SMITHSONIAN INSTITUTION
UNITED STATES NATIONAL MUSEUM
Bulletin 114
A REVISION OF THE KING SNAKES:
GENUS LAMPROPELTIS
BY
FRANK N. BLANCHARD
Instructor in Zoology, University of Michigan, Ann Arbor
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WASHINGTON
GOVERNMENT PRINTING OFFICE
1921
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ADVERTISEMENT.
The scientific publications of the United States National Museum
consist of two series, the Proceedings and the Bvlletins.
The Proceedings, the first volume of which was issued in 1878, are
intended primarily as a medium for the publication of original, and
usually brief, papers based on the collections of the National Museum,
presenting newly acquired facts in zoology, geology, and anthro-
pology, including descriptions of new forms of animals, and revisions
of limited groups. One or two volumes are issued annually and dis-
tributed to libraries and scientific organizations. A limited number
of copies of each paper, in pamphlet form, is distributed to specialists
and others interested in the different subjects as soon as printed.
The dates of publication are recorded in the tables of contents of
the volumes.
The Bvlletins, the first of which was issued in 1875, consist of a
series of separate publications comprising chiefly monographs of large
zoological groups and other general systematic treatises (occasionally
in several volumes), faunal works, reports of expeditions, and cata-
logues of type-specimens, special collections, etc. The majority of
the volumes are octavos, but a quarto size has been adopted in a few
instances in which large plates were regarded as indispensable.
Since 1902 a series of octavo volumes containing papers relating to
the botanical collections of the Museum, and known as the Contribu-
tions from the National Herbarium, has been published as bulletins.
The present work forms No. 114 of the Bulletin series.
William deC. Ravenel,
Administrative Assistant to tlie Secretary
In cTiarge of the United States National Museum.
Washington, D. C, July 19, 1921.
TABLE OF CONTEiNTS.
Page.
Introduction 1
Genus Lampropeltis 4
Description 4
Range 7
Habitat and habits 7
Variation 8
Subdivisions of the genus Lampropeltis 16
Key to the forms of Lampropeltis 18
Summary of structural characteristics of the forms of Lampropeltis 24
The getulus group 26
Splendida 26
Holbrooki 33
Niger 43
Getulus 49
Floridana 62
Brooksi 66
Yumensis 66
Boylii 75
Conjuncta 89
Californiae 94
Nitida 103
Summary 104
The calligaster group 115
Calligaster 115
Rhombomaculata 128
Leonis 138
Summary 138
The triangulum group 139
Polyzona 139
Micropholis 149
Nelsoni 155
Annulata 159
Crentilis 165
Amaura 172
Syspila 179
Triangulum 188
Elapsoides 206
Virginiana 217
Ruthveni 221
Multicincta 222
Pyrrhomelaena 231
Summary 237
Isolated forms 245
Mexicana 245
Alterna 247
Conclusion 249
Bibliography 255
Index 259
IV *
LIST OF ILLUSTRATIONS.
Page.
Fio. 1. Lampropeltis getulus splendida, fihowing normal arrangement of head
plates from above 5
2. Everted penis of Lampropeltis calligaster 7
3. Lampropeltis getulus splendida, showing normal arrangement of head
plates from the side 12
4. Lampropeltis getulus getulus, shovang extra supralabial scute inter-
polated l)etween normal second and third 12
5. Lampropeltis pyrrhomelaena, showing an extra labial in each series.. 13
6. Lampropeltis calligaster, showing position of the added labial in each
series 13
7. Lampropeltis elapsoides elapsoides, showing reduction of lower labials
from 9 to 8 13
8. Map showing locality records for Lampropeltis getulus splendida 28
9. Map showing locality records for Lampropeltis getulus holbrooki 37
10. Map showing locality records for Lampropeltis getulus niger 45
11. Diagram showing geographic variation in number of ventral plates in
Lampropeltis getulus getiilus 55
12. Map showing locality records for Lampropeltis getulus getulus 56
13. Map showing locality records for Lampropeltis getulus floridana 63
14. Diagram for comparison of numbers of ventral plates in Lampropeltis
'getulus boylii, L. g. yumensis, and L. g. splendida 70
15. Diagram for comparison of numbers of cross bands or rings in Lampro-
peltis getulus boylii, L. g. yumensis, and L. g. splendida 71
16. Map showing locality records for Lampropeltis getulus yumensis 72
17. Diagram showing geographic variation in numbers of ventral plates
in Lampropeltis getuhis yumensis and L. g. boylii 81
18. Diagram showing geographic variation in numbers of annuli in Lam-
propeltis getulus yumensis, and L. g. boylii 82
19. Diagram showing geographic variation in numbers of labial plates in
Lampropeltis getulus boylii, L. g. yumensis, and L. g. conjuncta... 84
20. Map showing locality records for Lampropeltis getulus boylii 85
21. Color pattern of Lampropeltis getulus conjuncta 90
22. Color pattern of Lampropeltis californiae cahfomiae from San Diego
County, California 96
23. Map showing locality records for I>ampropeltis californiae californiae. . 97
24. Map showing the distribution of the various forms of the getulus
group. 104
25. Diagram showing geographic variation in numbers of ventral plates
in the forms of the getulus group 106
26. Diagram showing geographic variation in numbers of rings or cross
bands in the various forms of the getulus group Ill
27. Typical color pattern of Lampropeltis getulus boylii To face 112
28. Typical color pattern of Lampropeltis getulus yumensis To face 112
29. Color pattern of Lampropeltis getulus splendida, showing approach
to the pattern of L. g. yumensis in the widened dark areas. . To face 112
30. Color pattern of Lampropeltis getulus splendida, showing the theo-
retically most primitive type of pattern in the getulus group. To face 112
31. Color pattern of Lampropeltis gelulus splendida, showing the common
pattern of this form To face 112
32. Typical color pattern of Lampropeltis getulus holbrooki To face 112
33. Typical color pattern of Lampropeltis getulus niger To face 112
V
VI LIST OF ILLTJSTEATIONS.
Fig. 34. Color pattern of Lampropeltis getulus getulus, showing approach to
pattern of L. g. niger To face 112
35. Typical color pattern of Lampropeltis getulus getulus To face 112
36. Typical color pattern of Lampropeltis getulus floridana To face 112
37. Typical color pattern of Lampropeltis getulus broolcsi To face 112
38. Diagram of the relationships of the forms of the getulus group 115
39. Typical pattern of Lampropeltis calligaster 118
40. Color pattern of Lampropeltis calligaster, showing the striped effect
commonly exhibited by dark individuals 119
41. Map showing locality records for Lampropeltis calligaster 122
42. Color pattern of Lampropeltis rhombomaculata 129
43. Map showing locality records for Lampropeltis rhombomaculata 131
44. Diagram showing variation in number of ventral plates in Lampro-
peltis polyzona and L. micropholis 144
45. Map showing locality records for Lampropeltis polyzona 146
46. Map showing locality records for Lampropeltis micropholis 150
47. Map showing locality records for Lampropeltis triangulum nelsoni 156
48. Map showing locality records for Lampropeltis triangulum annulata .
.
161
49. Map showing locality records for Lampropeltis triangulum gentilis. . 167
50. Diagram showing interrelationships between Lampropeltis triangulum
annulata, L. t. gentilis, L. t. amaura, and L. t. syspila 169
51. Map showing locality records for Lampropeltis triangulum amaura . . . 174
52. Lampropeltis triangulum syspila, showing typical color pattern of
anterior end of body 181
53. Map showing locality records for Lampropeltis triangulum syspila 182
54. Lampropeltis triangulum triangulum, showing typical pattern of
anterior end of body 191
55. Map showing locality records for Lampropeltis triangulum triangulum . 195
56. Diagram showing geographic variation in number of ventral plates in
Lampropeltis triangulum triangulum 198
57. Diagram showing geographic variation in number of dorsal blotches
in Lampropeltis triangulum triangulum 201
58. Lampropeltis elapsoides elapsoides, showing typical pattern of anterior
end of body 208
59. Map showing locality records for Lampropeltis elapsoides elapsoides. . 210
60. Map showing locality records for Lampropeltis elapsoides virginiana. . 217
61. Map showing locality records for Lampropeltis multicincta 224
62. Map showing locality records for Lampropeltis pyrrhomelaena 232
63. Map showing distribution of the various forms of the triangulum group . 238
64. Lampropeltis polyzona, typical color pattern To face 242
65. Lampropeltis triangulum nelsoni, typical color pattern To face 242
66. Lampropeltis triangulum annulata, typical color pattern To face 242
67. Lampropeltis triangulum amaura, a common color pattern of this
form To face 242
68. Lampropeltis triangulum syspila, typical pattern of this form. . To face 242
69. Lampropeltis triangulum triangulum, typical color pattern To face 242
70. Lampropeltis micropholis, typical color pattern in Ecuador—in
Colombia the heavily spotted intervals are narrower To face 242
71. Lampropeltis pyrrhomelaena, typical color pattern To face 242
72. Lampropeltis triangulum gentilis, typical color pattern To face 242
73. Lampropeltis elapsoides elapsoides, typical color pattern To face 242
74. Lampropeltis ruthveni, color pattern of type specimen To face 242
75. Lampropeltis multicincta, typical pattern To face 242
76. A diagrammatic presentation of the relationships of the forms of the
triangulum group 245
77. Lampropeltis mexicana, showing color pattern 246
78. Lampropeltis altema, showing style of color pattern 248
A REVISION OF THE KING SNAKES:
GENUS LAMPROPELTIS
By Frank N. Blanchard,
Instructor in Zoology, University of Michigan, Ann Arbor.
INTRODUCTION.
The snakes of the genus Lamprojpeltis have long been in a state of
great confusion from the systematic standpoint. Only about one-
third of the forms now recognized have been at all clearly defined;
of the rest some have been known from only a few specimens each,
some have been unnaturally divided among several forms, and
others have been included under a single name. The confusion has
been greatest among the forms here included in the 'Triangulum
group." In this group wide differences occur in color pattern
among forms exhibiting great similarity in structural features and
striking resemblance in pattern between others that are quite dis-
tinct in scutellation.
The reasons for this systematic confusion are mainly two: (1)
Many of the forms have been known from only a very few specimens,
and even these have not been assembled by any one reviewer. This
has made it difficult or impossible to correctly evaluate the differ-
ences exhibited, with the result that forms only distantly related
have been regarded as identical or only subspecifically distinct, and
others that are obviously identical have been considered distinct.
(2) Taxonomists have often disregarded the geographic probabilities.
Thus many absurd definitions and groupings have made identifica-
tion difficult and uncertain. Locality meant little; sometimes the
head of a specimen would belong to one name and the body to an-
other. With the systematic status of forms in such confusion, but
little can be done toward acquiring a knowledge of habits, life his-
tories, and ecologic and economic relations.
It was to bring order into this puzzling group of important and
economically valuable North American snakes and to put their
classification on a genetic basis that the present study was undertaken,
and the work is regarded as completed if the numerous forms have
been defined, our knowledge about them brought up to date, and
their genetic relationships traced out as well as the material at
present available will allow.
2 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
Every means has been taken to make the present review as accm-ate
and as complete as possible. Specimens have been borrowed from
many collections, both public and private, throughout the United
States. The assembling thus of a great many specimens not pre-
viously reported upon has helped greatly to a better understanding
of the distribution and relationships of the various forms. Every
specimen has been subjected to a careful examination of structural
and color pattern features. The former have of course been regarded
as of more significance than the latter in establishing relationships,
but, as a matter of fact, when the patterns of two specimens were
similar and the structural features different it was evident in nearly
every case that the resemblance in patterns was only superficial or
general, and that features easily overlooked were usually present to
confirm the fundamental differences.
It is fully realized that the present effort is but a preliminary at-
tempt at an understanding of the genus, and for this reason care has
been taken not to obscm-e features that may later prove to be of
importance. Nothing is to be gained by "lumping" of doubtful
forms; apparent constancy throughout a definite geographic range
has been deemed sufficient to warrant recognition by name, and inter-
gradation has been the criterion of subspecific distinction. If an
error has been made, the name will call attention to it, and wiU
result in ultimately determining the proper status of the form.
However, it is believed that more abundant material will increase
rather than diminish the number of forms now recognized, and,
where the material has suggested such a possibility, attention has
been drawn to the fact.
About 1,600 specimens have been assembled for examination.
This is but few, to be sure, when compared with the numbers often
available in other groups of animals, but very good for king snakes,
for it should be noted that the great bulk of this number is the result
of chance collecting of forms that are, in general, not plentiful any-
where, and that it is more than foiu" times as many as could ever
have been examined by any previous reviewer. Nearly all the type
specimens of the genus are in the United States National Museum,
in Washington, and the Academy of Natural Sciences of Philadelphia,
and these have been examined. One hindrance to the work has been
the occasional lack or ambiguity of information as to locality. Speci-
mens without locality labels or with vague or indefinite localities are
valueless, and those with incorrect labels are worse than useless, be-
cause they are apt to be misleading. Collectors can not be too care-
ful about attaching to their specimens exact information as to locality
as soon as possible after collecting, and if they wiU also attach brief
notes on habitat, they will add much to the value of the specimens.
It is hoped that the present work will emphasize the serious
inadequacy of our knowledge of the king snakes. The maps of distri-
REVISION OF THE KING SNAKES. 3
bution will show from what localities specimens are needed. But
above all there is great need of reliable information on the natural
history and ecological relations of all forms in the genus. Many
students of wild life already know far more about the habits of
some of these forms than they will find in these pages, and this fact
should encourage them to report any careful observations on all
phases of the natural history and ecological relations that they may
have opportunity to make.
Grateful acknowledgment is hereby extended to the numerous
museums and individuals who have assisted by loaning specimens
or supplying needed information on particular points. First of all
the writer desires to express his gratitude to Prof. Alexander G.
Ruthven, director of the Museum of Zoology of the University of
Michigan, under whose general direction the work has been carried
on, for suggestions and encouragement throughout its progress.
Much assistance has been rendered by Dr. Leonard Stejneger for
providing the opportunity to study the large collection at the United
States National Museum, and for placing every available facility
at the writer's disposal. Thanks for the use of large and important
collections are due es})ecially to Dr. John Van Denbm-gh of the
California Academy of Sciences, to Dr. Thomas Barbour of the
Museum of Comparative Zoolog}^, to Mr. Henry W. Fowler of the
Academy of Natural Sciences of Philadelphia, to Dr. J. O. Snyder of
Leland Stanford University, to Miss Mary C. Dickerson of the Ameri-
can Museum of Natural History, to Dr. Joseph Grinnell of the
University of California, to Dr. Lawrence E. Griffin of the Carnegie
Museum of Pittsburgh, to Dr. Albert H. Wright of Cornell University,
and to Mr. Carl L. Hubbs of the Field Museum of Natural History.
Smaller, but for the most part very important collections, have been
received from the Kansas State Agricultural College, the University of
Kansas, the BrookljTi Museum, Iowa State College, the San Diego
Natural History Society, Ohio State University, Wesleyan Univer-
sity of Connecticut, the Colorado State Teachers' College, the Univer-
sity of Colorado, the Colorado State Agricultural College, the Alabama
Natural History Museum, the University of Arizona, Smith College,
the Public Museimi of Milwaukee, Charles Mohr Natural History
Museum of Mobile, the Victoria Memorial Museum, the University
of Wisconsin, Dr. W. S. Blatchley, Dr. T. L. Hankinson, Mr. H. P.
Loding, Dr. E. R. Dunn, Dr. Thomas Van Aller, Mr. J. W. Mackel-
den, and Mr. W. R. Jones. Everything received has been of material
help. Often the receipt of only a few specimens has resulted in the
solution of an important problem.
In the hsts of specimens examined, which accompany the descrip-
tions of the various forms of the genus, the collections in which the
specimens may be found are referred to as follows: Acad. PMla.,
4 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
Academy of Natural Sciences of Philadelphia; Ala. Mus., Alabama
Museum of Natural History; American, American Museum of
Natural History; W. S. BlatcMey, private collection of Doctor Blatch-
ley of Indianapolis; Brooklyn Mus., Brooklyn Museum; Calif. Acad.
8d., California Academy of Sciences; Carnegie, Carnegie Museum of
Pittsburgh; Colo. S. Agr. Col., Colorado State Agricultural College;
Colo. 8. T. C, Colorado State Teachers' College; Cornell Univ.,
Cornell University; Field, Field Museum of Natural History; F. i?.
Jones, private collection of W. K. Jones of Satsuma, Alabama;
Kans. S. Agr. Col., Kansas State Agricultural College; Kans. Univ.,
Kansas University; E. P. Loding, collection of H. P. Loding of
Mobile, Alabama; M. C. Z., Museum of Comparative Zoology;
San Diego Mus., San Diego Natural History Society; Ohio Univ.,
Ohio State University; Stan. Z7mv., Stanford University; U.S.N.M.,
United States National Museum ; Univ. Ariz., University of Arizona
;
M. V. Z., Museum of Vertebrate of Zoology of the University of
California; Univ. Colo., University of Colorado; Univ. Mich.,
Museum of Zoology of the University of Michigan; T. van Aller,
collection of Dr. T. van Aller of Mobile; Wesleyan Univ., Wesleyan
University of Conneciticut.
References in the text are to author, year, and page, and are to be
found in full in the Bibhography. References in the synonomies,
which are inclosed in brackets have not been examined by the writer.
All of the outline and color pattern drawings are the work of
Miss Doris M. Cochran of Washington, District of Columbia, and all
are originals except figure 37 of Lampropeltis getulus IrooTcsi which
was prepared from a drawing kindly loaned by Dr. Thomas Barbour
of the Museum of Comparative Zoology.
In further explanation it should be mentioned that solid circles
on the maps of distribution indicate locahties from which specimens
have been examined, and hollow circles indicate reports that are
believed to be trustworthy. Only localities that are at least as
specific as counties are plotted, and a single circle may represent
several records near together.
Genus LAMPROPELTIS Fitzinger.
1843. Sphenophis FitzingeB', Sys. Kept., p. 25 (type, Coronella coccinea Schlegel;.
1843. Lampropeltis Fitzinger, Sys. Kept., p. 25 (type, Eerpetodryas getulus
Schlegel).
1853. Osceola Baird and Girard, Cat. N. Amer. Kept., pt. 1, p. 133 (type, Cala-
maria elapsoidea Holbrook).
1853. Ophibolus Baird and Girard, Cat. N. Amer. Rept., pt. 1, p. 82 (type,
Eerpetodryas getulus Schlegel).
1876. BellopMs Lockington, Proc. California Acad. Sci., vol. 7, p. 52 (type,
zonatus Lockington).
Description.—The genus Lampropeltis of Fitzinger, established upon
the Eerpetodryas getulus of Schlegel (1837, 198) and later defined as
REVISION OF THE KING SNAKES.
OpMholus by Baird and Girard (1853, 82), belongs to the family
Coluhridae. It was separated in 1860 by Cope (254) from the
CoroneUa of the Old World on the presence of the entire instead of
divided anal plate, and in 1862 (302) the same author noted the
further difference of two scale pits instead of one.
The genus may be diagnosed as follows: Maxillary teeth, 12 to 20,
solid, slightly increasing or slightly decreasing in size posteriorly,
subequal, or the last two a little enlarged; mandibular teeth decreasing
in size posteriorly; head not or but slightly distinct from neck; eye
moderate, with round pupil; scales
smooth, with two apical pits, in 17 to
27 rows; anal plate entire; tail mod-
erate, caudals in two rows.
Thecephalicplates are normal (fig.
1), consisting of paired parietals, pre-
frontals, internasals, and single fron-
tal ; the nostril lies between two na-
sals; the loreal is normally present
(commonly absent only in elapsoides
and virginiana); there is normally
one preocular, and two postoculars;
the temporals are normally two in
the first row, three in the second, and
four in the third (except elapsoides,
virginiana, and micropholis in which
there is usually one less in each row,
and aJtema in which there is one
more in each row) ; the upper labials
are normally seven, varying fre-
quently to eight only in Jloridana,
pyrrhomelaena, and calligaster; the
lower labials are normally nine, the
fifth largest, being commonly eight
only in elapsoides, virginiana, and
rhomhomaculata, and commonly ten
only in conjuncta, pyrrhomelaena, and calligaster. The first lower
labials meet on the median line behind the triangular mental plate at
the symphysis of the lower jaw, and are succeeded by two pairs of
parallel chin shields, the posterior of which are not often longer than
the anterior and may be only half as long and separated by one or
two small scales.
On the body there are from 17 to 27 longitudinal rows of dorsal
scales wider on the lower row or two and narrower above, all perfectly
smooth, and each provided with two pits near the posterior extremity.
On the abdomen is a single series of large transverse plates, the ven-
FlG. 1.
—
LaMPROPELTIS GETITLUS SPLENTDIDA
(U.S.N.M. NO. 1849). IJ X NAT. SIZE. Show-
ing NORMAL ARRANGEMENT OF HEAD PLATES
FROM ABOVE. /, FRONTAL; i 710, INTERNASAL;
I, loreal; no, nasal; pa, parietal; pf, pre-
frontal; po, postocular; pr, preocular; r,
EOSTRAL; <l, P, <', TEMPORALS OF THE FIRST,
second, and third rows, respectively; ul,
UPPER LABIALS.
6 BULLETIN 114, U]SriTED STATES l^TATIONAL MUSEUM.
trals, which vary in number, in the genus, from 152 to 254. These
are terminated posteriorly by a single anal plate (found divided in
only three specimens examined, one triangulum, one elapsoides, and
one californiae, and it is reported as divided in the type of CoroneUa
formosa anomala Bocourt). The plates under the tail, or caudals,
are normally in two rows, but occasional individuals have one or
several of these entire (particularly in micropJiolis, pyrrhomelaena,
and some forms of getulus) ; they vary in number in the genus from 27
to 79.
The head is but slightly distinct from the neck, except in a few
specialized or aberrant forms in which it is plainly wider at the
temples (pyrrhomelaena, alterna, mexicana, and micropholis) . The
body is in general slender, of nearly uniform diameter, cylindrical
above, the sides meeting the belly in a noticeable angle. The tail
is short, tapering rapidly to a horny tip, and varying from 0.09 to
0.18 of the total length; only a few aberrant forms have it commonly
longer than 15 per cent of the total length (mexicana, alterna, and
pyrrhomelaena). The sex of an individual may in a majority of
cases be told by the shape of the posterior end of the body. When
the body is viewed from the side, an abrupt slope on the dorsal
line from body to tail indicates a female, a gentle slope a male.
Viewed from below, the tail of a female is .much narrower at the base
than that of the male. The tail of the male must be larger at the
base in order to contain the copulatory organs, while the body of
the female must be larger near the end to hold the eggs; therefore the
sudden change in diameter in the female where the body meets the
tail, and the gradual change in the male.
No fundamental pattern of coloration holds throughout the genus,
but, except for two isolated forms of dubious aflSnities (mexicana and
alterna), all the styles of coloration may be put into three groups.
In one (the getulus group) the pattern is fundamentally an oval
white center in a black scale, and from this may be derived every
other arrangement in the group, including two other kinds of spotted
patterns, two of cross bands, one of rings, and one of longitudinal
stripes. In another (the calligaster group) the pattern is of numerous
black-edged brown blotches set on a ground color of lighter brown.
In the third (the triangulum group) the pattern is of encircling rings
of white or yellow, bordered with black, and separated by red. By a
joining on the ventral plates of the black borders of adjacent pairs
of black rings the red becomes restricted to dorsal saddles, and by a
migration upward, onto the dorsal scales, of these black borders the
red finally becomes restricted to dorsal blotches, and in this case,
smaller blotches are developed on the sides in alternation with those
above. In this group red is a fundamental and almost universal
color, but when it becomes restricted to dorsal blotches it changes to
REVISION OF THE KING SNAKES.
a brown or gray in the adult. In altema (fig. 78) the pattern is of
two kinds of transverse bands in" alternation on a uniform brown
ground color, one plain black, the other split or mixed with red.
In mexicana (fig. 77) the pattern is of dorsal blotches of red, but the
shape, scutellation, and pattern of the head forbid its association
with any of the other groups.
The penial characters are as follows (fig. 2), (terms as used by Cope,
1894, 831): Organ rounded, bilobed, or forked at end; sulcus sper-
maticus single; calyces, continuous across the end of the organ or
leaving there a small bare space, apical, few, with short processes
(fringes) which may be few—5 or 6 to 10,
or numerous—15 to 20; the latter passing
into spines which increase in size gradu-
ally toward the base; no spines distinc-
tively enlarged or separated from their
fellows; basal portion of organ, below
the large spines, smooth or with numer-
ous minute spines.
The dentition may be summarized as
follows: Maxillary teeth, 12 to 20, sub-
equal, the last two sometimes a little
stouter and longer than the preceding,
not separated from the latter by an inter-
space; mandibular teeth, 12 to 18, de-
creasing in size posteriorly; palatines, 8
to 14, subequal; pterygoids, 12 to 23,
a little smaller than the palatines, and
decreasing gradually in size posteriorly.
Range.—The genus Lampropeltis is
represented throughout North America,
south of the forty-sixth parallel east of
the Roclcy Mountains and the forty-third
west, including Florida, Yucatan, and Lower California, and extends
into South America west of the Andes Mountains as far as southern
Ecuador.
Habitat and habits.—^As will appear from the descriptions of the
various forms, surprisingly little is known of the life histories of
these snakes. It is hoped that the scarcity of information on this
subject will prompt those who have the opportunity, to make accurate
observations and report them.
There is great diversity in choice of habitat among the various
forms. Some belong in the deserts (yumensis, splendida), some in
the mountains and canyons of the- west (multidncta, pyrrhomelaena,
and perhaps gentilis), some on the prairies (calligaster, holbrooki),
one (triangulum) has become adapted to the deciduous forest region
Fig. 2.—Everted penis of Lampro-
peltis CALLIGASTER. (U.S.N.M. NO.
61726). About 2J x nat. size, ca,
calyx; /r, lobe of fringe; Zi, larger
lobe; Is, large shnes; ms, minute
spines; si, SMALLER LOBE; sp, SULCUS
spermaticus.
8 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
of the northeastern States, and three have adopted burrowing habits
(elapsoides, virginiana, and rhombomaculata) . They may all be
classed as truly land snakes; so far as known none seek their food
in the water. They have not the stout body of the snakes that
depend for safety upon inconspicuousness or poison, nor the attenu-
ated body and tail of the tree snakes and racers, but rather a well-
proportioned body and short tail, adapted to life on the ground and
more or less in the open. They seem in general to be fearless and
self-reliant. Their immunity to snake venom and their powerful
constricting ability render them truly kings among the reptiles of
North America. Much remains to be learned about the food prefer-
ences of the various forms, but in general it may be said that they
are the enemies of all small rodents and of all snakes and lizards;
fledgling birds are undoubtedly eaten on opportunity.
But little is known about hibernation and reproduction. The
fullest accounts will be found under the descriptions of triangulum
and getulus. Probably all are oviparous.
Variation.—Since so little careful study has been made of variation
in scutellation and color pattern in snakes, it is unsafe to generalize
on the systematic value of any of these points without a thorough
knowledge of the particular situation. Ruthven has shown, in his
excellent review of the garter snakes (1908), the extent and meaning
of variations in the genus ThamnopMs, but a comparison with the
present review will show that not the same value can be placed in
all instances upon variation in any particular characteristic. Thus
the forms of Thamnopliis exhibit great similarity in color pattern,
while in both series of labials there is much variation of specific value.
Lampropeltis, by comparison, exhibits the widest variation in pattern,
while there is remarkably little variation in the labials. This is but
one of numerous possible illustrations that color pattern may be less
variable than scutellation. It seems usually to be more so, but, as
said above, it is not safe to generalize upon the value of characters,
for what is true in one case may be quite otherwise in another. The
relative value of characters must be determined separately for each
group of genetically related forms. By a careful study of the extent
and meaning of variations in different characters in different genera
we may learn something more definite of the genetic relations between
genera, as we are now able to learn these relations between species
and subspecies. It is a lack of knowledge of the meaning of differences
between genera that results in errors and uncertainties as to their
relations. For instance, it is not now known what meaning to place
upon differences in method of change in number of scale rows. The
present review can not, of course, determine this, but it serves to
confirm the results obtained by Ruthven in the garter snakes to the
extent of showing the method of change in number of scale rows to
REVISION OF THE KING SNAKES. 9
be constant for the genus and to be distinctively, although not greatly,
different from that in Thamnophis.
As in any study of this kind, it becomes essential to distinguish
between se^'cral kinds of variation, particularly individual, sexual,
and geographic. Since in the present study a deficiency of material
has been a great handicap, usually not more than supportive reliance
can be placed upon statistical results. It is hoped, however, that
conclusions have been sufficiently conservative to allow for the
greater quantity of material that will be available to later workers.
Variation in dorsal scale rows.—The dorsal scales are arranged in
alternate series throughout the length of body, an even number of
rows on each side of a middorsal series. The number of rows varies
in the genus from a maximum of 27 to a minimum of 15; but in no
single individual is the variation less than 2 nor more than 6 (with
the exception of a few examples of elapsoides to be presently men-
tioned). If we take an average example, say, a hoylii, and c unt
the number of rows of scales behind the neck, we will find there are
23 ; if now we trace the sixth row backward (counting upward from
the ventral plates) we will find that at some point past the middle
of the body this row is lost or fused with the next row below, and
since this occurs almost simultaneously on both sides of the body,
the number of rows changes here to 21; if now we follow the fifth
row backward, we will find that this row, toward the posterior end
of the body, is either dropped or fused with the row below it, leaving
19; we call the dorsal scale formula in this case, therefore, 23-21-19.
If we take any other form in the genus that makes the change from
23 to 21 rows, we will find that this change always takes place by a
loss of the sixth row, due either to diminution and disappearance
or to fusion with the row below. Likewise any form or individual
in the genus that makes the change from 21 to 19 rows always does
so by loss of the fifth row or by its fusion with the fourth. We may
express these facts in figures as follows
:
23-21-19
6 5
in which the upper line denotes the number of scale rows around the
body, and the figures of the lower line denote which row, counting
from the ventral plates upward, is involved in the change indicated
by the figures above it. From a study of aU the forms in the genus
it is found that the change from any particular number of rows to
the next lower number always involves the same row, counting
upward from the ventrals. Expressing these results in the same
manner as above, we get the following:
27-25-23-21-19-17-15
7 6 6 5 5 4
10 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
A most interesting uniformity is revealed. It is only in a few indi-
viduals of a single form (calligaster) that 27 rows are ever found, and
only in a few individuals of two degenerate fornls {elapsoides and
virginiana) that 15 rows occur. Just as before the presence of 15
and of 27 rows was discovered, it was expected, from the table as
then worked out, that the change from 17 to 15 rows would involve
the loss of the fourth, and the change from 27 to 25 would involve
the loss of the seventh, so now we may expect if 29 rows should ever
be found, it would be by addition of a seventh row, and that a drop
to 13 would involve the fourth. The latter might not hold, however,
for we know that some snakes have the same number of rows through-
out the length of their body, and that such generally possess a small
number of rows, as 19, 17, or 15. It is not impossible that these
forms are the end results of degenerative evolution, involving diminu-
tion in number of scale rows. This is suggested by the case of
elapsoides, the most diminutive and degenerate form in the genus.
This form has clearly been evolved from the typical forms by degen-
erative evolution. In southern Florida it reaches its greatest reduc-
tion and it is here only, out of the whole genus, that individuals have
been found which possess the same number of scale rows throughout
the length of the body. Several individuals from here have been
noted with 17 rows of scales throughout the body. It is as if extreme
reduction leads to a minimum number of rows, uniform throughout
the length of the body. That an increase from 27 to 29 rows would
involve the addition of a seventh row is indicated by the genus
ElapJie. Examination of a few individuals of E. laeta, E. guttata, and
E. ohsoleta gives the following formula for change in number of scale
rows in this genus:
29-27-25-23-21-19
7 7 6 6 5
This, it will be noticed, is the same rule as derived from Lampropeltis.
The rule as derived by Ruthven for the genus TJiamnopMs is as fol-
lows:
23-21-19-17-15
5 5 4 4
This formula is the same in kind as for Lampropeltis and Elaphe, but
the lower line of figures has been moved one place to the left. The
rule for TJia7nnopMs appears, quite according to expectation, to hold
likewise for Natrix, but as to whether the similarity in formulae
between Elaphe and Lampropeltis indicates community of descent,
we can not say. Further investigation may show what value to
place upon similarity in method of change in number of scale rows,
in establishing relationships between genera.
The value of a knowledge of variations in scale rows in a study of
relationships within the genus lies not in which row is dropped, since
REVISION OF THE KING SNAKES. 11
that is a constant for the genus, but in the kinds of formulae pos-
sessed by any single form or group of forms. No individual ever
drops more than three rows on a side, and only certain formulae of
this kind occur. Thus while any individual that possesses a maxi-
mum of 25 rows may drop three rows on each side, making the formula
25-23-21-19, in only two forms (multicincta and pyrrhomelaena) is it
common for an individual with a maximum of 23 rows to have 17
rows at the posterior end, and in only three others has such a situa-
tion been observed to occur even rarely (as in polyzona, microjyholis,
and one specimen of iriangulum) . All others having a maximum of
23 rows possess the formidae 23-21-19, 23-21, or 21-23-21-19, or
21-23-21; those having a maximum of 21 rows never have a mini-
mum lower than 17; 15 rows is reached ohly by those that never
have more than 19.
It is very noticeable that some forms in the genus (elapsoides in
particular) are undergoing, or have undergone, a reduction in number
of scale rows concomitant with reduction in bodily size and in scutel-
lation. This shov.s that evolution in Lampropdtis may involve
reduction in scutellation, and that this reduction may be one evidence
that evolution away from the primitive type has taken place. On
the other hand, there is evidence from at least one instance (-floridana)
that evolution has taken place by increase in scutellation, including
increase in number of scale rows. From these definite instances of
evolution, involving change in number of scale rows, it is shown that
the formulae may be represented in descending order as follows
:
21-19
21-19-17
19-21-19-17
19-17
17-19-17
17-19-17-15
17-15
15-17-15
In some forms, between the formulae 21-19 and 21-19-17, there
occurs the formula 19-21-19, and then 21-19-17 may occur but
rarely or not at all (the getulus group in particular). Reduction
takes place by a shortening of the rows to be lost, proceeding first
from behind forward and later, in addition, by a drop in the region
behind the neck, approximately opposite to the twentieth to thir-
tieth ventral scute; from this point the loss proceeds in both direc-
tions. Exactly a reverse in this process is the means by which a
higher form.ula is attained, and is best illustrated hj floridana. The
added row enters at some point anterior to the middle of the body,
and extends its length from here in both directions.
186550—21—Bull. 114 2
12 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
FiQ. 3.
—
Lampropeltis getulus splendida (U.S.N.M. no.
22374). 2x NAT. SIZE. Showing normal akeangement
OF HEAD PLATES PEOM THE SIDE.
In nearly all forms the influence of sex upon the scale formula is
evident. While the extremes for the form are generally attained by
both sexes, more females possess the higher formulae and more
males the lower, and in some forms the sex can be told in m.any
cases by the formula alone. Thus in multicincta and pyrrliomelaena
the formula 23-21-19-17 appears to be possessed by males only, and
in some of the forms of triangulum the formula 21-19 is possessed
in the great majority of cases by females and 21-19-17 or 19-21-
19-17 by males. The ex-
act situation for each form
is includedwith its descrip-
tion. As a general rule it
is common for the male to
have one less row of scales
near the posterior end of
the body than the female.
This is undoubtedly corre-
lated with the fact that the
body of the female has a greater diameter in this region, due to the ne-
cessity at the breeding season for containing the eggs. This is another
illustration of the fact insisted upon by Kuthven that the number
of rows of scales is correlated with the size of the body.
The rule derived by Ruth-
ven for the garter snakes
that decrease in number of
scale rows is d ' to a dwarf-
ing of the body is, in a gen-
eral way, borne out by the
present study. Rhombomac-
ulata and elapsoides are ex-
cellent examples. However,
it does not appear that de-
crease in size of body is neces-
sarily accompanied by decrease in number of scale rows, or that the
latter is evidence of a decrease in size of body. Thus holhrooJci, niger,
and getulus are undoubtedly derived from splendida and their scale
formulae are decidedly lower, but it is certainly yet to be proved
that they are smaller snakes ; furthermore, conjuncta is in all proba-
bility a derivative of hoylii, but while its scale formula is the same as
that for hoylii, it gives strong evidence of being a smaller snake. It
seems quite evident that these facts are to be explained by a geo-
graphic change in shape of the dorsal scales. In the western forms
of the getulus group the scales are more elongate, in the eastern forms
relatively shorter and broader.
Fig. 4.—Lampropeltis getulus getulus (U.S.N.M. no.
14140). 1§ X NAT. SIZE. Showing extea sxn-EALABtAL
SCUTE INTEEPOLATED BETWEEN NOEMAL SECOND AND
THIRD.
REVISION OF THE KING SNAKES. 13
FiQ. 6.- lampropeltia pyrrhomelaena. about 2 x nat.
Showing an extra labial in each series.
A study of the whole genus indicates most strongly that the
primitive scale formula is 23-21-19. This and its simple derivative,
21-23-21-19, is characteristic of those forms of the two most wide-
spread and diversified groups (triangulum and getulus) which show,
in other structural
features, the most
primitive condition.
Higher and lower for-
mulae in the genus
Lampropeltis are be-
lieved to indicate spe-
cialization, or depar-
ture from the original stock of the genus.
Variation in labials.—-Both upper and lower labial series are
remarkably constant throughout the genus. The supralabials are
normally seven (fig. 3), but in very rare instances this number is
reduced to six by a fusion
of the sixth with the fifth.
Variation to eight occurs
occasionally in many
forms and in a few
(calligaster, pyrrhomelae-
na) it is rather common.
In all cases the presence
of eight supralabials is
due to the interpolation
4, 5, and 6), or to the
Fig. 6.—Lampropeltis calligaster (Univ. Mich. no. 44973).
li X NAT. SIZE. Showing position of the added labial in
each series.
of a scute between second and third (figs
division of the second.
The infralabials are more variable. Normally there are nine
(fig. 3), but in a few forms ten is common or usual (conjuncta, pyrr-
homelacna, calligaster, Jloridana, mexi-
cana, altema), and in a few others eight
is common or usual (micropholis, rJiom-
homacvlata, elapsoides, virginiana) ; those
that commonly have eight may rarely
have seven, and those that commonly
have ten may occasionally have eleven.
Reduction from nine to eight takes
place in the great majority of cases by
reduction of the seventh and its fusion with the eighth (fig. 7); in
some instances it seems to occur by loss of the ninth. The method
of increase to ten is less constant. Since this is commonly accom-
panied by increase to eight in the upper series, and since the latter
is due to the addition of a scute in front of the eye, it is most natural
to expect that the tenth lower labial would also be inserted in front
Fig. v.—Lampropeltis elapsoides elap-
soides (U.S.N.M. no. 28910). 2 X NAT.
size. Showing reduction of lower
lablals from 9 to 8.
14 BULLETIN" 114, UNITED STATES NATIONAL MUSEUM.
of tlie eye, and beneath tlie newly added labial in the upper row.
That this is normally the case seems evident from a study of pyr-
rhomelaena and caUigaster (fig. 6), but that the extra scute is often
acquired behind the eye must also be admitted. When the latter is
the case the added scute may be due to the division of the seventh
or to the addition of one between the seventh and the eighth (fig. 5),
or to the division of the ninth, or to a shortening of the last and a
migration into the labial series of one of the gular scales. The few
specimens that possess eleven lower labials usually show that this
is due to the addition of one before the eye and one behind. It is
not improbable that when the change from nine to ten is due to the
division of the seventh that this is an accident in place of the third,
and that the next division would then be due to occur before the eye
instead of behind. We may therefore express the normal change in
the lower labials as follows
:
8-9-10-11
7 4 8
in which the upper line refers to the number of labials and the lower
to the number of the scute that is involved in the loss or gain, as the
case may be. In case of increase it might be more accurate to replace
the second row above with the figures 6, 3, and 7, to indicate which
scute is divided to increase the number, since it more often looks like
a case of division than like an interpolation.
If we express the changes in the upper labials, as we have done for
the lower, we get the following:
8-7-8
6 3
This fits the scheme for the lower labials, and the two may be com-
bined thus:
6-7-8-9-10-11
6 3 7 4 8
We may express this in the following general rule, which seems to
hold throughout the genus, subject to the exceptions explained
above: In reduction—^when the number of labials is odd, a scute is
lost behind the eye, and when the number is even, one is lost in front
of the eye; in addition—when the number of labials is odd, a scute
is added in front of the eye, and when the number is even, a scute
is added behind the eye. The two labials of each series that lie
directly beneath the eye are never concerned in any change.
The actual method of change in the number of labials is of scant
importance in determining the relationships of the forms within the
REVISION OF THE KING SNAKES. 15
genus. It is included here only on the possibility that it may be of
value in establishing relationships with other genera. The real value
of the labial count in the present study is the sure indication it gives
of the degree of specialization of the form. An}^ marked change in
the normal 7-9 formula is, in the present genus, an earmark of
specialization. We may call attention here to the following forms
that show conspicuous departure from the 7-9 arrangement: Cun-
juncta, micropholis, pyn'JiomelaeTia, Jloridana, elapsvides, virginiana,
rhombomaculata, and calligaster. The descriptions and discussions
following will make it clear that these are the most specialized forms
in the genus, and in several instances occupy, geographically, the
most isolated portions of the entire range.
Variation in other features of scutellation.—The caudals and ventrals
vary geographically, sexuall}'^, and individually. A brood of eight
young of triangulum from Indiana shows a range in ventrals of from
200 to 208; of these, 5 are females and 3 males, and their averages
are respectively 205 and 203. Tliis expresses fairly well the average
difference between the sexes in the number of ventral plates for any
form in the genus. Sometimes a large series will show an average
difference of only half a plate in favor of the females. The influence
of sex on these averages is therefore negligible when they are to be
used to show geographic differences. And as the latter undoubtedly
occur the number of ventrals proves to be an important character
in the determination of relationships.
The caudals show a more distinct relation to sex than the ventrals.
The males always average to have a higher number, sometimes only
one or two, but usually four or five, more than the females. Geo-
graphic differences in their number can be satisfactorily demonstrated
only b}^ comparing averages of like sexes. As this always reduces the
dependability of the averages by reducing the number of specimens
upon which they are based, the caudals are much less valuable in
the present study than the ventrals. In subspecific determination the
number of caudals may be of more value than the proportionate tail
length, since the latter may remain the same throughout a series of
closely allied forms, while the ventrals and caudals are increasmg or
decreasing together.
