiring an effort which caused the body to appear to expand
54 NORTH AMERICAN FAUNA 67
slightly and sounding something like 6om-6om-6om-6om-6om-. The
purpose of his "booming" call is not known. It is not very loud, and
the females were not nearby when it was uttered, unless they were
well concealed.
Symbolic nest building
Symbolic nest building was observed at Stuttgart, Ark. In this
case a male King Eail was observed carrying nesting material into
a hole in a ricefield dike through which water was draining into a
roadside ditch. The dike was about 2i/2 feet in height, and the hole
was large enough for the bird to pass easily from one side to the other.
The light stream of water did not prevent an accumulation of nest-
ing material. However, the nest was not completed. Two days later
(April 2) the true nest was started about 10 yards from the hole in
the dike.
Copulation
Copulation usually takes place near the nest site, before and during
egg laying. Although no nests with eggs were found on the Arkansas
Grand Prairie before March 25, rails were observed copulating as
early as March 3, in 2 different years. Perhaps these birds nested
earlier than March 25.
On one nesting territory, the male came within 20 feet of the nest
(containing one egg), called, and was answered by the female who
left the nest and came to the male for copulation. The jupe-jupe-jupe-
jupe-jupe- call often precedes copulation during this period. Copula-
tion is performed with the female assuming a crouch and the male
mounting with legs and feet placed on the female's back.
NESTING PERIOD
The nesting period varies with latitude, being longer and starting
earlier in the southern part of the range. The nesting period in Florida
extends from late January, at least, until the middle of July, and in
Louisiana from early March to September. It is quite conceivable,
therefore, that in such States as Florida and Louisiana the breeding
season covers 7 to 8 months. Unlike the Bobwhite (Cottnus mrgin-
ianus), Redwinged Blackbird, and several other species which do not
nest much earlier in the gulf coast region than in the northern States,
the King Rail takes advantage of the long warm period, and nests
over a longer period of time. The long period of nesting in the South
should result in a greater total production of young, because of the
much greater opportunity for renesting and second broods. The nesting
season in the Middle Atlantic States is about 4 months; adults with
downy young have 'been observed in early August in Delaware.
NATURAL HISTORY OF THE KING RAIL 55
With the long warm period prevailing in the Deep South, time for
raising more than one successful brood would seem to be ample. A
breeding pair is busy with nesting activity for about 2 months (ap-
proximately 10 days for laying, 21-22 days for incubation, and 24—30
days with brood). As yet there is no evidence that the King Rail is
double-brooded ; however, no attempt has been made to determine this.
The closely related Clapper Rail in South Carolina is double-brooded
(Blandin, 1963, p. 66-67), and it is probable that some King Rails in
the Deep South also have more than one successful brood during a
season.
A nest found in a cypress pond in southern St. Johns County, Fla.,
in February 1933 contained 11 eggs in an advanced stage of incuba-
tion (Hallman, 1934, p. 18). Allowing a 21- or 22-day incubation
period plus 11 days for laying, it is conceivable that this nest was
started in January. D. J. Nicholson found a dead King Rail at Or-
lando, Fla., on February 16, 1925, with a hard-shelled egg ready for
deposit (Howell, 1932, p. 203). A single downy young King Rail was
seen on March 10, 1950, 2 miles west of 40-mile Bend, Dade County,
Fla., by J. C. Moore and D. B. Beard (U.S. National Park Service
files). W. B. Robertson, Jr. (personal communication), found several
young King Rails at Royal Palm Hammock, in the Florida Ever-
glades, on March 5, 1952. These last two examples indicate February
nesting.
Adults with six young approximately 2 weeks old were recorded
in Lee County, Fla., July 30, 1966, indicating that the nesting season
in Florida extends into July (Frederick H. Lesser, personal
communication) .
At Oakland Plantation, a few miles north of Charleston, S.C., a
brood of 10 young were seen by Francis Porcher on March 22, 1913
(Sprunt and Chamberlain 1949, p. 193). In this case nesting started
in early or mid-February. I found young 1 to 4 days old (egg tooth
present) and a nest of nine eggs in Jasper County, S.C., near Savan-
nah, Ga., on April 16, 1961, and young at Georgetown, S.C., on April
25, 1961.
At Grand Chenier, Cameron Parish, on the southwestern coast of
Louisiana on April 8, 1956, I observed a pair of King Rails feeding
2-week-old young. Back-dating about 38 days to cover the age of the
young and the incubation period, laying began about March 3.
Fifty miles north in the Louisiana rice country, nesting probably
gets underway a little later than on the coast. A nest of seven eggs was
found at Mamou, March 30, 1957. A late nest containing eight eggs
was found at Mamou, August 6, 1955.
On the Arkansas Grand Prairie, the important nesting months are
April, May, and June (table 8). The earliest indication of nest build-
56
NORTH AMERICAN FAUNA 67
ing. is in late March (March 25). During March 1952, nearly all King
Rails seen were in pairs.
In Maryland, most clutches are laid between May 15 and June 30.
A brood of chicks and their parents were seen by P. J: Van Huizen
near the Blackwater Kiver, Dorchester County, on May 21, 1965, giv-
ing evidence that some King Rails begin nesting in April.
In Central Ohio, Milton B. Trautman and R. Bales located 42 nests,
34 of which were found in May and 8 in June (M. B. Trautman, per-
sonal communication). Seven of the June nests were found during the
first half of the month, and 26 of the May nests were found during the
last half of that month. From this, the main nesting period seems to
extend from May 15 to May 31.
Table 8. — Nesting dates, clutch size, and habitat of King Rail nests at Stuttgart, Ark.
Nest
num-
ber
Date found
Number
of eggs i
Com-
plete Number
clutch hatched
size 2
Location
Dominant vegetation
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
40
41
42
43
44
45
46
47
48
49
Apr. 1, 1952
Apr. 10,1952
Apr. 15,1952
Apr. 16,1952
Apr. 21,1952
do
Apr. 24,1952
Apr. 25,1955
Apr. 26,1955
do
do
May 1, 1952
do.
May 2, 1952
do
May 5, 1962
May 6, 1952
May 9, 1950
do
May 10,1948
May 10,1952
do
May 11,1952
May 12,1962
May 13,1952
May 13,1962
May 15,1962
May 26,1952
May 26,1954
May 28,1952
June 3, 1954
June 4, 1952
June 9, 1952
June 13,1952
do
do
June 19,1952
June 25,1952
July 15,1950
do...
do
do
do
July 18,1951
Aug. 1, 1950
Aug. 9, 1951
Aug. 9, 1963
Aug. 10,1951
Aug. 29,1951
11 8 Ditch..
2d CanaL.
12 12 Ditch
11 9 do.
10 9 do.
14 14 do.
id. do.
do.
do.
do.
new do.
1
11
new
11
10
5
4
6
12
9
11
12
12
10
11
10
2
10
4
3
Id.
11
d.
11
do.
do....
do
do
do
11 9 do
5d Ricefleld
11 Ditch
12 .-.. Oat field ..
9 9 Ditch
11 10 Canal bank
12 10 Ditch
(?)_... .do
10 0 do
(?) do
(?)... ..do
2d Canal bank
10 Ditch
8 8 do
(?) do
eb Weedyfleld
10 10 10 Canal bank
7 Ricefleld
eb Pond edge
eb do
9 0 Ricefleld....
10 10 ...do
do
'. _...do
do..
do
.- do
Typha lati folia.
Juncus sp.
Typha latifolia.
Grass?
Juncus effusus.
Carex hyalinolepis.
Carex hyalinolepis.
Eleocharis sp.
Sedge.
Juncus effusus.
Juncus effusus.
.do.
.do.
_do.
.do.
.do.
9 Ditch
Typha latifolia.
Juncus effusus.
Sedge?
Carex stipata.
Juncus effusus.
Oryza saliva.
Juncus effusus.
Avena (van).
Juncus effusus.
Bromus secalinus.
Juncus effusus.
Carex stipata.
Juncus effusus.
Eleocharis palustris.
Carex stipata.
Grass?
Eleocharis sp.
Juncus effusus.
Grass? and sedge?
Avena (var.).
Oryza sativa.
Paspalum distichum.
Paspalum distichum.
Oryza sativa.
Oryza sativa.
Oryza sativa.
Oryza sativa.
Oryza sativa.
Oryza sativa.
Oryza sativa.
Oryza sativa.
Oryza sativa.
Oryza sativa.
Oryza sativa.
Oryza sativa.
Oryza sativa.
Echinochloa sp.
1 6=building; e6=eggs broken.
2 d= deserted.
3 Nest and eggs found; clutch size not recorded.
NATURAL HISTORY OF THE KING RAIL 57
Tanner and Hendrickson (1956, p. 54) reported that in Clay County,
Iowa, nesting begins soon after the arrival of the birds during the first
week of May.
In 1951 the nesting season extended for a period of 42 days from May 13, the
date that the first egg was laid, until June 28, the date that the last egg hatched.
NEST SITE AND MATERIALS
The usuial nest site is in the shallow- water part of a marsh. The
water depth was 4 to 18 inches at Clay County, Iowa (Tanner and Hen-
drickson 1956, p. 55), 2 feet at Buckeye Lake, Ohio (Trautman 1940,
p. 229), and 6 to 8 inches in Arkansas ricefields. In South Carolina,
Wayne (1910, p. 35) found nests in buttonbushes 8 to 18 inches over
water.
In a giant cutgrass marsh near Savannah, Ga., each of five nests
located was within 20 feet of the edge of the marsh, although the
vegetation density and other characteristics appeared uniform over
.extensive areas.
Occasionally a nest is placed on a dry-land site such as an oat or
wheatfield, or on a grassy embankment. In 1952, on Long Island, N.Y.,
Roy Latham (1954, p. 3-9) found a nest on the ground in a potato
field, 150 yards from the edge of a salt marsh where Clapper Rails
were nesting.
The nest site appears to be chosen by the male. On two occasions,
I have seen a male initiate nest-building.
Most King Rail nests are placed in fairly uniform stands of vege-
tation and are well concealed, but the shape of the nest canopy
(whether cone-shaped or round) sometimes disrupts the uniform pat-
tern of the vegetation and reveals the location of the nest to the hu-
man eye (figs. 22 and 23) .
The life form of some plant in the territory, such as a tussock of
grass or the stool of a rice plant, often determines the exact nest site.
A nest may be placed in a clump of grass or a sedge tussock, or be-
tween several clumps, parts of which are used in fashioning the canopy
and sides of the nest. The bases of most Arkansas nests were made of
wet decaying plants, and the platforms or cups were of dead dry
grasses, sedges, or rushes. These materials are obtained near the nest
site. The base of one nest found in Arkansas was made entirely of
mud and was 2% inches in depth.
Nest materials used in some Iowa nests consisted of one or two
species of plants (Tanner and Hendrickson 1956, p. 55). Most nests
in Arkansas ricefields were made of rice plants ; a few were made from
"weed" plants in the fields, such as wild millet. The completed nest
is a round, elevated platform with a saucer-shaped depression (figs.
23, 24, and 25). It usually has a round or cone-shaped canopy and a
ramp, and is nearly twice as large as that of the Virginia Rail or Sora.
348-693 O— 69 5
58
NORTH AMERICAN FAUNA 67
y;.' :'
MWm
-■ :;•;
Figxjre 22. — Canopy of King Rail nest in roadside ditch, Arkansas Grand Prairie.
In a uniform stand of vegetation the canopy is often quite conspicuous. Canopy
composed of spikerush {Eleocharis palustris) and smartweed (Polygonum
sp.).
Figure 23. — King Rail nest in roadside ditch near Stuttgart, Ark., May 30, 1952.
Nest constructed of softrush (J uncus effusus).
NATURAL HISTORY OF THE KING RAIL
Figure 24. — King Rail incubating in nest constructed of cattails (Typha lati-
folia) in roadside ditch, Arkansas Grand Prairie, April 1952.
•
-
#>
sd».
