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Full text of "North American fauna"

BOSTON PUBLIC LIBRARY 

illlL^ . 

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 "Rec d 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 2y 2 miles north of 
Georgetown. The mean tidal range at Georgetown is Sy 2 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. 163. 4 
. . 56. 0- 72. 5 65. 9 


147. 0-162. 154. 3 
60. 0- 70. 64. 4 


142. 5-159. 5 151. 1 
55. 0- 69. 64. 6 


135. 5-160. 146. 8 
55. 0- 69. 5 61. 9 


58. 0- 65. 5 62. 5 
.. 52.0-64.0 58.4 


50. 0- 63. 61. 9 
49. 5- 58. 54. 


55. 0- 69. 5 63. 3 
48. 0- 56. 51. 7 


53. 5- 67. 59. 6 
41. 0- 56. 48. 1 


.. 50.5-60.5 55.1 


46. 0- 56. 50. 8 


45. 5- 53. 5 48. 8 


40. 0- 52. 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 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 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 Valley 3 
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 4 1 / 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 iy 2 
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 dow r n 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 y 2 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, 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 form 3 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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NATURAL HISTORY OF THE KING RAIL 90 

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!00 NORTH AMERICAN FAUNA 67 

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NATURAL HISTORY OF THE KING RAIL 101 

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102 NORTH AMERICAN FAUNA 67 

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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 7y 2 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 &y 2 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). 



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