The temporals are normally two in the first row, three in the second,
and four in the third, expressed: 2+3+4 (fig. 3). Variation in some
forms is normally by increase of a scute in any row, more often in
the last row (getulus, pyrrhomelaema, calligaster), in others it is
normally by decrease {syspila, triangulum), but in only a few forms
has this formula been altered racially. In elapsoides, virginiana, and
micropholis the temporals are usually 1+2 + 3. Reduction in the
temporals takes place by a decrease in size of the upper scute in each
16 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
row and its final disappearance or its fusion with the next lower
scute. Usually the latter increases in size as the upper decreases,
but reduction in number is very evidently due to a flattening of the
head, and while the single anterior temporal is frequently larger than
either when both are present, it is generally narrower than the com-
bined width of the two, and in elapsoides the space between the upper
labials and the parietals has become decidedly narrowed (fig. 7).
The loreal plate in the great majority of the forms of the genus is dis-
tinctly longer than high and nearly quadrilateral in outline (fig. 3). It
may be absent by anomaly in any form and is commonly absent in elap-
soides, its place being taken by a downward extension of the pre-
frontal (fig. 7). In some its shape has been noticeably altered; in
getulus and hoTbrooki it is commonly nearly square or higher than
long (fig. 4); in pyrrhomelaena its length is increased and its lower
posterior angle is strongly acute; in micropJiolis its shape is dis-
tinctly atypical. The assumption is inevitable that the ancestral
type has this scute longer than high and with nearly right angles.
Variation in dentition.—Dental characters are of uncertain value
in a work of this kind. No relationships have been discovered by
their means that were not already evident from external characters.
The variability of the teeth is greater than that of the head scutes,
necessitating the examination of many skulls to determine the ex-
tremes and averages of the characteristics. The chief value in the
present study has been the confirmation derived from dental charac-
ters of the validity of the major groupings within the genus.
Variation in color pattern.—^There is no evidence of a fundamental
pattern from which all forms of the genus have been derived. On
the other hand, there are at least three styles of patterns, and, if we
include mexicana and alterna, probably two more. On the color
patterns alone the major groups in the genus can be recognized.
Two of the groups in particular {triangulum and getulus) show most
remarkable evolution in pattern. When we see the changes that may
take place in the color pattern within a single group of closely related
forms, we conclude that scant dependence can be placed upon differ-
ences and resemblances in pattern until these are supported by struc-
tural evidence. Furthermore, where structural differences are slight
or unreliable the color pattern may furnish conclusive evidence as to
relationships.
SUBDIVISIONS OF THE GENUS LAMPROPELTIS.
The many forms of king snakes fall naturally into three groups
(exclusive of the aberrant and little-known forms, alterna and mexi-
cana), recognizable on structural characters as follows:
REVISION OF THE KING SNAKES. 17
a}. Last two maxillary teeth usually not longer and stouter than those preceding;
spines of penis stout, extending one-third to one-half the distance to base of
organ.
6'. Penis bilobed or bifurcate; pattern in black, and white or yellow, in rings,
bands, or stripes, but fundamentally of a light spot on a dark scale.
getulus group.
b^. Penis scarcely bilobed; pattern of dorsal blotches of brown with black bor-
ders calligaster group.
a?. Last two maxillary teeth usually larger and stouter than those preceding; spines
of penia slender, extending half way or more than half way to base of organ;
pattern fundamentally of rings of black, red, and yellow triangulum group^
The greatest separation in the genus (still excluding altema and
mexicana) is undoubtedly between the sections a^ and a^ above.
The getulus group includes 10 very closely allied forms spread over
the southern two-thirds of the United States, Lower California, and
northern Mexico. This group is distinctly marked off by style of
color pattern from all other forms in the genus, but is closely allied
with the calligaster group in characters of skull, dentition, copulatory
organ, proportions, and scutellation.
The calligaster group includes but three forms, one of which is
known from only a single specimen. The other two are found only
east of the Rocky Mountains, one in the Mississippi Valley and the
other east and south of the Alleghenies. Both are more specialized
than any of the forms of getulus, and this together with the fact
that the group to which they belong is far less diversified than the
latter indicates that it is the result of an older radiation than the
getulus group.
In the triangulum group the anterior maxillary and particularly
the anterior dentary teeth are longer than the corresponding teeth in
the other two groups. The relative size of the last two maxillaries is
a-fairly, although not fully, distinctive feature. The penial characters
are likewise for the most part distinctive; the calyces are commonly
more numerous and have more and longer fringes, and the spines are
more slender, more numerous, and do not stop as abruptly.
The name mexicana has been applied to two specimens from San
Luis Potosi. The description will show that these can not be assigned
on structure or pattern to any of the above three groups, but, as the
genus is now defined, they must be assigned to Lampropeltis. Prac-
tically the same situation holds with respect to the specimen said to
have come from the Davis Mountains, Texas, and named by Brown,
altema. Penial characters can not be determined, since all three of
these specimens are females.
A fuller discussion of the three major groups is reserved for the
sunmiaries following the descriptions of their component forms.
The following key is expected to work for the great majority of
specimens, but difficulty may be expected from occasional individuals.
18 BULLETIN 114, UNITED STATES NATIONAL, MUSEUM.
Fuller knowledge of some forms that are now known from but few
specimens may be expected to reveal limits of variation that will
render invalid some of the distinctions now used, but this contin-
gency has been guarded against as much as possible.
KEY TO THE FORMS OF LAMPROPELTIS.
a^. Pattern of narrow cross bands of black, the alternate bands mixed or split with
red; ground color above, slate gray; head very distinct from neck. alterna, p. 247.
(Davis Mountains, Texas.)
a 2. Pattern not of narrow dorsal cross bands of black with the alternate bands mixed
or split with red; head usually only slightly distinct from neck.
6>. Color pattern without red ' and without dorsal blotches of brown or gray with
black borders getulus group.
c*. Scales chiefly black with sharply defined white or yellow spots (not light at
base shading gradually into a dark distal border); these yellow spots often
so grouped as to form 50 or more narrow cross bands on body and tail.
d^. Scale rows on middle of body 23 or 25; no light centers, dorsally, on the
scales between the cross bands; head mostly black splendida, p. 26.
(Southeastern Arizona to the ninety-seventh meridian; southern Texas,
and northern Mexico.)
d^. Scale rows on middle of body usually 21.
e ^ A yellow spot on practically all of the dorsal scales holbrooki, p. 33.
(Eastern Texas to southeastern Wyoming, east to eastern Illinois, and
south to the Gulf of Mexico.)
e 2. Scales between the dorsal cross bands without light centers or with only
a very few small ones niger, p. 43.
(Eastern Illinois to Ohio, south to central Alabama.)
c ^. Pattern in rings, cross bands, or stripes, or chiefly of scales white at base
shading gradually into a black distal border, but not chiefly of sharply
defined white or yellow spots on black scales.
/^ Posterior chin shields nearly as long and nearly as wide as anterior,
in contact or separated by not more than one small scale; pattern
neither of rings nor of longitudinal stripes.
g ^. Many dorsal cross bands of white or yellow.
h *. Cross bands less than 50; 21 (sometimes 23) rows of scales.
getulus, p. 49.
(From New Jersey to Mobile Bay and Central Florida.)
h ^. Ci'oss bands more than 50, or nearly indistinguishable; 23 (some-
times 21) rows of scales; scales between the cross bands usually
white at base .floridana, p. 62.
(Central to southern Florida.)
g ^. No dorsal cross bands distinguishable ; dorsal scales Light at base,
shading gradually into a dark distal border brooksi, p. 66.
(Extreme southern Florida.)
/^. Posterior chin shields generally much shorter and narrower than an-
terior and separated by one or two small scales; pattern of rings, or
of longitudinal stripes of white or yellowish,
i '. A dorsal longitudinal stripe, complete or interrupted.
j ^ Dorsal stripe white or yellow, sharply defined on a dark brown
or black ground color califomiae, p. 94.
(Fresno County, California to northern Lower California.)
1 The red fades to whitish in alcohol, but it is suflBcient, for the purpose of the key, to determine that
the pattern is in two colors instead of in three.
REVISION OF THE KING SNAKES. 19
j 2. Dorsal stripe poorly defined, of light brown or cinnamon on a
dark brown ground color; bellj'" uniform hxo'wn.. nilida, p. 103.
(Southern Lower California.)
i 2. Pattern of rings.
k . White ^ scales white to their bases, forming rings of uniform
white hoylii, p. 75.
(California, Nevada, southwestern Utah, northern and
western Arizona, and northern Lower California.)
k 2. White ^ scales mostlj' brown at their bases.
I '. White bars on prefrontals occupying less than half the
area of these scutes; frontal plate uniform black, or
with the white restricted to a narrow transverse bar at
its anterior end; no white on parietals; infralabials
usually 9 yumensis, p. 66.
(Southern Arizona, extreme southeastern California,
northeastern Lower California, and northwestern
Sonora.)
I 2. White bars on prefrontals occupying more than half the
area of these plates; frontal plate with prominent white
markings, or at least with a central spot of white; each
parietal with one or more white spots; infralabials
usually 10 conjuncla, p. 89.
(Southern Lower California.)
h 2. Pattern with red, or with dorsal blotches of brown, gray, or red, with black
borders.
m '. Pattern of black-edged dorsal blotches of brownish or
dark red, only narrowly in contact with the fifth row
of scales or extending no lov/er than the sixth or
seventh rows calligaster group.
n '. Blotches less than 40 leonis, p. 138.
(Nuevo Leon, Mexico.)
n 2. Blotches 45 to 80.
*. Scale rows 25 to 27 on middle of body; dorsal
blotches with concave anterior and posterior
margins; infralabials 9 or 10, rarely 8.
calligaster, p. 115.
(Western Texas to ^Mississippi, north to Indi-
ana) and northwest to Minnesota, thence south
to Texas.)
o 2. Scale rows 23 or 21 on middle of body; dorsal
blotches with straight or convex anterior and
posterior margins; infralabials 8, less often
9 rhombomaculata, p. 128.
(Mobile, Alabama, to Knoxville, Tennessee,
north to Maryland, south to Central Florida.)
m 2. Pattern in rings; or, if in blotches or saddles of brown,
gray, or red, these broadly in contact with the fifth
or a lower row of scales.
p '. Whitish cross bands on bodj' and tail less than
40; or, if more than 40, the snout not uni-
formly whitish.
' Specimens may be found which can be accurately identifled only by locality; in particular it should
be noted that young examples of yumensis and conjuncla may resemble boylii. See table, p. 77, com-
paring these three forms.
2 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
q ^ Whitish cross bands little if any widened on
the lower rows of dorsal scales, and the scale
rows more than 17 on the anterior portion of
the body,
r^ Red scales usually tipped with black,
s^. Snout black, with usually a light transverse
bar on or near the prefrontals; two tem-
porals in the first row; caudals usually
not less than 49 polyzona, p. 139
.
(Southern Mexico to Costa Rica.)
s^. Snout yellowish with transverse black spots;
a single, anterior temporal, or, if two, the
upper much the smaller; caudals not more
than -19 micropholis, p. 149
(Panama to Colombia and Ecuador.)
r*. Red scales not tipped with black.
i'. Whitish annuli usually more than 30;
snout black.
u^. Ventrals more than 200; black rings
often meeting across the red dor-
sally multicincta, p. 222.
(California.)
ii^. Ventrals less than 200; black rings not
meeting across the red dorsally.
ruthveni, p. 221.
(Southern Mexico.)
<'*. Whitish annuli less than 30.
v^. Dorsal red areas usually continuous
across the belly; snout whitish,
specked with black.
w^. Ventrals usually more than 210
(199 to 231) nelsoni, p. 155.
(Western Mexico.)
iv^. Ventrals usually less than 200,
(180 to 212) amaura, p. 172.
(Lower Mississippi Valley.)
v^. Spaces on belly between the yellow
rings filled with black; snout
totally black, or only very sKghtly
lightened on the top or sides.
x^. Yellowish rings 19 to 25; black
spaces on belly usually longer
than the intervening yellow
ones annulata, p. 159,
(Plateau region of southern
Mexico north to extreme south-
em Texas.)
x^. Yellow rings 25 to 40; black
spaces on belly usually shorter
than the intervening yellow
ones gentilis, p. 165
.
(Texas toSouthDakota, west
to Utah and Arizona.)
REVISION OF THE KING SNAKES. 21
a*. Dorsal whitish bands usually distinctly widened
on the first row of scales, or the scale rows
anteriorly not more than 17.
y^ Black practically uniform over
the head, except for the
snout region, which is more
or less lightened, at least on
the sides; scale rows an-
teriorly more than 17.
z'. Whitish annuli or cross
bands 25 to 40; black
often strongly encroach-
ing upon the red on the
middorsal line.
gentilis, p. 165.
z'. Whitish annuli or cross
bands 18 to 25; black show-
ing not more than a slight
tendency to encroach up-
on the red areas on the
middorsal line.
aviaura, p. 172.
(liower Mississippi Val-
ley.)
y*. Black of head practically re-
stricted to posterior portion,
or to various black-edged
light markings.
na^. Usually a single anterior
temporal; scale form-
ula usually 17-19-17,
rarely higher than 19-
17.
66'. Red areas continuous
across the belly.
elapsoides, p. 206.
(North Carolina
and Kentucky,south
to New Orleans, and
throughout Florida.)
66^. Red not continuous
across the belly, but
restricted to black-
bordered dorsal sad-
dles that extend up-
on the ventrals.
virginiana, p. 217
,
(Northern North
Carolina to Dele-
ware.)
22 BULLETIN 114, UNITED STATES NATIONAL MUSEUM,
Usually two anterior tem-
porals; scale formula
very rarely lower than
19-21-19-17.
cd. Whitish annxili or
cross bands 18 to
23; pattern of body
practically in rings.
ainaura, Tp. 172.
(Lower Missis-
sippi Valley.)
cc^. Whitish across bands
23 to 60; pattern of
dorsal saddles or
blotches of red or
brown.
dd'^. Infralabials 8 or 9,
rarely 10; tail
less than 16 per
cent of total
length; often a
dark band from
eye to angle of
month.
ee^. Dorsal saddles 35
to 60, reaching
down to the
fifth or third
row of scales;
often two rows
of lateral alter-
nating blotch-
es; a dark band
on posterior
portion of pre-
frontals; a
black-border-
ed light band
from the eye to
the angle of the
mouth; usually
a Y-shaped
light spot on
the back of the
head.
trian g ulum,
p. 188.
(Eas tern
United States
and southern
Canada.)
REVISION OF THE KING SNAKES. 23
ee''. Dores^l saddles 23
to 35, extend-
ing down to
the third row
of scales, or
lower; only
one series of
alter n a t i n g
spots; head
markings of iri-
angulum only
partially or not
at all devel-
oped.
syspila,
-p. 179.
(Southern
Indiana to
Minn e s o t a
,
south to cen-
tral Arkansas
and west to
central Kan-
sas.)
dd^. Infralaljials 10;
tail more than
.16 of total
length; a dark
l)lotch ])ehind
the eye.
VI exicana, J).245.
(San Luis Po-
tosi, Mexico.)
p'^. Whitish cross bands on body and tail more than
40; top of head black, snout uniformly white.
pyrrhomelaena, p. 231.
(Utah, Arizona, Western New Mexico, and
northern Mexico.
24 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
SUMMARY OF STRUCTURAL CHARACTERISTICS OF THE FORMS OF
LAMPROPELTIS.
Name. Ventrals Caudals.
Su-
pra-
labi-
als.
Infralabials.
Tem-
po-
rals.
Tail
divided
by total
length.
Pattern.
Num-
ber
of
bands
or
spots.
alterna
amaura
annulata .
.
boylii
brooksi ....
californiae.
caUigdster.
conjuncta .
elapsoides
floridana.
.
gentilis
getulus ....
holbrooki.
leonis
mexicana .
micropholis .
muUicincta
nelsoni
niger
nitida
polyzona
pyrrhomelaena. .
.
rhombomaculata.
ruthveni
splendida
syspila
triangulum
yumensis
virginiana
213
180-212
197-212
206-254
Similar
229-241
196-215
228-240
152-193
210-225
176-212
203-224
200-220
200
193-199
211-228
202-222
199-231
199-216
227
208-239
216-235
191-213
190
207-225
180-215
180-213
212-240
173-188
60
39-51
40-57
41-62
to floridana
47-60
38-57
48-54
32-48
42-55
41 (31)-53
38-58
38-55
50
55
40-49
45-61
42-59
41-53
56
42-61
61-79
31-55
50
43-56
40-54
29-54
44-57
36-44
7
7
7(8)
7(8)
7-8
7(8)
7
7(8)
7(8)
7(6-8)
7(6-8)
7
7
7(8)
7(8)
7(8)
7
7(8)
7-8
7
7-8
7(8)
7
7(8)
7(8)
7
10-11
I (8-10)
9(10)
9-10
3-1-4 0.170
2+3 .118-0.156
2+3 .123- .167
2+3 .107- .145
(8) 9-10
9-10
10 (9-11)
8 (7-9)
9-10
9 (8)-10
9 (8-10)
9 (8-10)
10
&-8 (10)
9 (8-10)
9(10)
9 (8-10)
2+3
2+3
2+3
1+2
2+3
2+3
2+3
2+3
2+3
2+3
[1+2
to
!2+3
2+3
2+3
2+3
9 (8-10)
9-10 (11-12)
8-9
9
9 (10-11)
9 (8-10)
9 (8-10)
9-10
7-9
2+3
2+3
2+3
2+3
2+3
2+3
2+3
2+3
1+2
Alternatingbands
Rings 118-26
do 19-26
do 2»-49
.113- .140
.110- .150
.115- .132
.118- .169
.107- .146
.115- .156
.100- .1.55
.096- .150
.16- .17
.112- .136
.131- .161
.120- .150
.110- .146
Stripes
Blotches
Rings
do
Cross bands.
Rings
Cross bands
.
do
Blotches
do
Rings
.124- .104
.153- .182
.100- .150
....do
....do
Narrow bands
Stripes
Rings
.100- .150
.114- .157
.100- .155
.090- .140
.129- .145
Blotches.-'.
Rings
Light crossbands.
Blotches
....do
Rings
Blotches
46-78
30-40
15-25
46-85
25-40
23-52
50-100
27
39
13-21
23-57
13-24
50-90
17-37
35-71
48-64
30
41-85
23-36
28-62
29-45
18-27
REVISIOlSr OF THE KING SNAKES. 25
Summary of structural characteristics of the forms of Lampropeltis—Continued.
Scale rows. Dentition. Total
num-
Name.
Extremes. Average. Maxillary.
Mandibu-
lar.
Palatine. Ptery-
goid.
ber of
speci-
mens
exam-
ined.
25-23-25-23-21.
21-23-21-19 to
19-21-19-17.
23-21-19-21 to
21-20-21-19.
23-25-23-21-19
to 21-23-21-
19.
Similar to flor-
idana.
23-25-23-21-19
to 21-23-21-
19.
25-27-25-23-21
to 21-23-21-
19.
23-25-23-21-19
to 21-23-21-
19.
19-21-19-17 to
15-17-15.
23-21 to 21-19.
23-21 to 19-17.
23-21 to 19-21-
19-17.
21-23-21-19 to
19-17.
13 14-15 12 1
21-19 27
21-19 13-15
13-14
13-15
14-15
9-11
8-10
15-20
13-17
12
23-21-19 168
californuie 23-21-19
23-2a-23-21-19.
23-21-19
17-19-17
21-23-21-19....
21-19
13-15
12-14
12-15
12-14
15
12-14
13(13-16)
13-14
14-15
13-14
12-16
12-14
16
12-16
14-15(17)
14-15(16)
9-10(11)
9-11
8-11
10-12
10
10-11
9(10)
9(8)
16-18
12-19
15-18
17-23
17-19
16-20
16-20
13-18
27
calligcuter 99
13
88
25
46
21-19 or 21-23-
21-19.
21-19 or 19-21-
19.
23
134
132
23-21-19
21-23-21-19 to
19-21-19-17.
23-2.5-23-21-19
to 21-19-17.
21-23-21-19 to
19-21-19-17.
21-23-21-19 to
19-21-19-17.
23
23-21-19
19-21-19-17....
21-23-21-19 or
21-19-17.
21-19-17
19-21-19
13
15-14
11-13
12-13
13
14-17
15
12-14
13-15
14-13
13
10-12
9-11
10-11
9
22
17-19
15-18
18-22
16-17
2
micropholis 24
49
15
33
21-23-21-19 to
19-17.
23-25-23-21-19
to 21-23-21-
19-17.
21-23-21-19 to
19-21-19-17.
21-23-21-19
21or23
23-21-19
23 or 21
13-15
13-20
12-13(12-15)
14-15
13-14(12)
11-13
12-13
13
13-16
13-15
14-18
12(12-16)
13-14
15
11-14
13(11-15)
13-15
14-17
10-12
11-14
9-10
11-13
9
10-12
9-11
10-11
9-10
16-22
16-22
13-16
21
15-19
17-20
17-23
19-22
12-19
48
pyrrhomelaena
rhombomaculata
33
58
1
23-25-23-21-19
to 21-23-21-
19.
21-23-21-19 to
19-17.
21-23-21-19 to
19-17.
17-19-17 to 19-
17.
23-25-23-21-19
to 21-23-21-
19.
21-23-21-19....
21-19
24
tytpila 02
21-19-17
17-19-17
23-21-19
404
virginiana 7
yutnemU 18
1,581
26 BULLETII^ 114, TJNITED STATES NATIOITAL. MUSEUM.
THE GETULUS GROUP.
LAMPROPELTIS GETULUS SPLENDIDA (Baird and Girard).
KING SNAKE.
Figs. 1, 3, 29, 30, 31.
1853. Ophibolus splendidus Baird and Girard, Cat. N. Amer. Rept., pt. 1, p. 83
(type locality, Sonora, Mexico; type specimen, U.S.N.M., no. 1726;
J. D. Graham, collector).
—
^Baird, U. S. and Mex. Bound. Surv., 1859,
p. 20, pi. 14; Pacif. R. R. Surv., vol. 10, pt. 3, art. 1, 1859, pi. 30, fig. 58;
vol. 10, pt. 6, a,rt. 4, 1859, p. 43.—Strecker, Baylor Univ. Bull., vol. 18,
no. 4, 1915, p. 39.
—
Lampropeltis splendida Cope, Proc. Acad. Nat. Sci.
Philadelphia, 1860, p. 255.
—
Van Denbtjrgh, Proc. California Acad.
Sci., ser. 2, vol. 6, 1896, p. 347.
—
Stejneger and Barbour, Check List,
1917, p. 89.
—
Coronella splendida Jan, Ai-ch. Zool. Anat., vol. 2, fasc. 2,
1863, pp. 238, 245.
1853. Ophibolus sayi Baird and Girard, Cat. N. Amer. Rept., p. 85 (Red. R.,
Arkansas), p. 159 (Eagle Pass, Tex.).
—
Marcy, Explor. Red R., p. 199,
1854, pi. 7.—Cope, Proc. U. S. Nat. Mus., 1888, p. 398 (San Diego, Texas),
1875. Ophibolus getulu^ splendidus Cope, Bull. U. S. Nat. Mus., no. 1, p. 37.
—
CouES, Rep. Geog. Geol. Explor. Sm-v. W. 100th Mer., vol. 5, chap. 5,
1875, p. 619.—Yarrow, Bull. U. S. Nat. Mus., no. 24, 1882, p. 93.—
GARMan, Mem. Mus. Comp. Zool., vol. 8, no. 3, pt. 1, 1883, p. 157.
Cope, Proc. U. S. Nat. Mus., vol. 14, 1891, p. 613; Rep. U. S. Nat. Mus.
for 1898, 1900, p. 918.
—
Lampropeltis getulus splendidus Wright, Proc.
Acad. Nat. Sci., Philadelphia, 1915. p. 148.
1901. Ophibolus getulus sayi Brown, Proc. Acad. Nat. Sci. Philadelphia, p. 77;
same, 1903, p. 550.
Description.—^This form is nearest to the ancestral stock of the
getulus group. Although represented by relatively few specimens,
these are well distributed over its range, thus presenting a fair
conception of the species. The scalation may be summarized as
follows: Ventral plates, 207 to 225; caudals, 43 to 56; supralabials, 7,
occasionally 8; infralabials, 9, sometimes 10, rarely 11; oculars,
1 and 2; temporals, 2+3+4; posterior chin shields shorter than
anterior, parallel, and separated from each other by one or two
small scales; loreal about as high as long, or longer than high; dorsal
scale rows usually 23-21-19, or 21-23-21-19, although 23-25-
23-21-19 is not uncommon in the western part of its range.
The proportions are the same as for the other forms of the getulus
group, namely: body cylindrical; sides meeting the belly at a
rounded angle; head scarcely distinct from neck; body tapering
slightly toward the head and toward the tail; tail short, tapering
to a homy tip, varying from 0.100 to 0.150 of the total length (males
average, 0.130; females, 0.120). Adults are commonly from 75 to
115 cm. in length. The largest examined was from Bexar County,
Texas, and measured 1,432 millimeters.
The color pattern is, briefly: Black, spotted on the sides with
white or yellow, one spot on each scale; across the back about 70
REVISION OF THE KING SNAKES. 27
(41 to 85) narrow dotted cross bands of white or yellow separated
b}^ unspotted areas; head, neck, and under parts mostly black.
The pattern is most typically developed in southwestern New
Mexico and northern Chihauhua. Specimens from this region
may be described as follows: Head, from between the eyes to about
a dozen dorsal scales beliind the parietals, black; neck and throat
black, except for a white spot, more or less developed, lengthwise
of each of the labial and chin shields and anterior gulars, and a
median light band on the anterior ventrals. On the frontal plate
two narrow transverse white spots anteriorly. On each prefrontal
and internasal, anteriorly, a similar white transverse band. Rostral
black at center and along posterior border. Nasals, loreals, pre-
oculars, and sometimes oculars with a central development of white.
Belly and caudals mostly black, with a development of white
on the ends of each alternate ventral plate and sometimes a median
wliite spot on ventrals that alternate with those bearing lateral
spots. Dorsal scales each with an oval white center, oriented length-
wise of the scale. These white centers occupy the most of the
scale on the first row and decrease in size dorsally, leaving three or
four rows of middorsal scales entirely black, except where the white
centers are continued across the back to form transverse white
bands, from two to four scales apart, throughout the length of the
bod3^ In distinction from holhrooTci the orientation of most of the
white centers of the cross bands is lengthwise of the scales and the
spaces between the bands are entirely unspotted, except rarely those
close behind the neck.
The penial characters as indicated by specimens from New Mexico
are as follows: distinctly bilobed; sulcus single, extending over the
side of the larger lobe, and ending in a small bare space, surrounded
by a few calyces; the latter extending far enough beyond the end of
the organ to be very evident in a lateral view; calyces with five to
ten fringes, which may be conspicuous or very short; latter passing
into spines which increase gradually in size to about one-third the
distance from the apex, here being replaced rather suddenly by a few
minute spines; remainder of organ smooth.
The dentition, as derived from examination of a few specimens,
is as follows: Maxillary teeth, 12 to 14, subequal, the last two scarcely
stouter; mandibulars, 15, third to sixth largest, decreasing posteriorly,
the last very small; palatines, 9, subequal, but slightly smaller than
the maxillaries; pterygoids, 15 to 19, subequal, smaller than the
palatines, decreasing a little in size posteriorly.
To distinguish splendida from Jiolhrooki and from yumensis, see
under these respective forms.
186550—21—Bull. 114 3
28 BULLETIN 114, U:N"ITED STATES NATIONAL MUSEUM.
Habitat and JiaUts.—^The o^ly reference that we have been able to
find concerning the natural history of this form is by Van Denburgh
(1897, 347): "Two specimens of this handsome snake were taken,
one of which was 'shot in a tree in a river bottom near Fort Lowell
(Tucson, Ariz.) May 28, 1893.'"
Range.—^The range of splendida extends east to about the 97th
meridian; northward to southern Oklahoma, and west as far as
Tucson. It is probable that it will be found of general distribution
in northern Mexico, although the only definite record for south of the
international boundary is for San Diego, Chihuahua. In New
Fig. 8.
—
Map showing locality eecoeds fob Lampeopeltis getulus splendida.
Mexico it doubtless extends considerably north of Fort Fillmore,
which is now the most northern record for that state.
The only published record for a locality not included in the list of
specimens examined is one for Eagle Pass, Maverick County, Texas
(Baird and Girard, 1853, 159). This specimen is listed as a variety
of OpJiibolus sayi, but as it occurs well within the rafige of splendida^
it is undoubtedly this species.
Variation.—Throughout the central portion of its range splendida
is a distinct and well-defined form, but at its east and west limits it
intergrades with holirooJci and yumensis, respectively. Only 23
specimens have been examined from the entire range, but these are
KEVISION OF THE KING SNAKES. 29
well distributed and indicate certain points which a fuller series may
confidently be expected to confirm. Thus the number of scale rows
is higher in the central and western portion of the range than in the
east. No specimen showed a lower maximum than 23 rows, but in
some of the specimens from central Texas this number was retained
for only a very brief space near the middle of the body, while in the
west it was present for a much greater distance, and some had 25
The table of scale rows brings this out to some extent.rows.
Approach to liolbrooki is shown also in the penial characters. That
form is characterized by having many minute spines below the large
ones, and this character is nearly as well developed in specimens of
splendida from San Diego, and Bexar County, Texas, as in typical
examples of Tiolbrooki; a specimen from Keeves County, in western
Texas, shows the minute spines present over nearly as great an area
as that occupied by the large ones, but they are only scarcely dis-
cernible, while specimens from Tucson, Arizona, show minute spines
present for only two or three millimeters below the large ones, thus
being close to yumensis in this respect. The ventrals and labials
seem to be much the same in one part of the range as in another.
Table of scale rows in splendida.
Formula.
Arizona. New Mexico andWestern Texas. Central Texas. Totals.
Male. Female. Male. Female. Male. Female. Male. Female.
23-25-23-21-19 2
1
1
1
3
2
4
4
1
5
5
23-21-19 1
1
2
2
1
1
1
421-23-21-19
Total 4 2 4 4 2 5 10 11
The most noticeable geographic variation occurs in the color pat-
tern, the full significance of which will be brought out in the discus-
sion of the evolution of the color pattern in the getulus group. Briefly
it is as follows: Westward from the central portion of the range the
number of dorsal cross bands is decreased perceptibly; they become
broader, the dorsal black areas become wider, and, on the sides,
opposite each dorsal white band, there is a fading out of the white
spots, tending to produce a series of lateral dark areas in alternation
with the dorsal series. Eastward from this central region, the wliite
centers of the cross bands tend to lose their symmetrical orientation
parallel to the long axes of the scales; the number of bands decreases
somewhat; the dorsal dark areas wdden very slightly; there is a dis-
tinct tendency for the development of a lateral alternating series of
dark areas, as in the western forms, sometimes also a second and
30 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
lower series alternating with the last, and extreme individuals begin
the development of white centers in the dorsal dark areas; the pro-
portion of white on the belly is increased, and near its eastern limits,
the head may become as speckled with white as a true Jiolhrooki.
Affinities.—^There can be no doubt, even from this brief summary,
that splendida bridges the gap between Tiolbrooki and yumensis, in
structural and color pattern features as well as in geographic position.
We will leave a fuller discussion of its relationships with these forms
to the section on the evolution of the group, and here offer only the
following diagram of affinities:
yumensis splendida liolbrooki.
REVISION OF THE KING SNAKES. 31
.9K
•spacq ssojo jo jeq
-nmu ejBtnpcojddv
So r- CO£ t« CO
papiAtp q:>3u9i iJ^Ji
m 00 cc ^ o «o
-inn ni q;3ii9i iB?Oi
00 CO o>c^
>0 OJ U5 CO U3 r-l CO
st^iodniex
^t< ^^ U5-* •^ u^ -^ -^ lO -^
+ + + + ++++++
CO CO COCO CO CO CO CO CO CO
+ + ++ ++++++
w M (NN CJMWC^^NM
++++ + ++++
CCCOtNCO CO COtTCOCO
++++ + +++ +
s i 2
CO
+
CO
+
CI
+ +
CO CO
+ +
—
y_• ^^ v-v-^ »
CSl (N WIN N N W N « MN N M C4 M c^ M M (M
•sicpioo '^ ^ -H^ '^ -*
__^
'^ -* ^^
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REVISION OF THE KING SNAKES. 33
LAMPROPELTIS GETULUS HOLBROOKI (Stejneger).
SPECKLED KING SNAKE, SAY'S KING SNAKE, KING SNAKE, GUINEA SNAKE,
SPOTTED KING SNAKE.
Fig. 32.
1842. Coronella sayi Holbrook, N. Amer. Herp., ed. 2, vol. 3, p. 99, pi. 22.
—
DuMERiL and Bibron, Erp. Gen., vol. 7, pt. 1, 1854, p. 619.
—
Lichten-
STEiN, Nomenclator, Mus. Zool. Berolineneis, 1856, p, 25, ("Mexico"!).
GuNTHER, Cat. Colubr. Snakes Brit. Mus., 1858, p. 41.
—
Jan, Icon.
Gen. Ophid., livr. 14, 1861, fig. 2, pi. 5; Arch. Zool. Anat., vol. 2, fasc. 2,
1863, pp. 238, 2Ah.—Coluber sayi De Kay, Nat. Hist. New York, pt. 3,
1842, p. 41.
—
Ophibolus sayi Baird and Girard, Cat. N. Amer. Rept.,
pt. 1, 1853, p. 84.—Baird, U. S. and Mex. Bound. Surv., 1859, p. 20
(Indianola, Texas); Pacif. R. R. Surv., vol. 10, pt. 3, no. 1, 1859, pi.
30, fig. 59.
—
Humphreys, Amer, Nat., vol. 15, 1881, p. 561.
—
Lainpro-
peltis sayi Cope, Proc. Acad. Nat. Sci. Philadelphia, 1860, p. 254.
Garman, S., Bull. Essex Inst., vol. 24, 1892, p. 10.—Beyer, Proc.
Louisiana Soc. Nat., (1897-1899), 1900, p. 14.
1875. Ophiboliis getulus sayi Cope, Bull. U. S, Nat. Mus., no. 1, p. 37.
—
Coues
and Yarrow, Bull. U. S. Geol. Geog. Surv. Terr., vol. 4, art. 11, 1878,
p. 269.—Cope, Bull. U. S. Nat. Mus., no. 17, 1880, pp. 23, 44.—Yarrow,
Bull. U. S. Nat. Mus., no. 24, 1882, p. 93—Garman, S., Mem. Mus.
Comp. Zool., vol. 8, no. 3, pt. 1, 1883, pp. 68, 156, pi. 5, fig. 4.—Davis
and Rice, Bull. Chicago Acad. Sci., vol. 1, no. 3, 1883, p. 29 (Wiscon-
sin).
—
Hay, 36th Annual Rep. Indiana State Board Agri. for 1886, vol.
28, 1887, p. 210; 17th Annual Rep. Dept. Geol. Nat. Res. Indiana, 1892,
p. 518.
—
Garman, H., Bull. Illinois State Lab. Nat. Hist., vol. 3, art.
13, 1892, p. 297.—Cope, Proc. Acad. Nat. Sci. Philadelphia, 1893, pp. 385,
387.—Hurter, Trans. Acad. Sci. St. Louis, vol. 6, no. 2, 1893, p. 255.—
Rhoades, Proc. Acad. Nat. Sci. Philadelphia, 1895, p. 392.—Cope,
Rep. U. S. Nat. Mus. for 1898, 1900, p. 911, fig. 226.—Strecker, Trans.
Texas Acad. Sci. for 1901, vol. 4, pt. 2, no. 5, 1902, p. 4.—Brown, Proc.
Acad. Nat. Sci. Philadelplua, 1901, p. 76; 1903, p. 550.—Branson,
Kansas Univ. Sci. Bull., vol. 2, no. 13, 1904, p. 397.—Ditmars, Reptile
Book, 1907, pp. 341, 358, pi. 107 (lower fig.); Reptiles of the World,
1910, p. 271.—VioscA, 6th Bien. Rep. Bd. Curators Louisiana State
Mus., 1918, p. 72.
—
Lampropeltis getulus sayi Stone, Proc. Acad. Nat.
Sci. Philadelphia, 1903, p. 542 (Sugar Loaf Mt., Oklahoma). Ruthven,
Proc. Iowa Acad. Sci., vol. 19, 1912, p. 207.
—
Wright, Proc. Acad. Nat.
Sci. Philadelphia, 1915, p. 148.
—
Coronella getula, var. sayi Boettger,
Kat. Rept.-Samml. Mus. Frankfurt, pt. 3, 1898, p. 71.
—
Lampropeltis
getula sayi Strecker, Proc. Biol. Soc. Washington, vol. 21, 1908, pp,
75, 89.
1880. Ophibolus getulus getulus Cope, Bull. U. S, Nat. Mus., no. 17, p. 23.
1894. Coronella getula (part )Boulenger, Cat. Snakes Brit. Mus., vol. 2, p. 198.
Johnson and Johnson, Reptile Life, 1907, fig. p. 64.
1903. Lampropeltis holbrooki Stejneger, Proc. U. S. Nat. Mus., vol. 25, p. 152
{=Coronella sayi Holbrook; tj^e locality, valley of the Mississippi).
Stejneger and Barbour, Check List, 1917, p. 188.
1905. Lampropeltis getula holbrooki Bailey, N, Amer. Fauna, no. 25, p. 47.
Lampropeltis getulus holbrooki Hurter and Strecker, Trans. Acad. Sci.
St. Louis, vol. 18, no. 2, 1909, p. 26.—Hurter, Trans. Acad. Sci. St,
Louis, vol. 20, no. 5, 1911, p. 185.
—
Strecker, Baylor Univ. Bull,
vol. 18, no. 4, 1915, p. 38.
34 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
1909. Lampropeltis getula holbrooMi Strecker, Baylor Univ. Bull., vol. 12, no.
1, 1909, p. 7 (Burnet, Texas).
1912. Ophiholus getulus, var. sayi Somes, Proc. Iowa Acad. Sci., 1912, p. 150.
As Stejneger has already pointed out (1902, 152), the name Coro-
nella sayi was originally used for this form by Holbrook under the
misapprehension that it was the species previously described by
Schlegel as Coluber sayi. As the latter is a very different snake,
PituopJhis sayi, the name sayi can not be applied to this form.
Description.—Holhroolci is nearly as well known as its represent-
ative on the Atlantic Coast. The scutellation as derived from about
132 specimens, is as follows: Ventral plates, 200 to 220; caudals, 38
to 55 (one specimen, 30); supralabials, 7, rarely 6 or 8; infralabials,
9, rarely 8 or 10; oculars, 1 and 2; temporals, 2+3+4, occasionally
varying by one in any row; posterior chin shields usually a little
shorter and a little narrower than the anterior, parallel, and generally
separated by one or two small scales; loreal usually as high as long
or higher than long, although sometimes longer than high; dorsal
scale rows commonly 21-19 or 19-21-19, occasionally as high as
21-23-21-19, and often as low as 19-21-19-17, very rarely 19-17.
The general proportions are the same as with splendida and getulus,
that is, head but slightly distinct from neck, belly meeting the sides
in a rounded angle; body cylindrical and of nearly the same diameter
except for tapering a little toward the neck and toward the tail,
latter short, tapering quickly to a homy tip, varying from 0.096 to
0.150 of the total length, the average for males 0.130, for females
0.120. The great majority of adult specimens measure between 75
and 95 cm.; the largest examined was from Galveston, Texas, and
measured 1,634 mm.
The general color above is blue-black with an oval white or yellow
or greenish-yellow spot near the center of each scale. At regular
intervals the white centers lose their normal orientation with the long
axes of the scales and become so grouped and oriented as to form
from 50 to 100 distinct, short, transverse bands on the back, which
are commonly less than half a scale in width and usually form a
dotted rather than continuous white band. Ventrally the white
centers increase in size so that the scales of the first row may be
described as white with black borders. The belly is checked with
black and white, the white usually predominating. The head like
the body is spotted with white. Transverse bars of white cross the
internasals, prefrontals, and supraoculars. There is a light spot on
each of the loreals, nasals, oculars, and temporals. The common
borders of the labials are black.
The copulatory organ may be described as follows: Bilobed or
slightly forked; sulcus single, extending over the side of the larger
lobe; calyces fairly numerous, apical, surrounding a small bare place
REVISION OF THE KING SNAKES. 35
at the tip; fringes of calyces few, fairly prominent, passing into
spines; latter extending a little less than half way to the base; not
particularly numerous or thick set (although apparently so if the
organ is not fully everted); basal portion with many conspicuous
but minute spines, extending downward for a distance about equal to
that covered by the major spines. This latter character appears to be
restricted to this form and niger. It is conspicuously well developed
in specimens examined from Missouri, Arkansas, and Louisiana, and
specimens from the following localities have these minute spines
developed over as much area but often more or less imbedded in the
tissue, and therefore less conspicuous: Galveston, Texas; Mobile,
Alabama; Kansas; Grinnell, Iowa; Reelfoot Lake, Tennessee; Kem-
per County, Mississippi. It is thus a characteristic of this form
throughout its range.
The dentition is as follows: Maxillary teeth usually 13, sometimes
14, the first very small, the rest large and subequal; mandibular
teeth, 14 or 15, rarely 16, the first very small, the next seven or eight
the largest, the remainder decreasing gradually to about the size of
the first; palatines usually 9, sometimes 8, subequal and about the
size of the anterior maxillaries; pterygoids, 13 to 18, usually 16,
smaller than the palatines and subsequal.
Intergradation between this form and splendida undoubtedly takes
place in Texas in the region of the ninety-seventh or ninety-eighth
meridians. It is impossible in some instances to refer a specimen
definitely to one form or the other. Distinctions which usually hold
are as follows: The head of holbrodki has many small yellow spots,
that of splendida is mostly black, with but few spots; in TiolhrooM the
dorsal spaces between the cross bands have a yellow spot on each
scale in adults (in young, few or none), in splendida there are no
spots here, or but few; JiolbrooH usually has no more than 21 rovs of
scales, while splendida has 23. To distinguish from niger and from
getulus see under these forms.
Habitat and habits.—Very Httle has been recorded on the natural
history of this form. Hurter (1911, 185, 255) states for Missouri
that it inhabits hilly places with sunny glades, occurring under rocks
and fallen trees. On the other hand, Mr. Percy Viosca informs the
writer that in Louisiana it is found in moist places throughout the
entire alluvial section of the State. Mr. H. P. Loding says that near
Mobile, ^Vlabama, they are turned up by the plow in the spring.
Like getuJus it is reported to be the enemy of the rattlesnake and
the moccasin. Coues (1878, 289) mentions this, and Humphreys
(1881, 561-2) gives a rather elaborate account of how one swallowed
a water moccasin (Ancistrodon piscivorous). Branson (1904, 397-8)
states it to be the enemy of all other snakes.