Figure 25. — King Rail incubating in open nest along roadside ditch, Mamou,
La., April 1957.
60 NORTH AMERICAN FAUNA 67
Dimensions of 11 eastern Arkansas nests were: average height from
ground to canopy, 43.0 cm. ; average height from ground to rim, 16.5
cm.; average exterior diameter, 28.0 cm.; and inside depth, 1.5 cm.
(Meanley,1953,p.265).
The height of the nest above water usually depends upon the depth
of the water. Eggs in most nests in Arkansas Grand Prairie ricefields
were less than a foot above the water level. In tidal marshes along the
lower Savannah Eiver, S.C., the eggs in two nests were about 2 feet
above the low tide mark and about 1 foot above the high water mark.
Nests in dry locations, such as oatfields, canal banks, or dry ditches,
are usually elevated very little, and the eggs may rest within an inch
or two of, or actually on, the ground. Nests placed above 2 or 3 inches
of water may be elevated as much as a foot during a heavy rain or
when a dry ricefield is being flooded.
After a heavy rain on the Arkansas Grand Prairie, an incubating
rail was observed working rapidly to build up its cattail nest above
the rising water in a roadside ditch. By reaching out with its bill all
around the nest and picking up materials (mostly cattail leaf frag-
ments), which it tucked beneath the eggs, and by using most of the
canopy for the same purpose, the bird managed to keep the eggs about
2 inches above the rising water. The ditch was nearly dry before the
rain, and the eggs were then 5 inches from the ground. At peak depth,
the water was 21 inches deep. On another occasion, a nest with eggs
2/2 inches from the ground was located in a ricefield that had been
temporarily drained. The next day, the field was flooded to a depth of
5 inches, and the eggs were raised to 7 inches from the ground. As the
water continued to rise, the incubating bird persisted in elevating the
eggs by tucking rice leaves from the canopy under them.
On one occasion I came upon a bird constructing a nest on a canal
bank, and watched the process only a minute or so before its mate
came to continue the work. The bird that was relieved left to feed in a
nearby ditch. I watched the newcomer for about 3 minutes, and then
collected it. Upon dissection it proved to be the male.
Apparently the male takes the more active part in nest building.
Males on three occasions were observed gathering nest material within
20 feet of the nest site. A captive male purred like a domestic cat con-
stantly as it carried nesting material to the nest site. The nest is shaped
as the bird (all observations were of males) sits in a clump of grass
or between clumps and semirotates its body. It later piles up dead vege-
tation, and shapes the cup. The canopy is formed by bending over
the tops of stalks of adjacent plants. One Arkansas nest was ob-
served under construction at 8 a.m. and 6 :30 p.m. of the same day,
another one at 12 :15 p.m. and 5 :15 p.m.
The nest is not always completed before the first egg is laid. While
driving along a paved road 5 miles south of Stuttgart, Ark., at 5 :30
NATURAL HISTORY OF THE KING RAIL 61
p.m. one day in May, I heard two rails uttering their characteristic
jupe-jupe-jiope-jupe-jupe call. One of the birds was standing in a
nearby ditch, and after about 3 minutes of watching, I saw the grass
move on the bank near the rail in view. As the same grass continued
to move, it was evident to me that the mate was building a nest.
Actually the bird was pulling in grass to form the sides and canopy
for a nest. After watching it for a few minutes I departed. The next
morning I found that construction was in the initial stage, but two
eggs had been laid on the bare ground and were surrounded by just
a few dead plant fragments.
A Purple Gallinule, another species of Rallidae, was reported by
Grimes (1944, p. 63) to have a nest platform 6 inches thick when the
first egg was laid. As the eggs began to hatch, the nest was built up
until it was 13 inches thick.
Similar nest building activity by Clapper Rails at Frogmore, S.C.,
was reported by Hoxie (1887, p. 181) :
The first time I found the nest it contained only one egg, and did not seem
wide enough to hold more than one more. ... As each new egg was laid they
added fresh material to the outside, until the nest was at least amply sufficient
to contain the full set of eight.
Several brood nests, usually without canopies, are constructed near
the egg nest.
EGG LAYING AND CLUTCH SIZE
Eggs were deposited daily at five Arkansas nests. In one Arkansas
nest, the eggs were laid between 7 p.m. and 7 a.m. In one South Caro-
lina nest, Wayne (1910, p. 36) noted that each egg was deposited after
11 a.m.
Parasitism or "dumping'' was recorded by B. H. Swales (1896, p.
142) in St. Clair County, Mich. On June 9, a King Rail was flushed
from a nest containing 17 eggs; nine were apparently laid by the King
Rail, seven by a Virginia Rail, and one by a Sora.
Clutch size
Clutches of 10, 11, or 12 King Rail eggs are most frequently found
(table 9). A smaller clutch may represent a replacement clutch, de-
pending upon when it occurs. On the Arkansas Grand Prairie, the
earliest clutch of eight eggs was found on May 28, approximately 2
months after the beginning of the laying season. In Maryland, in July,
I observed three complete clutches of six eggs each.
Description of eggs
Bent (1926, p. 261) gives the following description of the eggs:
They are ovate in shape and the shell is smooth and slightly glossy. The ground
color averages lighter than in eggs of the clapper rails, but not so light as in
62 NORTH AMERICAN FAUNA 67
those of the California species ; it is pale buff, varying from "cream buff" to "pale
olive buff." They are sparingly and irregularly spotted, mostly in small spots,
with various shades of "vinaceous drab," "army brown" and "vinaceous brown"
and sometimes with a few spots of brighter browns. The measurements of 56 eggs
averaged 41 by 30 millimeters, the eggs showing the four extremes measure
44 by 32, 38.5 by 28 millimeters.
I measured 20 eggs at Stuttgart, Ark. two from each of 10 nests.
The average measurement was 40.8 by 30.4 millimeters, with extremes
of 42.0 by 32.0 and 39.5 by 29.5.
Table 9. — Clutch sizes in King Rail nests at three locations
Number of clutches found
Clutches with— Stuttgart, Northern Delaware
Ark.1 and central Valley3
Ohio 2
8 eggs 3 1
9 eggs 113
lOeggs 3 9 3
11 eggs 7 11 2
12eggs__ 3 9 4
13eggs.._. .. 1 3
14 eggs - 111
Total... 16 37 14
Mean number of eggs 11.2 10.9 10.6
1 Meanley, unpublished.
2 Trautman, 1940, p. 229; R. Bales, ms.
3 Stone, 1937, p. 332; R. F. Miller, correspondence.
Weight of eggs
At Stuttgart, Ark., three eggs weighing 18.9, 20.3, and 18.8 grams
were marked on the day they were laid and were weighed on every
seventh day until hatching (table 10). The average weight loss was
0.47 gram during the first week, 0.83 gram during the second week, and
1.0 gram during the third week. The average total loss from laying
to hatching was 2.30 grams.
INCUBATION
The incubation period is about 21 or 22 days. Roberts (1936, p. 440)
stated that A. M. Bailey found the incubation period to be about 21
days. In Clay County, Iowa, Tanner and Hendrickson (1956, p. 55)
found it to be approximately 21 days. Incubation periods of four
Arkansas clutches were 21 days, 22 days, 22 or 23 days, and approxi-
mately 23 days.
One Arkansas nest was under daily observation from the time the
first egg was laid on April 1 until the last egg hatched on May 4.
Eleven eggs were in the completed clutch, and incubation started
with the laying of the 10th egg on April 10. At another nest in
Arkansas incubation began on April 22 or 23, and the eggs hatched on
May 13 and 14. A nest at Mamou, La., contained 9 eggs on June 9 and 10
hatching eggs on June 30.
NATURAL HISTORY OF THE KING RAIL 63
Both sexes incubate. To prove this, one night between 9 and 10 p.m.,
I placed white paint in a small can at the end of a long stick and
poured it on the backs of incubating birds at two nests. On subse-
quent visits to the nests, unmarked birds were often seen incubating.
Table 10. — Weight loss in three King Rail eggs during incubation
[In grams]
Weight on —
Day 1 Day 7 Day 14 Day 21
Egg 1 18.9 18.7 17.8 16.7
Egg 2 .. 20.3 19.5 18.8 17.9
Egg3._- 18.8 18.4 17.5 (>)
Mean.. 19.33 18.86 18.03 17.3
1 Clutch destroyed.
Later that season an exchange of sexes was observed at a nest
during the incubation period. At 5 :18 p.m. an incubating bird called
from the nest, whereupon its mate immediately came from the cattails
across the road to a point about 20 yards from the nest, and began
walking toward the nest until it was within 5 feet. The incubating
bird then left the nest and was replaced by its mate, which remained
on the nest for 17 minutes, when an exchange again took place.
In another instance, when one member of an Arkansas pair nesting
near a road was killed by an automobile, its mate continued to incubate
the eggs. An incubating bird caught on a nest at 5 :45 p.m. May 16,
was a male.
Incubating birds seldom flush until an intruder is within 10 feet or
less of the nest. As the hatching date approaches, they become more
tenacious. On several occasions I was able to band incubating birds,
but not without considerable resistance from them. On one occasion
when I approached a nest at hatching time, the bird flew from the nest
and struck me in the chest. On other occasions birds have struck at my
legs or have run to my feet where they remained with wings out-
stretched. Frequently they feigned injury by spreading the wings,
fluttering through the vegetation (fig. 26), and uttering a distress call
which might be written as a gutteral rack-k-k-, rack-k-k-, rack-k-k-,
sometimes varying to sound like chur-ur-ur-ur (the roll on the ur is
like the German "R"). Other scolding notes given by a rail flushed
from its nest are a resonant gip-gip-gip- and kik-kik-kik-.
In contrast to this type of behavior, the Clapper Rail is usually gone
before the intruder gets near the nest. At Chincoteague, Va., I have
examined some 200 Clapper Rail nests, and only on some half dozen
occasions has an incubating bird remained while approached to within
10 feet. This appears to be a striking behavioral difference between
these two closely related species.
64 NORTH AMERICAN FAUNA 67
I ■ fi \
gHHj
<>'{ k * Ia; • "*t
w * ' * > - i
V £■ *■**■- ,,<% Is^ A*i, _ *
Figure 26. — 'Distraction display of King Rail near neslt. This display is char-
acterized by feigning injury and emitting distress call.
HATCHING
Eggs in four Iowa nests hatched within a 24- to 48-hour period,
and were pipped from 24 to 48 hours before hatching (Tanner and
Hendrickson, 1956, p. 55).
Hatching was observed at Stuttgart, Ark., May 26-29, 1954. When
located at 12 noon on the 26th, the nest contained 10 eggs, only one
of which showed signs of hatching and had two small pip holes. By
4:30 p.m. the following day (May 27), 9 of the 10 eggs were pipped.
At 1 p.m. May 28, 3 eggs had hatched ; by 5 :30 p.m. that evening, 5
eggs had hatched; and by 10 a.m. May 29, all eggs had hatched. At
4 p.m. May 29, the entire brood and both parents were at the nest;
but at 9 :30 p.m. the entire family had deserted the nest and was prob-
ably spending the night in a nearby brood nest.
At one Arkansas nest the parent birds alternately participated in
brooding newly hatched young and hatching eggs. Toward the end
of the hatching period the nonbrooding parent was usually observed
within 25 feet of the nest, accompanied by several of the chicks.
As eggs hatch the shells are disposed of in several ways. One brood-
ing bird at an Arkansas nest ate most of an egg shell about 5 minutes
after the egg had hatched. Shell fragments were found in the stomachs
of several adult birds collected during the breeding season. Some shells
NATURAL HISTORY OF THE KING RAIL 65
remain in the nest, disintegrate, and eventually filter down into the
base of the nest. Shell fragments are found in virtually all nests that
have hatched young.
A pair may remain with their brood for more than a month after
hatching. I have collected three-fourths grown young rails that were
still traveling with an adult pair in August. In one instance a pair,
one of which was marked, and their 3-day-old young still spent most
of the day within 20 yards of their nest, and 19 days later were seen
only 10 yards from the nest ! Once I came upon a brood of young King
Rails approximately 3 weeks old traveling with three adult birds.
The call given by an adult with young chicks when all is well is a
soft continuous woof-ivoof-ivoof- (corresponding to the cluck-cluck-
cluck- of a barnyard hen). An alarmed parent with brood emits a
sharp gip-gip-gip-, which causes the young to scatter to a hiding
place.
NESTING SUCCESS AND SURVIVAL
In Clay County, Iowa, Tanner and Hendrickson (1956, p. 55) found
that four of six observed nests hatched one or more eggs each. Of 60
eggs in the six nests, 39 hatched.
Of 16 Arkansas nests I observed, 12 hatched one or more eggs each.
The average number of eggs hatched in each of these 12 nests was 9.9.
Of a total of 147 eggs in all 16 nests, 119 hatched.
An index of survival based on the number of young over 2 weeks
of age is difficult to obtain because complete broods are not always
seen. In Arkansas, I observed 10 broods with what I believe were full
complements. In each observation, the parent birds were unaware of
my presence as the family was crossing a road, feeding in a newly
sown ricefield, or moving about in some other comparatively open
spot. The number of young per brood ranged from two to nine and
averaged five. If my estimate of an average hatching success of 9.9 is
correct, then survival rate until 2 weeks of age was about 50 percent.
BREEDING STATUS OF FIRST-YEAR BIRDS
Although I know of no example of juvenile or immature birds being
marked and recaptured in breeding condition or in the act of nesting,
I collected a nesting bird in what appeared to be first-year plumage in
the Delaware Bay marshes. Only the lower throat and upper breast
regions of this bird were cinnamon, a whitish area covered most of the
lower breast and axillary regions, and the side of its head was con-
siderably paler than average for mature birds. The greater coverts
were heavily barred with whitish subterminal bars. The specimen, a
female, was extremely small. Measurements were as follows (with
adult female average in parentheses) : Wing 147.0 mm. (154.3) ; ex-
posed culmen 54.0 mm. (61.9); tarsus 50.0 mm. (54.0).
Development and Behavior of Captive Rails
DEVELOPMENT OF YOUNG
First-day chick
The newly hatched King Kail is very weak and wet. Contrary to
the statements of Audubon (1835, p. 28) and Howell (1932, p. 203),
it is unable to run about and follow its parents as soon as it is hatched.
Sometimes it emerges from the shell on its back and lies kicking and
struggling for some minutes before righting itself. A nest mate may
grab its toes or beak and so stimulate further activity. Most of the
chicks I have observed were more than an hour old before they were
able to go over the side of the nest and return. Chicks 15-20 minutes
old had considerable difficulty when we placed them in weeds and
water outside their nest, and they could not get back into the nest
under their own power.
As the down dries out, the young bird moves more actively about
the nest, the undeveloped wings assisting in this effort. As the rail
chick begins to gain strength, it sits on its tarsi and assumes a begging
display, with wings extended for balance.
The period of fluffing-out often takes half an hour or longer. It
took 41/2 hours for one of the chicks I observed. The fluffing-out proc-
ess may be necessary to produce buoyancy needed to enter the water
safely, as Gullion (1954, p. 389) suggests for the Coot (Fulica
americana) .
Chicks took food from their parents' beaks the first day, but I did
not see them picking up food from the ground until the second day.
The day -old chick has at least two calls : a loud begging call, chee-
wp; and a soft lower-pitched call of contentment, wee and wee-up.
The day-old chick is covered with black down that has a faint
greenish sheen or cast except in areas where it is most dense. The down
is very dense on the abdomen and sparse on the crown. The bill has
a pied pattern ; the basal half of the bill is grayish black, the narial
region is white, the distal portion is flesh-colored, and the egg tooth,
retained at the tip for 4-6 days after hatching is white. The legs and
feet are brownish gray although, at a quick glance, they appear to
be black. Eyes are grayish brown. A vestigial claw is present on each
wing.
66
NATURAL HISTORY OF THE KING RAIL
67
The young bird loses weight throughout the first day of life. One
female chick weighed 16.3 grams at hatching, 16.0 grams at 1 hour,
15.7 grams at 2 hours, and 13.2 grams at 24 hours (table 11).
One to thirty days
During the first month of life the major change in the appearance
of the King Rail chick is one of size and conformation (fig. 27). A
young captive male weighed 16.7 grams when he was 1/2 days old,
and 96.3 grams when he was a month old. His measurements at iy2
and 30 days, respectively, were: exposed culmen, 11.0 and 28.0 mm.;
tarsus, 20.0 and 42.0 mm. ; middle toe with claw, 22.0 and 50.0 mm.
(table 11.).
The thick natal down is present during most of the first month,
but during the fourth week there is evidence of development of the
juvenal plumage.
Table 11.
— Growth of four King Rails
Length of—
Age
(grams) Exposed
culmen
(mm.)
Middle toe
with claw
(mm.)
Tarsus
(mm.)
Bird A:
Hatched .
1 hour _ . .
2 hours _ .
1 day
8 days . . .
21 days..
60 days..
90 days . .
Bird B:
1 day
7 days..-
17 days..
21 days..
30 days..
45 days . .
Bird C:
1^ days . .
7-8 days . .
17-18 days.
21 days
30 days
45 days
60 days
BirdD:
1 J3 days . . .
7-8 days . . .
17-18 days .
21 days
30 days
45 days
60 days
U6. 3
16.0
15.7
13.2
14.0
40.0
202.0
265.0
15.6
18.9
37.0
50.7
75.4
177.9
16.7
25.7
46.5
63.8
'96.3
219.8
327.0
15.6
19.7
32.0
47.7
70.8
176.0
258.6
11.0
12.5
19.5
40.0
49.0
20.0
25.0
40.0
11.0
~2L0~
28.0
40.0
48.0
25.0
20.0
26.0
36.0
57.0
57.0
42.5
49.0
22.0
36.6"
50.0
61.0
61.0
43.0
20.0
21.5
29.0
53.0
53.0
38.5
52.0
20.0
32.6
52.0
56.0
58.0
36.5
iWet.
Toward the end of the first month the young rail begins walking
more deliberately and assumes the gait of the adult bird. When it is
seeking food, it tips its tail in typical adult fashion. Tail tipping was
observed in one 2- week-old chick.
68 NORTH AMERICAN FAUNA 67
Month-old King Eails have at least four calls: (1) seep-seep-seep-
( repeated) indicates general satisfaction and, particularly, acknowl-
edges the presence of others and notifies them of its presence; (2)
tah-eef tah-eef (repeated), very high pitched and progressively lower
in volume as sleep approaches, indicates relaxed comfort and sleep-
iness; (3) soo, tsoo {tsoo) indicates lonely dissatisfaction ; (4) keelp-
keelp-keelp-, a series of five or six hoarse notes in rapid sequence, ex-
presses protest.
Thirty to sixty days
During the second month the juvenal body plumage replaces the
natal down. The first juvenal feathers may be obscured by down until
the young rail is nearly a month old, but by the seventh or eighth
week the development of nearly all of the body feathers is complete.
The plumages of four captive King Rail chicks developed at about
the same rate through the first 6 weeks, but the rate varied consider-
ably thereafter.
The first evidence of change from the natal down plumage is the
appearance of white auricular tufts and pale juvenal feathers on the
underparts and flanks (figs. 27 and 28). Feather development in
these areas during the fourth week is as follows :
(1) The sternal region of the ventral tract : The pinfeathers are
pale buffy brown, and are tipped with natal down that is being pushed
out.
(2) The crural tract: Pinfeathers are whitish with black down at
the tips.
(3) The femoral tract: Feathers are approximately the same color
as those of the ventral tract.
By the latter part of the fifth week the juvenal plumage of most
young rails is developing in all body areas, but feathering of the
crown and back of the neck may begin slightly later in some individ-
uals. The abdominal region, axillary region, chin, and upper throat
are whitish and contrast rather sharply with the dusky upperparts,
particularly the lower back and rump. The dark brown feathers of
the upper back and humeral tract are well advanced, feathers of the
cervical region (lower throat) are approaching a cinnamon color, and
the thighs and flanks are faintly barred. The upper and undertail
coverts are making their appearance, and the anal circlet is sur-
rounded by short white feathers.
Quills began to appear the latter part of the fifth week on the
wings and tails of two of four captive birds. The primary and
secondary coverts developed more rapidly than did the primaries
and secondaries. The linings of the wings developed last.
By the sixth week the side of the head is whitish and faintly
washed with gray. A white supercilliary stripe is beginning to ap-
NATURAL HISTORY OF THE KING RAIL
69
Figure 27. — Downy young King Rail, 31 days old, with juvenal plumage begin-
ning to develop. White auriculars (of ear region) and white feathers of crural
tract are visible ; tip of bill and nares are white.
70
NORTH AMERICAN FAUNA 67
Figure 28. — Ventral view of 31-day-old King Rail showing development of white
juvenal plumage in sternal and abdominal region and crural tract.
NATURAL HISTORY OF THE KING RAIL
71
dm
1* Iff
- s»1 1
Figure 29. — Fifty-day-old King Rail with juvenal plumage nearly complete. Tail
and wings are undeveloped. This individual is slightly behind the average for
its age.
pear. The legs and bill approach flesh color, and the distal half of the
bill is darker than the proximal half. The eyes have brown irides and
black pupils.
By the seventh or eighth week the juvenal body plumage is almost
completed, and the young King Rail has a more browmish and less
dusky appearance (fig. 29). The cinnamon coloring of the lower
throat and breast approaches that of the adult. The juvenal feather-
ings of the crowm and nape are complete, and the flight feathers and
tail are well advanced on most birds.
There may be considerable variation in weight and size of young
rails during the second month of development. At two months a cap-
tive male and two captive females weighed 327.0, 258.6, and 202.0
grams respectively.
The juvenal plumage is nearly complete by the age of 60 days.
Remiges have developed enough so that some juveniles can make
short flights after the ninth week.
Ridgway and Friedmann (1941, p. 84) have presented a detailed
account of the juvenal plumage, as follows :
Above similar to adult, dark phase, but the dark centers of the feathers of
the back, etc., less fuscous, more dull black, the edges grayer and less well de-
veloped on the interscapulars and not at all developed on the lower back and
rump which are uniformly blackish, the long scapulars being the only feathers
with well-developed tawny-olive margins ; lesser and some of the outer greater
72 NORTH AMERICAN FAUNA 67
upper wing coverts tipped with white and crossed by another narrow white
band about 7 mm. anterior to the tip ; sides of head as in pale phase adult, but
the light ochraceous-buff areas cross-barred with narrow dusky lines (actually
the tips of the feathers) ; lower throat pale, light ochraceous-buff narrowly
barred with grayish hair brown to deep drab ; anterior part of breast more
heavily washed with pale ochraceous-buff; rest of breast and entire abdomen
white, crossed by broad, closely spaced, but somewhat broken bands of grayish
hair brown, the middle of the abdomen and lower breast unbarred; sides and
flanks dusky grayish olive-brown barred with white or buffy white; thighs and
vent like sides of breast but somewhat darker ; under tail coverts and under
wing coverts as in adult.
Call notes of young rails remain essentially the same during the
second month of life as they were during the first month, but the
voice becomes a little hoarser and deeper. During the second month
the young rail frequently exercises by jumping up and down, flap-
ping its wings at the same time.
The begging display may still be observed occasionally during the
ninth and tenth weeks, but it soon disappears.
A considerable change in calls occurs during the ninth and tenth
weeks. Some of them now approximate the calls of the adults. The
call most like that of an adult bird is a raucous crying squawk or cat-
like "meow." This call is made when a bird is separated from the
family group or is excited. The typical jupe-jupe-jupe-call of the
adult was not heard until the fifth month.
First winter plumage
For most individuals the first winter plumage is similar to that of
the adult. Some individuals, however, have whitish juvenal-like
plumage of the underparts and less distinct markings about the face.