36 BTJLLLiTIN 114, UNITED STATES IsTATIOFAL MUSEUM.
Mr. Mackelden of St. Louis writes that a large specimen of this
snake from St. Louis County, Missouri, which he kept in captivity
for five months, ate 21 Bascanion constrictors, some of which were
from a foot to 18 inches longer than the king snake and required
sometimes three days to swallow. It was often tempted with garter
and water snakes, but seemed to have an aversion to them as food.
Hurter (1911, 185) says that its food consists of mice, small birds,
lizards, and snakes. It seems to be much less amenable to life in
captivity than getulus. Most specimens kept by the writer refused
to eat, and long continued to vibrate the tail in nervousness or fear
whenever handled. Some individuals would always assume the
striking position whenever anyone approached the cage. Such as
could be induced to feed were observed to eat TJiamnopJiis hutleri
T. sirtalis, and dead mice. They constricted their prey exactly as
does getulus.
There are no records of breeding observations.
Range.—This form ranges from Mobile, Alabama, west to about
the ninety-seventh meridian in Texas. It extends north in the
Mississippi Valley probably to southern Wisconsin, west to south-
eastern Wyoming, southwest to the panhandle of Texas, and east to
eastern Illinois. It appears to be a common snake in eastern Kansas,
Missouri, southern Illinois, eastern Oklahoma, Arkansas, eastern
Texas, and Louisiana. Records are very few for Nebraska, Iowa,
and Mississippi.
Specimens have been examined from the following localities, in
addition to those represented by specimens in the United States
National Museum: Near Columbus Junction, Iowa; Duntanville
(Tuscaloosa County), University, Alabama Point (Mobile County),
and Mobile, Alabama; Cat Island and University, Mississippi; Man-
hattan, Twin Mounds, Mount Oread, Coal Creek in Coffey County?
Long Creek in Osage County, and Jefferson, Trego, Labette, and
Anderson Counties, Kansas; Dallas, Clifton, and Tule Canyon, Staked
Plain, Texas; Wister, Sugar Loaf Mountain, and Fort Supply, Okla-
homa; Arkadelphia, Greenway, Donaldson, Fayetteville, and Garland
and Jefferson Counties, Arkansas; Horseshoe Lake, Olive Branch,
Illinois; Reelfoot Lake, Samburg, Tennessee; Chastine, Louisiana;
Galena, Missouri.
Published records for other localities are as follows: Deming's
Bridge, Matagorda County, Texas (Garman, S., 1892, 10); Burnet,
Texas (Strecker, 1909, 7); Peoria, Illinois (Garman, H., 1892, 298);
Wisconsin (same); Montgomery County, Missouri (Hurter, 1911, 185);
and the following counties in Kansas (Branson, 1904, 398) : Mitchell,
Republic, Montgomery, Mami, Greenwood, Potawatomie, Franklin,
Lyon, Sumner, Scott, Logan, and Gove.
REVISION OF THE KING SNAKES. 37.
Variation.—^Minor variations of pattern appear to characterize
certain regions of the range. Thus specimens from southern Missis-
sippi, Louisiana, and southeastern Texas usually show the dorsal
cross bands plainly, but the spaces between and the sides are promi-
nently spotted with rounded or oval white spots (in specimens from
New Orleans the spots are a pale greenish white and the ground color
is velvety black).
In the area from extreme northeastern Texas to southern Kansas,
St. Louis, and the Mississippi River the cross bands are almost indis-
FlG. 9.—Map showing locality records for LAMPROPELTIS OETtrLtlS HOLBROOKI.
tinguishable; every scale has a large white or yellow spot on a ground
color of dark brown to bluish black (Hurter, 1911, 185), which, pro-
ceeding ventrally, occupies more and more of the area of the scale.
Often the only indication of cross bands is the asymmetrical orienta-
tion or atypical shape of certain of the dorsal spots, but the regu-
larity of their distribution indicates that they represent the original
bands. These spots may be unusually long and narrow and oriented
either lengthwise or crosswise of the scales, and, judging from recently
preserved specimens, they are while or pale greenish on a nearly
black ground color.
-38 BULLETIN 114, UNITED STATES NATIONAL. MUSEUM.
Specimens from Kansas are more like those from New Orleans in
respect to the distinctness of the dorsa bands and the prominence of
the spots between, but the spots are yellow (in alcohol)—Branson says
(1904, 398) pearly white or yellow.
The pattern of young individuals is deserving of attention. In all
the specimens examined the white crossbars are prominently devel-
oped and the scales between are unspotted, or the spots are small,
or present only midway between the bands. From the material at
hand it seems probable that, as suggested by Branson (1904, 398)
and by Hurter (1911, 185), the spots between the bands develop as
the young snake grows older.
The variation in scutellation and proportions needs very little dis-
cussion. In these characteristics the differences between this form
and its allies, splendida, niger, and getulus, are slight or not apparent.
This may be said of the labials, the caudals, the temporals, and the
proportionate tail length. The number of specimens is entirely inad-
equate for the demonstration of geographic tendencies in these char-
acters. It is probable, however, that geographic variation occurs in
the number of ventral plates. The average for this species is 210,
extremes 201 and 221; for splendida, 217, extremes 208 and 226;
ioT niger, 209, extremes 200 and 217; for getulus, 215, extremes 204 and
223. While it is apparent from the slight differences in the averages
for these species that but little reliance can be placed on variation in
the number of ventrals, it is nevertheless a fact that che lower numbers
of ventrals in Tiolbrooki are more common in the northern and north-
eastern parts of its range, and the higher numbers more frequent in
the western and southwestern portions. This means approach in this
character to niger and to splendida, respectively.
The scale rows are subject to considerable variation, as the table
shows, but the great majority of the specimens have one of the three
formulae—21-19, 19-21-19, and 19-21-19-17.
This form can thus be compared in lowness of scale formula only
with niger and getulus; the former is fully as much reduced as hd-
IrooJci, and the latter nearly as much so. Reference to the table will
show that the higher formulae characterize the western and south-
western parts of the range, and the lower formulae the northeastern
and the extreme southern. If hoTbrooki had a southwestern origin,
these regions would be the last reached, the former because it is the
farthest removed from the southwest, and the latter because it is but
recently elevated above the sea.
Affinities.—HoTbrooki is without doubt very closely allied to splen-
dida and to niger, and with both of these forms intergradation takes
place along the common boundaries of their respective ranges. We
will, however, leave a fuller discussion of afiSnities to the section on
REVISION OF THE KING SNAKES. 39
the evolution of the group, and leave the subject here with the fol-
lowing diagram of relationships:
splendida Jiolhrooici mger.
Scale/ormulae of holbrooki.
DISTRIBUTION BY SEX.
Formula. Male. Female.
21-23-21 1
8
23
16
1
10
21-23-21-19
. ..
21-19
19-21-19
21-19-17
19-21-19-17
19-17
Total
4
18
32
2
15
1
72 59
DISTRIBUTION BY LOCALITY.
Formula.
Oklahoma
and north-
ern Texas.
Elansas. Arkansas. Missouri. Louisiana.
Higher than 21-19
21-19
Lower than 21- 19
Total
5
6
6
2
17
7
3
9
7
1
1
13
2
7
25
17 26 19 15 34
40 BUIiLETIN 114, UNITED STATES NATIONAL MUSEUM,
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REVISION OF THE KING SNAKES. 41
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BEVISION OF THE KING SNAKES. 43
LAMPROPELTIS GETULUS NIGER (Yarrow).
Fig. 33.
1837. Herpetodryas getulus Schlegel, Phys. Serp., p. 198.
1882. OpMbolus getulus niger Yarrow, Proc. U. S. Na,t. Mus., vol. 5, p. 438 (type
locality, "^lieatland, Indiana; cotypes, U. S. Nat. Mus., no. 12149—two
specimens; Robert Ridgway, collector); Bull. U. S. Nat. Mus., no. 24,
1882, p. 93.—Hay, Journ. Cincinnati Soc. Nat. Hist., vol. 10, no. 2, 1887,
p. 64.—Gaeman, Bull. Illinois State Lab. Nat. Hist., vol. 3, art. 13, 1892
p. 299.—Hay, Batr. Rept. of Indiana, 1893, p. 111.—Cope, Rep. U. S.
Nat. Mus. for 1898, 19C0, p. 917, fig. 228.—Brown, Proc. Acad. Nat. Sci.
Philadelphia. 1901, p. 77.
1887. Ophibolus niger Hay, 36th Annual Rep. State Board Agri. Indiana for 1886,
vol. 28, p. 210.
1891. Ophibolus getulus sayi BlatchlIey, Journ. Cincinnati Soc. Nat. Hist., 1891,
p. 32; 24th Ann. Rep. Dept. Geol. Nat. Res. Indiana, 1899, p. 545.
1893. Ophibolus doliatus, var sayi Hay, Batr. Rept. State Indiana, p. 110.
1894, Ophibolus getulus Carman, H., Bull. Essex Inst., vol. 26, nos. 1, 2, 3, p. 35.
The status of this form has never been settled, and the specimens
avaihible for study are still too few. The \mter, however, believes it
to be entitled to the rank of subspecies, but should fuller material
prove it to be but a local or inconstant variation of TiolbrooTci, its name
will have to supersede the latter, a change by no means to be desired.
Description.—The scutellation of the 32 specimens examined is as
follows: ventral plates, 199 to 216; caudals, 41 to 53 (males, 46 to 53,
average 50; females, 41 to 47, average 43); supralabials, 7; infralabials,
9, rarely 8 or 10 ; 1 preocular, 2 postoculars; temporals usually 2+3 + 4;
posterior chin shields generally shorter than the anterior and separated
from each other by one or tv.'o small scales; loreal higher than long,
or about as high as long; scale formula usually 19-21-19.
The bodily proportions are the same as for JiolhrooTci and getulus.
The tail varies from 0.110 to 0.146 of the total length (males, 0.120
to 0.146, average 0.133; females, 0.110 to 0.128, average 0.120).
The largest specimen examined was the type—1,431 millimeters in
length.
The color pattern (fig. 33) is a reduction from that of holhrooH by
obUteration of the yellow centers on the scales of the dark areas on
the back, and contraction in size of all the other light spots. This
leaves the dorsal surface black, crossed by 50 to 90 very narrow cross-
bands of yellow, which tend to fork on the sides and there join an
alternating series of short, narrow transverse bars. Occasionally the
crossbands on the back may have nearly or quite disappeared, or
sometimes the white spots between the bands may be somewhat
developed. The lower rows of dorsal scales are more or less spotted
with yellow. The belly is checked with black and white or yellow,
the black sometimes predominating. The spotting on the head is
186550—21—Bull. 114 4
44 BULLETIN 114j UNITED STATES NATIONAL MUSEUM.
exactly like that of liolhrooTci except that the spots are fewer and
much smaller.
The penial characters seem to agree with those of Jiolbrooki, even
to the extent and prominence of the minute spines.
The dentition of a specimen from Wheatland, Indiana, is as follows
:
Maxillary teeth, 13-13; mandibular, 14-13; palatine, 9-9; ptery-
goid, 16-17.
This form may be known from TioTbrooki from the fact that the
adults have no white spots, or but very few, on the scales between the
dorsal crossbands; young examples may resemble TioTbrooki, but the
crossbands are very narrow, less than half a scale in width, and all
the white spots are smaller than in the latter form. It may be Imown
from splendida by the fact that the dorsal scales are in 21 instead of
23 rows; and from getulus it may be distinguished by the narrower
and dotted crossbands instead of wider and continuous ones, and by
their greater number, more than 50.
Habitat and habits.—^Almost nothing is recorded upon this subject.
Blatchley (1899, 545) says for Vigo County, Indiana, that it "fre-
quents rocky hillsides and the vicinity of streams," and mentions
finding one in the act of swallowing a Eutaenia sirtalis. The writer
found an adult at Henry, Tennessee, a little after simset, stretched
out at fuU length by the side of a road through farming country.
It had doubtless been concealed during the day in the thick bushes
between the side of the road and the open field adjacent. It offered
no resistance whatever to being picked up.
Range.—This form occurs from extreme eastern Illinois to southern
Ohio and south to northern Alabama.
The only published records for specimens not examined by the
writer are those of H. Garman (1894, 35), for Midland, Kentucky,
which was probably this form, and Blatchley (1899, 545) for Putnam
County, Indiana.
Three specimens in the collection of the Museum of Comparative
Zoology (no. 33), labeled ''Ohio," are almost typical of JiolbrooJci and
one in the National Museum collection (no. 12026) labeled "Mt.
Carmel, lUinois" is quite so. More specimens are much needed to
definitely settle the status of this form.
Variation and affinities.—The specimens named niger by Yarrow
were supposed to have been derived from typical getulus by increase
in black pigment at the expense of the yellow. It was evidently
overlooked that the crossbands were the same in character and in
number as in JiolbrooJci. The scale formulae and the penial characters
also ally it much closer with JiolbrooJci than with getulus. That it is,
however, closely comiected with the latter there can be no doubt.
A specimen from the Cherokee Nation in the moimtains of North
Carolina with 37 crossbands presents a strong contrast with one
REVISION OF THE KING SNAKES. 45
from so short a distance west as Knoxville, Tennessee, with 73, and
even though the former specimen shows the bands narrower than
usual for getulus, these forms might be supposed from these specimens
to be distinct even where their ranges meet, but specimens from
Georgia and Alabama practically prove intergradation. The speci-
40
35
Fig. 10.—Map showing locautt records for Lampropeltis getulus kiger.
mens from Marietta (fig. 34) and from Augusta, Georgia, cited by
Yarrow (1882, 91), have the cross bands very narrow, unlike any
typical getulus, but their number is 38 and 31, respectively, and one
would doubtless not hesitate to assign them to getulus. Specimens
from Ida and Gallant, Alabama, are decidedly niger, but one from
46 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
Anniston, Alabama, collected in 1919 by E. K. Dunn, is practically
transitional. This specimen shows the crossbands to be almost
obsolete, but where they can be made out they seem to be reduced
in number. The alternating light and dark areas on the sides of the
belly are about 33 in number—a getulus character, but in general
appearance it is nearer to niger. This enables us to determine
fairly closely the limits of the ranges of these two forms in the South
and the region where intergrades may be looked for.
That niger bridges the gap between liolbrooki and getulus there is
hardly room for doubt. Also that on the whole it is more closely
allied to TioTbrooki is evident from consideration of the characteristics
of all three. And from consideratiouxof the latter and the geographic
relationships, it becomes most logical to regard niger as a derivative
of lioTbrooki. This makes it necessary to consider niger as ancestral
to getulus. We may therefore express the probable relationships of
these three forms as follows:
liolhrooki—^niger—^getulus.
Further evidence for this view of their relationships will be found
in the summary of the getulus group.
ScaUformulae of niger.
Fonnula. Male. Female. Totals.
21-23-21-19 1
5
8
6
1
8
14
6
21-19 3
619-21-19
19-21-19-17
Total 20 9 29
REVISION OF THE KING SNAKES. 47
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REVISION OF THE KING SNAKES. 49
LAMPROPELTIS GETULUS GETULUS (Unnaeue).
kinqsn.vke; chain snake; cowsucker; black moccasin; horse racer; master snake; bastard horn
snake; thunder snake; thunder and lightning snake; "wamper; -wampum snake; rattlesnake
PILOT.
Figs. 4, 34, 35.
1766. Coluber getulus Linnaeus, Syst. Nat., ed. 12, vol. 1, p. 382 (type locality
Carolina).—Gmelin, Syst. Nat., vol. 1, pt. 3, 1788, p. 1106.—Lacepede,
Hist. Nat. Serp., vol. 2, 1789, pp. 84, 300.—Bonnaterre, Tabl. Encycl.
Meth., 1790, p. 45, pi. 18, fig. 33.—Sonnini and Latreille, Hist. Nat.
Rept., vol. 4, pt. 2, 1799, p. 174.—Daudin, Hist. Nat., vol. 6, 1800,
p. 314, pi. 77, fig. 1.—Shaw, Gen. Zool., vol. 3, pt. 2, 1802, p. 467.—Say,
Amer. Joum. Sci., vol. 1, 1818, p. 260.—Boie, Isis, 1827, p. 537.—
Harlan, Joum. Acad. Nat. Sci. Philadelphia, vol. 5, pt. 2, 1827, p.
358.—Peale, Contr. Macl. Lye, vol. 1, 1829, pi. 5.—Harlan, Med.
Phys. Researches, 1835, p. 122.
—
Lacepede, Hist. Nat. Quad. Ovip.,
vol. 1, 1836, p. 300.—De Kay, New York Fauna, 1842, plates, rept.
amph., pi. 10, fig. 21, pt. 3, p. 37.—Linsley, Amer. Jo\im. Sci., vol. 46,
1843, p. 43.—Hough, 5th Annual Rept. State Cab. Nat. Hist. New York,
1852, p. 23.—GiJNTHER, Cat. Colubr. Snakes Brit. Mus., 1858, p. 249.—
Pseudoelaps getulus Fitzinger, Neue Class. Rept., 1826, p. 56.
—
Coronella
getula Holbrook, N. Amer. Herp., ed. 2, vol. 3, 1842, p. 95, pi. 21.
—
Lichtenstein, Nomenclator, Mus. Zool. Berolinensis, 1856, p. 25.
Boulenger, Cat. Snakes Brit. Mus., vol. 2, 1894, p. 198 (part).
—
Ophi-
holus getulus Baird and Girard, Cat. N. Amer. Rept., pt. 1, 1853, p.
85.—Baird, Serp. of N. Y., 1854, p. 20; Pacif. R. R. Surv., vol. 10, pt. 3,
no. 1, 1859, pi. 31, fig. 65.—Coues, Proc. Acad. Nat. Sci. Philadelphia,
1871, p. 48.—Yarrow, Amer. Nat., vol. 12, 1878, p. 470.—Garman, S.,
Mem. Mus. Comp. Zool., vol. 8, no. 3, pt. 1, 1883, pp. 68, 156, pi. 5, fig.
3; Nat. Hist. Notes, Boston, 1887, p. 2.—Hay, 36th Annual Rep. Indiana
State Board Agri. for 1886, vol. 28, 1887, p. 210.—Nelson, Rep. State
Geol. New Jersey, vol. 2, 1890, p. 647.—Cope, Proc. Amer. Philos. Soc,
vol. 33, 1894, p. 221; Rep. U. S. Nat. Mus. for 1898, 1900, pi. 18, fig. 7.—
Brimley, Joum. Elisha Mitchell Sci. Soc, vol. 21, no. 4, 1905, p. 152.
DiTMARs, Reptile Book, 1907, pp. 341, 351, pi. 103, figs. 11, 12, pi. 108
(upper fig.).—Brimley, Proc. Biol. Soc. Washington, vol. 22, 1909, p.
134.—DiTMARS, Reptiles of the World, 1910, p. 267, pi. 59 (upper fig.);
Sci. Contr. New York Zool. Soc, vols. 1, 2, 1912, p. 222.—Brimley,
Joum. Elisha Mitchell Sci. Soc, vol. 30, no. 4, 1915, p. 202.—Engle-
HARDT, et al., Copeia, no. 17, 1915, p. 1.
—
Coronella getulus Dumeril and
BiBRON, Erp. Gen., vol. 7, pt. 1, 1854, p. 616.—Jan, Icon. Gen., livr. 14,
1861, pi. 5, fig. 1; Arch. Zool. Anat., vol. 2, fasc. 2, 1863, pp. 238, 244.—
Lampropeltis getula Cope, Proc Acad. Nat. Sci. Philadelphia, 1860, p.
255.—Abbott, Geol. of New Jersey, Appendix E, 1868, p. 802.—Xam-
propeltis getulus Abbott, Nat. Rambles, 1894, p. 476.
—
Loennberg,
Proc. U. S. Nat. Mus., vol. 17, no. 1003, 1894, p. 317.—Cory, Hunting
and Fishing in Florida, 1896, p. 131.—Ditmars, Proc. Linn. Soc. New
York, 1896, p. 12; 1902, p. 2.—Brimley, Amer. Nat., vol. 37, p. 263.—
Stone, Amer. Nat., vol. 40, no. 471, 1906, p. 167; Brimley, Joum.
Elisha Mtchell Sci. Soc, 1907, p. 145.
—
Fowler, Annual Rep. New
Jersey State Mus. for 1906, 1907, p. 178, text figs., pi. 45.—Brimley,
Proc. Biol. Soc. Washington, vol. 22, 1909, p. 134.—Hurter, Trans.
Acad. Sci. St. Louis, vol. 20, 1911, p. 186.
—
Brimley, Joum. Elisha
Mitchell Sci. Soc, vol. 30, no. 4, 1915, p. 202.—Dunn, Copeia, no. 18,
1915, p. 6; Bull. Amer. Mus. Nat. Hist., vol. 37, art. 23, 1917, p. 593,—
Stejneqer and Barbour, Check List, 1917, p. 88.
—
Brimley, Copeia,
no. 64, 1918, p. 97; no. 88, 1920, p. 100.
50 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
1875. Ophibolus getulus getulus Cope, Bull. U. S. Nat. Mus., no. 1, p. 37.
—
Yarrow, Bull. U. S. Nat. Mus., no, 24, 1882, p. 91.—Davis and Kice,
Bull. Illinois State Lab. Nat. Hist., no. 5, 1883, p. 33.—Cope, Eep. U. S.
Nat. Mus. for 1898, 1900, p. 914, fig. 227.—Eckel, Amer. Nat., vol. 35,
1901, p. 153.—Brown, Proc, Acad. Nat. Sci. Philadelphia, 1901, p. 77.—
EcEiEL and Paulmier, Bull. New York State Mus., no. 51, 1902, p. 375.
Henshaw, Occ. Pap. Boston Soc. Nat. Hist., vol. 7, 1904, p. 9.
—
Drowne,
Mon. Roger Williams Park Mus., no. 15, 1905, p. 12.
—
Lampropeltis
getuliis getulus Hay, Proc. Biol. Soc. Washington, vol. 15, 1902, p. 139.
Wright, Proc, Acad. Nat. Sci. Philadelphia, 1915, pp. 139 et seq., 168,
fig. 10.
Description.—^The eastern representative of this group is well
known and easily recognized. The scutellation as determined from
134 specimens is as follows: Ventral plates, 203 to 224; caudals, 38
to 58; supralabials, 7 (rarely, 6 or 8); infralabials, 9 (rarely, 8 or 10);
oculars, 1 and 2 ; temporals, 2 + 3 + 4 ; posterior chin shields about
equal in length to the anterior, and parallel; loreal as high as, or
higher than, long; dorsal scale rows commonly 21-23-21-19 or 21-19,
the higher formula being characteristic of the southern portion of the
range and the lower of the northern; rarely as high as 23-21 and
sometimes as low as 19-21-19-17.
This snake is somewhat stouter than the western representatives
of the group, and the head is distinctly narrower and higher. This is
reflected in the broader dorsal scales, the higher labials and loreal,
and narrower rostral. The tail varies from 0.100 to 0.155 of the total
length (males, 0.104 to 0.155, average, 0.127; females, 0.103 to 0,134,
average, 0,119). The largest specimen examined was from Gaines-
ville, Florida, and measured 1,752 mm,; the largest specimen from
the northern part of the range measured 1,434 mm,, and was taken
in Caroline County, Virginia.
The pattern (fig. 35) is formed of a series of narrow white or yellow
bands, 23 to 52 in number that cross the back transversely or ob-
liquely, bifurcate on the sides, and there join a series of quadrate
light spots. The latter overlap a little on the ventral plates, and
alternate with the dorsal bands. The belly is checked with a light
brown, and white or yellow, with a tendency for the dark to be
mostly opposite to the light areas on the sides.
The ground color of the head is dark. The rostral is white with a
black posterior border. The internasals, prefontals, and frontal
are dark, with each a transverse light bar anteriorly. There is a
medial light spot on each supraocular near the frontal suture. Each
parietal has a round light spot medially, near the common suture,
and an elongate light spot on the anterior-lateral border. There is a
light spot near the center of the nasals, loreals, oculars, and temporals.
The labials are white, with dark posterior borders; the chin and gular
shields are white. Close behind the head is a light spot about three
REVISION OF THE KING SNAKES. 51
scales long and two wide, or this may be elongated as the first trans-
verse band.
Normally the white is restricted as above, but in the soutliern por-
tion of the range there is commonly a development of white in the
centers of the scales of the dark areas of the back and sides. This is
usually most pronounced midway between the light bands.
The belly is checked with black or brown and white, or yellow and
may be nearly all dark or nearly all light.
The copulatory organ may be described as follows: Distinctly
forked; sulcus single, extending over the side of the longer fork; a
bare space at the apical end, surrounded by low calyces which extend
but slightly below the tips of the forks; fringes few, soon developing
into spines, which increase in size to about one-third the distance to
the base, and then stop suddenly, being succeded by a few minute
spines, or none at all; remainder of organ smooth. This description
holds for specimens from Baldwin County, Alabama, Indian River,
and Charlotte Harbor, Florida, Virginia Beach and Dunn Loring,
Vu'ginia, and Montgomery County, Maryland. This organ differs
from that of Tiolbrooki m the great restriction or entire absence of the
minute spines, the more distinctly forked character of the apical
end, the fewer caWces and their shorter fringes. It does not seem
to differ particularly from that of jloridana, except in having shorter
forks.
The skull is essentially like that of TwTbroold. Maxillary teeth 13
to 16, usually 13, subequal, the last one or two slightly stouter than
the preceding or actually a little smaller; mandibular teeth 14 to 17,
usually 14 or 15, the anterior a little the larger, the first and the last
five or six the smallest; palatines 9, sometimes 10, subequal; ptery-
goids 16 to 20, subequal, a little smaller than the palatmes and
decreasing in size posteriorly.
While getulus is a distinct and well-defined form, throughout its
range, at its southwestern limits it intergrades with niger, and m
central Florida v^iih floridana. The only character by which it may
be distmguished hi doubtful cases from uiger is the number of cross
bands, which are more than 50 in the latter and usually less than 40
in getulus. Where the range of getulus meets that oifioridana, again
the most valid separation may be made on the number of cross
bands; getulus generally has less than 50, and Jloridana more. The
totality of characters is the best guide, and yet there may still be
doubt about exceptional individuals from near the common boundary
of the ranges.
Habitat and h<ibits.—This species has received frequent mention in
the literature. Probably the first reference to it was that of Catesby
(1731, 52). He says: "It was the first one of the kind I ever saw, nor
was it known to any of the inhabitants I showed it to, therefore as
52 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
it wanted a name, the best I could think of was that of Chain-Snake,
from some resemblance to a chain that seems m many cases to en-
viron the body." Shaw (1802, 467-468) says: ''This snake is fomid
in Carolina, Virginia, etc., frequenting moist woods and shady places,
and preying on lizards, etc." DeKay says of it (1842, 37): "Not
uncommon in the pine woods of New Jersey, and is also found but
rarely in what are called the Brush Plains of Long Island." Ditmars
says with reference to habitat (1907, 359): "Specimens captured by
the writer were in rather dry patches of timber; some were taken
while basking in the sun of small glades in the forest; others were
found hiding under fallen tree trunks." Dunn has taken nimierous
examples in Nelson County, Virginia, in hay fields, near creeks, and
about farm buildings. Some of these would resent handling at first
by chewing at hand or arm, but usually they were perfectly tame
from the first. Wright says, for the Okefinokee Swamp (1915,
168-169): "Thirteen specimens of this fuie snake were taken and
many more seen. It is common throughout the drier parts of the
swamp and frequents the outskirts of the swamp as well. It keeps
to the islands and none were taken in other situations than the saw
palmetto or heath societies of the piney woods where it courses
through the low cover after its living prey or eggs. This species is
one of the most strikingly marked snakes of the swamp. The shining
black or brown with the contrasting white or yellow cross-bands
makes its appearance very attractive. In nature, it is mild, proves
an interesting and safe pet, and in no instance during our stay in the
swamp did it display any belligerency or suUenness toward any
member of the party."
This is a large, powerful, and fearless snake. It does not appear
to be secretive or to avoid an encounter. In fact (Ditmars, 1907,
360-361), "besides the promptings of its appetite this snake exhibits
a pugnacious interest in other serpents that may be considerably
larger than itself, engaging these creatures in a duel to the death,
during which, however, they are able to make but little resistance,
when encircled by the wonderfully strong constricting coils of the
enemy." Although sometimes (Ditmars, 1907, 363) "when first
caught, they strike vigorously, emitting a short hiss which sounds
more like a sneeze," they are frequently perfectly gentle and un-
afraid from the beginning. In fact, Loennberg (1894, 324-325)
writes that "when caught they never tried to bite, only one opened
its mouth, but they wind themselves round one's arm, showing
great muscular strength." For observation in captivity they are
probably the most desirable of all the North American snakes.
They feed upon other snakes, lizards, birds, and small rodents.
In this latter respect and because they are always victorious in an
encounter with a venomous snake, they are very useful to the agri-
REVISION OF THE KING SNAKES. 53
cultuiist and should receive full protection. When their prey is
large enough to struggle effectively, the King snake coils about it
in true constrictor style, but if it is small or dead it may swallow it
directly. Definite food records are few. Dunn has captured several
that contained mice. In captivity they have been observed to
eat ThamnopTiis lutleri, T. sirialis, Car'pho'pMs amoenus, house
mice, dead or alive, and even their relatives, L. getidus Tiolhrooki
and L. calligaster. Two of the latter kept by the writer were eaten,
one by accident (both starting to swallow the same mouse), and
the other apparently by intent, although they had both lived in
the same cage for more than a year. Hurter (1911, 186) records
the Black snake, Bascaniori constrictor, as having been eaten in
captivity, and Dunn mentions Fleterodon contortiix, Nairlx sipedon,
N. septemvittata, and the Chipping Sparrow.
By far the best account is given by Wright. Discussing the King
snake, as he found it in the Okefinokee Swamp, he writes (1915, 169):
The natives recognize its good nature and consider it harmless, though the king
of the snakes. They are aware of its usefulness as an enemy of moccasins and rattle-
snakes and report several combats which alwaj's resulted successfully for the King
snake, but these unlettered people, unlike many sentimental writers^ do not hold
that the King snake deliberately searches for the poisonous snakes in particular.
We, as they, believe it the enemy of every species of snake in the swamp, pre>ing of
course more on the terrestrial species of its own haunts. All the smaller snakes suffer,
and of the larger species, the blacksnake and spreading adder are the commonest
prey. It is surely a good "pilot" to the naturalist whenever one finds it digging,
for it almost invariably means other snakes, eggs, or some good capture. It will
seldom fail to react per schedule if you loosely hold it in one hand and a live black-
snake in the other. Almost before you can predict the outcome, the former may be
far within its captor—a demonstration we have tried more than once in the field.
It is especially fond of young snakes. One of our specimens had taken a newly
hatched Hderodon and the natives recounted several occasions when they had foxind
it working beneath a log for what proved a brood of young snakes. We do not doubt
but that it feeds on mice, rats, and other small mammals, but of such evidence we
found little in the swamp. Possibly, in early spring or in the fall these are more its
reliance. The principal food of this species is turtles' eggs, with snakes or their
^gs a second choice. Four of our specimens had eaten Florida cooters' (Chrysemys
floridana) eggs, which they dug out of the sand, and two had mud turtle {Cinosternum
pennsylvanicum) eggs in their stomachs. Mr. Francis Harper tells lis that he and
David Lee almost stepped on a King snake. After their recovery, what should
they find but a Kinosternon digging in sand, probably preparatory to lajdng, and the
King snake was close at hand. In fact, so addicted are they to this egg diet, that
the natives consider that it is a common happening to find the snake awaiting the
egg deposition. Unless it be the Florida bear, there is no form in the swamp which
eats turtles' eggs in such quantity as the King snake. It will take a whole nest of
eggs at one time, as many as 14 being found in the stomach of one snake.
This species, like the other King snakes, is o^iparous, and (Ditmars,
1907, 363) "deposits from 10 to 24 eggs, which require from five
to six weeks to complete the incubation." Wright (1915, 170)
says: "Mr. Harper reports a pair of them mating on May 19, 1912,
54 BULLETII^r 114, UNITED STATES NATIONAL MUSEUM.
[Okefinokee Swamp], and says another King snake was watching
the pair. One of our specimens (no. 6145), taken on June 15, 1912,
had 7 fair-sized eggs."
C. S. Brimley (1903, 263) says: "The eggs of the King snake
{OpJiibolus getidus) are long, oblong in shape, with a smooth, tough
skin and are more or less adherent to one another in clusters."
Range.—^This form is known to extend from central New Jersey
to central Florida, and west from the coast to the Alleghenies and
to Mobile Bay, Alabama. On the west side of Mobile Bay it appears
to be entirely replaced by Jiolbrooki, and in northern Alabama by
niger. The few specimens from the southwestern portion of the
range indicate that intergradation takes place in the region of a
line drawn from the point where the North Carolina-Tennessee
State boundary line meets that of Georgia southwest to Mobile.
Bay. (The locality of specimen no. 2294, United States National
Museum, labeled "Miss.," must be an error.) While there appear
to be no verifiable records north of New Jersey, numerous references
in the literature can not be overlooked. DeKay refers to it as
occurring in the "Brush Plains of Long Island" (1842, 38). Dit-
mars (1896, 13) says he has not heard of one being found in Long
Island or the vicinity of New York in the last five years. Baird
(1854, 21) says "it is quite maritime in its northern distribution,
being rarely found in the northern states except near the coast."
A specimen in the United States National Museum (no. 459), re-
ceived from the Jardin des Plantes, Paris, in 1858, bears the label
"New York." This may easily be an error. Hough (1852, 23)
reports this snake for Rossie, St. Lawrence County, New York, "of
common occurrence in this section of the state," and Eckel (1902,
376) refers to this record as one that "can not be neglected or sup-
pressed." Linsley (1843, 43) reports it at Milford, Connecticut, on
the word of a man whom he considers trustworthy.
Besides the localities represented by specimens in the United
States National Museum, specimens have also been examined from
the following localities: Bridgeton, Point Pleasant, Port Republic,
Stafford's Forge, Vineland, Lakewood, Lakehurst, and Beesley's
Point, New Jersey; WiUiams, Maryland; Seaford, Delaware; Midway
Mills, Nelson County, Virginia; Bluffton, Hampton County, and
Hilton Head, South Carolina; Raleigh, and Fort Macon (Carteret
County), North Carolina; Fargo, Savannah, and Billy's Island,
Okefinokee Swamp, Georgia; Orange Lake, Eustis, and Marion
County, Florida; Daphne, Baldwin County, Alabama.
Published records for localities from which no specimens have
been examined are as follows: Port Tobacco, Maryland (Yarrow,
1878, 470); Rossie, St. Lawrence County, New York (Hough, 1852,
REVISION OF THE KING SNAKES. 55
23); Milford, Connecticut (Linsley; 1843, 43); Nashville and Alapaha,
Berrien County, Georgia (Wright, 1915, 140).
Variation.—The averages for the ventral plates show distinct
differences with latitude. Thus the diagram (fig. 11) shows that
there is a progressive decrease in number of ventrals northward from
Georgia and an increase southward into Florida. The former is
much more gradual than the latter and is accompanied with other
gradual changes, which make a slight but not very distinctive differ-
ence in the appearance of the specimens from widely separated parts
of the range. The change into southern Florida is much less gradual
and is accompanied by other marked changes, particularly of pattern,
Ventral
plates.
222
220
218
216
214
212
210
208
200
204
202
Southeastern Northern Eastern Southeastern D. C. and New
Georgia. Florida. North Carolina. Virginia. vicinity. Jersey.
Localities. Eastern S. C.
Fig. 11.—Diagram showing geogkafhic variation in number of ventral plates in Lampropeltis
GETULUS GETULUS.
sufficient altogether to make necessary the recognition of a distinct
subspecies there (as jioridana)
.
The averages of the numbers of caudal plates are based upon too few
specimens to be of much value, but, such as they are, they support
the figures for the ventrals, that is, there is a slight decrease in number
northward, and increase southward. The same is true of the in-
fralabials. The higher numbers are more frequent in the south,
the lower in the north. Specimens from northern Florida show
about 15 per cent of increase to 10 infralabials; m fioridana this is
increased to about 30 per cent; north from Florida 10 is of less com-
mon occurrence, and the number is not infrequently reduced to 8.
The dorsal scale rows vary between the rather widely separated
formulae of 23-21 and 19-21-19-17. The extremes are uncommon,
Z4
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56 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
however. North of Georgia the usual formula is 21-19, sometimes
23 rows is attained and more often the formula is lowered to 19-21-19,
21-19-17, or 19-21-19-17. South of Georgia a maximum of 23
rows is more common than a maximum of 21, and of all the speci-
mens examined from Florida, no individual had a lower formula than
21-19. It seems almost certain that extensive series of specimens
Fig. 12.—Map showing locality records for Lampropeltis getultjs getulus.
will demonstrate a progressive decrease in number of scale rows with
increase in latitude.
The pattern, as already described, is fundamentally a series of
transverse dorsal bands which bifurcate on the sides to join there
an alternating series of light areas. This arrangement is subject
to much variation. The bands vary from J to 1| scales in width;
the forks are always narrower, and vary from interrupted spots, or
nothing whatever, to unbroken lines a scale wide; and the lateral
REVISION OF THE KING SNAKES. 57
white areas vary from squarish, about 2 scales by 2 to oblong and
about 4 or 5 scales long and 3 or 4 wide. In position the dorsal
bands may be transverse or oblique, and extend across from 9 to 15
or more rows of scales. The lateral alternating spots are always
located on the first few rows of scales; they are usually longer than
wide and nearly always overlap onto the ends of the ventrals. De-
pending upon the upward extension of these spots and the downward
extension of the cross bands, the forks may be horizontal or set at as
much as a 45 degree angle. It is rather more common for the white
scales to be all white, but frequently these scales, particularly in
the cross bands, are tipped, up to as much as half their area, with
deep black.
These variations seem to be all common and widespread, and
appear not to be of geographic significance. Of other variations the
opposite can be said.
From Georgia to southern Virginia the number of white cross
bands averages close to 30, northward this is increased to more than
40 near the District of Columbia and in New Jersey, and southward
into northern Florida it likewise is increased to 40. Correlated with
the change in number of bands northward is an intensification of the
white and a blackening of the ground color, and southward a lessening
of the distinction between the light and dark.
In southern Georgia and northern Florida it is by no means uncom-
mon for the scales of the dark areas to develop white centers. These
may extend to all the dark scales, but usually are best developed
midway between the white cross bands, fading gradually to disap-
pearance on either side. It seems likely that the potentiality for
the development of white centers is an inheritance from the form
Iholhrooki. Evolution into jloridana seems to have proceeded by an
enlargement basally, then distally, of these white centers, a softening
of their sharp outlines and a lightening of the dark parts, accom-
panied by an increase in number of cross bands and a decrease in
their distinctness, resulting in a light brown spotted appearance in
which the remains of the original pattern are to be distinguished
with difficulty. This reaches its logical outcome in extreme southern
Florida. In this phase, which has recently been described by
Barbour as hrooJcsi, the cross bands have completely disappeared,
and all the dorsal scales are yellow at the bases, shading to dark
brown at the tips.
Along with the general reduction in scutellation to the north is a
slight decrease in size of body, perhaps best realized by comparison
with the distinctly larger individuals from central and southern
Florida.
Mention must here be made of a most mteresting individual from
Occoquan Creek, Virginia (Cat. No. 59354, U.S.N.M.). This is a
58 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
variant in respect to pattern in every way similar to the specimens
of californiae from Fresno County, California. The belly is almost
an immaculate white on the anterior half of the body, but on the
posterior half it is mostly black, mixed with some white. The dorsal
pattern of cross bands is broken up into spots of varying shapes,
chiefly elongate, with a conspicuous tendency to form a dorsal white
stripe. It is to be hoped that more collecting in this vicinity will
reveal other examples. It indicates the inherent potentiality in
^etulus for the production of an Atlantic coast analogue of californiae.
Affinities.—It will doubtless already have been noticed that ven-
trals, caudals, labials, scale rows, and color pattern all indicate the
same thing, that is, that the region represented by Georgia is the
meeting ground of two diverging lines of evolution in getulus. That
one line has been southward there can hardly be a doubt. Floridana
is clearly an end form in all its characters, and it is certainly so in
geographic position. Similarly getulus, as it occurs in the northern
portion of its range, demonstrates itself to be a reduced form of the
southern type. Consequently evolution has proceeded from the
Georgia-Alabama region northward along the coastal plain, and
southward into Florida. In central Alabama and northern Georgia
getulus and niger intergrade in the region indicated above under
Range. The material at hand indicates that this is the only region
where intergradation occurs between these forms. The relationships
may therefore be expressed as follows:
getulus
(New Jersey)
niger getulus
(Georgia)
floridana
Iroolcsi
REVISION OF THE KING SNAKES. 59
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62 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
LAMPKOPELTIS GETULUS FLORIDANA Blanchard.
KING SNAKE.
Fig. 36.
1894. Lampropeltis getulus (part) Loennbeeg, Proc. U. S. Nat. Mus., vol. 17,
no. 1003, p. 324.
—
Ophibolus getulus (part) Ditmars, Reptile Book, 1907,
p. 359.
1904. Ophibolus getulus getulus Brown, Proc. Acad. Nat. Sci., Philadelphia,
p. 472.
1919. Lampropeltis getulus floridana Blanchard, Occ. Pap., Mus. Zool., Univ.
Mich., no. 70, May 5, p. 1, pi. 1, fig. 1 (type locality: Orange Hammock,
DeSoto County (northeast portion), Florida; U. S. Nat. Mus., no. 22368;
William Palmer, collector)
.
Description.—Characters in general like those of getulus. Ventrals,
210 to 225; caudals, 42 to 55; supralabials, usually 7, sometimes 8;
infralabials, 9 or 10; 1 preocular; 2 postoculars; temporals, 2 +3 +4;
posterior chin shields about equal in length to the anterior, parallel,
and sometimes separated by a scale; loreal rectangular, higher than
long or about as high as long (in one specimen fused with post-
nasal); scale rows 23 or 21, formulae, 23-21, 21-23-21-19, 21-19.
In bodily proportions this form is closely similar to getulus; it
appears, however, to be somewhat stouter and to grow to a larger
size. The tail varies from 0.107 to 0.146 of the total length. The
largest specimen examined bore the label "Florida," and measured
1,753 mm.
Pattern similar to getulus, except that the transverse light bands
on the back are about twice as many, varying as they do from 46 to
85, with an average of 66. In distinction from getulus, the forking
on the sides has usually disappeared, and the cross bands on the back
end abruptly at about the level of the seventh row of scales. Alter-
nating with the dorsal cross bands is a series of transversely elon-
gated light spots, about two scales wide, extending up from about
the second to the seventh row of scales. Alternating with these
latter is a series of light spots about two scales long which overlap
the ends of the ventrals and the first row or two of dorsal scales.
The scales of the dark bands are often sharply tipped with black.
Furthermore, in distinction from getulus, each scale of the black
areas has a development of white, beginning at its basal end and
spreading distally to cover one-half or three-quarters of its area.