Most rails in juvenal and first-winter plumage have some white bar-
ring on the wing coverts. This is also true of some adults.
MISCELLANEOUS NOTES ON BEHAVIOR OF YOUNG
Sleeping
From the time captive birds hatched until they were approximately
a month and a half old, the three to six occupying the same cage slept
together. Thereafter, as they assumed a somewhat different sleeping
posture, they usually slept separately, but sometimes still slept as a
group in the same part of the cage.
During the first 2 weeks after hatching, a warm quart-sized bottle
of water was placed in the box with the downy young. When sleepy
the chicks would huddle around the bottle, but not always in contact
with one another. However, if the bottle was removed, the chicks
huddled together when sleeping.
When just a few days old, chicks sleep in a prostrate position. They
simply flop dowrn on their bellies, usually with one side of their heads
NATURAL HISTORY OF THE KING RAIL 73
(cheeks) against the ground. Shortly thereafter, and until they
reach the age of V/2 to 2 months, they assume a sitting posture for
sleeping. Older young sleep more in the manner of adults, standing
on one or both legs, with the head turned around and the bill tucked
beneath the feathers of the back, or with the neck drawn in and the
bill pointed down to the ground.
Competition
Virtually no peck order was exhibited by captive King Rail chicks
during the first 2 weeks of life, when they fed together amicably in
one area. Thereafter, they competed for food, and after obtaining a
morsel from the common feeding site, would run away and ingest it
or run around the cage for several minutes before swallowing it.
Because of size variation of individuals older than 2 weeks of age,
there was an obvious peck order, but with little antagonism. A smaller
or more agile chick often would not hesitate to steal a morsel from a
larger competitor.
Bathing
When bathing, the King Rail assumes a partial squatting position.
By an up and down movement of the legs, the body moves in one
cadence, then the wings in another, and finally the head, dipping water
and flipping it over the back, in still another. The body feathers are
extended (somewhat ruffled), the closed wings are loose and moving,
and the head is immersed while cocked sideways, presumably for more
surface area, and hence functions better as a paddle in flipping water
over the body. Water also reaches the plumage through the up and
down action of the body and the movements of the wings. The bath-
ing operation usually takes 1 or 2 minutes. A captive bird evicts
another from the bathtub by pecking at its feet rather than at some
other part of its body.
WINTER BEHAVIOR OF CAPTIVE RAILS
During extended freezes or when there is a snow cover, water for
drinking is obtained by ingesting snow or small chunks of ice. Cap-
tive King Rails were observed ingesting snow and ice during periods
of heavy snowfall and during freezeups in a cage on Bluegill Pond at
the Patuxent Wildlife Research Center, Laurel, Md. One bird was
observed as it ingested a chunk of ice 3 inches in length and y2 inch
in width.
Captive King and Clapper Rails at Bluegill Pond preferred to
rest on the ice rather than in a more protected section of the cage pro-
vided with a windbreak and a bedding of straw (fig. 30) . During alter-
nating periods of freezing and thawing, spherical chunks of ice, up to
the size of a baseball, stuck to the tails of the Clapper Rails, and smaller
particles stuck to their breasts. Strangely, particles of ice virtually
never adhered to any part of the plumage of the King Rails.
348-693 0—69 6
74
NORTH AMERICAN FAUNA 67
0m ■■
Figure 30. — Captive King and Clapper Rails at Patuxent Wildlife Research
Center, Laurel, Md., January 1960.
Foods
Crustaceans and aquatic insects are the preferred foods of the King
Rail in most areas. Fish, frogs, grasshoppers, crickets, and seeds of
aquatic plants also have a high palatability rating with this species.
During the winter, particularly when the birds are hard pressed, con-
siderable quantities of grain or some other vegetable matter may be
consumed. In the southeastern Arkansas rice country, domestic rice
formed 30 percent by volume of the King Rail's winter food. A
stomach collected in December at Beaver Dam, Wis., was full of wheat.
In South Carolina, Audubon (1835, p. 29) examined a gizzard
crammed full of oats and collected King Rails in corn fields in autumn
near Charleston.
Some unusual foods found in gizzards include cherry {Prunus sp.)
seeds, skunk (Mephitis sp.) hair, feathers and vertebrae of a female
Red-winged Blackbird, King Rail eggshell fragments, a small water
snake (Natrix sp.), a mouse (Peromyscus sp.), a shrew (Sorex sp.),
fall army worms (Laphygma frugiperda), blackgum (Nyssa syl-
vatica) seeds, acorns (Quercus sp.) , and pine (Pinus sp.) seeds. A bird
collected near Fleming's Landing, Del., on September 30, 1961, had its
gizzard crammed with seeds of both waxmyrtle (Myrica cerifera) and
bayberry (M. carolinensis) .
The King Rail is more diversified in its choice of foods than its
salt-water relative, the Clapper Rail, as might be expected because of its
wider range and more variable ecology, which may find it feeding on
the edge of a salt marsh along the coast or in an oat field a thousand
miles inland.
Its adaptability to subsistence on a wide variety of foods in addition
to its usual diet of crustaceans and aquatic insects enables the King
to winter much further north than is generally realized. A King Rail
observed by I. W. Knight at Lome Park, Ontario, on December 26,
1960, remained in that locality until at least mid- January. It was
seen along an open stream where it was observed feeding on a frog and
the berries of "deadly" nightshade (Solanaceae) (Woodford and Bur-
ton, 1961, p. 326).
In some parts of its breeding range, particularly in brackish tidal-
river marshes of the Middle and South Atlantic coast, the King Rail
sometimes subsists largely on a 1-item diet, the red-jointed fiddler
crab.
75
76 NORTH AMERICAN FAUNA 67
From 1959 to 1961, several hundred observations were made during
the nesting season of King Rails feeding in brackish marshes along
the Delaware Bay between Fleming's Landing and Woodland Beach,
Del., where the red-jointed fiddler crab occurs in great abundance.
This little crab formed the main diet of the rail ; the only other item
of any importance was a clam (Macoma balthica) (fig. 31). Stomach
examinations confirmed field observations.
Table 12. — Principal foods of 118 King Rails from Arkansas ricefields
[ Volume = percent of total volume of stomach contents. Occurrence = percent of stomachs in
which found]
Collected in —
Winter Spring Summer Fall Annual
Food item (Dec.-Feb.) (Mar.-May) (June-Aug.) (Sept.-Nov.) volume,
33 stomachs 48 stomachs 16 stomachs 21 stomachs 118
stomachs
Vol- Occur- Vol- Occur- Vol- Occur- Vol. Occur-
ume rence ume rence ume rence ume rence
Animal:
Invertebrate:
Crayfish 7 18 61 81 22 25 3 5 23
Aquatic beetles 20 76 7 21 19 31 10 48 14
Land beetles 1 13 8 56 11 87 4 38 6
Grasshoppers 5 24 3 8 6 63 14 57 7
Aquatic bugs 5 24 1 15 10 44 6 29 6
Other insects (i) 21 2 17 8 75 5 38 4
Spiders (') 3 (') 2 (i) 13 1 10 (i)
Snails.. 3 12 2 8 0) 6 (») 10 1
Vertebrate:
Fish 7 30 1 21 8 19 26 43 11
Frogs 5 21 4 15 5 50 4 24 5
Miscellaneous 5 7 6 8 _. 3
Total 58 95 90 74 79
Plant:
Rice -_ 30 52 4 19 10 31 21 29 16
Ricefield weed seeds (i) 36 (') 4 (>) 56 2 38 1
Woody plant seeds 1 2 3 10 1
Tubers 12 21 3
Total. 42 5 10 .._ 26 21
» Trace.
There is considerable variation in food items taken by different
individuals in the same habitat and at the same time. Two birds col-
lected from a tidal marsh on the Choptank River in Maryland in
February 1961 present an interesting contrast. Bird A fed entirely
on fish, while bird B ate a wide and rather unusual assortment of
foods including the seeds of arrow-arum, hackberry (Celtis occiden-
talis), halberd-leaved tearthumb (Polygonum arifolium), dogwood
(Cornus florida), and grape (Vitis sp.). Bird B had also eaten cray-
fish and a snail (Gastropoda). The seeds of arrow-arum contain
calcium oxalate crystals and apparently are rejected by virtually all
water birds except the Wood Duck (Aix sponsa) . This was the only
time I encountered them during my studies of rail foods.
Only in the Arkansas ricefields has a fairly complete seasonal
survey of King Rail foods been made (Meanley, 1956, p. 252-258).
NATURAL HISTORY OF THE KING RAIL
77
Figure 31. — Foods of the King Rail in brackish bay marsh, Broadway Meadows,
Kent County, Del.: (1) Mud crab (Sesarma reticulation); (2) red-jointed
fiddler crab (Uca minax) ; (3) clam (Macoma balthica). (Photograph by
Frederick C. Schmid.)
Small series of stomachs have been collected from a few other local-
ities. Most of these were examined by John C. Jones of the U.S. Fish
and Wildlife Service.
ARKANSAS RICEFIELDS
In the Grand Prairie rice-producing area near Stuttgart, Ark., 118
stomachs were collected between 1950 and 1955 by Karl Kitler and
myself (table 12).
Animal life comprised 79 percent of the King Rail's annual diet.
It constituted 90 percent or more in spring and summer, dropped to
74 percent in the fall (the largest number of stomachs were collected
in November and may have made the figure lower than if there had
been better representation of the early part of this season), and was
still lower (58 percent) in winter.
The shift in feeding grounds from roadside ditches in the spring
to ricefields in summer and early fall, and finally back to natural
drainage ditches and small cattail marshes in winter, may account for
78 NORTH AMERICAN FAUNA G7
some of the seasonal variations in diet. However, seasonal fluctuation
in the abundance of aquatic animal life is apparently the basic
explanation.
The crayfish was the principal food of the King Rail in the rice
area, constituting 23 percent (by volume) of the annual diet; it
formed 61 percent in spring; 22 percent in summer; 3 percent in fall;
and 7 percent in winter. Since crayfish were available the year round,
it appears that consumption of this crustacean was influenced as
much by the availability of other favored foods as by the abundance
of crayfish. Possible seasonal variations in the size, agility, and/or
palatability of the crayfish according to age may have been factors
bearing upon the extent of seasonal use by the King Rail.
Another staple food available at all seasons was fish, which com-
posed 26 percent (by volume) of the diet in the fall when many fish
had become impounded in the shallow borrow pits of drained ricefields
and were easy prey for the foraging birds.
Aquatic insects were important foods, especially certain beetles and
waterbugs which were available the year round. Predaceous diving
beetles (Dytiscidae) furnished 19 percent of the winter diet.
Land beetles, chiefly ground beetles (Carabidae), scarabs
(Scarabaeidae), and snout beetles (Curculionidae) made up 6 per-
cent of the rail's annual food, while grasshoppers (Orthoptera)
constituted 7 percent.
A wide variety of other insects were taken in small quantities.
During the summer and fall they formed 8 percent and 5 percent,
respectively, of the food. Among these insects were dragonfly (Odon-
ata) nymphs, back-swimmers (Notonectidae), horsefly (Tabanidae)
larvae, fall army worms, rice water weevils (Lissorhoptrus simplex) ,
and rice stinkbugs (Solubea pugnax) .
Frogs accounted for about 5 percent of the annual diet.
The King Rail apparently is more of a vegetarian than its salt
marsh counterpart, the Clapper Rail. John Oney (1954, p. 23), in
studying fall foods of the Clapper Rail along the Georgia coast,
found that plant materials constituted only trace items of the Clapper
Rail's diet at that season. Martin, Zim, and Nelson (1951, p. 82) found
the volume of plant food in the Clapper Rail's diet to be 11 percent in
winter, 1 percent in spring, 0 percent in summer, and 3 percent in fall.
In the Arkansas area, vegetable matter in the diet of the King Rail
made up the following volumetric percentages during the 4 seasons :
42 percent in winter, 5 percent in spring, 10 percent in summer, and
26 percent in fall.
Cultivated rice seed was taken in larger quantities than any other
plant food, forming 16 percent of the annual diet. Increased consump-
tion of rice seed during fall and winter was due in part to the
abundance of waste grain left in the stubble. Kalmbach (1937, p. 60),
NATURAL HISTORY OF THE KING RAIL 79
in his study of the food of blackbirds in Louisiana, suggested that the
hard siliceous hulls of rice seed may be used in the gizzard for
grinding.
Kicefield weeds, abundant in all rail habitats, furnished some food
through the year. Seeds of jungle-rice (Echinochloa colonum), wild
millet, bullgrass, rice-cutgrass, beakrush, and smartweed (Polygonum
spp.) were found as traces in many stomachs in each season except
fall when they composed 2 percent of the contents.
The following seeds of woody plants were found in several
stomachs: blackberry (Rubus sp.), snowball (Styrax americana),
blackgum, and oak. Tubers of marsh plants, probably sedge
(Cyperaceae), were found in several stomachs, and one rail had eaten
tubers of an arrowhead.
TEXAS RICEFIELDS
Twelve stomachs were collected in ricefields at Eagle Lake, Colorado
County, Tex., during September 1938 by Valgene W. Lehmann.
Three items formed the bulk (63 percent) of the food and occurred
in at least half of the stomachs. The most important, the coneheaded
grasshopper (Neoconocephalus sp.), occurred in nine stomachs and
formed 30 percent by volume; dragonflies (Odonata) formed 20 per-
cent by volume ; and crayfish formed 13 percent by volume. An assort-
ment of insects accounted for most of the remainder. Rice seed was
the only plant food taken and comprised only 5 percent of the total
food consumed.
LOUISIANA RICEFIELDS
Nine stomachs were collected in ricefields in the gulf coast region of
Cameron and Vermilion Parishes in the summer of 1925 by
E. R. Kalmbach and in 1955 and 1965 by myself.
Crayfish were in seven of nine stomachs and were the major item
in six. Crickets (GryJlus sp.) were found in four stomachs and were
the most important items in three of those. Weevils were the only
other important food.
UPPER ST. JOHNS RIVER, FLA.
Six stomachs were collected in marshes in the Persimmon Hammock
area during the spring of 1905 by W. W. Worthington.
Crayfish were the major items in five of the six stomachs. Short-
homed grasshoppers (Acrididae) occurred in all of the stomachs,
but were important percentagewise in only one. Aquatic and land
beetles formed the balance of the food.
CURRITUCK SOUND, N.C.
Seventeen stomachs were collected, mostly in October, November,
and December, 1909 and 1910, at Church's Island by J. B. White and
W. L. McAtee.
80 NORTH AMERICAN FAUNA 67
Animal life formed 88 percent of the food with seeds of aquatic
plants forming most of the balance. Important animal foods were
sunfish (Centrarchidae) and perch (Percidae), grasshoppers and
locusts, and aquatic insects (mostly Belostomatidae, Hydrophilidae,
and Haliplidae) .
PATUXENT RIVER, MD.
Six stomachs were collected in fresh tidal-river marshes along the
Patuxent River in southern Maryland in early fall between 1923 and
1958, by O. J. Tremis, C. H. M. Barrett, and unknown rail bird
hunters.
An interesting assortment of materials was found in this small
series, including killifish (Fundulus heteroclitus) , crayfish, dragonfly
nymphs, snails (Amnicola sp.), grasshoppers, and crickets; leaves of
a bulrush (Scirpus sp.) and rice-cutgrass ; seeds of dotted smartweed,
halberd-leaved tearthumb, arrow-leaved tearthumb (Polygonum
sagittatum), burreed (Sparganium eurycarpum), water parsnip
(Slum suave), silky dogwood (Cornus amommm), and wild cherry.
BEAVER DAM, WIS.
Eleven stomachs were collected in various marsh types during
summer and fall, 1889-1908, by W. D. Snyder and C. F. Zimmerman.
Crayfish constituted over 90 percent by volume of the food in 6
stomachs and occurred in 9 of 11. Snails, soldier flies (Odontomyia
sp.), dragonfly larvae, a mollusk (Stagnicola paJustris), grasshop-
pers, and a fish (Etheostominae) were major items in the other four
stomachs.
CHICAGO, ILL.
Five stomachs were collected during spring (April-May), 1912-17,
by G. Eifrig and K. W. Kahmann. Crayfish were the major items
(50 percent plus by volume) in four of the stomachs, and dragonfly
nymphs were the major items (55 percent by volume) in the fifth.
Stalks of a bulrush (Scirpus sp.) were important (40 percent by
volume) in one stomach, and horsefly larvae were common (42
percent by volume) in another.
Two stomachs were collected during the summers (July) of 1878
and 1915 by S. A. Forbes and K. W. Kahmann. Frogs ( Rana sp.) were
the most important food (54 percent by volume) in one, and larvae of
soldier flies (94 percent by volume) in the other.
Crayfish were a major item (45 percent plus by volume) in each of
the three stomachs collected in the fall (September-October), one in
1913 and two in 1915 by K. W. Kahmann. Frogs (Rana sp.) were im-
portant (51 percent by volume) in one stomach.
Feeding Behavior
King Rails usually feed in areas concealed by plant cover or in com-
paratively open areas where they blend well with their surroundings
and are only a few steps from cover. Sometimes, however, they are
very conspicuous, as when feeding at low tide on mud flats or in
open roadside ditches. Dawson (1903, p. 443) observed such feeding
activity in Ohio :
In a region where they were in little fear of molestation, I have seen them
deploy upon an extensive mud flat in broad daylight and go prodding about in
company with migrant sandpipers, for the worms which riddle the ooze with
their burrows.
In tidewater areas, feeding probably occurs most frequently at low
tide. Whenever I visit the brackish marshes of the Delaware Bay at
ebb tide, I see King and Clapper Rails feeding in the tidal creek beds.
I suspect that King Rails do very little feeding at night, although
they are sometimes active during this period, as they are occasionally
heard calling, particularly during the courtship period. King Rails
that I kept in captivity in Louisiana and Maryland were relatively in-
active at night. In fact, some of them would habitually return to a
favorite spot in the cage each evening at dusk, sit down, and remain
quiet for long periods.
Generally King Rails forage in water so shallow that only the bill,
or part of it, disappears beneath the surface while food is sought.
However, on March 25, 1954, on the Arkansas Grand Prairie, I observ-
ed a pair of rails feeding in a roadside ditch where the water varied
from 6 to 12 inches in depth. Both of these birds immersed their entire
heads and necks in water, and several times their entire bodies disap-
peared beneath the surf ace, In fact, they occasionally fed by "tipping
up" like dabbling ducks.
Since King Rails are accustomed to procuring their food from the
water, if perchance they obtain a food item from land and are near an
aquatic environment they usually carry the morsel to the water and
immerse it before ingestion.
PELLET CASTING
Both the King Rail and the Clapper Rail, whose major food is crus-
taceans, reject most of the exoskeletal fragments of these animals
through the regurgitation of pellets (fig. 32) .
King Rail pellets examined in Arkansas and Maryland were com-
posed of crayfish and aquatic insect fragments. Nearly every pellet
81
82
NORTH AMERICAN FAUNA 67
Figure 32. — Regurgitated King Rail pellets from Dorchester County, Md. Note
the round gastroliths of the crayfish (Cambarus sp. ) in top and bottom pellets
at left. Pellets averaged 2 cm. in length by 1.5 cm. in width. (Photograph
by Frederick C. Sehmid.)
examined contained the hard cylindrical convex-shaped gastroliths of
crayfish.
In brackish marshes near Woodland Beach, Del., where King and
Clapper Rails occur together, pellets contained exoskeletal fragments
of the red-jointed fiddler crab and a clam (Macoma balthica) . As many
as 14 pellets were found on a single muskrat house.
FEEDING YOUNG
I have observed King Rails feeding their chicks within 2 hours after
hatching. Gross and Van Tyne (1929, p. 439) reported the same for
the Purple Gallinule, another member of the family Rallidae.
When the very small young are abroad, they follow one or both
parents about as food is caught for them. Larger food items such as
crayfish and large grasshoppers are dismembered and fed to the young
in pieces.
Sometimes, however, the young remain in a concealed place and
wait for the parent to bring them food. At Grand Chenier, La., April 8,
1956, I observed a pair of adult rails for over an hour as they kept
up a steady pace to and from a small pond catching fish and carrying
them to young hidden behind tussocks of grass 30 or so feet distant.
NATURAL HISTORY OF THE KING RAIL 83
Similar feeding was observed in a Louisiana ricefield where adults
brought crayfish to young that remained in the same spot.
As the young grow older, they not only accept food from their par-
ents but also begin to forage for themselves. An interesting example
of this dual feeding activity was observed near Woodland Beach,
Del., on July 29 and 30, 1959. An adult King Rail and three young
approximately 5 to 6 weeks of age were observed feeding on clams
at low tide in the bed of a creek. The adult bird dug in the mud for
the clams, usually inserting its entire head beneath the surface. It
would eat four or five clams and then carry one to the young. The
clams were swallowed whole. Sometimes one of the young, standing
next to its parent, would watch the digging operation and then start
digging for itself. The parent and its young were seen digging for
clams in the same place on both days. A raccoon also came to this
spot and dug many clams.
REGIONAL OBSERVATIONS
Arkansas ricefields
During March, April, and May 1952 it was not unusual to see 15
or 20 King Rails in the evening feeding along ditches bordering cer-
tain highways leading out of Stuttgart, Ark. The variety of rail food
available in these roadside ditches includes crayfish, tadpoles, frogs,
aquatic insects, small fish, and snails. Toward the end of May the rails
move out of the ditches and into ricefields, where they are found until
harvest. During the winter they are found about the network of rice-
field canals and natural drainage, often moving from place to place
along runways beneath matted vegetation.
The King Rail feeds almost exclusively in ricefields during the
summer. About the only time it emerges from this cultivated marsh
type is to move from one ricefield to another. When a field of nearly
mature grain is drained preparatory to harvesting, the rails move
over to a field of younger rice which is often contiguous to the dry
field. Some ricefields have a few low wet spots which prove attractive
to rails, even up to harvest time; but the last feeding place in nearly
all drying ricefields is along the "borrow" or ditch bordering the
levees.
This typical bird of the rice country performs a service to the rice
grower by consuming large numbers of crayfish that bore holes in the
ricefield levees. A single large crustacean is usually torn apart and
eaten in the course of several minutes; in one case the dismembering
operation was timed at 7 minutes. Small crayfish are ingested whole.
Delaware Bay marshes
In the brackish tidal marshes between Fleming's Landing and
Woodland Beach, Del., I have found King and Clapper Rails feeding
84 NORTH AMERICAN FAUNA 67
in the same tidal gut. An important food of the rails here is the red-
jointed fiddler crab, which is found further upstream than other
species of fiddler crabs, but not far above the brackish zone and, as
far as I know, not beyond tidewater in this area.
In some areas, fiddlers' holes or dens are concentrated mostly along
or just beneath the top of the embankment of a tidal gut, and at high
tide are inundated. Rails seem to feed mostly at low tide. When stalk-
ing fiddlers, rails are very slow and deliberate. When within striking
distance, a rail makes a quick thrust or stab at the crab. When a
fiddler is caught, it is often taken to some favorite feeding spot, such
as a muskrat house or pile of drift debris, for dismembering. The large
claw of the male crab is disengaged in the following manner, as
described by Oney (1954, p. 24-25) for the Clapper Rail:
The bird grasps the crab with its bill between the claw and the body. Then
holding the crab, it vigorously shakes its head. The claw goes one way and the
crab another. The bird then runs over and picks up the body and swallows it.
The female crab does not get the same treatment because both of their claws are
nearly equal size.
Some fiddlers, too large to swallow, are hacked to pieces and then
eaten bit by bit.