This basal lightening of the dark scales may be so pronounced as to
greatly obscure the pattern of alternating transverse bands of white.
The belly is checked with black and white. The actual colors vary
from white to creamy white to yellow, and from black to yellowish
brown.
The characters of the penis are as follows: strongly bifurcate,
the forks one-fifth the total length of the organ; sulcus single, ex-
REVISION OF THE KING SNAKES. 63
tending over the side of the larger fork, and down the inner side
of the V; distal surface smooth, succeeded proxiinally by a few
slightly fringed calyces; the short fringes rapidly becoming m-odified
into small spines which increase in size basally, and, one-half way
L
03 «/ 79
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63 6f 79
FlQ. 13.—Map SUOWINQ locality records job LAMPROPELTIS QETXn.U3 FLORIDANA.
dowTi the organ, stop abruptly; basal half of organ smooth except
for a few scattered minute spines, chiefly near the large ones; no
spines distinctly enlarged.
The skull is essentially like that of getulus. A specimen from
Orange Hammock, De Soto County, Florida (U. S. Nat. Mus., no.
64 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
22.367) has the following dentition: maxillary teeth, 15 on each side,
increasing very gradually in size posteriorly, the last three slightly
larger than those preceding; mandibular teeth 16 on each side,
somewhat larger anteriorly, decreasing in size posteriorly; palatines
strong, subequal, 10 on each side; pterygoids smaller than palatines,
subequal, 19 on the left side and 17 on the right.
Habitat and habits.—It is highly desirable that notes on the natural
history of this form be taken and reported, for comparison with the
typical form, as apparently there are no such observations in the
literature.
Range.—This form replaces getulus in Florida from about Orange
County south, probably to the Miami River in the southeast, where
it is in turn replaced by brooksi. In addition to the localities repre-
sented by specimens in the United States National Museum, specimens
belonging to other museums have been examined from Kissimmee
Prairie, Orlando, Gotha, and Vero, Florida.
Variation and affinities.—That this form is most closely allied to
getulus is plainly evident from its form, proportions, markings,
penis, skull, and scuteUation. Its pattern is obviously derived
from that of the latter by an obliteration of the narrow forks on the
sides, and an increase in the dorsal and lateral white markings. It
is uncertain just how this pattern was derived, but it is very possible
that it took place in the following way: new transverse bands were
developed from the white central scales midway between the original
bands; a lightening of each dark scale began near its base and ex-
tended from there basally, then distally, until all the area of these
scales, except the distal margin, was lightened. The isolation of
this form in the peninsula of Florida, the fact that no Lamprojpeltis
is known from the West Indies, and the juxtaposition of its range to
that of typical getulus allows of derivation from the latter species
only.
REVISION OF THE KING SNAKES. 65
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66 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
LAMPROPELTIS GETULUS BROOKSI Barbour.
Fig. 37.
1919. Lampropeltis getulus broohsi Barbour, Proc. New England Zool. Club,
vol. 7, June 5, p. 1 (type locality, 14 miles southwest of Florida City,
Dade County, Florida (near the Royal Palm State Park, formerly called
Paradise Key); Mue. Comp. Zool., no. 12456; W. S. Brooks and C. A.
Mosier, collectors).
Description.—The original description is as follows: "An adult,
1,350 mm. long. Similar to L. getulus jloridana Blanchard, in
squamation, but differing widely in coloration. Pattern so reduced
as to be almost everywhere undiscemible. Each scale dull chrome
yellow with a conspicuous, very dark brown apical spot."
Remarks.—The name hroolcsi, is based upon a single example taken
near Royal Palm State Park, and a badly crushed individual seen in
the highway near Homestead, Florida. Southern examples of
floridana show a distinct tendency toward the type of coloration of
hrooTcsi, so that the discovery in tropical Florida by Doctor Barbour
of this new race is most gratifying. It carries one step farther, and
to its logical conclusion the interesting series of pattern changes of the
getulus group in the southeast. It is to be hoped that more examples
will soon be found.
LAMPROPELTIS GETULUS YUMENSIS Blanchard.
KING SNAEK.
Fig. 28.
1861. Lampropeltis boylii conjuncta Cope, Proc. Acad. Nat. Sci., Philadelphia,
p. 301.
1865. Cornelia getulus splendida Jan, Icon. Gen. Ophid., Uvr. 12, pi. 6, fig. 1.
1901. Ophibolus getulus boylii (part) Brown, Proc. Acad. Nat. Sci., Philadelphia,
p. 78 (Yuma).
1912. Lampropeltis conjuncta Van Denburgh, Proc. California Acad. Sci.,
ser. 4, vol. 3, p. 164.
—
Grinnell and Camp, Univ. California Publ. Zool.,
vol. 17, no. 10, 1917, p. 187
—
^Hall and Grinnell, Proc. California Acad.
Sci., ser. 4, vol. 9, no. 2, 1919, p. 48.
1919. Lampropeltis getulus yumensis Blanchard, Occ. Pap., Mus. Zool. Univ.
Michigan, no. 70, May 5, p. 6, pi. 1, fig. 2, (type locality, 27 miles west
of Indian Oasis, Pima County, Arizona; type, U. S. Nat. Mus. no. 61318;
A. H. Howell, collector).
Description.—^The scutellation as indicated by 18 specimens is as
follows: Ventral plates, 212 to 240; caudals, 44 to 57 (males, 50 to
67, average 54; females, 44 to 51, average 48); supralabials, 7
(rarely 8); infralabials, 9, sometimes 10; 1 preocular; 2 postoculars,
temporals usually 2 + 3 + 4 ; posterior chin shields but little more than
half as long and half as wide as the anterior, and separated from each
other by two small scales; loreal longer than high; dorsal scale rows
usually 23-21-19, sometunes 23-25-23-21-19 or 21-23-21-19.
REVISION OF THE KING SNAKES. 67
This form seems not to differ in size or proportions from hoylii.
The tail varies from 0.090 to 0.140 of the total length (males, .121 to
.132, average, .127; females, .093 to .128, average .117). The
largest specimen examined was from the Graham Mountains, Arizona,
and measured, 1,204 mm.
The pattern is composed of white annuli separated by black, and
varying in number from 29 to 45 (average 37). It differs from
hoylii in that the most of the white scales are marked basally with
dark brown, which not infrequently extends irregularly over the
scales so as to greatly obscure the rings. The latter are narrow on
the back, 1 to 2^ scales in width, widening on the side to about 2
to 5 scales, and traversing the belly. They may be divided on the
midventral line and alternated instead of joined, and this may
occur less frequently above.
Characteristic of the group are the white bars across the prefrontals
and internasals, and the white centers on rostral, nasals, loreal,
oculars, and labials. Beneath the eye there is usually a rather con-
spicuous enlargement of the common dark border of the third and
fourth upper labials. Other markings on the head are infrequent.
There may be small white dots on supraoculars, anterior temporals,
anterior portion of frontal, and rarely on the edges of the parietals.
On the first scales behind the parietals there may be a small cluster
of white spots, the homologue of the first white band or ring, which,
in all members of the group, may at times be fairly well developed.
The belly is usually crossed by the continuations of the black and
white rings, and, like the dorsal white scales, the white ventral
plates are usually conspicuously edged basally with dark brown.
The skull is like that of hoylii. The dentition, as indicated by a
few specimens, is as follows: MaxiUary teeth, 13 to 16, the last two
somewhat stouter than those preceding; mandibular teeth, 14 to 17,
the anterior the larger; palatines, 9 or 10; pterygoids, 12 to 19.
The penial characters are very much like those of splendida. The
organ is bilobed, and has a small smooth area at the distal end. The
calyces are few, and their fringes may be fairly conspicuous or very
short. Minute spines are barely distinguishable over nearly as much
area as covered by the large spines.
To distinguish this form from conjunda and from hoylii, see the
table under the description of the latter. From splendida it may be
known by the fact that the white annuli are not joined on the sides,
but are completely separated by the black interspaces.
Hahitat and liahits.—Except for C. L. Camp's note for the vicinity
of Yuma, that this form "lives among cottonwoods and thick growth
along the river bottom," there are, apparently, no observations
available on the natural history of yumensis. Such observations
would be very desirable for comparison with those of splendida and
68 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
of hoylii. It would be interesting to know how severe desert condi-
tions it can withstand, and whether it differs in this respect from
hoylii. It may be a long time before the natural history of the various
forms of the getulus group is well known, but, when it is, we shall
expect to be able to trace courses of evolution in habits correspond-
ing with and bearing out the observations on the structural evolution
of the group.
Range.—This form inhabits the desert region of the southern third
of Arizona (except the extreme southeastern comer), the Colorado
Desert, and the area about the head of the Gulf of California.
Specimens are in the collection of the National Museum from Ash
Creek in the Graham Mountains, Arizona, at 3,200 feet elevation
(evidently the base of the mountains) and from Fort Grant and
Calva in Graham County. It will be interesting to learn whether
this form extends up the Gila River and its branches into New Mex-
ico, and to see if the divide between the Colorado drainage system
and the Rio Grande proves to be also the divide between this form
and splendida. If this proves to be the case, it will be a matter of no
small interest to see if these two forms are sharply distinct here.
This is to be expected, since none of the members of this group are
known to inhabit actually mountainous regions. Enough specimens
are available from the vicinity of Tucson to show that this is a region
of intergradation between splendid^ and yumensis. From the topog-
raphy of the country it niay be expected that yumensis ranges south
on the east coast of the Gulf of California to the region of the twenty-
eighth parallel, perhaps farther. West of the Colorado River, except
for the vicinity of Yuma, the only record is for Volcano Lake, Lower
Cahfornia, near the delta of the Colorado River, at the east base of
the Cocopah Mountains. West of the Yuma region it is assimaed that
the range of yumensis extends throughout the Colorado Desert, since
specimens from western San Diego County strongly suggest deriva-
tion from yumensis. South into Lower Cahfornia its range is probably
limited by the region where the mountains reach the east coast, at
about the thirty-first parallel. Specimens in the United States
National Museum from the northwest coast of Lower California and
from San Pedro Martir Moimtain, Lower California, are distinctly
hoylii.
Specimens have been examined from the following locaHties, in
addition to those represented by specimens in the United States
National Museum: Tucson, Fort Grant, and 10 miles below Cibola in
Yuma Coimty, Arizona; "Sonora" (southern Arizona); Pilot Ejiob
(Imperial County), California.
There seem to be no published records for other localities than
these.
REVISION OF THE KING SNAKES. 69
Variation and affinities.—The number of s})ecimens at present
available is too small to establish dependable averages. They are
well distributed, however, and a consideration of their structural
variations will be significant in relation to the intermediate or con-
necting position that this form occupies between boylii and splendida.
By reference to the accompanying table comparison may be made
of the extremes and averages of the numbers of ventral plates in
these forms for typical and for boundary regions. Southern New
Mexico and western Texas may be considered the region where
splendida is tyi)ical; the vicinity of Tucson, where this form inter-
grades with yumensis; Yuma, where yumensis is tyj^ical; and San
Diego County, Cahfornia, where boylii is typical.
From the graphic presentation of these figures (fig. 14) it will be
apparent that yumensis, in the eastern portion of its range, commonly
has numbere of ventral plates characteristic of splendida, and, in the
western portion, numbers characteristic of boylii. The nine speci-
mens from the Yuma region exliibit the ordinary variation and
usual average of boylii; those from the Tucson region exhibit a range
of variation and an average that is strongly suggestive of splendida,
and that in fact bridges the gap between the latter and boylii.
It is at Tucson as well that great variabihty occurs in the pattern,
and that all the transitional conditions are found between the pat-
terns of yumensis and splendida.
Table for comparison of splendida, yumensis, and boylii.
ipleiidida .
.
pnmensui.
boylii
Region.
I
Whole range
Southern Xcw Mexico and
v,-o^t("rn Texa?.
Tiicson and vicinity
(Tucson
i I '.raham County
[Yuma
f Wiiole range
\S&n I>iego County
Mum-
bor of
speci-
tnens
7
3
3
9
110
18
Ventrals.
Ex-
tremes.
207-225
207-22.5
210-220
212-229
2ir>-22.';
219-240
206-2.'>4
206-254
Aver-
age.
216
216
216
220
221
230
232
232
Cross bands
or rings.
Ex-
tremes.
41-85
69-85
41-71
33-37
30-35
29-42
28-49
29-41
Aver-
age.
69
A comparison of the numbers of rings or dorsal bands of these
related forms will be even more striking than the numbers of ven-
trals. From the table and diagram (fig. 15) it will be seen that the
normal lower limit of variation in cross bands of splendida (in southern
New Mexico) is above the upper limit of variation in rings of boylii.
But in the vicinity of Tucson the lower limit of splendida overlaps the
upper limit of boylii, and that, in specimens from this region in which
the pattern is so altered as to be no longer recognized as splendida,
but as yumensis, the number of bands—now changed to rings—has
70 BULLETIN U4, UNITED STATES NATIONAL MUSEUM.
i
<D OS 31
l.i^.te
"V*"
.
-
1^
/
p"
/
/
/
/
/
/
/
/
/
t
i
1
1
1
1
1
/
/
/
/
/
t
1
1
o H
% 3
S 3
REVISION OF THE KING SNAKES. 71
1
1
1
/
/
/
/
i
^
§
1
1
I
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1
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^j
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72 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
closely approximated the extremes and average of hoylii. This
range of variation in number of rings is not now altered throughout
the region occupied by hoylii, yumensis, and conjuncta.
The decisive change in the configuration of the pattern as well as
in the number of rings or bands occurs also in southeastern Arizona,
but striking as this change is, its steps are all easily recognizable,
and are explained in detail in the discussion of the evolution of the
group.
Its pattern differs from that of hoylii in that the white centers of
the dark scales, derived from splendlda, do not cover the whole of
the scale, but are mainly limited to the distal portion. Northv/ard
//p
3S
E V
//O
Fig. 16.—Map showing locality records fob Lampropeltis getulus yxxmensis.
this condition very soon changes into that typical of hoylii; that is,
the white spreads over all of the scale, producing rings of continuous
white. Westward, however, the pattern of yumensis is retained. In
the vicinity of Yuma the light rings are often heavily suffused with
brown, A specimen from near the mouth of the Colorado River, on
the west side (Volcano Lake), is still typical of yumensis.
It will be of interest, in connection with what is known of the
incomplete development of the pattern of the young of lioTbroolci, and
its possible ancestral significance, to learn whether the yoimg of
yumensis are entirely like the adults in the degree of basal shading
of the dark scales. Only three young specimens are now available.
One from Tucson looks more like hoylii, than any of the adults, but
REVISION OF THE KING SNAKES. 73
the white scales are nevertheless basally shaded with brown, and
this condition is evident on many of the ventral plates. A young
one from Fort Grant, Arizona, not far south of the range of typical
boylii, is indistinguishable from the latter species, and is identified as
yumensis only on locality. One from Fort Yuma is t>qncal of
yumensis in every respect. The evidence is of course inconclusive,
but favors the view that the young are like the jjarents. This will
be of interest in connection with the frequently noted fact that the
young of conjuncta are not like the adiUts, but like boylii. This is
in support of the distuictness of yumensis from conjuncta, and of the
derivation of the latter from boylii instead of from yumensis.
A dark subocular spot originates in this form on the upper labials.
In ty})ical splendida the head is nearly all black, but across the center
of each labial there is a narrow vertical band of white. Westward
this band widens until the labials appear white with dark mutual
borders. But beneath the eye there is a strong tendency for the
black pigment adjacent to the suture between the third and fourth
upper labials to be retained, or, if lost, to be recovered. The result
is the conspicuous dark subocular spot characteristic of yumensis,
boylii, califoi^iae, and conjuncta. Some specimens, however, from the
vicinity of Yuma have been subject to such a general increase in
pigment that this spot is not recognizable.
In brief, then, it appears that yumensis is not an mtergradational
condition between splendida and boylii, nor yet, as will be more
clearly brought out further on, is it identical with conjuncta of the
Cape Region of Lower California, but a recognizably distinct form
having a definite range that lies between the ranges of boylii and
splendida, and intergrading with both of these forms where their
ranges meet its own.
74 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
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REVISION OP THE KING SNAKES. 75
LAMPROPELTIS GETULUS BOYLU (Baird and GIrard).
unx snake; botle's kino snake; black and vhitk kino snake.
Fig. 27.
1853. OpMbolua boylii BArap and Girahd, Cat . N. Amer. Kept., pt. 1, Jan.,
ifr p. 82 (type locality,"lll Dorado County, California; type, U. S. Nat. Mus.,
noTiegS; Dr. C. C. Boyle, collector).—Baird, U. S. and Mex. Bound.
Surv., 1859, p. 20; Pacif. R. R. Surv., vol. 10, pt. 3, no. 1, 1859, pi. 30,
fig. 57; same, vol. 10 (Williamson's Route), pt. 4, no. 4, 1859, p. 11.
—
Cope, Proc. Acad. Nat. Sci. Philadelphia, 1866, p. 305.
—
Lampropeltia
boylii, same, 1860, p. 255.
—
Cooper, Proc. California Acad. Sci., vol. 4,
1873, p. 79.—Stejneger, N. Amer. Fauna, no. 7, pt. 2, 1893, p. 204.—
Van Denburgh, Proc. California Acad. Sci., ser. 2, vol. 5, 1895, p.
1006; Proc. California Acad. Sci., ser. 2, vol. 5, pt. 3, 1896, p. 1006;
Occ. Papers California Acad. Sci., vol. 5, 1897, p. 169.—McLain,
Coll. Rept. W. Coast U. S., 1899, p. 11.—Meek, Field Columb. Mus. Publ.,
no. 104, vol. 7, no. 1, 1906, p. 15.
—
Grinnell and Grinnell, Throop Inst.
Bull., no. 35, 1907, p. 41.
—
Grinnell, Univ. California Publ. Zool., vol. 5,
no. 1, 1908, p. 165.
—
Van Denburgh, Proc. California Acad. Sci., ser. 4,
vol. 3, 1912, p. 150.
—
Van Denburgh and Slevin, Proc. California
Acad. Sci., ser. 4, vol. 3, 1913, p. 415.—Camp, Univ. California Publ.
Zool., vol. 12, no. 17, 1916, p. 531.—Pemberton, Condor, 1916, p. 233.—
Grinnell and Camp, Univ. California Publ. Zool., vol. 17, no. 10, 1917,
p. 186.
—
^Bentlet, Copeia, no. 75, 1919, p. 90.
—
Coronella boylii Jan,
Arch. Zool. Anat., vol. 2, fasc. 2, 1863, pp. 238, 246.
1853. Coronella balteata Hallowell, Proc. Acad. Nat. Sci. Philadelphia, vol. 6,
p. 236 (type locality: California); Pacif. R. R. Surv., vol. 10, pt. 4, no. 1,
1859, pp. 14, 24, pi. 5.
1863. Coronella pseiuiogetulus Jan, Arch. Zool. Anat., vol. 2, faec. 2, pp. 238, 247.
1865. Coronella getulus pstudogetuhis Jan, Icon. Gen. Ophid., livr. 12, pi. 6,
fig. 2 (a-g).
1875. Ophibolus getulus boylii Cope, Bull. U. S. Nat. Mus., no. 1, p. 37.—Yarrow,
Rep. Geog. and Geol. Explor. Surv. W. 100th Mer., vol. 5, chap. 4, 1875,
p. 538.—CouES, same, chap, 5, 1875, p. 618.
—
Coues and Yarrow,
Bull. U. S. Geol. and Geog. Surv. Terr., vol. 4, art. 11, 1878, p. 283.—
Yarrow, Bull. U. S. Nat. Mus., no. 24, 1882, p. 92.—Cope, Proc. Acad.
Nat. Sci. Philadelphia, 1883, p. 29.—Garman, Mem. Mus. Comp. Zool.,
vol. 7, no. 3, pt.'l, 1883, pp. 69, 157.—Townsend, Proc. U. S. Nat.
Mus., vol. 10, 1887, p. 239.—Cope, Proc. U. S. Nat. Mus., vol. 14, 1891,
p. 613; Rep. U. S. Nat. Mus. for 1898, 1900, p. 919, fig. 230.—Brown, Proc.
Acad. Nat. Sci. Philadelphia, 1901, p. 78.—Ditmars, Reptile Book, 1907,
pp. 341, 363, pi. 103, fig. 9, pi. 1,08 (lower fig.); Rept. of World, 1910,
p. 271.
1894. Coronella getula (part) Botjlenger, Cat. Snakes Brit. Mus., vol. 2, p. 198.
1913. Lampropeltis boylei Atsatt, Univ. California Publ. Zool. , vol. 12, no. 3, p. 41.
1917. Lampropeltis boylii boylii Stejneger and Barbour, Check List, p. 87.
W
D
escription.—^The scutellation of the^Pacific Coast representative
of the getulus group may be defined as follows plates, 206Jo
254 ; caudars741 to 62; supralabials, 7, infrequently 8; infralabials 9,
often 10; oculars^ 1 and 2; temporals, 2+3+4; posterior chin shields
only about two-thirds as long^s the anterior, parallel, and separated
186650-21—BulL 114 C
76 BULLETIN lU, UNITED STATES NATIONAL MUSEUM.
by two small scales; loreal usually longer than higli, less often about
as high, as long; scale rows on middle of body, 23 or 25, formula
usually 23-21-19, or 21-23-21-19, often 23-25-23-21-19.
This is a somewhat smaller and slenderer form than the eastern
representatives of the group. In proportions it is close to splendida.
The taU varies from 0.107 to 0.145 of the total length (average for
males, .130, for females, .120). The largest specimen examined was
from Tehama County, California, and measured 1,280 mm.; the next
largest was 1,234 mm., from "California."
The pattern is of white rings, 28 to 49, often disposed more or less
obliquely, and separated by a ground color of black or dark brown; the
rings widen rather suddenly on the sides, from one to two scales
on the middorsal line to three to six on the first row of scales, and
cross the abdomen as broad bands. In distinction from yumensis
and conjuncta, the scales of the white rings are white to their bases.
However, the basal shading, characteristic of these forms, may
appear at almost any point in its range, and for identification in
such cases study should be made of the pattern and scalation of the
head (see Table of Comparisons, p. 77). The markings on the head
are typically like those of yumensis, but further white spotting may
be developed, particularly along the coast, and examples are not
infrequent in which there is more white on the head than in conjuncta.
The copulatory organ may be described as foUows: Bilobed;
sulcus single, extending over one of the lobes, and ending in a small
bare space; calyces strictly apical, fringes few, distinct or very short,
passing into spines; latter close set, somewhat larger toward the
base, covering from a third to less than half of the distal end of the
organ, suddenly replaced by a few minute spines which soon dis-
appear altogether, or which may not be present at aU. A specimen
from Washington County, Utah, shows a few pits as traces of the
minute spines, but these are not evident at all in one from Overton,
Nevada. Another from southern Nevada has the large spines pass-
ing rapidly, but not abruptly, into minute spines, and the latter
then soon disappearing. Specimens from Palo Alto show minute
spines only very close to the large ones.
The dentition of hoylii is as follows: MaxiUaries 13 or 14, subequal
except the last two or three which are slightly stouter; mandibulars,
14 or 15, longer in front, diminishing in size posteriorly; palatines,
8 to 10, usually 9, subequal, a little larger than the pterygoids; latter
13 to 17, diminishing posteriorly.
Although there is usually no difficulty in correctly identifying
specimens of hoylii, yumensis, and conjuncta, nevertheless puzzling
individuals will be found. Adult examples of yumensis and conjuncta
need never be confused, but specimens of hoylii can be foimd to
bridge all the distinctions. If the locality of a specimen is imknown.
REVISION OF THE KING SNAKES. 77
then reliance should be placed on the totality of characters. Adult
specimens which show no basal shading of the scales of the white
rings can be only hoylii. If basal shading is present, hoylii may be
distinguished from conjuncta by the fact that the infralabials are
usually 9 instead of 10, the frontal plate has usually a right or obtuse
posterior angle instead of an acute one, and its outline is distinctly
pentagonal instead of nearly trigonal. However, too much reliance
should not be placed upon these last two characters. If the number
of ventral plates is well above the average for yumensis and conjuncta,
the specimen is apt to be a hoylii. Adult examples of hoylii may
resemble yumensis, but if the head is much spotted posteriorly and
particularly if the wliite bars across the prefrontals are strongly
convex beliind and occupy more than half their area, it is a hoylii.
The accompanjdng table should prove useful in placing doubtful
examples, but since the distribution of these forms is fairly well known,
specimens may, with safety, be referred to their proper form by
locality only.
Table of comparisons.
Character. Yumenait. Boplii. Conjuncta.
Usually 9 Usually9 Usually 10.
Frontal Pentagonal More nearly trigonal.
Posterior angle
Postero-lateral angle .
.
Winterings:
Adults
Right or obtuse Usually distinctively
Obtuse
acute.
Nearly or quite 180 de-
grees.
Wliite scales always White scales only oc- White scales always dark
dark brown at base. casionally dark brown
at base.
No basal shading of
white scales.
brown at base.
Young Usually like adults Xo basal shading of white
scales.
Head: Parietals, frontals, White spots very re-
stricted or absent.
Usually like yumemif, Numerous wliite spots or
supraoculars, second except in southern marks on these plates.
and tliird rows of temp- California and north-
orals: Adults and young. ern lower CaUfornia,
where it is usually like
conjuncta.
Usually rounded or an-Wliite bars on prefrontals: Usually simple bars oc- Like bof/lii.
Adults and young. cupying less than half gular" posteriorly, and
the area of the scute. occupying more than
half the area of the
scute.
Habitat and Tiahits.—^Although this form has been known since 1853
there is very little recorded about its habits and habitat preferences.
Townsend (1887, 239) refers to it as "not uncommon on the Lower
McCloud, in damp places near the river." Hallowell (1859, 14) says
of it, "found in valleys and open prairies, very abundant, often
killed by travelers, and found lying on the roadside." A specimen
from Marin County, California, bears the note "by roadside in shade."
Merriam reports it (Stejneger, 1893, 204) as follows: "Found in the
valley of the Lower Muddy near an abandoned mill at Overton,
Nevada, where several were secured in dense thickets of Atriplex
torreyi. About dark they began to emerge from these retreats,
78 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
making a great noise in crawling over the dry leaves, and were soon
found in the open." Van Denburgh (1897, 171-2) says "The black
. and white king snake is most abimdant where the country is covered
with chaparral and where small streams are numerous." Grinnell
and Grinnell (1907, 41), reporting for Los Angeles County, California,
say "it is one of our least common snakes," and that they have
"met with it only in the upland regions bordering the foot hills."
That it occurs in the foot hills, along streams and small canyons
throughout its range, and not in the high mountains and on the open
deserts, seems evident from the material at hand and the few references
in the literature.
Its disposition Hallowell reports (1859, 14) as "timid, always
endeavoring to escape its pursurers." Van Denburgh (1897, 172)
says "it is usually very gentle, but sometimes fights its captor most
fiercely, rarely however, being able to draw blood with its small teeth."
Concerning its food the last-named author (1897, 172) writes
"I have twice found it swallowing the contents of quails' nests, and
once observed one crawhng along the ground and looking up into the
bushes for nests of small birds. Several times while I watched, its
quick eyes detected nests 3 or 4 feet above it, but although the snake
immediately chmbed up to these, it did not obtain a meal, for the
nests which it examined had been abandoned by their builders or
robbed by some earlier comer.
"While I was watching a man spade up a small plot of ground,,
he killed two gophers (Thomomys) and threw them a few feet away.
A few minutes later a snake of this species appeared, went directly
to the spot where the gophers lay side by side, and swallowed first
the adult and then the half-grown one. It took no notice of our
presence, and after completing its hearty m.eal disappeared in the
direction whence it had come."
A specimen in the collection of the University of California is pre-
served in the act of swallowing a rattlesnake about 2 feet long.
So far as known, therefore, hoylii, in disposition, food, and habitat
preferences, is very much like getulus, the only other member of the
group that is at all well known.
Range.—This form is known from Cape San Quintin and San Pedro
Martir Mountain in Lower California, north in California to the
forty-first parallel, and thence southeastward across western and
southern Nevada and southwestern Utah to the latitude of Phoenix
in Arizona. It appears to be generally distributed throughout the
range, except for the driest deserts and the mountains above 5,000
feet. Altitudes at which specimens haVe been taken are frequently
recorded on the accompanying labels and in the literature. The
highest records are for 4,500 feet; one specimen was taken by Mr.
Frank Stephens in the Providence Mountains, California, and another,
REVISION OF THE KING SNAKES. 79
in the Brooklyn Museum, was taken at Bellevue, Washington County,
Utah. Other altitude records are as follows: Hemet Lake, San
Jacinto Mountains, 4,400 feet; Wild Rose Spring, Panamint Moun-
tains, 4,060 feet; Hackberry, Arizona, 3,500 feet; Grand Canyon of
the Colorado, 3,000 feet; south of Mount Sanhedrin, Mendocino County,
California, 2,340 feet; base of the San Jacinto Mountains, 2,200 feet
and 1,700 feet; Santa Ana Canyon, west side of San Bernardino Moun-
tains, 2,000 feet; Eldorado County, 2,000 feet; Jarupa Mountains,
1,200 feet; Turtle Mountains, 1,000 feet; Sonoma County, California,
1 ,000 feet. No other specific records are for more than a thousand feet
.
Grinnell and Camp (1917, 186) record it as distributed "throughout
the southern and central parts of the State [California], except on
the high mountains (above 6,000 feet altitude)." But, as noted
above, the highest actual record is 4,500 feet. Since snakes are
found in California at much greater altitudes than this, notably the
coral king snake, multicinda, the absence of loylii from the moun-
tains must be considered as the expression of a habitat preference
or perhaps restriction, and this is quite in accord with what is
known of the other members of the group.
Concerning its ability to live in the deserts, less is known. From
the desert region between Yuma and Tucson only yumensis has been
taken. There is no record of either form from the Mohave Desert
or Death Valley. The Death Valley expedition of the United States
Biological Survey collected in these and other desert regions of the
vicinity but obtained hoylii only in the vicinity of streams and at the
bases of mountains.
Besides the localities represented by specimens in the United States
National Museum, specimens belonging to other museums have been
examined as follows: In California—Alameda County, Asuza and
Bairdsto\^Ti (Los Angeles County), Banning (Riverside County),
Beveridge Canyon on east slope of Inyo Mountains, Cabazon (River-
side Coimty), Camx^o (San Diego County), Cazadero and El Varano
(Sonoma (bounty), Forrest Lake (San Joaquin County), Gadwall,
(Merced County), near Guernevillo (Sonoma Coimty), Ilemet Lake
(San Jacinto Mountains), Irishes (Mendocino County), Jarupa Moun-
tains, La Puerta Valley (San Diego County), Los Angeles, near Mount
Hamilton (Santa Clara County), Mount Sanhedrin CNIendocino
County), Needles and Ontario (San Bernardino County), Owens
Valley (Inyo County), Pasadena (Los Angeles County), Piru (Ven-
tura County), Pleasant Valley (Mariposa County), Redondo Beach
(Los Angeles County), Riverside, Rumsey (Yolo County), San
Anselmo (Marin County), San Jacinto, Springs (San Luis Obispo
County), Tehachapai Mountains (Kern County), Tehama (Tehama
County), Turtle Mountains, Vacaville (Solano County), Vallecito
and Vallecito Creek (San Diego County), Vincent Creek, Wheatville
80 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
(Fresno County), Wild Kose Spring (Panamint Mountains); in Ari-
zona—Cave Creek (Maricopa County), Fort Verde (Yavapai County),
Cottonia (Mohave County) ; in Utali—Bellevue (Washington County) ;
in Nevada—Current school district in Nye County; in Lower Cali-
fornia—Cape Colnett, Cape San Quintin, San Salado, and the foot-
hills of San Pedro Martir Mountain.
Published records for other localities are as follows: Arizona
—
Date Creek, 50 miles south of Fort Whipple (Coues, 1875, 618), Gila
Desert (Baird, 1859, 20); California—^Tujunga Wash, east of San
Fernando (Grinnell and Grinnell, 1907, 41), Lower Santa Ana
Canyon (Grinnell, 1908, 165), and the following records by Van Den-
burgh (1897, 171), Oakland, Mount Tamalpais, Camp Taylor, Santa
Margarita, Healdsburg, Applegate; Nevada—St. Thomas (Clark
County) (Van Denburgh, 1897, 171), Pahranagat Valley (Stejneger,
1893, 204), "El Paso Creek" and "Bernicia" (Hallowell, 1859, 14).
In addition, the following localities in California are given on the
authority of Mr. Carl L. Hubbs: Bed of the Nacimiento River near
Jolon, Monterey Coimty; road between San Luis Obispo and Avila,
San Luis Obispo County; and near Fall Brook, San Diego County.
Variation.—In spite of similarity in general appearance throughout
its range, hoylii is very variable in all of its specific characters The
range includes very diverse conditions of topography and climate.
The long, high ridge of the Sierra Nevada extends from near its
northern limits southeastward for nearly 500 miles, splitting the
range into two sections—an east and a west. That this is a natural
barrier to its east and west migration in this region seems fairly cer-
tain, and the Mohave Desert, just south of the Sierras, may perhaps
continue the barrier where the mountains leave off—at least there
is no record of a loylii from this desert, although it was found by the
Death Valley Expedition in numerous other places.
Whether or not the Mohave Desert is or ever was a barrier to the
westward migration of hoylii, it is certain that this form must have
obtained entrance to the Great Valley of California, either from around
the south end of the Sierras or from the north. There is certainly
no reason for excluding its entrance from the south, as Imperial and
Riverside Counties offer no obstruction to yumensis from the lower
Colorado region, and, as has already been brought out m the dis-
cussion of the latter form, yumensis undoubtedly intergi'ades with
boylii in southern California.
In the north the situation is different. The Sierras continue to be
a barrier to very near the northern limit of the form. That it is not
adapted by nature to range much north of here is indicated by com-
parison with the northward distribution of its allies, Jiolbroolci and
getulus. The latter is not known north of the forty-first parallel, and
that the former extends somewhat farther north is due to a more
REVISION OF THE KING SNAKES. 81
favoring topography. The present distinctness of the east and west
sections of hoylii, the availability of the southward route to the Great
Valley, and the improbability of a northward one argue for the greater
probability of the former route to northern and western California.
This suggests the hypothesis that hoylii was differentiated from
yumensis in southern Arizona and spread from there north to Utah,
Nevada, and eastern California, and that it was likewise differen-
tiated from yumensis in southern California and from here migrated
north throughout California west of the Sierras, and south to Cape
Ventral
254
—
260
,,-- ' "-;^
—
246
/
/'
242
—
—
r—
1
1 .'
.^
—
238
(
1 /
i--''
--'
/''
234
/
_
J
,-''
.-- . -
^•'
' '^
230 —1
.„-
---
--"
'l
\
226
/''
1
222
-'"'
218
—
210
Tuc- Gra- Gen- Hack- Nee- Wild Inyo Over- Yu- San San Los Ava- Ft. Tu- Fres- San Shas-
son, ham tral berry, dies, Rose Mts.,ton, ma, Diego, Ja- An- Ion, Tejon, lare no, Fran- ta
Ariz. Mts., Ariz. Ariz, etc., Spring, Galif. Nev. Galif. Galif. cinto, geles, CaUf. Kern Co., Calif. Cisco, Co.,
3 Ariz. 11 1 Calif. CaUf. 1 4 and 18 Calif. Calif. 2 Co., CaUf. 15 Calif. Calif,
3 5 1 Grand Ariz. 11 13 Calif. 3 31 7
Localities, and numbers op Canyon, 9 3
SPECIMENS. Ariz.
1
Fio. 17.—Diagram showing geographic variation in numbers op ventrai, plates in Lampropeltis oetulus
YUMENSIS, AND L. G. BOYLH.
San Lucas. If there is any truth in this theory, there should be some
confirmation from a study and comparison of variations in scutellation
and pattern.
Examination of figures 17 and 18, on which have been plotted the
extremes and averages of the ventral plates and the white annuli for
numerous localities, brmgs out the following interesting relations:
(1) All examples of loylii from western and northern Arizona, Utah,
Nevada, and eastern California average much higher in number of
ventrals and almost distinctly higher in number of rings than exam-
82 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
N
•
y
1
1
1
1
1
1
1
N
1
1
s /
s
>
\
\
\
\
/
/
/
>
1
ofP—'"VT
1
1
•
II
ij''
\
\ *
.
»
il
•
\
^S
1 V
1
^^in
,
/
/
• /
>/
aa.
go**
o rv."
2 §.92
<"&!>-'
w^-^
a . B 2
to M
REVISION OF THE KING SNAKES. 83
pies from western and southern California, and that these higher
averages are bridged by a gradual change, through central Arizona,
from yumensis in the south. (2) Proceeduig west from Tucson there
is an increase in number of ventrals to the average and approximate
extremes characteristic of hoylii west of the Sierras, while the number
of rings, already the same as the average for hoylii as soon as yumensis
was differentiated from splendida, remains the same. The break
from Yuma to San Diego County is slight. (3) North from here
there is a progi'essive decrease in number of ventrals and increase in
annuli to the vicinity of Los Angeles. Specimens from this region
are sometimes distinguishable on sight by the evidently more nu-
merous and narrower white bands—about one scale wide dorsally,
and three on the first row of scales. It seems probable that these
specimens are also more often brown than black, but all those we
have examined were preserved. It should also be mentioned here
that two examples from Santa Cataltna Island are strikingly charac-
teristic of those from the adjacent mainland—^with a high number of
rings and a low number of ventrals. (4) The vicinity of Los Angeles
is doubtless west of the direct route of migration, as the normal
number of ventrals and rings is exhibited by specimens from
Kern County. (5) Thirty examples from the immediate vicinity of
San Francisco show that the form is characteristically developed
there, but leaves unexplained the high averages of the Fresno speci-
mens. (6) A small number of specimens from the vicinity of Shasta
County indicate that the tendency here is toward a decrease in
number of ventrals and an increase in rings (like Los Angeles speci-
mens). In fact, no specimen from here has even as many ventrals
as the average for San Francisco, while the lowest number found here
is not reached elsewhere in CaUfornia, except by a single aberrant
individual from San Diego County.
The situation with respect to the labials is shown by figure 19.
This indicates that on the west coast there is a uniform decrease in
niunber of infralabials from south to north. In this there may be
some significance. The change from Ymna to San Diego County is
too slight to be very significaiit, but from this region northward a
distinct retrograde tendency is evident attaining in northern Cali-
fornia the 7-9 formula characteristic of the genus; southward there
is an increase in the lower series that seems to have become distinctive
in individuals from the Cape Region. Specimens from Arizona are
too few to indicate the tendency northward from the range of
yumensis.
Further confirmation of the distinctness of the east section of
hoylii from the west is to be had by an examination of the pattern and
markings. Specimens from the east section retain the head markings
essentially as developed in yumensis. The white markings are
84 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
increased in size but not in number. The head thus has a clean-cut,
trim appearance with its well-defined black and white rings, and the
latter are nearly always at least a scale and a half wide on the mid-
dorsal line. The eastern section, on the other hand, is much more
variable, perhaps correlated with the much greater diversity in
environmental conditions. Basal shading of the white scales occurs
sporadically in all parts of its range, from Tehama County to San
Diego. Frequently numerous additional white markings are devel-
oped on the head. As already noted, the number and width of the
white rings is altered locally.
In brief, hoylii is characterized east of the Sierras by a high average
number of ventrals, by a somewhat greater number of rings, and by
constancy in head and body pattern; while west of the Range it is
characterized by a lower number of ventrals and a much greater
Number
of labials.
Infralabials 9
S
-. —
—-'
z c: c
Supralabials 7
6
h
-
z
:.
Shasta San Fresno, Los San Northern Cape
Co., Fran- Calif. Angeles, Diego Lower San
Calif. cisco, calif. Co.,
Calif.
Calif. Lacas.
Yuma. Tucson.
Fig. 19.—Diagram showing geographic variation in numbers of labl4x plates in Lampkopeltis
gettjlx7s botlii, l. g. yumensis, and l. g. conjctncta.
individual and local variation in other respects. It is not considered
that these differences are of subspecific value because both sections
of 'boylii were produced gradually by evolution from the same form
and in the same way, and there has been developed no recognizably
constant distinction in pattern, but it does seem to support strongly
the view that the Sierra Nevada Mountains form a natural barrier
that has forced hoylii to develop in two distinct sections and that the
Great Valley of California has been entered only from the south.
Affinities.—^The preceding discussion of the variations and their
significance leaves necessary here only a summary of such points as
indicate its affinities.
The eastern section of hoylii is related directly to yumensis, because
(1) the distinction in number of ventrals and number of rings is
REVISION OF THE KING SNAKES. 85
bridged in central Arizona; (2) the head pattern shows no develop-
ment not initiated in yumensis; (3) the change in coloration of the
white rings is effected in the vicinity of the Florence River, the
approximate boundary between the ranges of the two forms.
The western section of hoylii approaches yumensis in southern
California, and from here are developed new, although sometimes
Hg. 20.—Map showing locality records for Lampbopeliis getulus bovlii.
minor and local changes, northward and southward. Comparison
of variations and consideration of topography make it fairly evident
that the Great Valley of California has been populated from the
south and not from the east or from the north.
The relations of hoylii to conjuncta and to califomiae are discussed
under those forms.
86 BULUETIN 114, UNITED STATES NATIONAL MUSEUM.
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REVISION OF THE KING SNAKES. 87
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REVISION OF THE KING SNAKES. 89
LAMPROPELTIS GETULUS CONJUNCTA (Cope).
Fig. 21.
1860. Lampropeltis boylii Cope, Proc. Acad. Nat. Sci. Philadelphia, p. 255.
1861. Lampropeltis boylii conjuncta Cope, Proc. Acad. Nat. Sci. Philadelphia,
pp. 301, 305 (type locality. Cape St. Lucas, Lower California; cotypes (3),
U. S. Nat. Mus.. no. 5288; J. Xantus, collector).
—
McLain, Contr. Neotr.
Herp., 1899, p. 5.
—
Stejneger and Barbour, Check List, 1917, p. 87.
1875. Ophibolus getulus conjunctus Cope, Bull. U. S. Nat. Mus., no. 1, pp. 37,
92; no. 32, 1887, p. 78.
1882. Ophibolus getulus boylii (part) Yarrow, Bull. U. S. Nat Mub., no. 24, p.
92 (Cape St. Lucas, La Paz).
—
Belding, West Amer. Sci., vol. 3, no. 24,
1887, p. 98.—Cope, Rep. U. S. Nat. Mus. for 1898, 1900, p. 920.
1895. Lampropeltis conjuncta Van Denburqh, Proc. California Acad. Sci., eer.
2, vol. 5, pt. 1, p. 142.