Savannah National Wildlife Refuge
In early April 1960, I made observations of feeding King Rails
along an alligatorweed-choked canal on the Savannah National Wild-
life Refuge. Alligatorweed forms extensive mats upon which rails,
gallinules, coots, herons, and several species of ducks do much of their
foraging for aquatic insects, fish, tadpoles, frogs, and crustaceans.
While this vegetation type has no apparent value in a waterfowl
management program, it is of obvious value to birds that utilize its
growth form to facilitate food gathering. Some King Rail feeding
territories along the alligatorweed-choked canal were no more than
20 feet square, indicating the high rail-food productivity of these
aquatic mats.
The most frequently observed pair of rails defended their feeding
territory vigorously. Although they nested on the other side of the
dike some 40 yards distant in a sawgrass marsh, they consistently
returned to the same section of the canal for feeding.
The base of operations in the pair's feeding territory was a pile of
debris, possibly an old alligator nest, at the edge of a small clump of
giant cutgrass. From here the rails radiated out to feed on the mat of
alligatorweed. Whenever a crayfish or some other large morsel was
obtained, it was brought back to the pile of debris for "servicing"
and eating. Old earthen dikes still much in evidence throughout the
abandoned ricefield marshes of the Carolina Low Country are also
used for this purpose.
NATURAL HISTORY OF THE KING RAIL 85
Along the low banks of the canals, and sometimes partially sub-
merged in the alligatorweed, numerous alligators, some 5 to 6 feet
in length, sun themselves on warm spring days (fig. 14). They lie
motionless for several hours at a time, and if they move there is
simply a splash and complete submergence. It would seem that, to a
waterbird wading around in the canal, an alligator sunning along
the water's edge would look like another one of the many logs lying
half submerged in the alligatorweed. But this is not the case. Rails,
gallinules, and other birds feeding in the canal recognize the alligator
as an enemy and usually give it a wide berth. Sometimes, however,
they feed to within 2 feet of an alligator before circling the animal or
retreating.
SOME UNUSUAL OBSERVATIONS
A very unusual feeding performance was recorded by Earle McPeak
(Trautman, 1940, p. 230) at Buckeye Lake, Ohio. On June 11, 1929,
an adult King Rail was observed to uncover, break, and eat five eggs
of a painted turtle {Chrysemys picta), which on the preceding day
McPeak had watched the turtle lay in a hole and cover with earth.
Another unusual field observation concerned the capture and
devouring of a Semipalmated Sandpiper (Ereunetes pusilhis) by a
King Rail. E. D. Greaves (Chamberlain, 1960, p. 443) reported this
incident, which took place at Pea Island, N.C., May 22, I960 :
The rail darted out of the grass, picked the sandpiper from a feeding flock
and after stabbing it repeatedly, pulled it apart and devoured it.
In June 1960, at low tide in Taylor's Gut, Kent County, Del., I
saw a King Rail pursue and peck at a 3-foot-long water snake (Natrix
sp.) for a distance of some 50 feet. Finally, the snake stopped and
remained motionless for about 2 minutes as the rail continued to peck
at it. Eventually each took off in a different direction. Possibly the
rail was chasing the snake out of its nesting territory rather than
pursuing it for food.
At Grand Chenier, La., March 1956, I observed a rail catch a crab
in an open spot in the marsh. As the rail headed for cover to feed on
the morsel, it was harassed so much by a Boat-tailed Grackle (Cassi-
dix mexicanus) that it surrendered the crab to the blackbird.
Nauman (1927, p. 218) reported the following unusual feeding
activity which took place at his home in Iowa during a snow storm.
On April 16, 1921, when there were 8 inches of snow on the ground, a
King Rail was observed walking around on the porch picking up
bread crumbs. Until the snow melted, it returned to the porch on
numerous occasions to feed on crumbs.
At the Patuxent Wildlife Research Center, a captive 10-month-old
female King Rail attempted to eat a mouse and choked to death.
Mortality Factors
The most important factors in mortality of King Rails are (1)
striking (or being struck by) manmade objects and (2) predation.
In recent years, pesticides may also have become an important factor.
From time to time, hurricane-caused floods decimate coastal marsh
populations.
MANMADE OBJECTS
Since King Rails are nocturnal migrants, they strike various illu-
minated objects such as television towers, ceilometers, tall buildings,
and lighthouses. On their breeding grounds in the southern rice belt,
I have found dead King Rails under telephone lines and impaled on
barbed wire fences.
The automobile is an increasing hazard because of the network of
roads in the intensively cultivated rice country and the marshland
of the South Central States. Over a 3-month period (March 1-June 1,
1952), I found 24 dead adult King Rails along a 10-mile stretch of
paved road leading north from Stuttgart, Ark.
During floods in the gulf coast marsh country of Louisiana, King
Rails and other water birds are literally flushed out of the marshes
to the nearest high ground, which is often a well-traveled highway.
During one period of high water in the marsh bordering the highway
between the Intracoastal Canal and Creole, La., I saw 30 King Rails
(mostly adults with broods) walking back and forth across the road
in the face of heavy traffic. Many were being killed, particularly the
young.
Wherever muskrats are trapped, King, Virginia, and Sora Rails
become casualties since they use the runways where the traps are
placed. Whenever I have encountered muskrat trappers in the course of
my travels from New Jersey to Louisiana, I either have seen King
Rails removed from traps or have been told of the many that are
caught incidental to muskrat trapping. One trapper encountered in
Maryland caught 50 King Rails during the course of a single trapping
season (2^ months).
PREDATION
Judging from the many examples of predation in the literature,
the King Rail appears to have a wide variety of natural enemies. Fur
bearers are probably the most important, chiefly the raccoon, because
of its fondness for eggs, and its abundance in most marsh habitats.
NATURAL HISTORY OF THE KING RAIL 87
At a Lonoke, Ark., fish hatchery two rail nests found along the edge
of a pond had been broken up by raccoons as evidenced by numerous
tracks leading from the nest to a point in the open where the eggs had
been taken and destroyed.
The mink may also be an important predator of this rail. The Rev.
John Bachman, pioneer naturalist from Charleston, S.C., was quoted
by Audubon (1835, p. 29-30) regarding the fate of the King Rail as
follows :
Its feathers are frequently found lying on the banks of rice-fields, ponds and
lagoons, in places where the tracks of the minx plainly disclose the plunderer.
Similarly, on the Arkansas Grand Prairie, a mink trapper told me
of finding typical mink signs at freshly killed King Rails. In Cur-
rituck County, N.C., Kenneth Wilson (1954, p. 199-307) found six
species of birds taken by mink, including the King Rail. Wilson
also cited one case in which an otter (Lutra canadensis) ate one of
these large rails.
Predation by a bobcat (Lynx rufus) was noted by Bachman and
cited in Audubon (1835, p. 29-30) as follows:
. . . while placed on a stand for deer, I saw a wildcat creeping through a
marsh that was near to me, evidently following by stealthy steps something that
he was desirous of making his prey. Presently he made a sudden pounce into
a bunch of grass, when I immediately heard the piercing cries of the Marsh-Hen,
and shortly after came passing by me the successful murderer with the bird in
his mouth.
Tanner and Hendrickson (1956, p. 56) found a dead King Rail at
the den of a red fox ( Vulpes fulva) in Iowa :
During May and June 1951, the den of a red fox on the research area was
visited almost daily and the remains of prey brought to the den by the parent
fox examined. The only rail seen at the den was an adult King Rail found
May 12. Since this bird had not been present May 11, and obviously had been
dead several days, it seemed likely that it had been found dead by the foxes
and brought to the den as carrion. The carcass had not been mutilated.
Bachman (in Audubon, 1835, p. 29) also found remains of a King-
Rail in the stomach of a large moccasin (probably Agkistrodon
pisclvorus). Another ornithologist of the Charleston area, Arthur T.
Wayne, also cited an example of cottonmouth predation on the King
Rail (1910, p. 36) :
In the month of April, 1900, I was observing a nest of this species in a button-
wood bush, which was in a pond of water, and, about every other day, I waded
into the pond to see how many eggs were there. About the 8th of May, I judged
that the full complement of eggs would be completed, and upon visiting the nest
in the afternoon, which was very cloudy, I saw what I supposed to be the bird
incubating. But upon close inspection I was very much surprised to find that
what I took for the bird was a huge Moccasin (Ancistrodon piscivorus) , which
I promptly shot. This snake had eaten all the eggs and perhaps caught the bird
as the feathers were scattered around the nest.
88 NORTH AMERICAN FAUNA 67
John W. Aldrich (personal communication) collected a fox snake
(Elaphe vulpina) that had several King Rail eggs in its digestive
tract, in Little Cedar Point Marsh on Lake Erie in Ohio.
Alligators (Alligator mississipiensis) are known to take King Rails.
Kellogg (1929, p. 32) found remains of King Rails in 4 of 15 stomachs
of alligators taken in Cameron Parish, La., in 1926.
Coulter (1957, p. 18) examined the gastrointestinal tracts of 157
common snapping turtles ( Chelydra serpentina) from Maine marshes
and found bird remains in about one of every four.
Forty-two specimens contained evidence of a minimum of 52 birds including
25 ducks, 11 grebes, 3 rails and 13 unidentified birds.
No mention is made of the rail species taken. Birds the size of a
King Rail could easily be taken by a large snapper. As an example,
one 31-pound turtle consumed five birds including one Ring-necked
duck (Aythya collaris), one Common Golden-eye (Bucephala clan-
gula), and three Pied-billed Grebes (Podilynibus podiceps).
Hawks and owls take their toll of rails, gallinules, and coots. This
is to be expected since virtually all birds of prey hunt in marshes
as well as in uplands. Errington and Breckenridge (1936, p. 835)
found the remains of a King Rail in a Marsh Hawk (Circus cyaneus)
collected in August 1934 in the North Central States prairie region.
Errington (1932, p. 182) also found remains of a King Rail in a Great
Horned Owl (Bubo mrginianus) pellet collected in April 1930 in
southern Wisconsin.
The Fish Crow (Corvu-s ossifragus), well known as a plunderer of
Clapper Rail eggs, is also known to take eggs of other rails. Frank
C. Kirkwood and John Sommer found broken King Rail, Virginia
Rail, and Least Bittern egg shells beneath an active Fish Crow nest
at Gum Swamp Island, Blackwater Marsh, Dorchester County, Md.,
on June 16, 1929 (from Kirkwood's field notes, 1929) .
HURRICANES
An interesting account of King and Clapper Rail behavior during
a hurricane in the Louisiana prairie marsh country was given by
Robert J. Newman (1957, p. 409), from information based on eye
witness accounts of H. W. Belknap, graduate student from Louisiana
State University, and J. H. Sutherlin, Manager of Sabine National
Wildlife Refuge. Hurricane Audrey was Louisiana's worst coastal
storm of the present century and struck with its greatest force along
the southwest Louisiana coast on the morning of June 27, 1957. New-
man's report was as follows :
Rafts of marsh debris, ranging in size from 10X10 to 20X100 feet, went float-
ing by (HWB, JHS). On them huddled a strange company of water moccasins,
nutria, rails, and gallinules — sometimes as many as 20 birds to a raft. Occa-
sionally as the great rollers surged forward, the mats of vegetation would buckle
NATURAL HISTORY OF THE KING RAIL 89
and disintegrate. Then, according to Sutherlin, the Purple Gallinules would
drown, but the supposedly less aquatic Clapper or King Rails would swim
adeptly away through the rough water and flying spindrift.
The rain-drenched lawn at Sabine headquarters became a sanctuary within
a sanctuary. The wildlife congregated there included water snakes, a marsh
deer, a skunk, armadillos, and 200 to 250 King or Clapper Rails.
348-693 O— 69-
The King Rail as a Game Bird
Although a prized game bird, this large rail is seldom hunted be-
cause of the difficulty of maneuvering in its habitat, and the unlikeli-
hood of finding concentrated numbers. Hence there are few localities
where the King Rail can be considered an important game bird. In
the gulf coast domestic rice producing areas of Texas and Louisiana,
where King Rails are most numerous, they are probably shot in greater
numbers than anywhere else; however, here considerably less than 1
percent of the local population is shot. In the Middle Atlantic States,
the largest numbers are taken by hunters shooting Sora in the wild-
rice marshes of the Delaware Valley and Chesapeake Bay. However,
in this area, some hunters will gun the wildrice marshes all season
without seeing a King. Elsewhere in its range, the King Rail is shot
only incidentally. Quail hunters in the South frequently encounter
them and take a few.
Probably the best known hunting grounds in the Delaware Valley
are in the wildrice marshes of Salem and Cumberland Counties, N.J.
The tidal marshes of the Maurice and Cohansey Rivers in Cumber-
land County are famous rail hunting areas. The Patuxent River
marshes near Upper Marlboro are the most important areas in the
Maryland part of Chesapeake Bay ; while in Virginia Tidewater, King
Rails are hunted mostly along the lower James River in the upper
reaches of some of its tributaries such as the Chickahominy ; and
along the Pamunkey and Mattaponi, famous Sora shooting rivers
which downstream form the York. One of the most popular spots for
King Rail shooting in Texas ricefields is the Eagle Lake area in Colo-
rado County.
A few King Rails are killed by Clapper Rail hunters in coastal salt
marshes. Hunters Tom Reed and Gordon Clark killed 3 Kings and
50 Clappers in 2 days of gunning at Chincoteague, Va., during Sep-
tember 1961.
In the open piney woods of Central Louisiana it seems odd to see a
King Rail during the winter in some wet spot in the bluestem range.
I know of several hunters in that locale who take one or two in each
quail and woodcock season.
METHODS OF HUNTING
Contrary to popular opinion, the King Rail is not always an easy
mark for the hunter despite its alleged weak flight. Characteristics
90
NATURAL HISTORY OF THE KING RAIL
91
of its habitat and other circumstances govern the manner in which it
flies, and I have seen several fly at a fast clip for 300 yards.
The Marsh Hen, as this bird is sometimes called, tends to be secretive,
but at times it is found in completely open situations near cover. It is
not readily flushed, usually escapes by running, and flies only as a last
resort. Flight begins with the legs dangling, but as the bird levels
off, it flies in a straight line close to the ground with its legs extended
straight back beyond the tail and its neck stretched forward.
Methods of hunting King Kails in the wildrice marshes of Chesa-
peake Bay and the Delaware Valley are quite different from those in
the domestic ricefields of Louisiana and Texas. In the Chesapeake Bay
and Delaware River tidal marshes, rails are hunted mostly from long,
narrow, flat-bottom boats that are poled through the flooded wildrice
beds by a "pusher" (figs. 33 and 34). When tides reach peak level, the
rails flush as the boat approaches them.
Patuxent River, Md.
On the Patuxent River in the Southern Maryland tobacco country,
rail hunting has been a sport enjoyed by the tobacco growing aristoc-
racy since Colonial times. Perc Blogg (1944, p. 67-68), famous Mary-
land sportsman of the "good old days," presents an account of the
hunting of Soras and King Rails on the Patuxent in his delightful
Figure 33. — Method of hunting railbirds in Patuxent River wildrice marshes in
Maryland. Boat is poled through marshs at flood tide ( September 1958) .
92
NORTH AMERICAN FAUNA 67
Figure 34. — After the hunting season : railbird boats tied up at dock in Decem-
ber. Low tide on old wildrice marsh on the Patuxent River in Maryland.
little book There Are No Dull Dark Days. He begins his essay with a
poem about the Marsh Hen :
Give me a gun and some old Marsh
Where the pusher's voice calls mark right !
As the king rail springs from the ditch beyond
Then as suddenly drops out of sight.
"Dah he! Mark left!" What a thrill as the excited "pusher" calls the first
bird on a beautiful September morn. This is the moment for which the gunner-
man has waited many a month.
We are on the Patuxent. Everything has clicked ; the wind is southeast and
gentle, the day warm but not hot. . . . high tide at 7 :30 a.m. As far as the eye
can see on both sides of the river, artistic stalks of wild "oats" stand. Over on
the higher marshes, a solid mass of brilliant yellow blossoms, called butterweed
by the natives, greets the eye.
Rails, being in good requisition for the table, have been extensively hunted,
particularly on the marshes of the Delaware and Chesapeake bays. Most sought
after of the rails is the little sora or Carolina rail. The Virginia and king rail
often add variety to the bag, however.
It is next to impossible to make these birds take wing when they are able to
run. Because of this, rail are hunted only when the tide is so high that the flooded
marshes afford no shelter and make it impossible for them to run. While the
pusher poles a small skiff over the flooded marsh, the hunter stands in the bow,
gun in hand. Every now and then a bird will jump, sometimes almost from under
the boat, flying away with apparent feebleness, just over the tops of the foliage.
As it flies, its legs dangle awkwardly. This ruse, however, is merely to prepare
it for the sudden drop which often leaves the surprised hunter drawing a bead
NATURAL HISTORY OF THE KING RAIL 93
on empty space. Oftimes the gunner doesn't even see the rail announced by
Mark right, Mark left, or simply "Dah he goes."
On the Patuxent marshes in the 1940's, I would flush about 2 Vir-
ginia and 1 King to each 100 Sora.
Eagle Lake area, Tex.
The method of hunting King Rails in the Texas ricefields was
described by Dev Klapp (1961, p. 14) :
On the Gulf Coast the rail is found in the great ricefields where it seeks its
food. The close-knit rice stalks afford the bird ideal cover, and any attempt to
hunt such terrain in an orthodox manner is next to useless. The hidden rail
would merely sneak away as the hunter approached and never be seen.
So Texans waste no time slopping around through ankle-deep gumbo mud
chasing a bird that refuses to flush. They patiently wait until rice harvest time,
then, by the hundreds they take to the fields.
Beginning about October 1, reapers appear in the fields to gather the grain.
The machines take positions at the outer perimeter of the fields and, as the sun
rises above the horizon, go into action. Snorting and rocking, they circle the
fields time and again, gradually working toward the center. Running side by
side, usually in pairs, they cut a swath some 30 feet wide.
The hunters wait until all but a 30-foot swath of rice is cut, then hurriedly
take stands to each side of the uncut grain. In this narrow strip, they know, all
the rail in the field are crowded. Reluctant to fly, the birds have crept through
the standing grain, just ahead of the oncoming machines, until forced into this
bit of cover.
As the reapers get about half-way through this swath the birds begin to panic.
That's when the fun starts. A hunter suddenly shouts, "There's one!" as the
first rail rises from cover, almost straight up. Before the bird has set its course
it is brought down.
The combine (fig. 35) has replaced the reaper in most areas, but the
method of hunting is still the same.
Other areas
Milton B. Trautman (personal communication) reported that be-
tween 1907 and 1918 King Rails were hunted in the marshes of Indian
Lake, Logan County, Ohio, and that bags of half a dozen birds a day
were not uncommon. M. G. Vaiden, an ornithologist from Rosedale,
Miss., told me that while Bobwhite hunting in the delta, January 30,
1945, he shot five King Rails and three Bobwhites; and on January
27, 1946, he shot four King Rails. The Rails were flushed from boggy
spots in growths of ragweed (Ambrosia sp.) .
In the days of the market gunner, rails were shot in much greater
numbers than at present, and were sold in the markets of most of the
large cities along the eastern seaboard. To the epicures, the King
Rail of the fresh-water marsh was far superior to the Clapper Rail
of the salt marsh. Charles S. Westcott ("Homo") writing in Forest
and Stream magazine in the 1880's, said (Stone, 1937, p. 332) :
Many of the latter, however, carefully plucked were palmed off for King Rails
on those less expert in identifying them.
94
NORTH AMERICAN FAUNA 67
Figure 35. — Method of hunting railbirds in southern riceflelds. As rice is
harvested the hunter walks beside the reel of the combine which flushes the
birds. (Arkansas Grand Prairie, September 1953.)
Summary
Audubon described the King Rail. Alexander Wilson and, for a
while at least, Audubon considered it to be some form3 perhaps an
adult, of the Clapper Rail. Audubon and Bachman, staunch friends,
spent a lot of time together in the Charleston area, where Bachman
was able to show Audubon that the large rufescent rail was associated
almost entirely with the fresh- water marshes and ricefields, while the
"ash-coloured" rail was a denizen of the salt marshes. By so doing,
Bachman apparently convinced Audubon that the two were distinct
species.
The King Rail is essentially an inhabitant of fresh and brackish
marshes, while the Clapper Rail is more an inhabitant of salt marshes.
In some transition areas, however, particularly in the lower reaches
of brackish river marshes, both species occur and sometimes inter-
breed. Viable eggs resulting from a mixed mating are known to occur,
and some specimens taken in areas of interbreeding appear to be hy-
brids. Further study is necessary to understand more precisely the
relationship between King and Clapper Rails.
Rallies elegems elegans is restricted mostly to the humid section of
North America, east of the 100th Meridian. The Cuban form, R. e.
ramsdeni, is restricted to Cuba and the Isle of Pines. A third form,
R. e. tenuirostris (sometimes considered to be a race of Rallus longiros-
tris), is restricted to the fresh- water marshes of the Valley of Mexico.
In the United States the King Rail is found in greatest numbers in the
South Atlantic and Gulf Coastal Plain provinces, especially in coastal
fresh and brackish marshes and in ricefields.
While it has been shown that the King Rail is migratory, so few
birds have been banded that little is known about the time, distance,
and routes of migration. Apparently the major routes are along the
Atlantic Coastal Plain and through the Mississippi Valley. Most birds
probably arrive on the breeding grounds in the north during April
and May and depart in August and September. Although most King
Rails migrate to the South Atlantic and Gulf Coastal Plains in the
fall, a few individuals may be found wintering almost anywhere
within the geographic range of the species. Since there are numerous
records of birds wintering in the middle and northern latitudes, it is
not known whether birds seen in the spring are recent arrivals from
the south or birds that have wintered in the area. Rails are very secre-
tive in winter, becoming quite vociferous with the onset of the breed-
ing season; thus spring arrival dates may be based on sighting or
hearing of birds that have been present in the area for some time prior
96 NORTH AMERICAN FAUNA 67
to the recorded date. During the summer King Rails may restrict
their activities to a relatively small area for as long as 3 months, dur-
ing which time the adults are nesting and molting, and the young,
flightless for about 2 months after birth, are. growing.
The 4,000,000-acre Louisiana coastal marsh is the largest block of
breeding habitat in the range of the King Rail. The southern rice-
fields are a good example of an optimum breeding habitat. The breed-
ing density of a King Rail population in a South Carolina river marsh
was 25 males per 100 acres; in an inland Florida marsh the density
was 30 males per 100 acres. King and Clapper Rails were found breed-
ing in the same brackish marshes in Louisiana, Maryland, and
Delaware.
King Rail sexes appear to be alike in plumage. The male averages
larger than the female. Immature birds apparently can be externally
distinguished from adults during the first autumn by the color of the
soft parts. Most rails in juvenal and first-winter (immature) plumage
have some white barring on the wing coverts. This is also true of some
adults. The light-phase adult plumage described by Ridgway and
Friedmann (1941) is probably typical of hybrids. The small sample
of weights and measurements given in this report indicates that the
King Rail averages slightly larger than the Clapper Rail.
Molt is not well understood. Apparently all individuals molt after
the nesting season, but some also molt during it.
King Rails are known to return to the same section of the same
marsh for several consecutive years to breed. Territories are estab-
lished and maintained by aggressive behavior, primarily that of the
male. The mating call, given by the male, presumably serves the same
purpose as the song of a passerine bird on its territory, namely to at-
tract a mate and to repel other birds of the same sex (but also, in the
case of the King Rail, to maintain contact after pairing) .