Description.—Ventrals, 228 to 240; caudals, 48 to 54 (males, 53
to 54; females, 48 to 51); supralabials, 7; infralabials usually 10,
sometimes 9 or 11; oculars, 1 and 2 ; temporals normally 2 + 3 + 4
;
posterior chin shields about half as long and half as wide as anterior,
separated from each other by two small scales; loreal longer than
high; frontal and parietals somewhat more elongate than in hoylii
and yumensis; scale rows on middle of body 23 or 25.
In size and proportions this form is nearest like loylii; it seems,
however, to be somewhat smaller and more slender, and to have a
more elongate head. The tail varies from 0.115 to 0,132 of the total
length (males, 0.122 to 0.132, average 0.126; females, 0.115 to 0.122,
average, 0.119); more specimens may show that these limits are
exceeded, and that the averages are practically the same as for the
rest of the getulus group. The largest specimen examined measured
965 mm.
The pattern is in general like that of hoylii and yumensis, that is,
white annuli on a dark brown or black ground color (fig. 21). Like
yumensis, the white scales are edged basally with dark brown, except
that the basal darkening is not always evident on some of the scales
of the first one or two rows. None of the specimens examined
shows any tendency for the brown to overspread all of the white,
like some examples of yumensis from the Yuma region. Young
specimens show no basal shading of the white scales. The rings are
generally complete on the belly, but may be more or less imperfectly
alternated; sometimes dark spots are developed on the belly in the
white areas. Unlike yumensis and the great majority of specimens of
hoylii, the head, behind the prefrontals, is much lightened with white
spots. The frontal usually has a roughly triangular white mark, or
some fragments of this mark, located somewhat anteriorly. The
supraocular has anterior and posterior white dots, which may be
joined. The parietals have a lateral white dash, and on ten of the
eleven specimens examined there is a white dot on each parietal near
90 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
the common suture. There is a large white dot on each temporal of
the first two rows and often on some of the temporals of the third
rows. The light spots on the dorsal scales close behind the parietala
are more extensively developed than is usual with hoylii or yumensis.
The anterior headplates have large light marks, as in hoylii, and
the mutual borders of the labials are dark; like this form, too, the
subocular dark spot is well developed. Since the young specimens,
as well as the adults, show these head markings, they may thus be
Fig. 31.—Color pattekn or Lampeopeltis getulus conjtjncta (U.S.N.M. no. 5288, onb or 9W»
COTYPES). About IJ x nat. size.
usually distinguished from the young of hoylii and yumensis, and the
presence of 10 infralabials would be confirmatory.
The dentition is as follows: Maxillaries, 12 to 15, usually 13 or
14, subequal, the last two barely stouter than the preceding; man-
dibular, 14 to 16, usually 14, longer in front, decreasing in size pos-
teriorly; palatines, 8 to 11, usually 9, larger than the pterygoids;
latter 15 to 18, diminishing in size posteriorly.
The copulatory organ seems to be essentially like that of hoylii.
Minute spines are present only as traces just below the large ones;
the fringes are very short, and perhaps slightly more numerous than
in hoylii.
REVISION OF THE KING SNAKES. 91
Habitat and habits.—Nothing is recorded on the natural history of
this form.
Range.—Conjunda is at present known only from the "Cape
Region" of Lower California, from La Paz to Cape San Lucas.
No published records for other localities than those given with the
list of specimens have been noted.
Variation.—Little variation is to be expected in this form as at
present known, and, as brought out in the description, the speci-
mens examined are homogeneous in every respect. It will be noted
that in general the scutellation is close to the mean for boylii, but dif-
ferences are to be seen in the much greater frequency of ten infralabial
plates than nine, in the more acute posterior angle of the frontal
plate, in the more elongate parietals, in the narrower head, in the
numerous light spots on the posterior portion of the head, and in
the basal shading of the white scales, and it is on these points that
this form is regarded as distinct.
Affinities.— Conjunda has hitherto been considered as identical
with what we have called yumensis. Cope first noticed the basal
shading of the white scales (1860, 255) and later (1862, 301) called
attention to the similarity between specimens from Cape San Lucas
and Fort Yuma. He then based the name conjunda on the speci-
mens collected by Xantus at Cape San Lucas, and since that time
they have been regarded as practically identical with those from the
Yuma region. Tl is perhaps already plain why the separation has
been made, but it may be well to summarize the situation. It is
here held as certain that yumensis is derived directly from splendida,
and that boylii is likewise derived from yumensis. It is believed, for
the following reasons, that conjunda is more closely allied to boylii
than to yumensis.
1
.
The head markings of conjunda are rather frequently developed
in part, and sometimes in entirety, in specimens of boylii in various
parts of its range, but especially in the San Diego region and in other
localities along the coast, but not in any specimen of yumensis yet
examined.
2. The ranges of conjunda and yumensis apparently do not meet,
but are separated in northern Lower California by a southward, and
perhaps somewhat eastward extension of the range of boylii.
3. The similarity in the basal shading of the light scales of conjunda
and yumensis is not necessarily an indication of close afiinity. Indi-
viduals of boylii from most diverse localities occasionally show this
shading in as marked a form as it appears in conjunda or typical
yumensis. In boylii it must be regarded as an instance of reversion,
a situation that is duplicated frequently in other members of the
getulus group.
186550—21—Bull. 114 7
92 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
4. The young of yumensis, with possible exceptions near the
northern and western limits of its range, are like the adult, and if
there had never been a loylii-stsige between this form and conjunda,
the young of this latter would also be like the adult. But young
examples of conjunda are like hoylii in the character of the white
rings, their scales only with age attaining the basal shaduig, while
in head pattern and in shape of frontal they are like a variation of
hoylii that is not uncommon in some coastal portions of its range,
but not like any variations of yumensis.
The basal shading of the white scales of conjunda, and the same
condition as an individual or possibly local variation in loylii, is there-
fore to be regarded as a case of reversion to an ancestral type—analo-
gous to the development of white centers or bases on the dark scales
oijloridana.
5. If, as noted elsewhere, nitida attained the Cape Region from
southwestern California, there is then no reason why conjunda may
not have done so, too.
From the above considerations the following hypothesis is therefore
presented to explain the derivation of conjunda. Yumensis retained
its identity in the desert region of southern Arizona and the head of
the GuLf of California, but northward, westward, and perhaps south-
westward, it developed into hoylii. The latter migrated south into
the desert of Lower California west of the mountains, and became
altered, somewhere in the peninsula, into conjunda, partially by the
development of new characteristics, or the accentuation of certain
variations, partially by reversion to the pattern of its desert ancestor,
yumensis.
REVISIOIT OF THE KIN"G SNAKES. 93
S 8
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94 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
LAMPROPELTIS CALIFOENIAK CALIFORNIAE (Blainville).
Fig. 22.
1835. Coluber (Ophis) californiae Blainville, Nouv. Ann. Mus. d'Hist. Nat.,
Paris, vol. 4, p. 292, pi. 27, figs. 1, la, 16 (type locality, California; M.
Botta, collector).
—
Baird and Girard, Cat. N. Amer. Kept., pt. 1, 1853,
p. 153.
—
Coronella californiae Dumeril and Bibron, Erp. Gen., vol. 7,
pt. 1, 1854, p. 62S.—Ophibolm californiae Cope, Bull. U. S. Nat. Mus.,
no. 1, 1875, p. 37; Rep. U. S. Nat. Mus. for 1898, 1900, pi. 19, fig. 8.—
Lampropeltis californiae Van Denburgh, Occ. Pap. California Acad.
Sci., vol. 5, 1897, p. 172.—McLain, Crit. Notes Coll. Kept. W. Coast
United States, 1899, p. 11.
—
Van Denburgh, Proc. California Acad.
Sci., vol. 3, 1912, pp. 149, 151.—Grinnell and Camp, Univ. California
Publ. Zool., vol. 17, no. 10, 1917, p. 187.—Stejneger and Barbour,
Check List, 1917, p. 87.
—
Hall and Grinnell, Proc. California Acad.
Sci., ser. 4, vol. 9, no. 2, 1919, p. 54.
1861. Coronella getuliLS californica Jan, Icon. Gen., Livr. 14, pi. 5, fig. 3.
1863. Coronella californica Jan, Arch. Zool. Anat., vol. 2, fasc. 2, pp. 238, 246.
1882. Ophibolus getulus eiseni Yarrow, Proc. U. S. Nat. Mus., vol. 5, p. 439 (type
locality, Fresno, California; type, U.S.N.M., no. 11788; G. Eisen, col-
lector); Bull. U. S. Nat. Mus., no. 24, 1882, p. 94.
1883. Ophibolus getulus californiae Garman, Mem. Mus. Oomp. Zool., vol. 8^
pt. 1, p. 157.—Cope, Proc. U. S. Nat. Mus., vol. 14, 1891, p. 614; Rep.
U. S. Nat. Mus. for 1898, 1900, p. 922, fig. 231.—Brown, Proc. Acad.
Nat. Sci. Philadelphia, 1901, p. 78.—Ditmars, Reptile Book, 1907,
pp. 341, 363.
1894. Coronella getula (part) Boulengbr, Cat. Snakes Brit. Mus., vol. 2, p. 198.
Description.—The scutellation of this form differs from that of
hoylii only in that the range of variation appears to be less. It
may be described as follows: ventrals, 229 to 241, average 233;
caudals, 47 to 60 (males 55 to 60, average 57; females 47 to 53, aver-
age 50); supralabials, 7, rarely 8; infralabials, 9 or 10, sometimes 8;
oculars 1 and 2 ; temporals, 2 + 3 + 4 ; posterior chin shields much
shorter and narrower than the anterior, and separated from each
other by two small scales; loreal longer than high (rarely fused with
the prefrontal); scale rows commonly 23-21-19 (although sometimes
25 rows are developed near the middle of the body, and less fre-
quently a row is dropped on each side for a brief space anteriorly).
In size and proportions this form does not differ from hoylii.
The tail varies from 0.113 to 0.140 of the total length (males, 0.128
to 0.140, average 0.133; females, 0.113 to 0.123, average 0.120).
The largest specimen examined was 1,233 millimeters long and was
from Julian, San Diego County, California.
The pattern is very different from that of any other king snake.
The ground color above is dark brown or black. A bright yellow
or white stripe, about two scales wide, extends along the middorsal
line from a few scales behind the head to the tip of the tail. This
stripe is sometimes interrupted, most often close behind the head,
in the region of the vent, and on the tail. The scales adjacent to
REVISION OF THE KING SNAKES. 95
the middorsal stripe are wholly dark brown or black, and this con-
dition may obtain as low as the second or first row of scales, but
the scales of the lower rows nearly always have white centers which
are smaller dorsally and larger ventralty. These white centers
may extend as high as the seventh row of scales, or no higher than
the first. Usually the scales of the first row or two are all white.
The belly is usually white or yellow with a development of black,
laterally, on the posterior edges of the ventral plates. This black
is increasingly pronounced posteriorly. The underside of the tail
is almost constantly a uniform black or dark brown; when white
occurs there, it is usually as a tapering midventral stripe extending
back from the vent for a short distance.
The head markings are like hoylii, except that numerous white
spots are developed on the parietal, supraocular, and frontal plates,
and on the anterior temporals, a condition which is not, however,
uncommon in specimens of hoylii from southern California, and
which is developed likewise in conjuncta of the "Cape Region" of
Lower California. The upper and lower labials are white or yellow
with their common borders dark, except that, below the eye, on the
third upper labial, or bridging the suture between it and the fourth,
is the dark blotch characteristic of the western representatives of
the getulus group.
Specimens from San Diego County usually have this typical and
striking pattern (fig. 22) : The conspicuous and often perfect dorsal
stripe, the regular white spotting of the lower rows of scales, the
nearly immaculate belly, and the uniformly black caudals. North
of here the most constant feature of the pattern is the uniform dark
color underneath the tail; the rest is extremely variable. This
variation and its significance is discussed farther on.
The penial characters are practically identical with those of
hoylii. The following characters are derived from examination of
alcoholic material: Apparently sHghtly bifurcate, the sulcus single,
and extending over one of the lobes; calyces, with low fringes,
restricted to the extreme distal end; the fringes soon becoming
modified into spines which increase gradually in size toward the
base; no spines unusually enlarged; about halfway toward the base
of the organ the spines suddenly replaced by a few very small ones,
which soon disappear entirely, leaving the basal portion smooth
(or, in preserved specimens, wrinkled).
The dentition is as follows: Maxillary teeth, 13 to 15, subequal,
the posterior not enlarged; mandibular teeth, 14 to 15, subequal;
palatines, 9 to 10 (abnormally 11); pterygoids, 16 to 18.
Habitat aiul habits.—Practically nothing is recorded on the natural
history of this form. One specimen from Santa Ysabel, San Diego
County, California, is recorded as having been taken "along a small
96 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
stream," C. L. Camp refers to it as usually found in "brushy
localities." It is highly desirable in connection with its alleged
"mixing" with loylii, that all possible information bearing on its
life history, habits, and habitat preferences be obtained and reported.
In particular it would be desirable to obtain records and notes on
complete broods of yomig. Breeding experiments, if they could
be carried out, should yield valuable information.
Range.—This form is now kno^vn only from Fresno County, and
from Los Angeles County south through San Diego County and
riG.i22.—COLOB PATTEEN OF LAMPEOPELTIS CAUFOKNIAE CALIFOROTAE (U.S.N.M. NO. 54363) FROM SAN
Diego County, Califoenia. About IJ x nat. size.
just south of the international boundary. Further collecting will
doubtless show its range to extend well south into Lower California,
where an area of intergradation with nitida may be expected.
Published records for localities not included in the list of speci-
mens examined are as follows: Waterman's Canyon, and Cuyamaca,
California (Van Denburgh, 1912, 149, 151).
Variation.—Specimens are too few and variation too sHght for
the recognition of geographic differences in any characteristics
except pattern. The latter, as already remarked, is very constant
and distinctive in aU specimens examined from San Diego County,
but farther north it is extremely variable. The pattern of these
EEVISION OF THE KING SNAKES. 97
more northern specimens is a variable mixture of the jDatterns of
hoylli and typical californiae. Frequently complete rings occur for
a short distance behind the head and on the tail; more often these
rings are continuous on the scales only; more commonly still they
are reduced to short dorsal transverse bars. All stages occur be-
tween the latter and irregular or elongate middorsal blotches, short
rounded spots, short and interrupted stripes, long and somewhat
as-
/zo if^
f20 U^
Fig. 23.—Map showing locality records for Lampropeltis caluorniae califoenlae,
irregular strij^jcs, and the continuous smooth stripe symmetrically
placed on the middorsal line. Even in the last and most perfect
case there is generally an enlargement at the anterior end of the
stripe, not infrequently detached from the rest, which is obviously the
homologue of the first complete ring of hoylii. Most of the north-
em specimens have the underparts a uniform brown, although here
too the ventral pattern of hoylii may often be observed. Along
98 BULLETIN 114, UI^TITED STATES NATIONAL MUSEUM.
with the progressive development of the dorsal stripe, the lateral
pattern of californiae is gradually formed by a breaking up into
spots of the lateral portion of the white rings, and the development
of white centers in the dark scales between these areas.
The development of the coloration beneath, in californiae, appears
to take place first by the formation of dark spots in the white portions
of the belly. These enlarge until the whole ventral surface except
for the lateral tips of the ventrals is a uniform dark brown or
black. While the belly is becoming thus overspread with black,
the white portions of the rings on the first row or two of scales are
becoming extended laterally into a continuous stripe bordering the
black of the belly. In the southern specimens, however, the belly
is generally immaculate, except for the ends of the ventral plates.
That this condition has been produced by a secondary development
of a midventral white line, beginning anteriorly and increasing in
length and width, is indicated by a specimen from San Jacinto
(Stanford University collection. No. 1216), and is borne out by the
majority of specimens from San Diego County. A few specimens,
however, indicate that the immaculate condition of the belly has
sometimes been produced by an extension onto the ventrals of the
continuous white of the first row or two of the dorsal scales. The
black in these cases disappears last from the midventral line, and the
ventrals show no darkening of their postero-lateral borders.
Recently a most interesting confirmation was obtained of the
specimens collected at Fresno in 1879 by G. Eisen. In the spring
of 1918 two specimens were foimd by Van Denburgh and Evermann
in Fresno Coimty, at Jameson and Firebaugh, respectively. The
Jameson specimen (California Acad. Sci., no. 41668) is as uniformly
deep brown on the beUy as if the color had been painted on with a
brush; the white rings are nearly all complete on the dorsal scales;
on the first row, and overlapping the ends of the ventrals, the
white rings have fused laterally into a continuous stripe on each
side. In the Firebaugh specimen (CaHfomia Acad. Sci., no. 41700)
the belly is only a little less uniformly dark; but dorsally only the
first five white rings are continuous, the rest being broken into
lengthwise stripes that are mostly short, except for the middorsal
stripe which is prominent and practically continuous over nearly
two-thirds of the posterior portion of body and tail. From Los
Angeles County only a single specimen is known, and this is fully
as aberrantly marked as any of the examples from Fresno County.
Of the three from San Jacinto, one is rather aberrant, and the other
two are as perfect as those from San Diego.
We may therefore summarize the situation by saying that (1)
typical ioylii is found throughout the region inhabited by cali-
REVISION OF THE KING SNAKES. 99
forniae, or its aberrant representatives, (2) typical californiae is
found only south of Los Angeles County, (3) in San Diego County
all the individuals are typical, (4) in Riverside Coimty some are
typical and some partly aberrant, and (5) north of Riverside Coimty
aU are aberrant.
The specific identity of this form has been often questioned on
accomit of (a) its identity in structural features with boylii, (b) the
remarkable variation in its pattern and its evident approach in these
variations to the pattern of boylii, (c) its alleged rarity and apparently
sporadic distribution, (d) its occurrence wholly within the range of
its nearest relative.
In support of its specific identity it may be remarked that (a)
color pattern differences alone are often used in the differentiation
of species, (b) There can be no doubt of its approach in pattern to
boylii in the northern ]>ortion of its range, but typical boylii occui*s
here while typical californiae does not, indicating that this is not an
instance of intergradation ; and in the southern part of the range,
both forms are common in the same localities and no aberrantly
marked individuals are found. If interbreeding produces aberrant
individuals, why is californiae not found in typical form in Fresno
County, if boylii is; and if interbreeding takes place in San Diego
County and Mendelian segregation is to explain the occurrence here
of only the typical patterns of both forms, why should aberrant
patterns be found in some other portion of the range ? (c) Its rarity
in the region of its typical development is not apparent; where it
does seem to be rare it is variable, (d) The fact of its range lying
wholl}^ within that of its nearest relative is an unusual instance and
in need of explanation.
Affinities.—As brought out in the discussion of variation, the
affinities of this form are all with boylii. The only other member of
the getulus group which ranges anj^where near californiae is yumensis.
To assume yumensis to be the ancestor of californiae raises a question
more difficult to answer than the one raised by assuming boT/lii to be
the ancestor. It would then have to be explained how one species
gave rise to two other species, which occupied the same region to-
gether, in competition, and both survived. Yumensis is however,
excluded from direct relationship by the fact that the only region
where its range is near that of boylii is the region where californiae is
constant in its specific characters, and by the fact that the abberra-
tions of californiae are toward boylii and are in a region that could
never have been occupied by yumensis.
We have then the peculiar condition of a species being differen-
tiated within the range of its parent. If we say it is a mutation,
we must explain why it is constant in pattern in the southern portion
100 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
of its range and variable toward hoylii in tlie northern. If we sug-
gest that it was evolved from loylii by adaptation to a different
habitat, we consider a possibility that may have no proved counter-
part elsewhere, and that in this case has, as yet, no support from
observation in the field.
The mutation theory, even though it rests upon no direct evidence
in this case seems at present to offer the best exp anation of the
origin of this form. On the basis of this theory the origin may then
be hypothecated as follows: Californiae originated from hoylii by
mutation, somewhere in the Great Valley of California. It spread
southward west of the Sierra Nevada Mountains and the deserts of
southwestern California becoming more different from boylii toward
southwestern California. From here it extended its range into
Lower California to Cape San Lucas At some point in this penin-
sula, probably pretty well south, it became modified into the color
variety nitida.
It will be recognized that more information is essential to an ade-
quate explanation of californiae.
REVISION OF THE KING SNAKES. 101
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REVISION OF THE KHSTG SNAKES. 103
LAMPROPELTIS CALIFORNIAE NITIDA (Van Denbureh).
1887. Ophibolus califomiae Cope, Bull. U. S. Nat. Mus., no. 32, p. 79.
1895. LampropeUis nitida Van Denburoh, Proc. California Acad. Sci., aer. 2,
vol. 5, pt. 1, p. 143, pi. 14 (type locality, San Jose del Cabo, Lower
California; type, California Academy of Science collection. No. 800;
Gustav Eisen, collector).
—
Stejnegeb and Barbour, Check List, 1917,
p. 88.
Description.—Since no specimen of this form has been examined
by the writer, the original description is quoted in full:
Allied to L. califomiae, but with the gastrosteges, urosteges, and upper surfaces of
head and snout, entirely brownish black.
The head is slightly distinct, considerably depressed, its plates normal; 1 loreal;
1 pre- and 2 postoculars; scales in 23 rows, smooth, with 2 apical pits; postgeneials
very small; anal entire; seven superior labials, the third and fourth entering the orbit;
227 gastrosteges; 56 pairs of urosteges.
The back and sides are blackish brown; the former, with a rather indistinct longi-
tudinal line composed of cinnamon colored spots upon the centers of the scales of
the median series, and upon the inner edges of those forming the first row on each
side of this series; the latter, Avith a few scales of the first and second rows dotted,
centrally, with cinnamon or yellowish white. A band of cinnamon crosses the nape.
The gulars, geneials, and inferior labials, are blackish brown mth paler centers.
The plates on the top and sides of the head are brownish black, with faintly indicated
dots of raw umber upon the loreal, pre-, and postocular plates, and near the posterior
edges of the supraoculars and parietals. There are 6 cinnamon colored blotches on
the upper surface of the tail. The gastrosteges and urosteges are entirely brownish
black, with the exception of the first 10 gastrosteges, which show faint cinnamon
colored dots.
Total length 965 mm. Tail 125 mm.
A small specimen (290 mm.) has, on the sides, rather numerous cinnamon colored
blotches or enlargements of a similarly colored longitudinal line. This line is of
about the width of one row of scales, and occupies the tips of the gastrosteges and the
lower half of each scale of the first aeries.
Remarks.—Only the two specimens described above are known.
They are of great interest, however, for they prove the occurrence in
the Cape Region of an ally of califomiae paralleling in every way the
case of hoylii and conjuncta. Since there is good reason to believe,
from its present distribution, that califomiae never reached southern
Arizona, its derivative, riitida, must have attained the Cape by way
of the pensinsula of liower California, and not by way of Sonora and
the islands of the Gulf of California. This is an argument in favor of
the feasibility of the former route for conjuncta, thereby favoring
its derivation from hoylii instead of from yumensis; and it also
favors the specific distinctness of califomiae from hoylii, as it shows
the same situation to hold at Cape San Lucas with respect to these
two forms as obtains in the vicinity of San Diego County, California.
104 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
SUMMARY.
From the preceding descriptions it appears that this group is
represented by a series of forms arranged in linear order across the
United States and northern Mexico (fig. 24), and that each member
Fig. 24.—Map showing the distribution of the various forms op the getulus group.
of the group is most closely related to the form inhabiting the con-
tiguous region. Cdliforniae offers the only exception, and this is
discussed separately in the section devoted to it.
REVISION OF THE KING SNAKES. 105
Since the forms are thus linearly arranged and related, evolution
may have proceeded from either extreme toward the opposite one,
or from some intermediate point in two diverging directions. Flori-
dana, hroolcsi, and conjuncta must be excluded from further con-
sideration as ancestral forms, for reasons already discussed, chief of
which are that they belong to regions that have received their faunas
from the mainland to which the peninsulas are attached and which
have thus not been centers of dispersal, and that they are very
clearly derivatives of the adjacent forms, getulus and hoylii, respec-
tively. Also, californiae and nitida are too obviously derived from
hoylii to receive consideration in this connection. There are left
therefore six forms to which to look for an approach to the ancestral
condition of the group.
A comparison of the characteristics of these various forms may
give some clue as to whether evolution has proceeded from one coast
to the other or from some intermediate point in two directions.
Certain structural features will be considered first.
Scale formulae of theforms of the getulus group.
Form. Maximum.
conjuncta
j
23-25-23-21-19
hoylii 23-25-23-21-19
yumensis ' 23-25-23-21-19
splendida
;
23-25-23-21-19
holbrooki 21-23-21-19
niger 21-23-21-19
getulus
i
23-21
floridana i 23-21
Muiunum.
21-23-21-19
21-23-21-19
21-23-21-19
21-23-21-19
19-17
19-21-19-17
19-21-19-17
21-19
Average.
23-21-19
23-21-19 or 21-23-21-19
23-21-19
23-21-19 or 21-23-21-19
21-19 or 19-21-19
19-21-19
21-19 or 21-23-21-19
21-23-21-19
The table giving maximum, minimum, and average scale formulae
shows that the forms from splendida westward are identical in the
matter of scale rows, and that those east of splendida have a decidedly
lower range and average, with the exception of Jloridana. As ex-
plained in the discussion of getulus, the drop to 17 rows posteriorly
is rather common from Virginia to New Jersey, and since this form
must have reached this portion of its range from the south where
17 rows at the end is almost unknown, this must be considered a
secondary and not a primitive character. Seventeen rows occur most
frequently in holbrooki in the vicinity of New Orleans, and in niger in
the northern portion of its range, both places that would be more
recently populated whether the origin be placed in the southeast
or in the southwest. This indicates that in these places a reduction
in number of scale rows is taking place, and strongly suggests that
the formula 19-21-19 is also a result of reduction from a higher
formula. That evolution has taken place in this group in Jloridana,
106 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
from a lower number of rows to a higher can not be denied, and it
may therefore be admitted as possible to regard getulus or Jiolhrooki
as ancestral. But if we consider that a maximimi of 23 rows and a
minimum of 19 is characteristic of 5 of the 8 forms and conmion at
widely separated localities in the 3 (which, as will appear later, are
\
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\
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s
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1
1
1
1
1
1
/
/
1
1
•
•
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/
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/
1
1
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1
"3 o
evidently specialized in other structural features), it looks as if this
were a fundamental character of the getulus group, and as if forms
possessing a lower maximum and a lower minimum were distinctly
removed from the ancestral condition.
A glance at the diagram (fig. 25) of extremes and averages of ven-
tral plates shows that the eastern forms differ decidedly in this respect.
REVISION OF THE KING SNAKES. 107
Again, it will be noticed ihat Jloridana has changed from getulus by
an increase in number of ventrals. Since evolution by increase must
therefore be considered as a possibility, we can base no argument for
direction of evolution upon ventral plates alone. We may simply
note, for later reference, that the forms exhibiting the lowest scale
formulae also have the smallest number of ventral plates.
Variation in number of caudals and labials is of little use in itself
in this discussion, but it is noticeable that the lowest numbers occur
in the forms having the fewest ventral plates and the lowest scale
formulae. Changes in the skull and dentition have been too slight
to be of much use in tracing relationships within the group. This
very constancy will, however, be of value later in connecting this
group closely with the calligaster group.
The loreal shield is markedly different in the extremes of the group.
On the east coast it is square or decidedly higher than long, while on
the west coast it is usually longer than high, the dimensions here
rarely approaching equality. The intermediate forms are inter-
mediate in respect to the shape of the loreal, but TiolhrooJci is more like
getulus (fig. 4) and splendida (fig. 3) is more like hoylii. If the getulus
group is correctly associated with the other forms of Lampropeltis,
the shape of the loreal in the eastern forms is a decidedly speciahzed
instead of primitive condition. This is a very good reason for look-
ing west for the origin of the group.
Like the loreal, the relative length of the posterior chin shields is
decidedly different in the extremes of the group. In the eastern
forms they are approximately as large as the anterior shields and
usually in contact with each other, while m hoylii and its allies they
are hardly more than half as large as the anterior ones and are
separated usually by two smaller scales. Splendida here occupies the
intermediate position. Its western individuals are like hoylii and
its eastern show a decided approach to the getulus condition. The
nearest allies to this group are calligaster and rhomhomacuJota, and
here the posterior chin shields are usually a little shorter than the
anterior and tend to be separated by about one scale, an arrangement
that is true as well of the triangulum group. Both coast forms
therefore represent extreme modification in the shape of the posterior
chin shields, and it is splendida that presents the closest approach in
this character to the other groups of the genus.
186550—21—Bull. 114 8
108 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
Comparison of the coast forms with respect to the shape of^^the
head and its scutes shows considerable divergence. Such compari-
son may be tabulated as follows:
Character. boylii. getulus.
Snout More blunt More pointed.
Less than twice as wide asRostral About twice as wide as high
(proportion of height to
width averages distinctly
less than in getulus).
Usually longer than high
Increased
Loreal
high.
Usually higher than long, or
as high as long.
Reduced.Internasal suture
Eye Larger—its diameter gener-
ally greater than the great-
est height of the 4th upper
labial.
Usually fails to reach the
posterior limit of the pari-
etal.
Averao'e lower
Smaller—its diameter gener-
Upper temporal of third
row.
Supralabials
ally less than the greatest
height of the 4th upper
labial.
Usually extends to the pos-
terior limit of the parietal.
Average deeper.
About as long and nearly as
wide as anterior, and in
contact or separated by not
more than one small scale.
Posterior chin shields. .
.
About half as wide and half
as high as anterior, and
separated by two smaller
scales.
Briefly interpreted, getulus has a higher and more pointed head
to which the deeper loreal and supralabials contribute, and hoylii
has a lower, flatter, and blunter head. The shape of head in Jiol-
hrooTci is decidedly like getulus; yumensis and conjuncta are decidedly
like hoylii; it is in splendida that we find a mean between these ex-
tremes, and the closest similarity with the other groups of the genus,
and in the forms east and west that we find speciahzation.
The penis is essentially the same throughout the group. In hol-
hrooki and niger, however, the basal half of the organ is beset with
minute spines. It would be easy to say that this condition is primi-
tive, and that the others have been derived by reduction of these
small spines. But primitive conditions generally show more per-
sistency than this. Most of the other structural characters show a
marked degree of permanence; reduction or increase is the excep-
tion, not the rule. In the calligaster group, the one nearest related
to this, the copulatory organ has the minute spines, when present
at all, usually restricted to the immediate vicinity of the large ones.
This is the situation that holds in the triangulum group and else-
where in the getulus group, except that there may be a slight ten-
dency toward extension of these spines in yumensis and hoylii. Thus
these small spines of the penis, instead of favoring holhrooTci or niger,
as ancestral, are distinctly against it, for the reason that they rep-
resent a speciaHzed condition not foimd elsewhere in the genus.
In getulus, too, this organ is decidedly specialized. The bilobed
condition, noted in the descriptions for holhrooTci and probably for the
REVISION OF THE KING SNAKES. 109
other more western forms, is intensified in getulus into distinct forks,
a character that is even more strongly developed in Jloridana. Since
this feature is unique for the genus, we must consider it a strong reason
for excluding these forms from the list of ancestral possibilities.
In connection with the positive evidence of specialization in the
forms east of splendida derived from the characteristics of the loreal,
chin shields, and copulatory organ, we must now consider that the
reduced scutellation of these forms is also a result of specialization.
We are forced to conclude, therefore, on structural grounds, that
Jloiidana, getulus, niger, holbrooTci, and probably the forms west of
the Rockies, must be excluded from consideration as possible ancestors
of the group.
Turning now to the pattern, we find that its variations are truly
remarkable, ranging as they do from cross bands, to small spots,
to rings, and stripes, and to various combinations of these. In fact,
the pattern is the most variable of all the characters upon which
specific value is placed. It may therefore be expected that pattern,
if anything, will give a clue to relationships.
Since the head pattern is the least variable, we may consider that
first. One of its strikingly constant characters is the pair of white
bars on the prefrontal and internasal plates. In the Pacific coast
forms these are broad, occupying most of the area of the scutes, and
usually so broadened as to largely lose their transverse character;
in getulus and Jloridana they are narrow and usually close to the
anterior and antero-lateral borders of the scutes; often they are
curved, or broken in the middle, or reduced to spots; in Tiolhrooki
they are pretty constantly narrow transverse bars, the chief varia-
tion being to reduction, breaking into spots, and curving; in yumensis
they are strong, symmetrical transveree bands on the anterior por-
tions of the scutes; in splendida they vary from the yumensis form
at the extreme west to the Tiolhrooki condition in the east; in the
central portion of its range these bars are narrow, rather irregular,
sometimes broken, or somewhat bent or curved. These bars, there-
fore, reach their extremes of diversity in the east and west coast
forms, and are developed most perfectly and symmetrically in Arizona
and in the Mississippi Valley. If we feel it necessary to assume as
the starting point the perfect and symmetrical condition, we must
explain why, starting with either liolhroolci or yumensis, evolution has
proceeded in one direction to an extreme condition (the nearer coast
forms), and in the other through a reduced condition {splendida),
to a perfect condition again, and then to another variable extreme.
To the writer the only logical explanation is to consider the condi-
tion presented by splendida in southern New Mexico as the starting
point. Here we have the beginnings of the white bars, imperfectly
formed as symmetrically placed lightened areas on a black head; evo-
110 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
lution, resulting from radiate migration, has perfected the marks in
central Texas and in Arizona; farther migration has resulted in
further change in these bars by which they lose their sim.ple form and
become modified in two different ways independently—in the west
becoming broadened, in the east more or less broken into spots, and
bent, or curved. We see then, in these prefrontal and internasal
bars, their inception, perfection, and diverse modification.
These white bars are not the only parts of the head pattern requir-
ing mention. White markings, spots, and bars of various shapes,
are considerably developed in all forms of the group except yumensis,
the Great Basin section of hoylii, and splendida. The first two are
alike with white markings symmetrical, clean-cut, and limited to
the snout and sides. In splendida, in the central portions of its range,
the only white markings present occupy the same positions as in
yumensis, but are much more restricted; the rest of the head is
black. In the western portion of its range these markings attain
the same development that they show in yumensis; in the eastern
portion it develops the markings of JioTbrooki. It is not necessary
to explain the variations of these markings in further detail; the
case is the same as with the prefrontal white bars; in s'plendida we
have the inception of two diverging lines of head markings, the east
and the west.
One other portion of the head pattern may be mentioned. In all the
forms west of S'plendida the labials are white with narrow dark mutual
borders, except for the suture between the third and fourth upper
labials, the dark mutual border of which is so much widened as to
form a conspicuous circular spot or blotch on the upper lip beneath
the eye. In all the forms east of splendida the labials are light with
dark mutual borders, but there is no widening of the mutual dark
border of the third and fourth supralabials. Only as an occasional
individual variation is there any increase in pigment on the upper
labials beneath the eye—a variation as uncommon as the occasional
great reduction of the subocular spot in the west coast forms. Con-
necting these two divergent sections of the group we have splendida,
in the New Mexican portion of its range, with an almost totally black
head, its labials lightened only by a naedian narrow white bar on
each scute which often is not even continuous across its middle.
The condition in the other forms of the group has undoubtedly been
produced by a widening of these narrow median white bars; while
eastward they all widened, westward the dark pigment was retained
on the upper series beneath the eye. This seems to be the only
logical explanation of the peculiar distribution of this subocular
dark patch.
Perhaps of even greater value is the evidence from the body pattern.
Here the diversity is apparently much greater. If we except cali-
fomiae (its relationship to hoylii has been sufficiently discussed, along

HPII
iiovLii (Univ. ('Ai.ipdUNiA No.fiM;i, viidm Tuhhk Mount-
AiNB, Bah DicitHAuniNo County, CAuroimu]. Aiiout
isaMo-ai. (Tot«wi«mouio
Km. aa.-TTriCAJ. COLOR rATTLit;
ULUS NU)KR (rROU A Sl'SCIUEN
MUSaVU, FROU Tallai>koa
About HAT. suK.
FlO. »—TWICAL COLOR PATTERN Of LaUPROPRLTIS OBTO-
LUS OEIULUS (O.S.K.M. NO. 22S56, VlROWU BBaCH,
VttGiKiA). About NAT. SIZE.
PlO.36,—TYTICALCOLORPATTKBNOP LAMPEOfELTiaOETUUja
ruiRiDANA {U.S.N.M. NO. 22368, ttpe, Obanoe Ham.
MOCK, DeSoto County, Florida). About nat. flia.

Fig. 27.—Typical coloe pattern of lyAMPROPELTis
;
BOYLii (Univ. California no. 5543, from Turtl
AiNS, San Bernardino County, Californla.)
NAT. SIZE.
- (ETULUS 3PLEN-
MZ0NA),SH0'W-
YITMENSIS IN
SIZE.
Fig. 32.—Typical color pattern of Lampeop,lusget-
GETULUS HOLBEOOKI (U.S.N.M. NO. 13041, NE^W ) SHOW-
LEANS, LOUISIANA). About NAT. SIZE. About
186550—21. (To face page 111.)
K
Fig. 30—Color"
DIDA(U.S.N.]I
SH0"WING THE
TEEN IN THE G-
FiQ. 35—Typical
LUS QETULUS (
ViEQINU.). AB
KEVISION OF THE KING SNAKES. Ill
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112 BULLETIlsr 114, UNITED STATES NATIOFAl, MUSEUM.
with its description), the various forms of the getulus group all have
a pattern that is fundamentally one of narrow transverse white bands
on the back, separated by a ground color of black (figs. 27 to 37).
The numbers of these crossbands or rings, as the case may be, have
been averaged and plotted, for each form, in the accompanying
diagram (fig. 26). Two contrasted conditions are shown—the high
numbers of splendida and Jiolbrooki, and the low numbers of getulus
and the west coast forms (the situation with floridana has already
been explained under the discussion of that form) . It is noteworthy
that the extremes of the group—the coast forms—^have approxi-
mately the same variation and average in number of bands. The
fundamental difference in pattern between these forms is that the
bands in the west coast forms (figs. 27 and 28) are continued around
the body as rings, while in getulus (fig. 35) they fork on the sides and
the connections across the belly are imperfect; from above both body
patterns look almost identical. It seems to the writer quite out of
reason to suppose that any of the west or east coast forms could have
changed into the spotted condition of splendida and Jiolbrooki and
then emerged with the pattern exhibited on the other coast. The
fact that the coast forms have patterns so strikingly similar suggests
that, like the diverse modifications in the pattern of snout and labials,
they have been produced by independent evolution from a similar
beginning. Then, of necessity, either lioTbrooki or splendida must be
examined for ancestral possibilities. The former is, however, suffi-
ciently rejected on structural grounds.
The pattern of splendida as exhibited by examples from southern
New Mexico and northern Chihuahua briefly is this: Ground color
black; back crossed by about 50 or 60 narrow white bands formed of
black scales with oval white centers; sides with a white oval center on
each scale, the white occupying more area toward the belly; latter
black, except that the ends of about two gastrosteges, opposite the
dorsal white bars, are white. The important thing to notice is that
the white centers are all very approximately oval and oriented the
long way of the scales. Figure 30 shows a typical example from
southern New Mexico in which the black spaces on the back are very
small and the white centers are all very nearly uniform in orientation.
Going eastward in the range of splendida we find that in western
Texas the white centers are often angular or long and narrow and
tend to lose their symmetrical orientation. This is shown by a
specimen from Reeves County, Texas (fig. 31) . In central and south-
ern Texas this change is more marked, and there is a decided tendency
for some of the white centers on the sides opposite the crossbands to
fade, causing a series of small dark areas to appear on the sides, in
alternation with the dorsal dark areas. This is the beginning of the
chain pattern of niger and getulus. It is obscured, however, in
Jiolbroolci by the appearance of an oval white center in each scale of
REVISION OF THE KING SNAKES. 113
the dark areas. This is well shown by a specimen from New Orleans
(fig. 32). In this the symmetrical oval spots occupy only the areas
that were without white spots in the specimens from central and
southern Texas, and the spots which were present in these specimens
are present here too, and with much the same unsymmetrical orienta-
tion as was shown initiated in figure 31. That is, Jiolbrooki has
retained the pattern of splendida, as developed in Texas, but has
added something to it.
The form niger is developed from holbrooki by a fading of the newly
derived white spots between the bands and a contraction of the white
in the crossbands and sides (fig. 33). The exact derivation of getulus
(fig. 35) from nige?' (fig. 33) is uncertain, due to scarcity of specimens
from the critical region. A specimen from Anniston, Alabama, sug-
gests that the transition to getulus was accomplished by a disappear-
ance of alternate cross bands on the back and possibly by fusions of
the light areas on the ends of the ventrals. In this specimen the
crossbands are scarcelv distinguishable but seem to be reduced in
number. A specimen from Marietta, Georgia (fig. 34), has the pat-
tern of getulus, but the crossbands are very narrow. The change
from this to the typical pattern was accomplished by a widening of
the white parts, thus making the crossbands conspicuous and bringing
out the chain pattern that was initiated in the splendida of central
and southern Texas and remained inconspicuous in holbrooki. The
pattern of such a typical getulus is shown in figure 35.
Floridana was developed from getulus by a basal lightening of each
dark scale, and an increase in the number of crossbands (fig. 36).
Here, as in Tiolbroolci, we have a spotted pattern that is at the end of
an evolutionary series, but it is a decidedly different sort of spotted
pattern from that of splendida (fig. 30) in that it bears the vestiges
of an earher pattern while that of the latter apparently does not.
In extreme southern Florida this series of patterns is carried one step
farther and to its logical conclusion in hroolcsi (fig. 37), in which even
traces of the crossbands appear to be lost. Numerous structural
features, however, mark it as a specialized type.
Yumensis is formed from splendida even more simply. It occurs
as follows: Westward from southern New Mexico splendida becomes
somewhat altered. The essentials of the process are fundamentally
the same as the changes eastward. The white centers lose their sym-
metrical form and arrangement and assume various shapes. Small
dark areas develop on the sides opposite the crossbands (fig. 29).
The doi-sal dark areas increase in length and breadth. At this point
we have the fundamentals of another getulus pattern, but, apparently
by chance, rings are produced instead of a chain pattern. The inter-
mediate forms upon which this outline is based show that the white
spots separating the lateral dark areas from the dorsal ones are
114 BULLETIN 114, UlSTITED STATES NATIONAL MUSEUM.
weaker on one side of the crossband than on the other, so that the
dorsal dark areas become fused with the lateral on one side of the
crossband, and on the other the white is strengthened. Similarly the
lateral dark areas fuse with a ventro-lateral series, and thus become
complete on the belly. It now only remains for these newly formed
rings to be perfected. This occurs by a whitening of the white rings
and a fading of the white that may still form ragged edges projecting
into the dark areas. All these transitional stages may be observed
in specimens from the vicinity of Tucson. With the perfection of
the pattern of yumensis (fig. 28), the only change necessary to pro-
duce the pattern of hoylii (fig. 27) is an extension of the white in the
white rings to the whole of the scales in these rings.
Thus the patterns of the coast forms of the getulus group result
from the tendency of the pattern of tj^pical splendida to become
broken into an alternating series of black and white areas.
That splendida may well be the parent form of the getulus group
is indicated by its present distribution. It appears to be confined
to the region between central Texas and eastern Arizona. Its north-
ern limit is unknown, but is doubtless the mountains of central or
northern New Mexico. That it extends south into Mexico is as cer-
tain as anything can be, but how far we can only surmise. For
southern Mexico, where considerable collecting has been done, there
is no record of splendida, nor of any form allied to it. We ma}^ there-
fore say that the arid plateau region of the southwest is its home.
This has often been commented upon as being a favorable center for
preservation and for dispersal of reptilian life. The present diversity
of forms may be considered the result of an east and west dispersal,
with consequent modification in each major environment.
The foregoing discussion may be sunoimarized as follows : '
1. Since each member of the getulus group is nearest and evidently
directly related to the form inhabiting the adj acent region, and since
the forms are in linear arrangement geographically, one of them
must therefore present a condition from which all the others may
be derived (californiae and nitida are ruled out of consideration here
for reasons discussed under the former)
.
2. Of the eight forms entering the discussion, two
—
conjuncta and
jioridana—are eliminated because their geographic position and the
evidence for their derivation from the adjacent continental forms
is too strong to allow them to be considered as ancestral possibilities.
3. Comparative study of loreal and chin shields indicate that the
east and west forms present specializations in these scutes.
4. The proportions of the head and its scutes indicate that splendida
is the most closely allied to the other forms of the genus, and that
both the east and the west forms of the getulus group are specialized.
5. Getulus, jioridana, niger, and liolbrooki are excluded from con-
sideration as possible ancestral forms on account of the unique
REVISION OF THE KING SNAKES. 115
specializations in the structure of the copulatory organ. Since they
are specialized in this respect we must regard the low averages in num-
bers of scale rows, ventrals, and labials as also due to specialization.
-6. Splendida is the only member of the group not eliminated on
structural grounds.
7. There are no structural peculiarities that can not be more easily
explained by assuming splendida as the ancestor.
8. That splendida, as represented in southern New Mexico and
northern Chihuahua, possesses the only pattern from which all the
others may be simply and naturally derived is definitely indicated
by the variations and geographical distribution of (a) the white bars
californiae< boyliu yumensis^ splendida—-^holbrooki >niger >getulii8
I
floridana
nitida conjuncta I
brooksi
Fig. 38.—Diagram of the relationships of the forms of the getulus group.
on the prefrontal and internasal scutes, (6) the subocular black spot,
(c) the other head markings, and (d) the body pattern.
9. The region inhabited by splendida is the one theoretically most
favorable for being the center of preservation and dispersal of a
plains form of land snake.
THE CALLIGASTER GROUP.
LAMPROPELTIS CALLIGASTER (Harlan).
YELLOW-BELLIED KING SNAKE; EVANS KING SNAKE.
Figs. 6, 39, 40.
1827. Coluber calligaster Harlan, Journ. Acad. Nat. Sci. Philadelphia, vol. 5,
pt. 2, p. 359 (type locality, Missouri); Med. Phys. Researches, 1835,
p. 122.
—
Lampropeltis calligaster Cope, Proc. Acad. Nat. Sci. Philadel-
phia, 1860, p. 255.
—
Strecker, Proc. Biol. Soc. Washington, vol. 21,
1908, p. 75.—HuRTER and Strecker, Trans. Acad. Sci. St. Louis,
vol. 18, no. 2, 1909, p. 26.—Hurter, same, vol. 20, no. 5, 1911, p. 187.—
Stejneger and Barbour, Check List, 1917, p. 87.
—
Ophibolus calligaster
Cope, Bull. U. S. Nat. Mus., no, 1, 1875, p. 37.—Smith, Geol. Surv. Ohio,
vol. 4, 1882, p. 689.—Yarrow, Bull. U. S. Nat. Mus., no. 24, 1882,
p. 94.—Cope, Proc. U. S. Nat. Mus., vol. 14, 1891, p. 610.—Garman,
H., Bull. Illinois State Lab. Nat. Hist., vol. 3, art. 13, 1892, p. 293.—
Hay, Annual Rep. Indiana State Board Agric, vol. 28, 1887, p. 210;
Batr. Rept. State Indiana, 17th Annual Rep. Indiana Dept. Geol. Nat.
Resources, 1892, pp. 515, 590; Batr. Rept. Indiana, 1893, p. 182.—
Hurter, Trans. Acad. Sci. St. Louis, vol. 6, no. 2, 1893, p. 255.—
Blatchley, 24th Ann. Rep. Dept. Geol. Nat. Res. Indiana, 1899, p.
545.—Cope, Rep. U. S. Nat. Mus. for 1898, 1900, p. 905, fig. 223, pi.
18, fig. 6.—Brown, Proc. Acad. Nat. Sci. Philadelphia, 1901, p. 80.—
Strecker, Trans. Texas Acad. Sci. for 1901, vol. 4, pt. 2, no. 5, 1902,
p. 4.—Ditmars, Reptile Book, 1907, pp. 341, 355.—Somes, Proc. Iowa
Acad. Sci., 1912, p. 150.—Strecker, Baylor Bull., vol. 18, no. 4, 1915,
p. 38.
—
Coronella calligaster Boulenger, Cat. Snakes Brit. Mus., vol. 2,
1894, p. 198.
116 BULLETIN 114, tTNITED STATES NATIONAL MUSEUM.
1842. Coluber eximus (part) Holbeook, N. Amer. Herp., ed. 2, vol. 3, p. 72.
1856. Ablabes triangulum, var. calligaster Hallowell, Proc. Acad. Nat. Sci.
Philadelpliia, p. 244.
1859. OpMholus evansii Baird, Pacif. R. R. Surv,, vol. 10, no. 4, p. 43 (quotation
from Kennicott's forthcoming manuscript).
1859. Ophibolus evansii Kennicott, Proc. Acad. Nat. Sci. Philadelpliia, p. 99
(type locality, central Illinois; cotypes, U.S.N.M., no. 1593; 3 specimens;
R. Kennicott, collector).
—
Coronella evansii Jan, Arch. Zool. Anat,,
vol. 2, fasc. 2, 1863, pp. 237, 243; Icon. Gen. Ophid., livr. 17, 1866,
pi. 2, fig. 3.
1863. Coronella tigrina Jan, Arch. Zool. Anat., vol. 2, fasc. 2, pp. 238, 244.
1883. Ophibolus triangulum calligaster Garman, S., Mem. Mus. Comp. Zool.,
vol. 8, no. 3, pt. 1, pp. 66, 155.
1892. Lampropeltis rhombomaculatus Garman, S., Bull. Essex Inst., vol. 24,
p. 9.
The first description of this form appeared in 1827 in the Genera
of North American Reptiles and a Synopsis of the Species, by R.
Harlan (p. 359). The description is applicable to the present form,
but is very indefinite. Harlan attributed the name calligaster to Say,
probably because the latter found the specimens, recognized them
as new, and proposed the name. But it was Harlan who drew up
and pubHshed the description. Thus Holbrook, in 1842 (vol. 3,
p. 72), said: ''Say seemed to consider the serpent he observed in
Missouri as new; but I am not aware that he described it as such.
Harlan, however, gave a description of it from specimens in the
Philadelphia Museum, and under the name calligaster, from the
beautiful arrangement of colors on the belly." That Harlan had
the present form before him when he drew up the description, while
not beyond doubt, yet appears fairly certain, for Cope in 1861 (1860,
255), says, in reference to the specimens from which Harlan
prepared his description, ''One of these, a stuffed skin, presented
to the Academy by Doctor Holbrook, and labelled by Doctor Hal-
lowell 'original specimen,' is now before us. We can assert its
identity with Ophiholus evansii of Kennicott both from his descrip-
tion and from comparison with specimens collected by Doctor
Hammond in Kansas, and described by Hallowell (1856, 244). They
all have 25 rows of smooth scales." One of the specimens, however,
described by Hallowell (1856, 244) is probably an Elaj^Tie since it
has 65 caudal plates and a divided anal. We have examined three
of Doctor Hammond's Kansas specimens and there is no doubt
about their identity, and two of these were evidently used by Hal-
lowell in his description and discussion of caUigasier, just referred
to above. Kennicott, m 1860 (1859, 99) thought that Harlan had
before him a specimen closely related to what we now know as
Elaphe laeta, and so attributed Harlan's Coluber calligaster to the
genus ScotopUs {ElapTie). Kennicott undoubtedly had before him
specimens of ElapTie laeta which is considered as somewhat different
REVISION OF THE KING SNAKES. 117
from true E. laeta and to be the equivalent of Harlan's 0. calligaster.
Kennicott therefore gave a new name to the specimens in the col-
lection of Northwestern University, which were unquestionably
the Lam-propeltis calligaster that we now loiow, calling them OpTii-
holus evansii.
Granting that Harlan had our form before him, the name calli-
gaster has priority over evansii of Kennicott.
Description.—The scutellation is as follows: Ventral plates 196 to
215, average 205; caudals, 38 to 57 (males 44 to 57, average 51;
females 38 to 52, average 46); supralabials, 7, sometimes 8; infra-
labials, usually 9, often 10, occasionally 11; oculars, 1 and 2; temporals
normally 2+3+4; posterior chin shields usually a little shorter than
anterior, close together or separated by 1 or 2 small scales; loreal
usually longer than high, but sometimes as high as long; suture
between the inteniasals only about half, or less than half, as long as
that between the prefrontals; scale rows on middle of body usually
25, sometimes 27, very rarely no more than 23. (For the scale
formulae, see under Variation.)
The body is rather slender, and uniform in diameter; the head
is scarcely distinct from the neck, the snout is generally somewhat
longer and more pointed than in rhomhcmaculata, the tail is rather
short and tapers rapidly, varying from 0.110 to 0.150 of the total
length (males, 0.123 to 0.150, average, 0.137; females, 0.110 to 0.145,
average, 0.123). The largest specimen examined measured about
1,355 mm., and was taken at Lawrence, Kansas,
The pattern (fig. 39) is composed of about 60 (46 to 78) trans-
versely elongate quadrate brownish or greenish blotches (reddish
in the young), 2 to 3 scales long and 8 to 12 across, narrowly margined
with black, set on a ground color of lighter brown, and extending
down on the sides to the seventh or sixth row of scales. These
blotches are more or less concave before and behind and are occa-
sionally divided on the median line; on the tail they may become
narrowed to transverse black bands, indistinct and irregular near
the end. On the sides, alternating with the dorsal row, there is a
similar series of smaller dark-edged blotches, roundish for the most
part, but often elongated anteriorly. On the first row or two of
scales, and overlapping the ventrals, is a third series of spots, in
alternation with the last, and tending, posteriorly, to fuse with them.
Each scale, usually including the ends of the ventral plates, is minutely
mottled with brown.
The head is somewhat variously spotted, but certain markings are
usually evident. Transversely across the posterior portion of the
prefrontals and not overlapping the frontal plate is a brown bar,
edged with darker; extending forward from behind the parictals
there is a dark-edged brown band which forks on the parietals, and
118 BULLETIN 114, UK-iTED STATES NATIONAL MUSEUM.
extends thus forward to between the eyes, where the forks may-
unite again. From behind each eye a long black-bordered brown
band about two scales wide extends backward on top of the neck.
From the eye to the angle of the mouth there is a narrow dark band.
Thus there is a strip of the ground color from the upper postocular
to the last supralabial. The other head and throat plates are mostly
light-colored. The belly may be faintly or heavily checked with
Fig. 39.—Typical pattern of Lampropeltis calligaster (U.S.N.M. no. 1593, cotype of 0phibolu3
EVANSn Kennicott from Central Illinois). About 1| X nat. size
small quadrate brownish or yellowish blotches, or nearly immaculate
except for the ends of the ventrals.
This pattern is well defined in young individuals, but in many
adults the markings, particularly of the head, lose their distinctness
and become greatly obscured by a darkening of the ground color.
Such darkening appears to be accompanied by an alternate length-
wise lightening and intensification of pigment, producing a rather
prominently striped effect (fig. 40). Throughout the middorsal
line there is a light stripe; bordering this on either side is a dark
band which passes through the lower portions of the dorsal blotches.
Below this, and between the dorsal and lateral series, is a light band;
then below this last and passing through the upper lateral series of
blotches is another dark band. The two or three lowest rows of
REVISION OF THE KING SNAKES. 119
scales are lighter and the beUy is apt to be veiy hght, due to a fading
of the small quadrate dark spots. This darkening and striping seems
to progress with age. Its beginnings may be detected in specimens
that still show the normal pattern slmrply defined. These darkened
individuals seem to come chiefly from the region from eastern Mis-
souri to Indiana.
The copulatory organ (fig. 2) is bilobed; the sulcus single, extend-
ing over the longer lobe and ending in a smooth area on the distal
end of the shorter lobe; calyces best developed on the longer lobe,
strictly apical, barely showing in a side view of the fully distended
FiQ. 40.—Color pattern of Lampeopeltis caluqaster (U.S.N.M. no. 61726, Jerseyville, Jerset
County, Illinois), showing the striped effect commonly exhibited by dark individuals. About
li XiNAT. SIZE.
organ; fringes few and short; spines short and stout, increasing
gradually in size to a little more than a third of the way to the base,
stopping suddenly; minute spines may succeed the large ones for a
few millimeters; basal portion of organ smooth.
The skull is very similar to that in the getulus group, and to that of
rhomhomaculata. Maxillary teeth, 12 to 14, usually 13 or 14, sub-
equal, the anterior and posterior ones slightly smaller than those
in the middle; mandibular teeth 13 or 14, the posterior smallest,
the fourth to seventh largest; palatines subequal, larger than the
pterygoids, 9, 10, or 11 in number; pterygoids 12 to 19, most com-
monly 16.
120 BULLETIN U4, UNITED STATES NATIONAL MUSEUM.
While there is usually no trouble whatever in distinguishing this
form from rJiombomaculata, every individual test may be expected to
fail in exceptional cases. The possession of 25 or 27 rows of scales
will distinguish calligaster in nearly every instance; yet a specimen
from southwestern Missouri has the formula 23-21-19-17-19, and a
specimen from University, Mississippi, has the formula 23-21-19. So
far no calligaster has been found with only eight infralabials and no
rThomhoraaculata with ten, but both may have nine. The shape of the
head is not reliable, nor are the markings on the head . In calli-
gaster the dorsal blotches tend to have concave anterior and posterior
margins, while in rJiombomaculata these are usually shghtly convex or
straight, and in the latter these blotches are narrower and tend to
come to a point on the sides instead of being more trimcate, as in
calligaster.
Haiitat and Jiahits.—^There have been recorded but few observa-
tions on the natural history of this form. For Illinois Garman
writes (1892, 294) that it "occurs on prairies throughout the State.
Not very common." Blatchley (1899, 545) records the capture of a
specimen in Indiana "in an open woods 2 miles east of Terre Haute,
at a point where the prairie meets the upland. It was crawling
slowly over the ground and did not quicken its speed when pursued,
though it struck rather viciously when caught."
An excellent accoimt is given by Branson (1904, 396-7) in his paper
on The Snakes of Kansas. He says:
This species is qmte numerous tliroughout the eastern part of Kansas and occxirs
in all parts of the State. In the collection of snakes at the Beloit High School there
are eight specimens of Ophibolus calligaster and not more than seven of any other
species. This collection is representative of Mitchell County and indicates that
0. calligaster is as numerous as any other species foimd there. 0. calligaster lives
upon mice, frogs, small fish, etc. I kept three specimens in this laboratory last
suimner. Birds, toads, lizards, mice, insects, and smaller snakes were placed in the
cage with them. They paid no attention to the birds, toads, lizards, and insects,
but attacked the mice as soon as they saw them. They would attempt to swallow
dead mice that were placed in the cage, but always seized them by the middle of
the body. They could not swallow them without beginning at the head and even-
tually gave it up. I captm-ed a specimen in Gove County in August, 1893, that
had just swallowed a mouse. From these observations I conclude that mice are its
principal food. This snake is not often foimd far from water. The ones that I kept
in captivity last suromer stayed in the water most of the time. One of these snakes
molted three, one four, and one two times during their five months' captivity. The
one that molted four times always became cross just before molting and would strike
me when I attempted to handle him. He did not strike hard enough to do any
injury.
The conclusion reached by Branson with regard to mice is fully
confirmed by Mr. Mackelden, of St. Louis, who writes that
—
On Easter Sunday, 1917, I collected 38 Lampropeltis calligaster along two ravines
in Jersey County, Illinois, and every one of them when caught disgorged from 1 to 8
REVISION OF THE KING SNAKES. 121
mice, mostly young mice. We saw a great many more, but did not attempt to catch
them. I consider the Evans King snake a valued asset to the farmer, for it comes
out of hibernation earlier than most other species and lives largely upon mice, rate,
and moles. I have never been bitten by the Evans King snake nor had one attempt
to bite me, although I have caught hundreds of them. My experience has been
that this snake does most of its himting for food between sun-up and 10 o'clock. It
is seldom that I have caught one out in the open in the afternoon.
Seven specimens of caUigaster were taken in an open field along a
small ravine in Jersey County, Illinois, early in April, 1919, by Mr.
Mackelden and Mr. Froman A. Beach. One specimen disgorged
three nestlings of the field mouse, Microtus acrogaster, about five
hours after being caught. The mice showed no evidence that diges-
tion had begun.
Eleven eggs of caUigaster were plowed up early in August, 1918,
by Mr. Beach. These all hatched about a month later.
Range.—This form is Imo'svn from western Indiana south to Mis-
sissippi and Louisiana, westward to western Texas, and northeast to
Minnesota. It appears to be v/ell known only from Illijiois, Missouri,
and Kansas. The limits of its range can at present be only sur-
mised. It should be expected in western Kentucky and Temiessee.
The first record for Mississippi rests upon a specimen only recently
sent to the Museum of Zoology of the University of Michigan from
University, Mississippi, and the two records for Louisiana (Cornell
Univ. no. 7154, Jennmgs, Jefferson Davis County, Louisiana, October,
1906, A. G. Hammer, collector; no. 7153, Chastme Natchitoches
County, Louisiana, Apr. 2, 1915, K. P. Schmidt, collector) have not
heretofore been recorded. It has several times been said to occur in
Wisconsin, but no definite record has been cited. A specimen in the
Museum of the Academy of Natural Sciences of Philadelphia bears
the label "Minnesota." There is no record for Iowa or Nebraska
and for Arkansas only the extreme northeastern corner of the State,
Greenway. Writing for Texas, Strecker (1915, 38) says, "this beau-
tiful King snake has been reported from only a few scattered locali-
ties. It is found in the neighborhood of Waco, but is extremely
rare. In addition to the localities represented by specimens in the
United States National Museum, specimens from other museums have
been examined from the following localities: "Minnesota;" Wme-
mac, and Vigo County, Indiana; Calhoun, Charleston and Cooks
Mills, Illmois; Manhattan, Wathena, Salt Creek, La^vrence, Rock
Creek, and Osage and Labette Counties, Kansas; Alva, Oklahoma;
Jennings and Chastme, Louisiana; University, Mississippi.
Published records for other localities are as follows : Pekin, Illinois
(Garman, H., 1892, 293); Deming's Bridge, Matagorda County,
Texas (Garman, S., 1892, 9); Greenway, Arkansas (Hiu'ter and
Strecker, 1909, 26) ; and the foUowmg counties in Kansas (Branson,
122 BULLETIN 114, UNITED STATES NATIOISrAL MUSEUM.
1904, 376), Mitchell, Lyon, Franklin, Republic, Shawnee, Scott,
Gove, Logan, Geary, Sumner, Miami, Pottawatomie, and Neosho.
This form has been reported from Sapulpa, Oklahoma, by Mr. Karl
P. Schmidt (1919, 71), but examination by the writer of the specimen
upon which this record is based shows it to be a very different snake,
Fig. 41.—Map showing locautt eecoeds foe Lampeopeltis calugastee.
but one which is frequently confused with calligaster, that is, Ela-
pJie laeta.
It is reported for Lancaster, Fairfield County, Ohio, by W. H.
Smith (1882, 689), but, from the description, the identification is
obviously mcorrect. This is doubtless the basis of Morse's report of
later date (1904, 130).
There is a specimen m the collection of the American Museum of
Natural History, labeled "west coast of Mexico," Frank Trubaudt,
collector, but it seems to be impossible to verify this record, or to
learn anything more definite as to the locality. It is not at all im-
REVISION OF THE KING SNAKES. 123
possible that its range may actually extend to the west coast of
Mexico, but the record here cited is too indefinite to bear any reliance.
It may be remarked as bearing upon the unreliability of this record
that there is also at the American Museum a specimen of holhroohi
(no. R4298) labeled ''West Coast of Mexico, Frank Trubaudt,
collector."
Summary of certain structural characteristics of calligaster.
Ventral Dorsal Tail divided hy i-
plates. blotches.
"3
1
total length.
"
.
Locality.
a
i 0) i
"c a
sK 5y^ > X > g
H t> > 3
» < W < M w <! w < :?;
Texas 196-209 205.2 61-69 65 7.08 9.33 0.143
0.121- .137 0.128
49-50
38-52
53
45
Male...
Female.
3
3
Kansas 196-213 204.0 57- 78 68 7.05 9.40 .124- .150
.115- .135
.138
.123
49-57
41-48
52
45
Male...
Female.
12
9
St. Louis and vi- 195-214 205.2 46-68 57 7.08 9.23 .124- .149 .140 46-56 52 Male... 15
cinity. .110 .125 .122 40-47 44 Female. 9
Illinois and west- 201-212 206.0 53-69 60 7.11 9.22 .123 .143 .136 44-53 50 Male... 17
ern Indiana. .112- ,145 .126 42-46 45 Female. 10
Whole range 195-214 205 46-78 62 7.08 9.28 .123- .150
.110- .145
.1.37
.123
44-57
38-52
51
45
Male...
Female.
47
31
Scaleformulae of calligaster.
Locality.
25-27-25-
23-21 25-23-21
23-25-2a-
21
23-25-21-
19
23-25-23-
21-19 Total.
Male. Fe-
male. Male.
Fe-
male. Male.
Fe-
male. Male.
Fe-
male. Male.
Fe-
male. Male.
Fe-
male.
Texas 2
2
2
1
1
1
3
1
1
1
1
5
1
2
3
1
2
1
1
1
2
1
......
3
1
7
7
14
......
4
3
4
12
15
13
4
Kansas 1 9
St. Louis and vicinity .... 11
Illinois and Indiana 1 1 9
Total 1 7 6 8 7 5 5 6 29 8 44 33
One male, Jasper County, Missouri, has the formula, 23-21-19-17-19.
One male, Umversity, Mississippi, has the formula, 23-21-19.
Variation and affinities.—The number of specimens and their dis-
tribution throughout the area inhabited by this form are entirely
inadequate for the demonstration or even detection of geographic
variations. The form appears to be very homogeneous throughout
its range. It is not unlikely, however, that large series of specimens
would reveal geographic tendencies or local differences that may not
now be even suspected.
The accompanying tables of structural characters summarize the
situation as well as can be done with the material at hand.
The table of scale formulae is built upon so little data that it can
do no more than suggest the likelihood that specimens from the region
186550—21—Bull. 114 9
124 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
from St. Louis to Indiana may have a lower average formula than
those from farther west and southwest.
Sex influence on the number of scale rows seems evident. The
large majority of individuals that attain the maximum number of
rows, 27, are females; of those possessing the lowest formula, three-
fourths are males ; of the individuals having the formula next to the
highest, the majority are females; and of those possessing the inter-
mediate formulae, 23-25-23-21-19 and 25-23-21-19, about half are
males and half females.
This form is not at present known to intergrade with rTiomboTnacu-
lata, but this may be due to our lack of knowledge of the situation in
eastern Mississippi and western Alabama and Tennessee. The close
relationship between calligaster and rhombomaculata is more fully
brought out under the discussion of the latter form.
REVISION OF THE KING SNAKES. 125
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REVISION OF THE KING SNAKES. 127
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128 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
LAMPROPELTIS RHOMBOMACULATA (Holbrook).
molecatcher; beown king snake.
Fig. 42.
1840. Coluber rhombomaculatus Holbrook, N. Amer. Herp., ed. 1, vol. 4, p. 103,
pi. 20 (type locality, Georgia and Alabama).
—
Coronella rhombo-maculata
Holbrook, N. Amer. Herp., ed. 2, vol. 3, 1842, p. 103, pi. 23.—Jan,
Arch. Zool. Anat., vol. 2, fasc. 2, 1863, pp. 237, 243; Icon. Gen. OpMd.,
livr. 17, 1866, pi. 2, figs. 1, 2.
—
Ophibolus rhombomaculatus Baird and
GiRARD, Cat. N. Amer. Rept., pt. 1, 1853, p. 86.—Baird, Pacif. R. R.
Surv., vol. 10, pt. 3, no. 1, 1859, pi. 30, fig. 60.—Cope, Bull. U. S. Nat.
Mus., no. 1, 1875, p. 37.—Yarrow, Bull. U. S. Nat. Mus., no. 24, 1882,
p. 94.
—
Davis and RicE, Illinois State Lab. Nat. Hist. Bull. no. 5, 1883,
p. 34; Bull. Chicago Acad. Sci., vol. 1, no. 3, p. 29 (Southern Illinois—
probably an en-or).-Cope, Proc. U. S. Nat. Mus., vol. 11, 1888, p. 381;
same, vol. 14, 1891, p. 610.
—
Garman, H., Bull. Illinois State Lab.
Nat. Hist., vol. 3, art. 13, 1892, p. 296.—Cope, Rep. TJ. S. Nat. Mus. for
1898. 1900, p. 903, fig. 222, pi. 18, fig. 5.—Brown, Proc. Acad. Nat.
Sci. Philadelphia, 1901, p. 79.—Miller, Proc. Biol. Soc. Washington,
vol. 15, 1902, p. 36.
—
Hay, same, p. 90.—Brimley, Journ. Elisha
Mitchell Sci. Soc, vol. 21, no. 4, 1905, p. 152; same, 1907, p. 146.—
DiTMARS, ReptUe Book, 1907, pp. 341, 354, pi. 103, figs. 5, 8, pi. 104.—
Brimley, Journ. Elisha Mitchell Sci. Soc, vol. 30, no. 4, 1915, p.
202.
—
Lampropeltis rhombomaculata Cope, Proc. Acad. Nat. Sci., Phila-
delphia, 1860, p. 255.—Stejneger, Proc. TJ. S. Nat. Mus., vol. 14;, 1891,
p. 503.—DxJNN, Copeia, no. 18, 1915, p. 6.
—
Stejneger and Barbour,
Check List, 1917, p. 89.
—
Lampropeltis rhombomaculatus Hay, Proc.
Biol. Soc. Washington, vol. 15, 1902, p. 138.—Schufeldt, Guide to
Natiu-e, vol. 5, no. 9, p. 269, fig. 2.
1883. Ophibolus triangulus rhombomaculatus Garman, S., Mem. Mus. Comp.
Zool., vol. 8, no. 3, pt. 1, p. 156.
1894. Coronella calligaster (part) Boulenger, Cat. Snakes Brit. Mus., vol. 2,
p. 199.
Description.—The scutellation is as follows: Ventral plates, 191
to 213; caudals, 31 to 55 (males 37 to 55, average 45; females 31 to
44, average 41); supralabials, 7; infralabials usually 8, often 9;
oculars, 1 and 2; temporals 2+3+4; posterior chin shields a little
shorter than the anterior or about equal, parallel, only occasionally
separated from each other by a small scale; suture between the
internasals only about haK as long as that between the prefrontals;
loreal usually longer than high, sometimes as high as long; scale
rows on middle of body 21 or 23, formula usually 21-19 or 21-23-21-
19.
In bodily proportions this form is very similar to calligaster; the
snout is generally shorter and blunter, and apparently the whole
animal averages somewhat shorter and more slender. The tail
varies from 0.100 to 0.150 of the total length (males, 0.111 to
0.150, average 0.124; females 0.100 to 0.131, average 0.119). The
largest specimen examined was from Raleigh, North Carolina, and
REVISION OF THE KING SNAKES. 129
measured 1,137 mm., although Hay records a specimen 1,150 mm. iu
length (1902, 90).
The pattern is very similar to that of calligaster, but in adults it
is, as a rule, so greatly obsciu-ed that an ordinarj' individual presents
a nearly uniform brown appearance. In the yoimg, however, and
in some adults, the markings are well defined.
The body pattern consists of about 55 (48 to 64) transversely
elongate dorsal blotches, about 1^ to 2 scales long, and 8 to 11 scales
wide, that become narrower on the sides, and extend down to the
FiQ. 42.—Color pattern of Lampropeltis rhombomaculata (U.S.N.M. no. 16832, Brooklanp, Dis-
tEicT OF Columbia). About ij x nat. size.
sixth or fifth row of scales (fig. 42). On the tail they become much
narrower. The two series of lateral alternating blotches are narrow,
irregular, and commonly fused into a single series in alternation
with the blotches of the dorsal row. Small dark spots are often
present, posteriorly, between the spots of the lateral series. The
belly is white or yellowish, and checked, scantily or heavily, with
dark brown. Large dark individuals often exhibit a lengthwise
striping precisely like that described for calligaster (fig. 40), and first
observed by Miller in a specimen from Alexandria, Virginia (1002, 36).
130 BULLETIN 114, UNITED STATES E-ATIONAL MUSEUM.
The head markings are like those of calligaster, but much less dis-
tinct. The prefrontal bar may be just visible; the forked mark on
the parietals is evident; the elongate bands on the neck, and the
dark diagonal from the lower postocular to the last supralabial are
nearly as well developed as in calligaster.
The groimd color is a light or dark brown or greenish, lighter on
the sides. The dorsal blotches are a darker brown or reddish, nar-
rowly margined with black. In young individuals the blotches are
a dark red, and the belly is yellowish.
The penis is closely similar to that of calligaster. It may be de-
scribed as bilobed; the sulcus single, and passing over the end of the
larger lobe where it is continuous with a narrow smooth area which
widens on the distal end of the smaller lobe; this smooth area bor-
dered by a few calyces, most numerous on the larger lobe, but still
so few that they are scarcely visible in a side view of the fully dis-
tended organ; fringes extremely short, becoming modified into short,
stout spines which increase in size basally; less than half way down
from the end of the organ the large spines replaced abruptly by a
few scattered minute ones; basal portion smooth.
The skull is essentially like that of calligaster. The dentition is
as follows: Maxillaries 12 to 15, usually 12 or 13, subequal, but dis-
tinctly decreasing in size anteriorly and posteriorly; mandibular
teeth 12 to 16, usually 12, decreasing in size before and behind, the
third to sixth largest; palatines, 9 or 10, subequal; pterj^goids usually
14, varying from 13 to 16, subequal, smaller than the palatines.
Habitat arid JiaMts.—^This form was rare in collections until about
1888 when it was found near the District of Columbia, on the Virginia
side of the river. Since then many examples have been taken in
this vicinity on both sides of the river. Its apparent rarity is un-
doubtedly due to its secretive and burrowing habits. It is sometimes
found in the open. Ditmars (1907, 355) reports finding one in Fair-
fax County, Virginia, "that lay stretched upon a grassy bank, enjoy-
ing the warm rays of a spring sun. When captured it defended
itself vigorously for the moment, but soon became quiet. Having
no receptacle in which to place the snake, the writer carried it several
miles coiled quietly about his hand. Its only symptom of anger was
an occasional shaking of the tail." One taken near Mobile, Alabama,
last spring by Mr. W. R. Jones was found "crossing a dry road in
the latter part of the afternoon." Mr. Jones describes the situation
as "cut-over pine land with more or less scrub oak. Not much
brush, as it is frequently burned over. Light soil, sandy loam;
about 600 yards from nearest water." Frequent notes on labeled
specimens indicate that it is most commonly found in the ground,
and turned out by the plow or in excavations. Mr. E. R. Dunn
thinks they can be readily found by following a plow in any upland
REVISION OF THE KING SNAKES. 131
field near the District of Columbia, and reports seven plowed up in
one 12-acre field in middle Virginia.
In regard to its food in captivity, Ditmars (1907, 355) says it
"feeds upon the young of other snakes and upon lizards as well, but
seems to prefer small rodents and birds," and to be especiallj^ fond
of "very young birds. Some specimens refused these tempting
morsels, and ultimately starved to death. Generally speaking, these
snakes are sluggish and uninteresting in captivity." One example
Fig. 43.—Map showing locality records for Lampropeltis riiombomaculata.
found by Mr. Dunn in Nelson County, Virginia, disgorged a mouse.
In captivity they will feed upon dead mice.
Accurate observations on the natural history of this form are very
much to be deshed.
Eange.—This species is at present known from Montgomery
County, Maryland, south through the Atlantic States to northern
Florida, west to Mobile County, Alabama, and northeast to Knox-
ville, Tennessee. Until about 30 years ago it was considered a rare
snake, and was known only from the Carolinas to Alabama. Since
that time numerous examples have been taken from Ealeigh, North
Carolina, from Nelson County, Virginia, and from the vicinity of
Washington, District of Columbia. Now that we know better how
132 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
to look for it, we should be able to extend our knowledge of its
range.
It is highly desirable to learn how far west it occurs in order to
see if there is any approach in structural characters to calligaster.
It will undoubtedly be found farther north in Maryland, perhaps
even in New Jersey.
Published records for localities not included in the list of speci-
mens examined are as follows: Illinois (Davis and Rice, 1883, 34),
undoubtedly an error; Groveton, Virginia (Ditmars, 1907, 345).
Furthermore Mr. Dunn reports finding examples at Warminster,
Norwood, Midway Mills, and Wingina, in Nelson County, and at
Bent Creek in Appomattox County, Virginia.
Variation and affinities.—The study of variation in this form can
not be undertaken at present with any satisfaction, due to the totally
inadequate series of specimens available, and their unequal distri-
bution over the area inhabited by the species. The accompanying
table has been prepared as a summary of the situation as it now
stands.
Summary of certain structural characteristics of rhomhomaculata.
Region.
Knoxville, Tennessee.
.
Alabama, Georgia, and
Florida.
Southern Virginia to
South Carolina.
District of Columbia
and vicinity.
Whole range
Ventral
plates.
202-205
}l97-208
|l96-213
|l91-211
191-213
203.5
202.4
203.7
200.3
201.5
Dorsal
blotches.
f^
51-55
50-58
4&-60
48-64
48-64
W
7.00
7.00
7.03
7.00
7.01
8.50
8.50
8.38
8.13
8.29
Tail divided by-
total length.
r
0. 150
t
.128
r0.127- .139
.
121-
. 148
.
112-
. 123
. Ill- . 135
.
101-
. 130
150
131
/ .111 .]
\ .101- -1
0.133
.132
.118
.124
.117
.124
.119
Caudals.
55
44
47-52
4:
43-50
42-43
37-48
31-43
37-55
31-44
Male..
Femalej
Male..
Feraalej
Male..
Female
Male..!
Female! 13
Male.. 37
Female 18
Upon comparison with the similar table for caUigaster, it will be
noticed (1) that the averages are strikingly similar for everything
except the labials, (2) that they are, in nearly every instance, slightly
lower than the corresponding figures for the other form, (3) that the
ventrals, infralabials, and caudals reach their lowest averages at the
most northerly locality, and that here this form is farthest removed
both structurally and geographically from calligaster.
Not only in its externally visible features of scalation, pattern,
and bodily proportions, but also in its penial and dental characters
does it demonstrate its close relationship with calligaster, and with
or through calligaster its near relationship to the members of the
getulus group. Its closest relatives, therefore, are all forms that get
REVISION OF THE KING SNAKES. 133
their living chiefly above ground. But it appears that this is largely
a burro\\dng form. Certain degenerative tendencies may therefore
be expected. The infralabials furnish one instance, the averages for
which, although admittedly based on a very inadequate number of
specimens, can hardly be without significance. This decrease in
number of infralabials is probably to be correlated with a shortening
of the snout, and the latter is very likely associated wdth burrowing
habits. One of the characteristic differences between this form and
caUigaster is the shorter and blunter snout, a difference that may
usually be observed but that is not easy to measure.
The number of specimens is too small to reveal geographic tend-
encies in the number of scale rows, but, for comparison with the
condition in caUigaster, it should be noted that along with the dis-
tinctive difference in numbers of labials and the somewhat smaller
average size of body, there is a decided decrease in scale formula,
an average difference of two rows on each side of the body.
A sexual difference in scale formula is fairly evident. The figures
in the table may be summarized as follows: (1) About half of the
individuals of each sex have the formula 21-19; (2) nearly all the
rest of the females have a higher foiinula, 21-23-21-19 (one of those
having the formula 19-21-19, U.S.N.M. no. 17444, is a highly
aberrant individual in other matters of scalation), and most of the
males have a lower; (3) no female has 17 rows at the end, while this
is the case with about one-fourth of the males.
In pattern as well as in scutellation and proportions, this form shows
itself to be a reduced caUigaster. Moreover, the differences are but
slight. The markings on the head are in every way similar, but
are less well defined. The smaller average number of dorsal blotches
may be accounted for by the shortening of the tail, but these blotches
are different in shape. Their anterior and posterior borders are
generally slightly convex, instead of concave, as in caUigaster, and
they taper nearly to a point on the sides, instead of being truncate
or blunt, as in the latter form. The fact that they extend down on
the sides commonly to the fifth row of scales instead of to the seventh
or sixth is due to the loss of one or two rows on each side of the body.
This lower extension of the dorsal blotches may account for the fact
that the two lateral series of alternating spots, characteristic of
caUigaster, tend to fuse into a single series. Degeneracy is indi-
cated by the contracted and transversely zigzag shapes that tliis
lateral series assumes. Then, too, the tendency for the pattern to
be indistinct seems to be an accentuation of the condition illustrated
by the caUigaster of the region from eastern Missouri to Indiana.
All the differences between rhomhomaculata and caUigaster are
traceable to reduction or degeneration and not to any further de-
velopment of the characteristics of the latter form.
134 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
We may therefore summarize the situation with regard to rhom-
homaculata by considering it a derivative of calligaster by (1) a
shortening and strengthening of the head, developed in response
to burrowing habits, and particularly noticeable in the shape of the
snout; (2) a reduction of one in the number of infralabials, correlated
with the shortening of the head; (3) an average reduction of one tooth
in the maxillary, palatine, and mandibular series, and of two in the
pterygoid series; (4) a reduction in number of scale rows, amounting
to about two on each side of the body; (5) a shght reduction in
numbers of ventrals and caudals, and, correlated with the last, a
decrease in the proportionate tail length; (6) reduction and fading
of the pattern; and (7) a slight decrease in body length and girth.
Scale formulae of rhombomaculata.
Locality.
21-23-21-19 21-19 19-21-19
19-21-19-17-
21-19-17
Male. Fe-
male. Male.
Fe-
male. Male.
Fe-
male. Male.
Fe-
male.
District of Columbia and vi-
cinity 2
1
4
3
10
11
2
1
7 3 2 4
3
1
1
Southern Virginia to South
Carolina
Georgia, Florida, and Ala-
bama 2
Tennessee
Total 3 7 24 9 3 2 9
Total number of males, 39.
Total number of females, 18.
REVISION OF THE KING SNAKES. 135
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136 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
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REVISION OF THE KING SNAKES. 137
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138 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
LAMPKOPELTIS LEONIS (Giinther).
1893. Coronella leonis Gunther, Biol. Cent. Amer., Rept. and Batr., p. 110,
pi. 39, fig. A (type locality, Nuevo Leon, Mexico; type in British Mu-
seum; W. Taylor, collector).
—
Boulenger, Cat. Snakes Brit. Mus.,
vol. 2, 1894, p. 199.
Description.—^As this form is known only from the type speci-
men, the original description is here quoted in full
:
Scales in 23 rows, without pit, smooth; head similar to that of Coronella laevis;
anterior frontals not quite half the size of posterior; vertical 5-8ided, with the lateral
margins convergent; 1 praeocular not reaching the vertical; 2 postoculars; loreal
longer than deep; 7 upper labials, the third and fourth entering the orbit; temporals
2+3, of the 2 anterior only the upper one is in contact with the postoculars. Ventrals
200; anal entire; subcaudals 50. Body pale olive-color on the back, with 27 salmon-
colored incompletely black-edged spots, some being of a transversely oval shape, but
the majority presenting the appearance of being formed of 2 rounded portions. On the
tail the spots lose their light center and appear merely as brown spots. The lateral
spots which are so conspicuous in most variations of Coronella triangulum are here
nearly entirely absent. Vertical and each occipital with a black spot, red in the
center; abdomen with only a few blackish blotches irregularly scattered. A black
band along the middle of the lower part of the tail. The single specimen measured
23^ inches, the tail 3f inches. This snake may be considered to be one of the aberrant
forms of Coronella triangulum.
Remarlcs.—The description is incomplete in some respects, and
the type specimen has not been available for examination. If, as
stated above, scale pits are actually absent, in which case other
distinctive differences will almost certainly be found, the specimen
can hardly be a Lampropeltis, but an error may easily have been
made on this point. Further examples must be obtained before the
status of this form can be stated with assurance. But from the
description and excellent figure, it appears to be a distinct form
closely allied to the other members of the calligaster group. Further
discussion, however, is hardly profitable at present.
SUMMARY.
The handicap of insufficient material has been more than usually
acute in the study of this group.
The intimate relationship existing between rhorribomaculata and
calligaster and the fact that the former is a derivative by reduction
of the latter need not here be more than recalled, after the discus-
sions aheady giVen under Variation and Affinities of these forms.
That leonis is a member of the group must be allowed, pending
further knowledge of the form.
We may therefore express the relations of the forms of the caUi-
gaster group, diagramatically, as follows:
leonis calligaster ^rJiomoomaculata
The Atlantic States—the ''Southeast"— may therefore be excluded
from consideration as a possible center of origin or dispersal of this
group.
REVISION OF THE KING SNAKES. 139
Of the present known range of calligaftter, the northeast has points
distinctly against its consideration as the required center, namely,
(1) this portion is north of the southern limit of glaciers, (2) the
pattern appears to be degenerate (indistinct) here, (3) the number of
scale rows is somewhat diminished in the northeast, (.4) the infra-
labials are here less often 10. We must therefore look south of the
Missouri River and west of the Mississippi for the center of the
group. Of this region we loiow very little of the situation outside
of Kansas, and this is too far north to be a center of reptilian preserva-
tion and radiation.
We are forced therefore to look to the "Southwest"
—
perhaps
Texas or northern Mexico—for the center of dispersal of this group,
and to await fuller information before being more specific.
THE TEIANGTJLUM GBOUP.
LAMPROPELTIS POLYZONA Cope.
COEALILLO.