The display of the male during prenuptial courtship consists mostly
in walking about with tail uplifted and white undertail coverts ex-
tended. After pairing, other forms of display and a repertoire of
subdued calls are used to maintain the pair bond.
Copulation takes place near the nest site, before and during egg
laying.
The nesting season of the King Rail is one of the longest among
birds in the South. In Florida, there is evidence of nesting from
January to July ; and in Louisiana, from March to September. In the
middle and northern latitudes the nesting season is usually about 3
to 4 months long.
Since the Clapper Rail in South Carolina is known to be double-
brooded, it is possible that the King Rail in the southern part of its
range may also have more than one successful brood; however, this
has not yet been established.
NATURAL HISTORY OF THE KING RAIL 97
Clutch size is large, averaging 10 or 11 eggs. There appears to be
no geographic variation in clutch size.
Nesting success appears to be high in most areas. In one Arkansas
sample, success was 75 percent; and in one Iowa sample it was 67
percent. Such success is probably due in some measure to the incu-
bating birds' pugnacity toward would-be offenders. Survival of young
until 2 weeks of age was about 50 percent in the Arkansas rice belt.
Downy young of the King Rail are black. A change from the downy
plumage begins at about 1 month. Juvenal plumage is obtained in
about 60 days, and wings are developed enough for short flights after
the ninth week.
Usually chicks are more than 1 hour old before they can go over
the nest and return. During the first month six different calls were
recorded.
The King Rail occurs in a wider range of habitats and feeds on a
greater variety of foods than most other North American rallids.
Aquatic animals, particularly crustaceans, are its main food. Plant
food items are taken more under emergency conditions. When the
King Rail occurs in the same environment as the Clapper Rail, it may
subsist mostly on a 1-item diet like that species.
In most areas King Rails feed mainly in shallow water where the
depth is usually 2 or 3 inches. In Delaware Bay marshes, King Rails
fed almost entirely on mud flats, exposed at low tide, and on the
Arkansas Grand Prairie, in summer, they fed almost exclusively in
ricefields.
In some areas King Rails were observed to have feeding territories
to which they returned regularly at times other than the breeding
season.
Most food items are ingested whole, but larger crustaceans often are
dismembered before eating.
Mortality of King Rails apparently is due mainly to birds coming
in contact with manmade objects, and to natural predation, especially
the destruction of eggs by raccoons. Studies to determine the effects
of pesticides on this rail have not been made; however, its favorite
food, aquatic animal life, is highly susceptible to these chemical agents.
In at least one intensive agricultural area, the Arkansas rice belt,
where ecological conditions have not changed during the last 20 years,
the King Rail has shown a marked decline.
The King Rail is one North American game bird that certainly is
not overhunted. This is so primarily because the population is gen-
erally scattered and its habitat is usually difficult for hunters to work
through. The only time that I found them concentrated and fairly
easy to shoot was during the harvest in the Louisiana ricefields. A
few are shot incidentally in the course of hunting Sora in the Middle
Atlantic States, and Bobwhite in the deep South in damp piney woods.
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Appendix 1— Methods of Capturing for
Banding
Most of the King Rails captured for banding have been taken in
traps or with long-handled dip or clap nets. A few downy young
have been caught by hand. Some King Rails are inadvertently caught
in the course of trapping ducks.
TYPES OF CAPTURING DEVICES
Long-handled dip or clap net
This device is very effective in capturing King Rails on nests. Traps
equipped with drift fences and placed in breeding territories will
also capture rails during the nesting season as the birds wander
about their territories. However, in such situations, where only one
or two birds are involved, long-handled nets are more efficient because
of the time required to install traps. Most incubating King Rails
can be approached closely enough to be caught on a first attempt. The
long-handled net I use has a bamboo handle 7y2 feet in length, a
hoop 2 feet in diameter, and a net 3 feet in depth.
Nest traps are effective in catching incubating King Rails, but
are time-consuming to construct and are probably no more effective
than long-handled nets. Blandin (1936, p. 62-63) described a nest trap
for the Clapper Rail, a species that flushes more readily from the
nest than does the King Rail.
All-purpose or cloverleaf trap
As nearly as I can ascertain, this is the type of trap most often
used for capturing King and Clapper Rails (figs. 36 and 37). Seth
H. Low (1935, p. 16-20) originally designed this trap for catching
shore birds for banding. He used long leads or drift fences placed
at right angles to the trap and running up to the trap entrance. Robert
E. Stewart (1954, p. 1) caught nearly 1,000 Clapper Rails with this
type of trap at Chincoteague, Va., and designed a very effective gather-
ing cage that prevents rails from getting back into the trap (fig. 37).
The effectiveness of Stewart's design is due to a ramp that begins at
the opening at ground level and runs toward the top and rear of the
cage. When a rail reaches the top of the ramp it drops down into a
small chamber where it is well contained and easily retrieved.
The all-purpose trap used by Stewart and other Patuxent Wildlife
Research Center biologists is about &y2 feet in length. Each of the two
103
104
NORTH AMERICAN FAUNA 67
Figure 36. — All-purpose or cloverleaf trap and drift fence in shrub swamp at
Patuxent Wildlife Research Center, Laurel, Md.
LEAD OR
ENTRANCES
DRIFT FENCE
_GATHERING
CAGE
1 FUNNEL
BAFFLE
FUNNEL
RAMP
Figure 37. — All-purpose or cloverleaf trap.
NATURAL HISTORY OF THE KING RAIL 105
cells is about 3 feet by 3 feet ; flattened out, each cell section is approxi-
mately 9 feet in length by V/2 feet in width. The top is 7 feet by 4 feet.
The trap is best made of hardware cloth or welded wire. A less
expensive and also less durable trap can be made of 1-inch poultry
mesh (chicken wire). A trap made of poultry mesh should have a
hardware cloth gathering cage.
The lead wire or drift fence should be a minimum of 1 foot in height.
Two feet is better. One-inch poultry mesh is an ideal size for the drift
fence. Downy young of banding age can probably get through a 2-inch
mesh.
The length of the lead or drift fence depends upon the trapping
situation. In marshy impoundments at the Patuxent Wildlife Research
Center near Laurel, Md., I used some half-dozen different trapping
designs at various times.
The trap is placed in that part of a marsh known to be inhabited by
rails. Their presence is determined by hearing a bird call regularly
from the same area during the course of several consecutive days, by
locating a nest, or by hearing the cries of young that have been sep-
arated from their parents. The general area should be checked for
tracks, droppings, and fragments of crayfish or other rail foods, and
when tracks along a well-worn runway or path are found, the strategy
then is to block it off with the drift fence.
Rails often feed along the edge of a marsh on a mud flat bordering
open water. In such a situation, the drift fence should extend a foot
or so out into open water and landward 10 or 15 feet (or more) to the
trap. At low tide, in cordgrass marshes in Delaware, King and Clapper
Rails feed along the exposed muddy bottoms of narrow creeks. In such
a situation, a drift fence leading to a trap on either bank can be placed
across the creek. Thus the rail's passage is cut off, and it tends to
follow the fence toward the trap and often goes into it.
In a 10-acre shrub swamp-marsh mixture at the Patuxent Wildlife
Research Center, the most select area for rails was in a 1-acre cattail
patch. By placing four all-purpose traps about 30 feet apart in a row
and connecting them with drift fences, I was able to block off an exten-
sive area and succeeded in catching many rails (King, Virginia, and
Sora). Numerous Common Snipe (Capella gallinago), Red-winged
Blackbirds, Rusty Blackbirds (Euphagus carolinus), Swamp Spar-
rows and Song Sparrows were also captured. A few Wood Ducks,
Woodcocks, American Bitterns (Botaur-us hntiginosus), and Green
Herons (Butor'uJes virescens) were taken as well. The trap placed at
the pond end of the cattail patch caught more rails than did the three in
the interior of the patch.
TENDING TRAPS
The number of times a day that traps should be tended depends
somewhat upon the size of the population in the area of interest and
106 NORTH AMERICAN FAUNA 67
the frequency of catch. In high population areas, such as some Clapper
Rail marshes, traps should be checked four or five times a day. In areas
of low population, two or three times a day is adequate, as too frequent
disturbance will frighten rails from trapping sites.
In areas of high predation by raccoons, opossums (Didelphis virgm-
iana) , minks, and domestic rats, it is well to leave traps open at night.
When operating on a 24-hour basis it is important to check the trap
at dusk or shortly thereafter. Rails left in traps at night in areas of
high predator populations will almost without exception be destroyed.
This is especially so because paths made by the operator seem to become
natural highways for mammals.
Between dusk and dawn I virtually never caught rails (King, Vir-
ginia, or Sora) in traps at the Patuxent Wildlife Research Center.
Most species of North American Rallidae are not as active at night as
during the day. Some rails are vociferous at night especially during
the breeding season; however, at that time they call mainly from
territories.
When I first started trapping rails, I expected to encounter trouble
with muskrats getting into traps. They do get into traps occasionally,
but burrow out quickly, and I have never had to remove a single animal.
AGE FOR BANDING KING RAIL CHICKS
Downy young can be banded at 2 weeks of age, a few individuals as
early as 10 days. The band size is number 5.
NEED FOR BANDING DATA
In view of the paucity of information on movements and popula-
tion dynamics of rails, a large number of these birds should be banded.
To accomplish this, it is necessary to know which methods are best
for capturing rails, where to place traps, and in what geographic areas.
The all-purpose or cloverleaf trap with drift fences is the best known
device for capturing rails for banding.
The best areas for trapping are usually determined by noting calls,
reading "sign" (tracks, droppings, piece of discarded crayfish), and
locating nests.
Localities in which I have found high populations are as follows : In
Louisiana marshes near the intersection of the Intracoastal Waterway
and the road to Pecan Island ; just below the intersection of the Water-
way along the road to Creole, and in a silted-in canal near Dulac ; in
marshes across the river (north) from Savannah, Ga.; and in marshes
on the Savannah National Wildlife Refuge near Savannah.
Appendix 2— Local Names
The King Rail is probably better known in life to marsh hunters and
trappers than to most ornithologists, and to these hunters and trappers
it has its own special name depending upon locality. I picked up many
of these names while working in the marshes and ricefields of the
South.
Great Red-breasted Rail Audubon (1835, vol 3, p. 27).
Fresh- Water Marsh Hen ___ Audubon (1835, vol. 3, p. 27).
Rale de Prairie Creole hunters of Louisiana ac-
cording to Audubon (Arthur,
1931, p. 235).
English Rail Hunters on Arkansas Grand Prai-
rie, Ark. (author).
Slash Guinea Hunters on Arkansas Grand Prai-
rie, Ark. (author).
Sage Hen Southeastern Arkansas (Chicot
County) rice farmers (author) ;
also muskrat trapper, Rappa-
hannock River, Va. (author).
Rice Guinea Northeast Arkansas rice belt
around Weiner and Hickory
Ridge (author).
Rice Chicken Northeast Arkansas rice belt
around Weiner and Hickory
Ridge (author).
King Sora Potomac River, Va. (Kirkwood,
1895, p. 278) ; Powhatan River,
Va. (author).
King Ortlan Patuxent River, Md. (author).
King Water Rail Muskrat trapper, Choptank River,
Md. (author).
Marsh Pullet Allen's Fresh, Wicomico River,
Md. (author).
Mud Hen New England (Forbush, 1925, vol.
1, p. 352).
Injun Hen Raleigh, N.C., area (Brimley, 1887
p. 201).
107
108
NORTH AMERICAN FAUNA 67
Okankiskoki (ruddy raven). Big Cypress Seminole Indians
(Mikasukis), Florida (Gilles-
pie, 1956, p. 123).
Oklani kahki (ruddy raven) _ Cow Creek Seminole Indians
(Muskogee), Florida (Gilles-
pie, 1956, p. 123).
Marsh Hen Many localities.
Stage Driver Lake Erie marshes, Ohio (author).
U.S. GOVERNMENT PRINTING OFFICE: 1968 O — 348-693
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