Fig. 64.
1858. Coronella doliata, var. A Gunther, Cat. Colubr. Snakes Brit. Mus., p. 42.
—
Salvin, Proc. Zool. Soc. London, 1861, p. 227 (Vera Paz, Guatemala).—
Ophibolus doliatus Duges, La Naturaleza, 1876, vol. 3, pp. 222-226, fig.
1860. Lampropeltis polyzona Cope, Proc. Acad. Nat. Sci. Philadelphia, p. 258
(type locality, Quatupe, near Jalapa, Mexico; type specimen, Acad. Nat.
Sci. Philadelphia, no. 9770; Mr. Pease, collector); same, 1861, p. 302
(Mirador, Vera Cn\z).—Ophibolits polyzonus Cope, Proc. Amer. Philos.
Soc, vol. 11, 1869, p. 162.—SuMicHRAST, Bull. Soc. Zool. France, 1880,
p. 181.—Ferrari-Perez, Proc. U. S. Nat. Mus., 1886, p. 187.
1861. Lampropeltis micropholis Cope, Proc. Acad. Nat. Sci. Philadelphia, p. 302
(Minatitlan, Mexico).
—
^McLain, Contr. Neotr. Herp.,1899, p. 5.
—
Coronella
micropholis Boulenger, Cat. Snakes Brit. Mus., vol. 2, 1894, p. 203, vars.
B, C, E, F.—Werner, Rept. Batr. Centralamerika, Chili, 1896, p. 10.—
BoETTGER, Kat. Rept.-Samml. Frankfurt, pt. 2, 1898, p. 72. ^Werner,
Abhand-Akad. Wiss. Munchen, vol. 22, pt. 2, 1903, p. 347.—Gadow, H.,
Proc. Zool. Soc. London, 1905, vol. 2, p. 196 (var. B, Chilpancingo, San
Luis Allende).
—
Barbour and Cole, Bull. Mus. Comp. Zool.,vol. 50,
no. 5, 1906, p. 152 (Chichen Itza, Yucatan).
—
Gadow, H., Zool. Jahrb.,
Abt. Syst. Geog. Biol., vol.31, 1911, pp. 6, 24, pi. 1, figs. 6, 7 (Carrizal,
S. Michoacan; Chilpancingo, Guerrero).
—
Sternfeld, Sitz. Gesell. Nat.
Freunde, 1913, pp. 105, 106, 115, pi. 10, fig. 4.
1861. Coronella doliata, \B,r.formosa Jan (part). Icon. Gen. Ophid., livr. 14, Dec,
pi. 4, fig. B (no locality given); Arch. Zool. Anat., vol. 2, fasc. 2, 1863,
pp. 237, 241 (Mexico).
1883. Ophibolus triangulus zonatus Garman, S., Mem. Mus. Comp. Zool., vol. 8,
no. 3, pt. 1, p. 67 (Acapulco, Mexico).
1886. Coronellaformosa Bocourt, Miss. Sci. Mex., p. 612, pi. 39, figs. 3, 3a to 3c.
1886. Coronellaformosa polyzona Bocourt, Miss. Sci. Mex., p. 615, pi. 39, figs. 7,
7a-7d (Oaxaca).
1886. Coronella formosa anomala Bocourt, Miss. Sci. Mex., p. 614, figs. 4, 4a-4c
(type specimen in Paris Museum; tj'pe locality Haute Vera Paz, Guate-
mala).
186550—21—Bull. 114 10
140 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
1886. Coronellaformosa oligozona Bocourt, Miss. Sci. Mex., p. 614, pi. 39, figs. 8,
8a-8c? (Tehuantepec, west slope of Guatemala) (two cotypea in Paris
Museum; type locality, Tehuantepec),
1887. Ophibolus doliatus cocdneus (part) Cope, Bull. U. S. Nat. Mus., no. 32,
p. 78 (Guatemala, Zacualtipan).
1887. Ophibolus doliatus polyzonus (part) Cope, Bull. U. S. Nat. Mue., no. 32,
p. 78; Proc. U. S. Nat. Mus., vol. 11, 1888, p. 382; vol. 14, 1891, p. 608;
Amer. Nat., vol. 27, 1893, p. 1067.
—
Osceola doliata polyzona Cope, Rep.
U. S. Nat. Mus. for 1898, 1900, p. 898.
1903. Coronella micropholis arcifera Werner, Zool. Anz., vol. 26, p. 250 (type
specimen in Naturhistorischen Museum in Briissel; type locality, Mex-
ico).
In the same paper in which the original description of polyzona
appeared Cope described a specimen from Panama under the name
of micropholis. He later decided that it was identical with polyzona,
and thereafter called both by the latter name. Other authors
have likewise considered the two names synonymous, but have
generally used the name micropTiolis in preference to polyzona, be-
cause it appeared a page earlier than the latter. The present
review, however, has disclosed the fact that Cope really did have two
very distinct forms before him. The great variability and relative
abundance of specimens of the northern one and the scarcity of
specimens of the southern form have hitherto obscured the distinct-
ness of the latter.
The paper containing the original descriptions of polyzona and mi-
cropolis was read before the Academy of Natural Sciences of Phila-
delphia in June, 1860, and was probably published before the end of
that year in one of the quarterly issues of the Proceedings. The
volume for 1860 bears the date 1861. It seems to be impossible to
determine the exact date for the whole volume or for the quarterly
portion containing Cope's paper, but it in all probability antedates
Jan's figure of Coronella doliata, y&t.formosa of December, 1861 . Jan's
formosa, however, was a composite, as a study of his figures B and B*
shows, and the writer therefore, in accordance with his privilege as
first reviser, designates as the type of formosa figure B* (Jan. 1861,
livr. 14, pi. 4), thus making the type locality definite and fixing the
name formosa Jan as a synonym of micropholis Cope.
Description.—In Mexico and Central America this is the best known
form of the genus. The following summary of scutellation is based
upon the examination of 48 specimens and published records of about
a dozen more: Ventral plates, 208 to 239; caudal, 42 to 61; 1 pre-
ocular; 2, very rarely 1 or 3, postoculars; supralabials, 7, rarely 8;
infralabials, 9, sometimes 8, less often 10; temporals usually 2 + 3+4;
loreal usually present, and longer than high, sometimes about as
high as long, occasionally absent, its place being then occupied
chiefly by a backward extension of the posterior nasal; posterior chin
REVISION OF THE KING SNAKES. 141
shields usually a little shorter than anterior, in contact with each
other, or occasionally separated by 1 or 2 small scales. The maxi-
mum number of scale rows is nearly always 23 or 21, becoming
posteriorly 19 or 17, but examples from Costa Rica and Yucatan may
not have more than 19 rows.
The proportions are nearly the average for the genus. The head
is widest at the temples and tapers toward the snout, which is blunt;
it is but slightly distinct from the body; the latter is cylindrical,
fairly stout, and of approximately uniform diameter; the tail is
short and stout, but averages a little longer than for most forms
of the group; its length in proportion to the total length varies
from 0.124 to 0.164 (males, 0.130 to 0.164, average 0.144; females
0.124 to 0.140, average 0.131). The largest specimen examined was
1,580 mm. in length, and was from Nicaragua; however, a dried
skin of one from the same country measured 1,610 mm.
The body pattern is made up of from 17 to 37 paired rings of
black bordering narrow annuli of white or yellow and separated
by broader ones of red. The dorsal scales of the red areas are
usually strongly tipped with black, but these black tips are some-
times very small or absent altogether. The white rings are ^ to 1^
scales in width, widening but little if any on the first row of scales;
the black rings are from 1| to 3 scales wide on the middorsal line,
and, rarely, excluding the red altogether from some of the annuli.
The red rings, although not infrequently narrower, are usually
wider than the groups of black and yellow rings that separate them.
The scales of the yellow rings may be uniform in color, but more
often they are strongly tipped and mottled with black, and infre-
quently the latter color may almost obliterate the yellow.
All of the annuli are usually continuous across the belly, but some
specimens from the State of Vera Cruz and from Yucatan Peninsula
show the red bands to have widened on the sides so far as to exclude
the black and yellow annuli altogether from the belly. The yellow
then appears in dorsal semicircles, bordered with black. (Further
change in this direction probably accounts for Boulenger's variety
E, 1894, 205.) In a few specimens the belly is much overspread
with black, but usually the black is conspicuous only as a blotch
in the yellow rings, and it may be absent even there.
The head is black except for a more or less perfect cross band of
white or yellow on the snout, chiefly on the prefrontal plates. It is
sometimes much reduced, and rarely absent. On top of the head
the black extends back to the posterior portion of the parietal plates,
on the supralabials it extends as far back as the fifth. The first black
band is close behind the parietals, or involves the tips of these plates,
and in the great majority of cases is continuous across the throat.
142 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
Polyzona may be distinguished from annulata and from nelsoni
by the presence of black tips on the red scales; by the snout, which
usually has a transverse light band in polyzona, is entirely black in
annulata, and is mottled with dark and light in nelsoni. From
micropholis it may be known by the white cross band on the snout
instead of the white snout with black transverse marks, and by
the fact that the yellow annuli are narrow, and their scales plaia
or more or less mottled with black as well as black-tipped, instead
of being wide with the scales heavily tipped with black and not
mottled, and by the presence of two anterior temporals instead of
usually only a single large one.
The dental characters are as follows: Maxillary teeth, 13 to 15,
usually 14, subequal except that the last 2 or 3 are distinctly, although
not strongly, enlarged; mandibulars, 13 to 15, commonly 14, the
third, fourth, and fifth or the fourth, fifth, and sixth, decidedly the
largest, those posterior much smaller and decreasing slightly in
size; palatines, 10 to 12, commonly 10, subequal; pterygoids smaller,
decreasing, varying from 16 to 22.
The penial characters, as indicated by preserved specimens, are
as follows: Sulcus single; a small space at the distal end without
calyces or with only ridges across it; immediately adjacent to this
a few calyces with only a very few fringes; succeeding calyces with
numerous fringes, gradually changing to spines; latter closely set
together, numerous and slender, increasing in size to about the
middle of the organ, then rapidly decreasing to very small spines,
and disappearing altogether; basal third of organ smooth (or ridged
or ribbed in preserved specimens); no spines distinctly enlarged
or set apart from the others.
Habitat and Tidbits.—Sumichrast (1880, 181) says ''Cette espece
vit dans les terres chaudes et temp^rees des deux c6tes du Mexique.
Sa coloration, analogue h. celles des Elaps, la fait injustement accuser
de venin par les habitants qui J'appellent corallUo."
Cope (1900, 900) quotes, as from the manuscript notes of Sumi-
chrast, as follows
:
Among the numerous Mexican snakes which are called "coralillas, " this one
attains the largest dimensions. It is distributed throughout the warm and temper-
ate regions, but disappears in the alpine region, where, at least, I have never observed
it. This snake prefers shaded localities, as plains covered with tall herbs, and along
rivers. Although of a very harmless disposition, it is not easily caught, since on
being alarmed it glides swiftly through the vegetation and is not long in disappearing
in the gallery excavated by some other animal. It also lives in the enormous nests
of the ant, Orcodoma mexicana, on which it warms itself in the sun. Although entirely
inoffensive, it does not escape the charge of being poisonous, as all the coraUllas are
supposed to be by the natives.
REVISION OF THE KING SNAKES. 143
Range.—This form is apparently the only representative of the
genus in Central America, It is known from Costa Rica to southern
Mexico; in the latter country it occurs at least as far north on the
east side as Tuxpan in the Province of Vera Cruz, and, on the Pacific
side, it is known from Oaxaca, Guerrero, and southern Michoacan.
According to Cope (1900, 900) Sumichrast says of it in his manu-
script notes: "It is distributed thi-oughout the warm and temperate
regions, but disappears in the alpine region, where, at least, I have
never observed it." Apparently the only record at variance with
this distribution is one for the "City of Mexico" (Boulenger, 1894,
204), and this may easily be an error.
PubUshed records for other localities than those listed are as follows
:
Haute Vera Paz (Salvin, 1861, 227; Bocourt, 1886, 614), Retalhuleu
(McLain, 1899, 5), and Cohan, Guatemala (Werner, 1903, 347);
Teziutlan (State of Puebla), and Misantla (State of Vera Cruz)
(Ferrari-Perez, 1886, 187); Acapulco, Mexico (Garman, S., 1883, 67);
Oaxaca (Bocourt, 1886, 616); Carrizal, South Michoacan, west of the
lower Balsas River (Gadow, 1911, 6 and 24); Zamora, Michoacan
(Duges, 1876, 222); Chilpan-cingo, San Luis iUlende (Gadow, 1905,
196) ; in Boulenger's Catalogue of Snakes (1894, 204) are the follow-
ing records: Tierra Colorado and Amula, Guerrero; Teapa, Tabasco;
Huatuzco; Belize; Duenas, Guatemala ; Irazu, Costa Rica; Chiriqui;
City of Mexico.
Variation.—The necessity of a knowledge of the variations in
a form is well illustrated by the present case. Without regard for
normal variation and geographic probabilities, numerous names
have been applied to specimens from Mexico and Central America
with the result that quite distinct forms have been lumped together
and individual variations of the same form, even from the same
locality, have been granted distinction. Confusion is the only end
attained, and that was the situation in the beginning. It is by no
means impossible that fuller series of specimens may demonstrate
the necessity for further subdivision, but it is hoped that a natural
group has been distinguished in polyzona as here defined, so that no
radical rearrangements will be necessary.
The diagram of extremes and averages of ventral plates for certain
regions (fig, 44) shows that throughout all the central portion of the
range, from Nicaragua to southern Mexico, the numbers are high and
fairly constant. North through Tehuantepec and Vera Cruz the
decrease is distinct but not great. The material from Yucatan and
Guerrero is too scanty to be more than suggestive, but differentiation
by isolation is indicated in the former locahty and either that or
approach to nelsoni in Michoacan is the inference in the latter. Costa
Rica and Panama show a definite passage to the low average of
micropholis.
144 BULLETIN 114, TJlsriTED STATES NATIONAL MUSEUM.
The present material is too poorly distributed to show anything
about variation in caudals or tail length, since the numbers for aver-
ages are too much reduced by the necessity of treating the sexes
separately.
rjs>™
^9-
^r^-
m ^ oa
hi o «i*^ Pl<N
S 3
lo-l"'
5-^ ».
2 H -^
3 § S
The number of yellow annuh of the color pattern is so variable that
Httle can be said beyond calling attention to the fact that the averages
for a few localities vary in the same sense as the averages for the
ventrals—Vera Cruz, 25; Guatemala, 31; Nicaragua, 24; Costa Rica,
26; Panama to Ecuador, IS.
REVISION OF THE KING SNAKES.
Scaleformulae of polyzona.
145
Locality.
21-23-21-19 21-19
21-19-17
r.n'l
19-21-19-17
19-17
Male. Female. Male. Female. Male. Female. Male. Female.
Vera Cruz 6
3
7
1
10 4
1Guatemala and Chiapas . .
.
Yucatan
1 2
1
1
i 1
Hondiuras and Nicaragua . . 2 2
Costa Rica 1 1
Total 9 10 3 4 10 6 2
The table of scale formulae proves nothing, but points of interest
indicated by it are (1) that the two higher formulae are evenly
divided between the sexes, and, of the individuals having the two
lower formulae, the majority are males; (2) that the great majority
of the individuals from Nicaragua to southern Mexico have the higher
formulae; (3) that the lower formulae are much commoner at the
extremities of the range; that is, Yucatan, Costa Rica, and Vera
Cruz. Boulenger records (1894, 204) the highest number of scale
rows, 25, from Tabasco, and the lowest number, 19, from Costa Rica.
His two specimens from Guerrero with 21 rows are in keeping with
the suspected approach to nelsonl here.
We may safely say then that in the structural characteristics,
ventral plates, and dorsal scale rows, polyzona shows a]:)proach to
micropholis in the southern portion of its range and to nelsoni on the
west coast of Mexico. Discussion of other structural characters can
not be profitably undertaken here because the specimens at hand are
too few to prove anything by figures. It can onl}'' be remarked that
there appears to be nothing to refute the testimony of the ventrals
and scale rows.
The striking characters of the pattern of polyzona are (1) the black
head with white crossband on the snout; (2) the black tips on the
yellow and the red scales; (3) the narrow yellow annuli not widening
on the sides.
The white snout band may be V-shaped, pointing backward, and
divided on the middle line, represented only by a few dots in the
loreal region, of irregular outline, or prominent, well defined, and con-
tinuous across the snout. It appears to be most often poorly devel-
oped in Vera Cruz; in two from Yucatan it is absent; from Chiapas
to Nicaragua it is usually weU developed; two specimens from Costa
Rica show it broad and transverse. This seems to be a change in the
extreme south toward micropholis, and in the north appears like a
primitive or an extreme condition, certainly not a typical one.
146 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
The black tips on the red scales are exceedingly variable. Nearly
all individuals show them to some degree, but they are absent in a
few. The latter is true of a specimen from Chiapas and of another
from Guatemala. The black tips on the yellow scales are often
accompanied by a dark mottling of the rest of the scale. This is
rarely excessive. Occasional specimens show no black on the yellow
scales. Vera Cruz specimens show no approach to annulata, and the
relationship to nelsoni can not be told for lack of specimens. The
two examples from Costa Kica show no approach in black spotting
of the yellow rings to micropliolis.
The narrowness and uniform width of the yellow wings characterizes
annulata and nelsoni as well as polyzona, and is probably evidence of
relationship. The two Costa Rican specimens both have these rings
Fig. 45.—Map showing locality kecords for Lahpropeltis polyzona.
very narrow. The transition to micropholis is shown by a specimen
from Panama and one from Darien, both of which have the yellow
rings wider than in polyzona but decidedly narrower than Ecuadorean
examples of micropliolis. These have the scales of the yellow rings
strongly black-tipped but less so than is typical of the latter form.
Affinities.—That all the Mexican and Central American represen-
tatives of the triangulum group are closely related hardly needs
argument. It has not been doubted. The difficulty has lain in
correctly defining the several forms and determ.inmg just what the
relation o± each to each is. The evidence from variation and geo-
graphic probabilities is that polyzona is directly related to micropholis
and that it is distinct from annulata on the Gulf side of Mexico. The
nature of its relationship to nelsoni, on the Pacific side of Mexico,
must be learned at a future date.
REVISION OF THE KING SNAKES. 147
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REVISION OF THE KING SNAKES. 149
LAMPROPELTIS MICROPHOLIS Cope.
Fig. 70.
1860. Lampropeltis micropholis Oope, Proc. Acad. Nat. Sci. Philadelphia, p. 257
(type specimen originally at Acad. Nat. Sci. Philadelphia, now appar-
ently lost; type locality, Panama; Dr. John L. LeConte, collector).
—
Fowler, Proc. Acad. Nat. Sci. Philadelphia, vol.65, 1913, p. 168.—Coro-
neZZa ?mcrop/8oKsPERACCA, Bull. Mus. Torino, vol. 11, 1896, p. 9 (Panama);
same, vol. 12, 1897, p. 17 (Cuenca, Ecuador); same, vol. 19, 1904, p. 13
(Vinces, Ecuador); Voy. Exp. Sci. Colombie, 1914, pp. 96-111.
—
Des-
PAX, Bull. Mus. Nat. d'Hist. Nat. Paris, no. 7, 1910, p. 370; Kept. Batr.
de I'Equateur, 1911, p. 28.
—
Griffin, Mem. Carnegie Mus., vol. 7, no.
3, 1916, p. 176 (Cacagualito, Colombia).
1861. CoTonella doliata formosa Jan. Icon. Gen. Ophid., livr. 14, Dec, pi. 4,
fig." B* " (this specimen from Colombia, now probably at Vienna, may
be taken as the type).
—
Boulenger,Bu11. Zool. Soc. France, 1880, p. 44
(Quito, Ecuador).
—
Coronella doliata, var./ormosa Boulenger, Ann. Mag.
Nat. Hist., vol. 9, 1882, p. 458 (Guayaquil).
1887. Ophibolus doUatus polyzonus Cope, Bull. U. S. Nat. Mus., no. 32, p. 78
(Panama, Darien).
Description.—This is the most southern representative of the genus
known. Its scalation is as follows: Ventral plates, 211 to 228, aver-
age about 218; caudals, 40 to 49 (males, 43 to 49, average about 46;
females, 40 to 44, average 42), frequently several entire; suprala-
bials, 7, rarely" 8; infralabials, 9, sometimes 8, rarely 10; one pre-
ocular, two postoculars (fused in one specimen); temporals usually
1 + 2 + 3, varying to 2 + 3 + 4 ; posterior chin shields in contact, shorter
than anterior, or about as long; loreal longer than high or about as
high as long, its upper posterior angle obtuse and its lower posterior
angle acute; scale rows on middle of body 21 or 23, the formulae
commonly 21-23-21-19, or 21-19-17, or 19-21-19-17.
In proportions this form differs somewhat from its nearest rela-
tive, polyzona. The body is fairly stout and of nearly uniform
diameter throughout; the head is more distinctly set off from the
neck; the tail is distinctly shorter and blunter than is usual in poly-
zona, varying from 0.112 to 0.136 of the total length (males, 0.114
to 0.136, average about 0.123; females 0.112 to 0.129, average about
0.118). The largest of the few adults examined measured 1,232 mm.,
and probably came from Ecuador.
The pattern (fig. 70) is conspicuously different from that of poly-
zona, although built upon the same plan. The homologues of the
white or yellow rings, 13 to 21 in number, are much widened and the
dorsal scales of these rings, instead of being tipped or mottled with
black or entirely without black, have each a conspicuous oval black
spot on the distal end, which often occupies more than half the area
of the scale and may sometimes cover the whole scale. Further-
more, the black on the head has so receded as to be continuous only
from the anterior end of the frontal plate to the posterior portion of
150 BULLETIIsr 114, UNITED STATES NATIONAL MUSEUM.
the parietals, and laterally as far as the eyes. The temporals,
labials, and anterior head plates have each a black blotch on the
posterior portion. On the upper labials, beneath the eye, there is a
large black spot.
The rings vary greatly in width, but the red ones are commonly
about as wide as the black and white together. All the annuli are
generally completed on the belly, but the red bands here are often
®rSstw;v.
Fig. 46.—Map showing locauty recoeds foe Lampeopeltis micbopholis.
checked or even completely filled with black, and, opposite the whit-
ish bands, there is often a large black blotch. The dorsal scales of
the red areas are usually black-tipped, but the black may be greatly
reduced or entirely absent.
The colors can not be satisfactorily determined from alcoholic
material, but some of that at hand is in excellent condition. It is
quite evident that the homologues of the red areas of polyzona are
red here also, and that this red may extend across the belly; further-
REVISION OF THE KING SNAKES. 151
more, tliat the homologues of the white or yellow rmgs may be
strongly suffused with red (confirming Cope's original description of
a specimen from Panama) , and that this red may extend across the
belly. The specimens at hand indicate that these rings may, on the
other hand, often be white or yellowish instead of red.
The dental characters from a few specimens are as follows: Max-
illary teeth, 15 or 14, subequal, the last two or three only slightly
larger; mandibular teeth, 15, the anterior distinctly larger, decreas-
ing in size posteriorly; palatines, 10 to 12, more commonly 10, sub-
equal; pterygoids smaller than the latter, decreasing posteriorly, 17
to 19 in number.
Habitat and habits.—^Apparently nothing is recorded upon this
subject.
Range.—^This form is at present known only from Panama, and
from Colombia and Ecuador west of the Andes. In the latter coun-
try it has been taken at 4,200 feet elevation (Fowler, 1913, 168). Its
range meets that of polyzona in western Panama or southeastern
Costa Rica. If intergradation occurs it takes place in this region,
but two specimens from Panama and two from Costa Rica are nearly
typical of their respective forms.
Published records for other localities than those listed are as follows
:
Quito, Ecuador (Boulenger, 1880, 44); Cuenca, Ecuador (Peracca,
1897, 17); Vinces, Ecuador (P. acca, 1904, 13); Nanegal, Ecuador
(Despax, 1910, 370); Angelopolis, Colombia (Peracca, 1914, 96-111).
Variation.—Since che specimens examined are nearly all from
Ecuador, nothing definite can be said about geographic variation.
Enough of individual variation is included in the description.
Affinities.—On geographic grounds polyzona is the only form from
which microplwlis can be derived, but from the description it will
appear that it is very distinct from the latter in numerous structural
as well as color pattern characters. The differences between the two
are, however, only such as can be best explained on the assumption
of the above relationship.
The most conspicuous changes, and how they may be accounted
for, are as follows: (1) The reduced number of temporals has re-
sulted from a reduction in size of the upper scute in each row and an
accompanying increase in the size of the lower plates; (2) the ventrals
have imdergone a slight decrease in number, a change often paralleled
by other forms in the genus; (3) the caudals have decreased dis-
tinctly, a change that is greater than but not different in kind from
that illustrated by other forms; (4) the paired caudals frequently fuse,
particularly near the tip of the tail; such fusion occurs sporadically
throughout the genus, but is more noticeable here than in any other
form; (5) the homologues of the yellow rings of polyzona, here more or
less red, have decreased in number, accompanying a decided increase
152 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
in width; (6) the black tips on most of the yellow scales of polyzona
have here become much accentuated; (7) the black tips on the red
scales have become reduced in area or have disappeared entirely;
(8) the black of the head has receded, leaving a black spot on the
upper labials below the eye, and leaving the snout and temples free
from black except for a spot on the posterior portion of each scute,
and occasional minute mottlings; (9) the number of dorsal scale
rows averages decidedly lower.
The pattern changes represent only a further development of the
pattern of polyzona. The structural changes represent normal varia-
tion ia polyzona and all other forms of the genus, but they are changes
that in other forms, notably elapsoides, are associated with speciali-
zation. In fact, in every way micropJiolis gives the impression that
it is an end form
—
quite beyond consideration as an ancestral type.
It must then be looked upon as derived from polyzona and not as
ancestral to it.
EE\1SI0N OF THE KING SNAKES. 153
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154 BULLETIN U4, UNITED STATES NATIONAL MUSEUM.
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REVISION OF THE KING SNAKES. 155
LAMPROPELTIS TRIANGULUM NELSONI Blanchard.
Fig. 65.
1887. Ophibohis multistratus Cope, Bull. U. S. Nat. Mus. no. 32, p. 78 (Guana-
juata).
1887. Ophibolus doliatus coccineus (part) Cope, Bull. U. S. Nat. Mus., no. 32, p.
78 (Guadelaxara; Colima).
1894. Coronella micropholis, var. A. Boulenger, Cat. Snakes Brit. Mus., vol. 2,
p. 204.
1899. Lampropeltis micropholis oligozona Stejneger, N. Amer. Fauna, no. 14,
p. 70 (Maria Madre Id).
—
^Van Denburgh and Slevin, Proc. California
Acad. Sci., ser. 4, vol. 4, 1914, p. 149.
1920. Lampropeltis triangulum nelsoni Blanchard, Occ. Pap., Mus. Zool.,
Univ. Mich., no. 81, p. 6, fig. 1 (tj^je locality, Acambaro, Guanajuato,
Mexico; type specimen, no. 4(5552, United States National Museum;
E. W. Nelson, collector.)
Description.—This is the west coast representative of the Mexican
Lampropcltes allied to triangulum. Its scutellation mar be sum-
marized as folios: ventral plates, 199 to 231; caudals, 42 to 59;
supralabials 7, infrequently 8; infralabials 9, sometimes 10; pre-
oculars, 1, postoculars, 2; temporals usually 2 + 3 + 4; posterior chin
shields usually in contact, and generally shorter than the anterior;
loreal longer than high, sometimes as high as long; scales rows on
middle of body 21 or 23, the commonest formulae being 21-23-21-19
and 21-19-17.
The bodily proportions are practicall}' the same as for the rest of
the group. The head is only slightly distinct from the neck, the
body cylindrical and of about the same diameter throughout, and
the tail tapers uniformly. The latter varies from 0.120 to 0.150 of
the total length (males, 0.128 to 0.150, average, 0.140; females, 0.120
to 0.137, average 0.129). The largest specimen examined was from
Maria Madre Island, and measured 1070 mm.
The pattern of this form (fig. 65) lilce that of the other Mexican
members of the group, is made up of pairs of black annuli, 13 to
24 in number, bordering narrow rings of white or yellow and separated
by bands of red. It differs from polyzona chiefly in the fact that
the scales of the red areas are never tipped with black, and the pairs
of black rings average fev'er in number.
The black and yellov*^ rings are complete on the belly, and this is
generally true of the red ones, also. Opposite the latter on the
belly, however, there is frequently a blackish mottling, and toward
the interior of Mexico, this space may be as completely filled with
black as in annulata. The yellow rings are of nearly imiform
diameter or may widen a little on the sides. They are sometimes
mottled with darker laterally, and a black spot may or may not be
present within their area on the belly.
186550—21—Bull. 114 11
156 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
The black rings tend to extend their width dorsally at the expense
of the red, and to become narrowed on the sides. They may com-
pletely obliterate the red toward the end of the taU, but on the body
are usually widely separated.
The head is black, except for the region anterior to the frontal
plate, which is much lightened or mottled with dark and light. In
some individuals from the interior this condition is restricted to the
extreme anterior end. It is in contrast with the black snout with
light cross band of polyzona, and the entirely black snout of annulata.
FlQ. 47.—Map showing locality EECOEDS foe LAMPEOPELTIS TEIANGULXm NELSONI.
The dental characters, as indicated by the examination of a few
specimens, are as follows: maxillary teeth, 12 or 13, subequal,
except that the last two are somewhat enlarged; mandibulars, 13
to 15, decidedly larger anteriorly, decreasing posteriorly; palatines,
10 or 11, subequal; pterygoids, 18 to 22, smaller than palatines and
decreasing posteriorly.
Habitat and Imbits.—Nothing is recorded on this subject for this
form.
Range.—At present nelsoni is known only from western Mexico,
from Acambaro in the state of Guanajuato to southern Sinaloa and
south to Colima, including the Tres Marias Islands. Its southern
limit may be looked for in Michoacan or Guerrero, where its range
REVISION OF THE KING SNAKES. 157
meets that of polyzona. It is probable that it intergrades with
annvlata somewhere on the Mexican plateau, perhaps in the region of
the divide between the Gulf and Pacific drainage. Its range north-
ward can only be surmised, but it should be expected in Sonora
west of the Sierra Madre.
Published records for localities not included in the list of specimens
examined are as follows (Boulenger, 1894, 204): Mazatlan; Presidio,
near Mazatlan; Mezquital del Oro.
Variation and ajjinities.—The material at hand indicates that this
is a fairly homogeneous form, characterized by a low number of
pairs of black annuli separated by broad red interspaces, absence
of black tips on the red scales, and by a light colored snout mottled
with darker. These characters apply equally well to the Tres
Marias Islands and to the adjacent mainland, but the numbers of
ventrals and scale rows is distinctly higher on the islands. Using
Boulenger's figures (1894, 204) for Forrer's specimens from the Tres
Marias, the average for 4 specimens is 231, the extremes 229 to 232;
this contrasts rather strongly with the average of 214 for 19 specimens
from the mainland. The extremes for these, 200 to 221, do not even
reach the numbers for the islands. All of the Tres Marias specimens
attain 23 rows of scales, while from the mainland this number is
possessed by only three, most of the others having the formula
21-19-17. Since the pattern and other structural features seem to
be the same in the island as in the mainland forms, it is not desirable
to make a specific distinction, on the basis of the specimens now on
hand.
While the color pattern seems to be constant on the west coast,
it is very noticeable that toward the interior of Mexico some speci-
mens have the spaces on the belly opposite the dorsal red areas
partially or completely filled with black, presenting in this a striking
approach to annulata. The snout, too, may be blacker and the red
interspaces between the pairs of black rings may be much narrower
and strongly encroached upon by the latter.
Since, in other respects, this form is like annulata, it is believed
that these similarities in pattern toward the interior of Mexico are
evidence of close affinity between those forms, and, in fact, are
sufficient evidence of intergradation.
If the writer's conclusions as to the stem-character of annulata,
stated further on, with respect to the forms north of it be accepted
then the most natural inference with respect to nelsoni is that it, too,
is a derivativ^e of annulata.
158 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
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REVISION OF THE KING SNAKES. 159
tAMPROPELTIS TRIANGULUM ANNULATA (Kennicott).
Fig. 66.
1860. LampropeUis anmdata Cope, Proc. Acad. Nat. Sci. Philadelphia, p. 257
(nomen nudum; advance reference to Kennicott's forthcominc: article).
1860. LampropeUis annulata Kennicott, Proc. Acad. Nat. Sci. Philadelphia, p.
329 (type locality, Matamoros, Mexico; tj-pe, Acad. Nat. Sci. Philadel-
phia, 3613;* Lieut. Couch, collector).—Stejneger, Proc. U. S. Nat. Mua.,
vol. 14, 1891, p. 503; Proc. Biol. Soc. Wasliington, vol. 31, 1918, p. 99.—
Coronella annulata (part) Gunther, Biol. Cent.-Amer., 1893, p. 109, pi.
38, figs. A, B, C.
1861. Coronella doliata, var. conjunci.a Jan, Icon. Gen. Ophid., livr. 14, Dec,
pi. 4, fig. c (type locality, Brazil—probably through error; type, Milan,
Mus.; from the collection of Prince Max de Neuweid); Arch. Zool. Anat.,
vol. 2, fasc. 2, 1863, pp. 237, 242; Blenco sist. degli Ofidi, 1863, p. 46.—
BocouRT, Miss. Sci. Mex., pt. 3, vol. 2, 1886, p. 611, pi. 39, figs. 6, 6a
to M.—Ophiholus doliatus eonjunctm Cope, Proc. TJ. S. Nat. Mue., vol.
14, 1891, p. 608; Amer. Nat., 1893, Dec, p. 1067.
1875. Ophibolus dolialus annulatus Cope, Bull. U. S. Nat. Mus., no. 1, 1875, p.
36.—Yarrow, Gcog. Geol. Explor. Surv. w. 100th mer., vol. 5, chap. 4,
1875, p. 537.—(part) Yarrow, Bull. U. S. Nat. Mus., no. 24, 1882, p. 90
(Matamoros).—Cope, Proc. U. S. Nat. Mus., vol. 11, 1888, p. 382; vol.
14. 1891, pp. 608, 609; Amer. Nat., vol. 27, no. 324, 1893, p. 1067, fig. 9;
Rep. U. S. Nat. Mus. for 1898, 1900, pi. 36, fig. 10.—Brown, Proc. Acad.
Nat. Sci., Philadelphia, 1901; Jan., p. 7b.—Osceola doliata annulata Cope,
Rep. U. S. Nat. Mus. for 1898, 1900, p. 895, fig. 219.
1882. Ophibolus dolintus gentilis (part) Yarrow, Bull. XJ. S. Nat. Mus., no 24, p.
90 (Brownsville, Texas; Caderita, Nuevo Leon, Mexico; Mexico).—
Strecker, Baylor Bull., vol. 18, no. 4, 1915, p. SS.—CoromUa doliata,
var. gentilis (part) Bocourt, Miss. Sci. Mex., pt. 3, vol. 2, 1886, p. 611.
1883. Ophibolus triangulalus annulatus Garman, Mem. I^Ius. Comp. Zool., vol.
8, no. 3, pt. 1, 1883, p. 156.
1888. Ophibolus doliatus cccipilalis Cope, Proc. U. S. Nat. Mus., vol. 11, p. 382
(neither type nor locality designated); same, vol. 14, 1891, p. 609; Rep.
U. S. Nat. Mus. for 1898, 1900. p. 883.
1894. Coronella micropholis Eoulenger, Cat. Snakes Brit. Mus., vol. 2, p. 203,
var. D.
Kennicott's description of annulata, which was road before the
Academy of Natural Sciences of Philadelphia in August, 1860, un-
doubtedly ^ras published before the api)earance of Jar^'s cor.jui-cta
in December, 1861. The type locality is thus fixed satisfactorily,
as it would not have been had the name conjuncta appeared first.
The latter is recorded from Brazil and Caracas. It seems impossi-
ble that these localities can be correct, as the figure is exactly like
» The location of the type specimen has only recently been learned. Kennicott gave it as number 4293
of the United States National Museum collection. This number at present belongs to a specimen from
the same locality and collector as the type, but which was transferred at some unknown time to this number
from the number 1857. Comparison with the original description proves that this is not the type. The
latter has, however, been found in the collection of the Academy of Natural Sciences of Phila<lelphia,
number 3613. It answers to Kennicott's description and bears the original parchment label ol the Smith-
sonian Institution with the number 4293 and the measurements and scalation in the same handwriting as
the original entry in the record book ol this Jfuseum. When it went to Philadelphia is not known.
160 BTJLLETIIsr 114, UNITED STATES NATIONAL MUSEUM.
the form from northeastern Mexico (that the black on the belly is
not solid between the yellow rings is because it is the anterior end
of the body that is shown; this is the usual condition with typical
specimens). If we are correct in judging Jan's conjuncta to be the
equivalent of Kennicott's annulata, then the occipitalis of Cope is
also a synonym of annulata, for he expressly states (1891, 609) that
his name is a synonym of conjuncta Jan, having, in 1888, when he
proposed the name occipitalis, apparently overlooked the name
conjuncta.
Description.—The scutellation based upon twelve specimens is
as follows: Ventral plates, 197 to 212; caudals, 40 to 57; supralabials,
7, rarely 8; infralabials, 9, sometimes 10; a single preocular, 2 postocu-
lars; temporals generally 2 + 3 + 4; posterior chin shields usually in
contact with each other, sometimes separated by a small scale,
shorter than, or about as long as, the anterior; loreal longer than
high; scale rows usually 21-19 or 21-23-21-19.
The bodily proportions are much the same as for the rest of the
group. The head, however, is less distinct from the neck, and the
tail, in the vicinity of the type locality, is rather long. For this
region the proportion of the tail to the total length ranges from 0.138
to 0.167 for eight females, averaging 0.151; two males have the pro-
portions 0,150 and 0.156; and for a female from Puebla, Mexico, it
is 0.123. The largest specimens examined were from Montemoreles
and Peubla, Mexico; each measured 836 mm.
The pattern (fig. 66) is composed of from 19 to 26 white or yellow
rings, from head to tip of tail, bordered by black and separated by
broader areas of red. The white rings are about 1^ to 2 scales wide,
uniform m diameter or a little widened on the first row of scales,
completed upon the belly or partially interrupted there with black,
and usually mottled on the sides with darker. The black rings are
generally widest on the middorsal line, sometimes even confluent
here across the red, and narrowest on the first row of scales. They
are contmuous across the belly. The red areas, except the first ones,
are not completed across the belly, except partially so in individuals
from the northern limits of the range, but are here replaced by black.
The belly thus normally presents a series of large quadrate black areas
separated by much narrower bands of white or yellow.
The head is normally black from the tip of the snout back to the
posterior portion of the parietals and the fifth to seventh upper
labials. In the northern portion of the range, lightening of the snout
begins in the loreal and nasal region and the lower side of the rostral
plate. The chin is more or less mottled with black and white. The
first black ring is usually complete on the neck.
Dental characteristics, as exhibited by five specimens, are as fol-
lows: Maxillary teeth, 13 to 15, subequal except the last two which
BEVISION OF THE KING SNAKES. 161
are enlarged; mandibulars, 14, 13, or 15, the anterior enlarged,
diminishing posteriorly, a little greater space between the fourth
and the fifth; palatines, 11, 10, or 9, subequal; pterygoids 15 to 20,
smaller than the palatmes, and dimuiishing posteriorly.
The penial characters could not be satisfactorily determined,
since all the typical specimens were females.
That this form has suffered so at the hands of herpetologists is
doubtless due to the fact that it inhabits a region where but little
collecting has been done, and that it is consequently rare m museums.
Its name has usually been included in tlie synonymy of its northern
Fig. 48.—Map showinq locauty records for Lampropeltis Triangulum annulata.
or southern relatives. It appears, however, to be a well marked form.
From gentilis, its nearest relative on the north, it differs (1) in hav-
ing the snout nearly or entirely black mstead of lightened on the
end; (2) in having the white rings of about the same width on sides
and belly as on the middorsal line, instead of widened there; (3) in
having the belly marked with broad areas of black, separated by
narrower bands of white, instead of nearly equal areas of black and
white; and (4) in having a distinctly higher average number of ventral
plates. From polyzona it may be distinguished, (1) by the entire
absence of black tips on the red scales, (2) by the absence of the white
band across the snout, and (3) by the smaller number of ventrals.
162 BULLETIN U4, UITITED STATES FATIOSTAL MUSEUM.
From nelsoni it is ciiiefly distinguished by the entirely black snout,
and the completely black areas on the belty, betv/een the white
rings, and the narrower red areas.
Habitat and Jiahits.—Of these nothing is recorded.
Range.—This form is best known from extreme southern Texas
and northeastern Mexico. One from Peubla, Mexico, is referable to
annulata, and one in the British Museum, labeled "Tehuantepec,
"
belongs probablj^ to this form (Boulenger, 1894, 205), It should
be looked for in the plateau region of southern and eastern Mexico;
westward it apparently intergrades Vvath nelsoni. Since the gentilis
of Arizona and New Mexico is most closely related to this form, the
latter may be expected to range pretty well north in Mexico, east
of the Sierra Madre. The onl}^ reliable record, aside from those
included in the list of specimens examined, is that for Tehuantepec
(Boulenger, 1894, 205).
*
Variation and affinities.—This form has never been collected in
numbers, and the great majority of specimens obtained have come
from northeastern Mexico and extreme southern Texas. Future
collecting will undoubtedly extend its knov»Ti range and increase the
limits of its variation. Specimens from Cameron Comity, Texas,
and the adjacent country in Mexico are typical and homogeneous;
they need not be confused with the gentilis from farther north nor
with the Mexican forms of the triangulum group. The ventrals here
range from 197 to 210—a number that is low for polyzona, a little
lower than nelsoni, and a little high for gentilis and amaura. A
specimen from San Antonio, which has been referred to ge^aiilis, has
193 ventrals, and one from Puebla has 213. Boulenger's specimen
(1894, 205) from Tehuantepec, which is probably close to annulata,
has 218 ventrals. This indicates that on the plateau region of
southern Mexico annulata may be characterized b}^ a higher average
number of ventrals than it possesses in the northern part of its range.
The same may be true of the dorsal scale rows. The Fuebla specimen
has the formula 23-21-19, while of those from the type region, only
3 out of 10 have the formula 23-21-19, the rest having it 21-19. This
is significant of the reduced scutellation that will be noticed to charac-
terize the forms of the triangulum group northeast and east from
here.
The labials are, as usual, 7 and 9, but 10 in the lower row is attained
by occasional specimens from the type region. The temporals are
usually 2+3 + 4, v/ith mfrequent drops of one in the second and
third rows. The posterior chin shields are a little shorter or about
equal to the anterior, and the loreal is oblong. The tail is about the
same length as that of folyzona, a little longer than that of the major-
REVISION or THE KING SNAKES. 163
ity of forms in the genus. The important thing to note in all these
matters of structm-e is that annulata is typical of the co^iservative
forms thi'oughout the genus. It is not specialized by having a very
short tail (jaicroyliolis) or ver}'' long one (pyrrJiomelaena) , in havijig
the temporals reduced in number (elapsGides, microplwUs) or irregular
(tnangulum), in havmg a high number of labials (j^yrrhomelaera,
conjuncta) or a low number {elapsoides , rhomhomaculata) , in havmg
unusually short postgenials (h&i'lii, califorrnae), in having a low scale
formula (elapsoides, micropholis, holbrooJci), or in having a low number
of ventrals (elapsoides, gentilis in Utah). It is near to the normal or
average for the genus.
The pattern, too, is one from which the patterns of all the northern
members of the group may be derived. This will be made m.ore
plain later on, but it will be well here to describe the pattern of the
Puebla specimen (tig. 66), since this is theoretically the most primi-
tive in the group (no. 9555, Mus. Comp. Zool., Puebla, Mexico). It
has 19 white (m alcohol) rings on the body and 4 on the tail. These
are 1| to 2 scales in -v\ idth, are a little mottled with darker, chiefly on
the sides, and encircle the body. The space between the white rmgs
may be described as black, more or less split with red. The red never
extends onto the belly beyond the tips of the ventrals, and dorsally
it narrows toward the median line and is frequently excluded there-
from by a Vv'idening of the black borders. The tail is ringed with
black and white. The head is black as far back as the middles of the
parietals, anterior temporals, sixth supralabials and fifth infralabials,
except for small irregular spots of lighter on the prefrontals and
anterior part of frontal, and in the loreal region. This last is sug-
gestive of the light cross band on the snout of polyzoiM, but it is too
much to say now what its significance may be. It is expected to be
shown that all the forms of the group north and west of ammlata may
be easily derived from such a pattern as this.
About all that can be definitely said of the relationshi]) of this form
to those mhalnthig adjacent ranges is that it is a direct relati\'e of
nelsoni, and of gentilis. It is a fairly safe conclusion that mter-
gradation takes place with both of these form.s. Its relationship to
polyzona., judging from the material now at hand, is not at all clear,
but it must be close.
164 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
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REVISION OF THE KING SNAKES. 165
LAMPROPELTIS TRIANGULUM GENTILIS (Baird and Girard).
Fig. 72.
1853. Ophibolus gentilis Baird and Girard, Cat. N. Amer. Kept., pt. 1, p. 90
(type locality: North Fork, Red Tdver, near Sweetwater Creek, Wheeler
County, Texas; type, U. S. Nat. Mus. no. 1853; Capt. Marcy, collector).—
Marcy, Ex. Doc. Ho. of Rep., 33d Congress, Ist sess., 1854, p. 200, pi.
8.—Baird, Pacif. R. R. Surv., vol. 10, pt. 3, no. 1, 1859, pi. 30, fig. 64.—
Coronella gentilis (part ?) Boulenger, Cat. Snakes Brit. Mus., vol. 2,
1894, p. 201.
1860. Lampropeltis doliata (part) Cope, Proc. Acad. Nat. Sci. Philadelphia, p.
256 (Kansas).
1860. Lavipropeltis muUistriata Kennicott, Proc. Acad. Nat. Sci. Philadelphia,
p. 328 (type locality, Fort T>ookout, South Dakota; type specimen, U. S.
Nat. Mus., no. 1842; Lieut. Warren and Dr. Hayden, collectors).
1875. Ophibolus muUistratus Cope, Bull. U. S. Nat. Mus., no. 1, p. 36; Proc. U. S.
Nat. Mus., vol. 14, 1891, p. 611.
—
Ophibolus multistrata Coues and Yar-
row, Bull. U. S. Geol. Geog. Surv. Terr., vol. 4, art. 11, 1878, p. 284.—
Cope, Rep. U. S. Nat. Mus. for 1898, 1900, p. 909, fig. 225.—Lavipropeltis
multistrata Stejneger, Proc. U. S. Nat. Mus., vol. 14, 1891, p. 502.
1882. Ophibolus doliatus gentilis (part) Yarrow, Bull. U. S. Nat. Mus., no. 24, p.
90,—Brown, Proc. Acad. Nat. Sci. Philadelphia, 1901, p. 75.—Bran-
son, Kansas Univ. Sci. Bull., vol. 2, no. 13, 1904, p. 402, figs. 25, 25a.—
DiTMARS, ReptUe Book, 1907, pp. 340, 348, pi. 106 (upper fig.).—Cock-
erell, Univ. Colorado Studies, vol. 7, 1910, p. 131 (footnote).—Ellis
and Henderson, Univ. Colorado Studies, vol. 10, no. 2, 1913, p. 91
pi. 4, fig. 2S.—Osceola doliata gentilis Cope, Rep. U. S. Nat. Mus. for 1898
1900, p. 894.
1882. Ophibolus doliatus annulatus (part) Yarrow, Bull. U. S. Nus. Mus., no. 24
p. 90 (Apache, Arizona).
1883. Ophibolus triangulus gentilis Garman, Mem. Mus. Comp. Zool., vol. 8
no. 3, pt. 1, pp. 66, 155.
1893. Ophibolus doliatus sysputus Cope, Proc. Acad. Nat. Sci. Philadelphia, p
387 (Hennessy, Oklahoma).
1903. Lampropeltis pyrrhomelaena celaenops Stejneger, Proc. U. S. Nat. Mus.
vol. 25, p. 153 (type locality, Mesilla Valley, New Mexico; type specimen
U. S. Nat. Mus., no. 22375; H. B. Lane, collector).—Stejneger and
Barbour, Check List, 1917, p. 89.
1917. Lampropeltis triangulum gentilis Stejneger and Barbour, Check List
p. 90.—Stejneger, Proc. Biol. Soc. Washington, vol. 31, 1918, p. 90.
Description.—Forty-six specimens of this form have been available
for study, and from these the following summary of the scutellation
has been derived: Ventral plates, 176 to 212; caudals, 31 to 53
(males, 41 to 53, average 48; females, 31 to 49, average 45); supra-
labials, 7, rarely 8; infralabials, 9, rarely 8 or 10; one preocular;
two postoculars, rarely one; temporals usually 2 + 3+4, occasionally
one less in any row; posterior chin shields shorter than the anterior or
nearly as long, in contact with each other or separated by one or two
small scales; loreal distinctly longer than high; scale rows usually
21-19, often 21-19-17, on middle of body sometimes 23.
166 BULLETIIT 114, UNITED STATES NATIONAL, MUSEUM.
Head somewliat distinct from the neck; body moderately stout,
cylindrical; tail tapering, 0.115 to 0.156 of the total length (males,
0.124 to 0.156, average about 0.141; females, 0.115 to 0.148, average
about 0.134). The largest specimen from a typical portion of the
range (Wheeler County, Texas) is 713 mm. in length. Some exam-
ples from central Kansas are larger than this.
The pattern (fig. 72) on body and tail is composed of 25 to 40
whitish annuli from one to three scales wide dorsaliy and from two
to five in width on the first row of scales. Between these rings are
pairs of black annuli separated by red. The black and white rings
are continuous across the belly, but the space there between the
black rings and opposite the dorsal red areas is usuallj^ filled v/ith
black. The latter color tends to encroach upon the red, and, on
the middorsal lines the black rings of adjacent pairs are often con-
fluent. The whitish rings are usuallj'' mottled on the sides with
darker; the belly is generally crossed by approximately equal bands
of black and white, but the white areas may be blotched with darker,
the black may be partly broken up, and rarely there is little or no
black below.
The head is generally a uniform black on the posterior half, and
this black may extend forward over the snout. The latter is, how-
ever, usually lightened, at least on the sides. The labials show
varying proportions of dark and light, corresponding with the extent
of black on the head.
The actual colors of the whitish annuli, according to Ellis (1913, 91),
are light gray to bright j^'ellov/, while the red varies from slate brown,
through brick red, to scarlet. A specimen from Blue River, Nebraska,
bears the note: "Original color: Deep orange, lemon yellow, and
black." The red and yellow fade to white in alcohol.
The dentition, as revealed by the examination of a few specimens,
is as follows: Maxillary teeth, 12 to 14, most often 13, the last 2
slightly larger; mandibulars, 12 to 15, the third and fourth largest, a
little greater space between the fourth and fifth; palatines, 10 or 11,
oftener 10; pterygoids, 16 to 20.
This form may nearly always be immediately recognized by its
black head with snout mottled with red and black, its conspicuous'
black rings which encroach on the red dorsaliy and are separated by
not more than 40 nor less than 25 whitish annuli on body and tail.
Specimens examined from west of the Rockies, where the color pat-
tern is confusingly similar to that of vyrrJiomelaena and 'niuliicincta,
have less than 200 ventrals, while the latter forms have several to
many more. Its distinction from araaura and from syajnla is given
under these respective forms.
Habitat and liabits.—^Apparently nothing has been recorded on the-
natural history of this form.
REVISION OF THE KING SNAKES. 167
Range.—As here defined, gentilis occurs from western Utah and
southeastern Arizona east to the ninety-seventli meridian, north to
southern South Dakota, and south prohahly to northern Mexico.
Besides the localities included in the accompanying list of specimens
Ellis (1913, 92) has recorded a few other stations for Colorado; Yuma;
Orchard; Clear Creek, near Golden; Beulah; and Baca County.
Fig. 49.—Mj\j saowiNG locauty uscords for Lampkopeltis tkiangulum gsntilis.
Variation.—Considerable variation with locality is indicated by
the ventral plates. The extremes and average for Kansas are about
the same as for sys'pila in that region ; the number is lower in Colorado,
and the few from west of the Rockies indicate a still greater reduction.
It will be interesting to see whether future specimens bear out the
unusual lack of variation in the four Utah specimens now on hand.
Subsequent proof of the existence of a distinct race west of the Rockies
would not be surprising, but the present material certainly would not
justify any separation.
168 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
Table of scaleformulo.e of gentilis.
Formula.
23-21
21-23-21
23-21-19
21-23-21-19
21-19
19-21-19
21-19-17
19-21-19-17
19-17
Total
^lale. Female.
1
1
1 ..
1 3
9 9
1 1
5 . .
1 1
1
20 15
Geographic differences on scale rows can not be detected from the
scanty material now available. A notable point is the great varia-
bility in formulae. Like the southern forms of the group, 17 rows
toward the posterior end of the body is much less common than 19,
and this number is possessed almost exclusively by males. The com-
monest formula is 21-19.
Variation in other points of scalation may be considered as only
normal for the group. Reduction in the temporals below 2 + 3 + 4
occurs frequently in the third row and less often in the first two.
There is no evidence that the lower labials are becoming reduced
to 8. In fact, it may be said that reduction in scutellation from the
mean of the group has not proceeded in gentilis quite as far as in
syspila and triangulum.
In spite of the scarcity of specimens certain significant differences
that depend upon locality may be noted in the color pattern, chief of
which is perhaps the increase in dark pigment westward. Eastern
specimens as a rule have more red on the snout, while examples from
west of the Rocky Mountains may have the whole head black with
only a very little lightening in the loreal region and at the end of the
snout. Likewise on the body, the black pigment increases at the
expense of the red, and frequently excludes the latter altogether from
the dorsal line, thus presenting a striking resemblance to multicincta of
the Pacific region. These differences in pigment are, however, only
average ; occasional specimens from east of the Rockies are as heavily
pigmented as those west. The latter are definitely ringed, while those
on the east side of the mountains show a variation toward syspila.
Here the black on the belly opposite the dorsal red areas is sometimes
split lengthwise in the middle or is continuous across the belly but
much narrower. Thus specimens from Nebraska and Kansas may
have the red restricted to wide dorsal saddles, but there are usually
no lateral alternating spots.
REVISION OF THE KING SNAKES. 169
Affinities.—Very few specimens are available from the southern
portion of the range, but the indication is that no constant differences
will be found to separate gentilis from annulata. Their close affinity
is indicated in every characteristic. Annulata becomes gentilis when
the yellowish annuli widen on the sides and restrict the black on the
belly to an area no greater than that of the yellow, and when the black
snout becomes lightened with reddish. Intergradation certainly takes
place with amnaura and with sijspila where their respective ranges
meet its own. The relationship held to exist between these forms
and discussed at greater length in the summary of this group may be
expressed in the following diagram
:
gentilis syspila
(Utah, Col.) /
gentilis > amaura
(Texas)
T
annulata
FiQ. 50.—Diagram showing mTERBELATioNsmps between Lampropeltis trungulum annulata
L. T. gentilis, L. T, AM\tJRA, AND L. T, SYSPILA.
170 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
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REVISION OF THE KING SNAKES. 171
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186550—21
172 BULLETIN 114, UNITED STATES NATIOlfrAL MUSEUM.
LAMPROPEXTIS TRUNGULUM AMAURA (Cope).
Fig. 67.
1853. Ophibolus doliatus Baird and Girard, Cat. N. Amer. Rept., pt. 1, Serpents,
r p. 89.—Baird, Pacif. R. R. Surv., vol. 10, pt. 3, no. 1, pi. 30, fig. 63.—
Coronella doliata (part) Dumeril and Bibron, Erp. Gen., vol. 7, pt. 1,
1854, p. 621.—Gunther, Cat. Colubr. Snakes Brit. Mu3., 1858, p. 41,
var. B.
—
Lampropeltis doliatus Garman, S., Bull. Essex Inst., vol. 24,
1892, p. 9.
1860. Lampropeltis amaura Cope, Proc. Acad. Nat. Sci. Philadelphia, p. 258
(type locality, "unknown"—Original entry of type U. S. Nat. Mus., no.
5282 reads " ? Mississippi").
1863. Coroneda gentilis Jan, Arch. Zool. Anat., vol. 2, fasc.|2,?pp. 237, 241.
1866. Coronella doliata gentilis Jan, Icon. Gen., livr. 17, pi. 1, fig. 2, (Fort Towson,
Oklahoma).
—
^Bocourt, Miss. Sci. Mex., 1886, p. 610 (part).
—
Osceola
doliata gentilis Cope, Rep. U. S. Nat. Mua. for 1898, 1900, fig. 218 (Cal-
casieu Pass, Louisiana).
1875. Ophibolus doliatus amauru^ Cope, Bull. U. S. Nat. Mus., no. 1, p. 36.
1882. Ophibolus doliatus doliatus Yarrow, Bull. U. S. Nat. Mus., no. 24, p. 89.
—
Strecker, Baylor Univ. Bull., vol. 18, no. 4, 1915, p. 38.
—
Osceola doliata
doliata Strecker, Trans. Texas Acad. Sci. for 1901, vol. 4, pt. 2, no. 5,
1902, p. 3. Lampropeltis doliatus doliatus Strecker, Proc. Biol. See.
Washington, vol. 21, 1908, p. 75.
1888. Ophibolus doliatus cocdneus Cope, Proc. U. S, Nat. Mus., vol. 11, p. 382;
vol. 14, 1891, pp. 608, 609; Amer. Nat., vol. 27, 1893, p. 1067, pi. 28, fig.
10; Rep. U. S. Nat. Mus. for 1898, 1900, pi. 36, fig. 10.—Stone, Proc.
Acad. Nat. Sci. Philadelphia, 1903, p. 542 (Limestone Gap, Oklahoma).
Strecker, Baylor Univ. Bull., vol. 18, no. 4, 1915, p. 38.—^VioscA, 6th
Bien. Rep. Louisiana State Mus., 1918, p. 72.
—
Osceola doliata cocdnea
Cope, Rep. U. S. Nat. Mus. for 1898, 1900, p. 896 (part).
1888. Ophibolus doliatus parallelus Cope, Proc. U. S. Nat. Mus., vol. 11, pp. 383,
385 (type locality, unknown; type specimen, U. S. Nat. Mus., no. 10544);
vol. 14, 1891, pp. 608, 609; Amer. Nat. vol. 27, 1893, p. 1067 pi. 27, fig. 8;
Rep. U. S. Nat. Mus. for 1898, 1900, pi. 35, fig. 8. Osceola doliata parallela
Cope, same, p. 893, fig. 217.
—
^Wright, Proc. Acad. Nat. Sci. Philadel-
phia, 1915, p. 140.
1917. Lampropeltis triangulum amaura Stejneger and Barber, Check List, p. 90.
As here defined, amaura occupies a very limited range in the lower
Mississippi Valley. Nearly every reference in the literature includes
with it iudividuals here referred to elapsoides, syspila, gentilis,
nelsoni, and perhaps others. The indications are that it is a fairly
well-defined form, intergrading, however, on the north with syspila,
on the west with gentilis, and probably with annulata on the south-
west. The type is typical, and undoubtedly came from a central
portion of this range, very likely from Mississippi, as the original
entry suggests.
Parallelus Cope is included in the synonomy with some hesitation.
The pattern of the head is like that of syspila, but the body pattern,
which is imlike any example of syspila yet examined, occurs in some
specimens of amaura and of virginiana. The scutellation excludes
REVISION OF THE KING SNAKES. 173
the latter, so that there is the least difficulty in referring it to amaura.
Where it came from is unknown. Originally only one specimen bore
the United States National Museum number 10544. This was en-
tered as from Gainesville, Florida, James Bell, collector. Later
another was found with the same number—10544. The original
name in the record book is erased and " ? Ophiholus doliatus" written
over it. This was probably meant for the specimen which later
became the tyi^e of 'parallelus. This " ? Ophiholus doliatus'' has a
line through it and above is written " ? Osceola elapsoidea;" this was
doubtless the specimen that originally bore the number 10544. This,
too, has a line through it, and above it is written "OpTiiholus doliatus
parallelus." Then when it was noticed that two different specimens
bore the number 10544, one was given a new number, 20137, and the
type of parallelus was left with the old number. Specimen number
20137 is a perfect example of elapsoides, and is without doubt the one
received from James Bell. This leaves parallelus with no locality
and no way of finding it out.
Description.—^This form has the following scutellation : Ventral
plates, 180 to 205; caudals, 39 to 51 (males, 41 to 51, average about
45; females, 39 to 47, average about 44); supralabials, 7, infralabials,
9, occasionally 8 or 10; 1 preocular, 2 postoculars; temporals, 2 + 2 + 3
or 2+3+4, sometimes 1+2 + 3; posterior chin shields shorter than
anterior, and frequently separated by one or two small scales; loreal
distinctly longer than deep; scale rows usually 21-19, occasionaly as
low as 19-21-19-17, and as high as 21-23-21-19.
This is a small snake, but not as small as elapsoides. It is rather
slender; the snout is a little more pointed and projecting than that of
gentilis; the tail varies from 0.118 to 0.156 of the total length (males,
0.133 (0.118) to 0.156, average 0.143; females, 0.125 to 0.143, average
0.136). The largest specimen examined measured 629 mm., and
came from Jefferson County, Texas.
The pattern is typically in rings of black, yellow, and red. There
are 18 to 26 yellow rings on body and tail bordered with black and
separated by red. The yellow rings widen on the lower rows of
scales, and traverse the belly, but are sometimes more or less inter-
rupted here with black blotches. The red areas may be continuous
across the belly or interrupted by the ventral junctions of the black
borders of adjacent pairs of rings (fig. 67). The head is usually
black and the snout red, but the red, in a few cases extends back
onto the parietals.
This form may be distinguished from elapsoides by the greater
number of ventral plates (nearly 200), a maximum of 21 instead of
19 rows of scales, the usually greater extent of black on the head,
the fact that the snout is usually more or less mottled with black
instead of being a uniform red; from syspila it may be distinguished
174 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
b3^ the fact that ( 1) the pattern is in rings, or, when in blotches of
red, there is no ventro-lateral series of dark spots alternating with
the dorsal blotches, (2) the head is black with a red snout, instead of
red with a posterior black band and various dark-edged light markings
between the eyes, and ( 3) the number of transverse yellow bands is
not commonly above 23; from gentilis it is best known (1) b}/ the
smaller number of yellow cross bands, usually not more than 25,
(2) by the fact that the black rings show but slight tendency to
encroach dorsally upon the red, and (3) the black of the belly in
Fig. 51.—Map showing locality records foe Lampkopeltis trlvngxtlum amauea.
gentilis is usually concentrated between the black rings opposite the
dorsal red areas, instead of being divided v/ith red, as in amaura.
HoMtat and Jiahits.—Of the natural history of this form nothing
is recorded.
Range.—-As here defined, amaura occupies the southern portion of
the Mississippi Valley, from eastern Mississippi to southern Arkansas,
and west to about the 97th meridian.
Tnere are no apparentl}^ reliable localit}^ records aside from those
included in the accompanying list of specimens.
Variation and affinities.—^With this form we must confine ourselves
to the discussion of individual variation, since the demonstration of
geographic variation is rendered unsatisfactory by the insufficiency
REVISION OF THE KING SNAKES. 175
and poor distribution of the material, and, exceptfor pattern charac-
ters, b.y the small extent of the range. In the matter of scale rows
it should be noted that the common formula is 21-19. In this, and
the small percentage of specimens with 17 rows at the end, amaura is
like gentilis, and probably like annulata (but the undue proportion
of females among the specimens of this form lessens the value of
figures based upon characters tliat vary v/ith sex). The infrequent
occurrence of a lower formula, as 19-21-19-17, foreshadows a situa-
tion that is common in syspila and still more common in triangulum.
The distribution of scale formulae between the sexes in amaura is as
follows
:
Formula. Male. Female.
21-23-21-19 3
21-19 fi 7
19-21-19 3 1
19-21-19-17 3 1
19-17 1
Thus the formula 21-19 is about as common to one sex as to the
other, and may be considered the normal. In accordance with the
general rule, the higher formulae are mostly possessed by females,
and the lower by males.
The number of veutrals in the majorit}' of cases lies between 188
and 199, therefore averaging a little less than annulata in north-
eastern Mexico. With the exception of local variation, this number
of ventrals is liardly different from that of syspila and triangvlum.
Variation in infralabials and temporals is the same in kind and
apparently about the same in frequenc}" as in these same two
forms.
The pattern is rather variable, and to properly interpret this
variation many more specimens must be secured. We may say that
amaura is characterized by a black head, reddish snout somewhat
mottled with darker, and about 22 pairs of black annuli bordering
yellow, and separated by red. The latter color may encircle the
body or be restricted to wide saddles bordered below with black.
Wlien the latter is the case we may have the ventral borders of the
saddles near together on the belly, or, at the northern edge of the
range, they may be indistinct and only just reach the ventrals.
This is a close approach to syspih. When, added to this, there are
developed alternating spots near the ends of the ventrals, and the
whitish bands have increased to more than 25, and the black of
the head has receded or become broken up, then we have syspila.
The change to gentilis is not as well shown for lack of specimens, but
westward, where the paired black rings increase to more than 25,
176 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
and the black spreads more or less over the red, in particular above
the dorsal line, we have gentilis.
On the basis of this single form and its variations, it would not be
safe to assert that evolution had proceeded in one direction and not
in another, for proof that it did not occur in any one of several ways
would be difficult to produce. We will therefore leave to the sum-
mary of the group our hypothesis of its actual relations to its nearest
relatives, and the reasons therefore, and, with the briefest summary,
leave the subject at this point.
It can hardly be doubted that the closest affinities of amaura are
with syspila on the north and with gentilis on the west, and with
these forms intergradation certainly occurs. With annulata its
relationship is very close, but whether direct or through gentilis is
uncertain. It is the closest relative of elapsoides, but between the
two there is apparently a slight but distinct structural gap, strongly
suggesting differentiation of the latter by isolation, followed by a
rejoining of the two ranges.
REVISION OF THE KING SNAKES. 177
1 IS
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178 BULLETinsr 114, LTNITED STATES NATIONAL MUSEIJ]\T.
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REVISION OF THE KING SNAKES. 179
LAMPROPELTIS TRIANGtLlJM SVSPILA (Cope).
Red Snake; Red MaK Snake
Figs. 52, G8.
1861. Caronella doliata Jan, Icon. Gen. Ophid., livr. 14, pi. 4, fig. A (Illinoifl
meridional); Arch. Zool. Anat., vol. 2, fasc. 2, 18G3, p. 237.—Bocourt,
Miss. Sci. Mex., pt. 3, vol. 2, 1886, p. 009, vol. 3, pi. 39, fig. 2.—Ophibolus
dolialus Smith, Geol. Surv. Ohio, vol. 4, 1882, p. 691.—Blatchley,
Journ., Cincinnati Soc. Nat. Hist., 1891, p. 32.—Hurter, Trans. Acad.
Sci. St. Louis, vol. 6, no. 2, 1893, p. 255.
—
Lampropeltis doliata McLain,
Notes Coll. Rept., 1899, p. 3.
—
Lampropeltis dolialus Hurter and
Strecker, Trans. Acad. Sci. St. Louis, vol. 18, no. 2, 1909, p. 26.—
Hurter, Trans. Acad. Sci. St. Louis, vol. 20, no. 5, 1911, p. 183.—
RuTHVEN, Proc. Iowa Acad. Sci., vol. 19, 1912, p. 207.
1882. OpUholm dolialus dolialus (part) Yarrow, Bull. U. S. Nat. Mus., no. 24,
p. 89.—Hay, 36th Annual Rep. State Board Agric. Indiana for 188G, vol.
28, 1887, p. 210; Journ. Cincinnati Soc. Nat. Hist., vol. 10, no. 2, 1887,
p. 64 (New Harmony, Ind.); 17th Annual Rep. Indiana Dept. Geol. Nat.
Res., 1892, p. 515.—Cope, Proc. Acad. Nat. Sci. Philadelphia, 1893, p.
385; Rep. U. S. Nat. Mus. for 1898, 1900, pis. 34, 35, figs. 5 to 7.—Branson,
Kansas Univ. Sci. Bull., vol. 2, no. 13, 1904, p. 400.—Osceoia doliata
doliata Morse, Proc. Ohio State Acad. Sci., vol. 4, pt. 3, spec. pap. no. 9,
p. 130.
—
Lampropeltis dolialus dolialus Strecker, Proc. Biol. Soc.
Washington, vol. 21, 1908, p. 89 (Hot Springs, Arkansas).
1888. OpUholus dolialus syspilus Cope, Proc. U. S. Nat. Mus., vol. 11, p. 384
(type locality, Richland County, Illinois; type specimen, U. S. Nat.
Mils., no. 13380; Robert Ridgway, collector); vol. 14, 1891, pp. 608, 609.—
Hay, 17th Annual Rep. Dept. Geol. Nat. Res. Indiana, 1892, p. 517.—
Cope, Amer. Nat., vol. 27, 1893, p. 1067, pi. 27, fig. 7.—Osceola doliata
syspila Cope, Rep. U. S. Nat. :Mus. for 1898, 1900, p. 891, fig. 210.
1892. Ophibohis triangulus dolialus Garman, H., Bull. Illinois State Lab. Nat.
Hist. vol. 3, art. 13, p. 295.
1898. Coronella triangulum, var. collaris Boettger, Kat. Rept.—Samml. Frank-
furt, pt. 2, p. 72.
1903. Lampropeltis dolialus coccineus Stone, Proc. Acad. Nat. Sci. Philadelphia,
p. 542 (Petit Jean Mountain, Arkansas; South McAlester, Oklahoma).
1919. Lampropeltis triangulum syspila Ruthven, Occ. Pap., Mus. Zool., Univ.
Mich., no. 60, p. 3.
1919. Lampropeltis triangulum amaura Schmidt, Copeia, no. 73, p. 72 (Sapulpa,
Oklahoma).
This form has usually been Inown as doliatus Linnaeus, but this
name, as Stejneger has pointed out (191S, 99), can not be definitely
identified with any known form of Lampropeltis. The name syspilus
Cope is the next one to be considered. This is a composite, as a
glance at the list of specimens accompanying Cope's original descrip-
tion shows, but the type is definitely indicated, and as this belongs to
the present form its name is available. An error in the locality of
the type occurred in the original entry and was copied by Cope in
his list. The entry in the record book of the National Museum is
Richland, Illinois; this was undoubtedly intended for Richland
180 BULLETIN U4, UNITED STATES NATIONAL MUSEUM.
County, Illinois, as Mr. Eidgway has collected extensively here but
not at Richland, which is in Sagamon County. Richland County
is in much better agreement with the specimen; its pattern is not
typical for syspila, as here defined, but is closely approached by some
specimens from southeastern Illinois and southern Indiana. Saga-
mon County is close to the range of triangulum, and an individual
like the type of sys'pila is not to be expected there.
Description.—The scutellation, as exhibited by nearly a hundred
specimens, is as follows: Ventrals, 180 to 215; caudals, 40 to 54
(males, 43 to 54, average 48; females, 40 to 50, average 47); supra-
labials, 7; infralabials, 9, rarely 8 or 10; 1 preocular, very rarely 2;
2 postoculars, rarely 1; temporals commonly 2+2 + 3, or 2 + 3+4,
occasionally 1+2+3; posterior chin shields usually shorter than the
anterior, sometimes about as long, in contact with each other or
separated by a small scale; loreal distinctly longer than high; maxi-
mum number of scale rows commonly 21, often 23, very rarely 19,
beginning with 21 or 19 and ending with 19 or 17.
In size and proportion this form is nearest like triangulum. The
head is scarcely distinct from the body, is tapering, and blunt; the
body is fairly stout, and cylindrical, except that the belly meets the
sides at something like a right angle; the tail is typically short and
tapers quicldy to a horny tip. Of the total length, the tail varies
from 0.114 to 0.157 (males, 0.130 (0.116) to 0.156, average 0.137;
females, 0.114 to 0.146 (0.157), average 0.131). The largest speci-
men examined was from St. Clair County, Illinois, and measured
1060 mm.
The pattern (fig. 68), although apparently in rings when viewed
from above, has definitely changed to the blotched or saddled type.
The whitish cross bands, varying from 23 to 36 (one specimen, 43) in
number, are IJ to 3 scales wide above and from one to several scales
wider on the lower rows. They are usually strongly mottled on the
sides with darker, and are narrowly bordered with black. The black
borders of adjacent pairs meet usually on the first or second row of
scales to inclose the dorsal saddles of the predominating color—red,
or brownish, or gray. These dorsal saddles vary much in width
but are usually decidedly broader than the groups of white and
black bands that separate them. The belly is generally strongly
checked with black and white. On the sides, usually overlapping
ventral and dorsal scales, is a series of small black blotches, often
inclosing some of the ground color, and alternating with the dorsal
saddles. These alternating spots are nearly always present (rarely
fused into a single midventral series), except in transitional individ-
uals from the western and southern limits of the range, where inter-
gradation occurs with the ringed forms, gentilis and amaura.
The head typically shows a narrow black band across the posterior
extremities of the parietals, followed by a whitish half collar on the
REVISION OF THE KING SNAKES. 181
neck (fig. 52), Anterior to the black band the head is reddish or
brownish to the snout, or the latter may be grayish. The markings
on the top of the head vary much with locality; in southern Missouri
and Arkansas there is much black, lightened by streaks or vague
spots of the ground color; in eastern Kansas the black may be
restricted to the tips of the parietals while the rest of the head is
without markings, or the black of gentilis may be more prominent;
eastward the markings typical of tnangvlum appear in varying com-
binations and perfection. Light, black-bordered superciliary spots
are generally present (fig. 52) and often there is a black line from the
eye to the angle of the mouth.
Syspila is bounded on three sides by closely allied forms, from which
it is not separated by well-defined physical barriers. Consequently
difl5.culties in identification must be expected. From gentilis it is
best distinguished by the restriction of black on the head to the
posterior portions of the parietal plates, and the narrower black cross
bands on the body which do not tend to overspread the red areas
Fig. 52.—LAMPEOPELTia TBiANQTn.UM SYSPILA (U.S.N.M. NO. 61680, Jefferson County, Missouri).
About li x nat. size. Showing typical coloe pattern of anteeioe end of body.
dorsally; from amaura it may best be known by the greater number
of transverse whitish bands, usually 25 or more, the presence of
ventro-lateral alternating spots, and the breaking up or practical
absence of black on the frontal and anterior portion of the parietal
plates; from triangulmn it may be distinguished by the smaller
number of dorsal saddles—35 or less—the presence of only one series
of spots in alternation with the dorsal saddles, the yellow half collar
behind the head, the incompleteness of the head pattern of triangulum,
and the generally red color of the dorsal saddles as contrasted with
the usually brown or gray in triangulum (See also under Variation in
triangulum,, p. 200).
The dentition, as indicated by a few specimens, is as follows:
Maxillaries, 11 to 13, the last 2 a little larger; mandibulars 11 to 14,
the anterior the largest; palatines, 10, 11, or 12; pterygoids, 17 to 20.
Range.—As here defined, syspila is found from the ninety-seventh
meridian north to northwestern Iowa, south to southeastern Olda-
homa, and east to western and southern Illinois, southern Indiana,
and extreme southwestern Ohio. It also may be expected in western
182 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
Kentucky, western Tennessee, and northern iiississippi, and it should
be looked for in southern Minnesota.
Specimens hare been examined from the following localities in
addition to those represented by specimens in the United States
National Museum: New B!armony and Vigo County, Indiana;
Charleston, Grand Chain (Pulaski County), St. Clair County, and
Horseshoe Lake at Olive Branch (Alexander County), Illinois; Ames,
Boone, Fort Des Moines, Grinneli, and vicinity of Sioux City, Iowa;
Galena, Missouri; Petit Jean Mountain and Fort Smith, Arkansas;
Fig. 53.—Map sho-wxng locauty becoeds fob Lampeopeltis Triangulum syspila.
South McAlester and Sapulpa, Oklahoma; Manhattan, Douglas
County, Anderson County, and Labette County, Kansas; and near
Belleview (Davidson County), Tennessee.
Habitat and Jiahits.—^According to Hurter (1911, 184) this snake
makes its home around spring houses so as to be near its food—rats
and mice. He mentions finding one hiding under the loose bark of
a heavy rotten log. " I placed it in my collecting bucket with a lizard,
Eumeces fasciaius. On looking into the bucket a little later I found
only a small end of the lizard sticking out of the snake's mouth, and
the wriggling tail, which had been broken off in the struggle, at the
REVISION OF THE KING SNAKES. 183
bottom of tho bucket." According to the same authority this
snake "is a cannibal, swallowing its own kind and other small ser-
pents and lizards."
Branson (1904, 401-402) says that "these snakes are ver}' gentle.
They no^'er bite unless very much aggravated. When they do bite
they do not strike like other snakes, but take the offending object
in their mouths and shut down on it. Their teeth are so small that
they can do little injury."
Mr. Mackelden of St. Louis ^\Tites that "nearly all the milk snakes
I have caught were found under rocks, boards, or logs, etc., in the
early part of the day. I have found them in the open field and also
in the densest Vv^oods. The stomach contents of those I examined
revealed earthworms, June beetles, spiders, and smaller ii\sects. I
recall having caught one tliat was in the act of sv.^allo"wing a young
blue racer."
This form is so closely related to triangulurn and so similar in size
and structure that its habits must be much the same as those of the
latter.
Variation and ojfinities.—Examination of the table showing geo-
graphic variation in number of ventral plates, brings out plainly the
fact that the lo\yest numbers occur in the southern portion of the
range.
Table of ventral plates in syspila.
I.oralirv.
Arkansas and Oklahoma
Southern Missouri
East central Missouri
Southeastern Illinois and
southern Indiana
Iowa
Eastern Kansas
s'^Sens! J^-^r^™*^- i -^-«'^««^-
o
21
12
6
5
18'-] 93 188
180-198 11)0
190-213 1!)9.
104-205
203-213
202-213
201.1
208. 7
208
Table of scaleformulae in sysjnhi.
Formula.
21-23-21-19
21-19
19-21-19
21-19-
19-21-19
19-1
Total.
171
-17/
Male.
3
7'JiJ
2
34
1
47
Female.
2
13
40
184 BULLETIN 114:, UNITED STATES NATIONAL MUSEUM.
Arkansas and Oklahoma with an average of 188, and southern Mis-
souri with 190 show a condition closely like amaura, the average for
which throughout its range is 193. In east central Missouri, and in
southern Indiana and the adjacent portion of Illinois the averages are
200 and 201 respectively. This is closely like the averages for triangu-
lum in Indiana and Ohio, and in Michigan, which are respectively 201.6
and 202. The figures for syspila in Iowa and eastern Kansas are 208
in each case. These are high like triangulum in northern Illinois and
Wisconsin (202), and gentilis in Kansas (202). The point to be noted
is that on the three sides of its range where syspila meets a closely
allied form it shows in the number of its ventral plates a definite
approach to these respective forms.
Since the dorsal scale rows vary somewhat with sex, and the
specimens available are very limited in number, geographic variation
in this character cannot be conclusively shown. The totals for the
whole range, however, are of interest for comparison with alhed forms.
The table shows that the general expectation of a greater proportion
of high formulae in females holds here. While in amaura about half
of each sex has the formula 21-19, in syspila only about one-seventh
of the males have this number; and, whereas, in the former only a
tenth of the females and half of the males have 17 rows at the end,
in this form about one-third of the females and three-fourths of the
males have this number at the end. This indicates a distinct dif-
ference between amaura and syspila and a decided approach to
triangulum.
Differences in other matters of scalation between this form and
its near relatives are too slight to demonstrate change with locahty.
In its color pattern, as in the number of its ventral plates, syspila
proves its close relationship to its three nearest neighbors. The
line of separation on the south is arbitrary at best. Here it is the
change from a ringed pattern to a saddled pattern that marks the
change from amaura to syspila, while on the north and east it is the
change from a saddled to a blotched pattern that indicates the change
from syspila to triangulum. Whatever may be our conception of
the origin of these forms, we must agree that syspila gives every
evidence of being an intermediate or connecting link between amaura
and triangulum. On the south, near the range of the former, it is of
smaller size, tends to partake of the ringed pattern of amaura, has
more black on the head, and only scant indications of the complicated
head pattern of triangulum. In Missouri the body is larger, the
saddles of red well-defined, long, and broad, with usually a ventro-
lateral series of smaller spots alternating with them, the black is
restricted to a black band across the posterior portion of the parietal
plates, forming an anterior border to the first whitish cross band,
REVISION OF THE KING SNAKES. 185
while on the rest of the head various portions of the elaborate pattern
of triangulum appear, commonly a dark-edged light spot over each
eye, and more or less of a light band from behind or over the eye to
the side of the neck, bordered below by a dark band from the eye to
the angle of the mouth. As the range of triangulum is approached
the head markings characteristic of that form attain greater perfec-
tion, and some specimens show a tendency to a dorso-lateral fusion
of the dark borders of the first whitish cross band, to inclose an oval
or Y-shaped spot of white or gray within the ground color of the head
on its posterior portion. This and the shortening and narrowing of
the dorsal saddles are the slowest of the characteristics of triangulum
to appear, and when they have been attained, we say that we have
triangulum.
Thus, syspila is central both geographically and structurally to
the three forms, triangulum, amaura, and gentilis, and with each of
these forms intergradation takes place in the region of the common
boundary of the respective ranges.
186 BULLETIN 114, UISTITED STATES NATIONAL MUSEUM.
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REVISION OF THE KING SNAKES. 187
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188 BUUIjETIN 114, UNITED STATES NATIONAL MUSEUM.
LAMPROPELTIS TRIANGULUM TRIANGULUM (Lacgpede).
MILK snake; spotted adder; checkered adder; hotjse moccasin; house snake.
Figs. 54, 69.
1789. Coluber triangulum Lacepede, Hist. Nat. Quadr. Ovip. Serp., vol. 2, tabl.
meth., pp. 86, 331 (type locality, America).—Bonnaterre, Tabl. Encycl.
Meth., 1790, p. 18.
—
Sonnini and Latreille, Hist. Nat. Kept., vol. 4,
part 2, 1799, p. 132.—Daudin, Rept., vol. 6, 1800, p. 322.—Boie, Isis,
1827, p. 537.—Lacepede, Hist. Nat. Quadr. Ovip., vol. 1, 1836, p. 307.—
Ablates triangulum Dumeril and Bibron, Erp. Gen., vol. 7, pt. 1, 1854,
p. 315.
—
Lampropeltis triangula Cope, Proc. Acad. Nat. Sci. Philadelphia,
1860, p. 256.—Verrill, Proc. Boston Soc. Nat. Hist., vol. 9, 1863, p. 197.—
Abbott, Geol. New Jersey, Append. E, 1868, p. 802.—Allen, Proc.
Boston Soc. Nat. Hist., vol. 12, 1868, p. 180; vol. 13, 1870, p. 262.—
OpMbolus triangulum Goode, Proc. Amer. Ass. Adv. Sci., Aug. 1873,
p. 1S2.—OpMbolus trianx/ulus Smith, Sup. to Sci. News, 1879, p. 6; Geol.
Surv. Ohio, vol. 4, 1882, pp. 636, 639, 689.—Garman, S., Mem. Mus.
Comp. Zool., vol. 8, no. 3, pt. 1, 1883, pp. 65, 155, pi. 5, fig. 1; Nat. Hist,
notes, 1887, p. 2.—Higley, Trans. Wisconsin Acad. Sci., vol. 7, 1883-87,
pp. 166, 174.—Garman, H., Bull. Essex Inst., vol. 26, 1894, p. 35.—
Rhoades, Proc. Acad. Nat. Sci. Philadelphia, 1895, p. 392.—CoroncZfa
triangulum Boulenger, Cat. Snakes Brit. Mus., vol. 2, 1894, p. 200.—
Boettger, Kat. Rept.-Samml. Frankfurt, pt. 2, 1898, p. 72 (Milwaukee
Wisconsin).-Sternfeld, Sitz. Ges. Nat. Freunde, 1913, p. 105.—
Lampropeltis triangulum Abbott, Nat. Rambles about Home, 1894,
p. 300.—Dunn, Bull. Amer. Mus. Nat. Hist., vol. 37, art. 23, 1917, p.
630.
—
Lampropeltis triangulus Dunn, Copeia, no. 25, 1915, p. 63.
1827. Coluber eximius Harlan, Joum. Acad. Nat. Sci. Philadelphia, vol. 5, pt. 2,
p. 360 (type locality, Pennsylvania).—Hitchcock, Cat. Animals Plants
Massachusetts, 1833, p. 552, ed. 2, p. 534.—Harlan, Med. Phys. Re-
searches, 1835, p. 123.—KiRTLAND, 2d Annual Rep. Geol. Surv. Ohio, 1838
p. 167.—Storer, Rep. Fishes, Rept., Birds Massachusetts, 1839, p. 227.—
HoLBROOK, N. Amer. Herp., ed. 2, vol. 3, 1842, p. 69, pi. 15.—DeKay,
New York Fauna, 1842, pt. 3, p. 38, plates, Rept. Amph., pi. 12, fig. 25.—
LiNSLEY, Amer. Joum. Sci., 1843, p. 43.—Hough, Cat. Rept. Fishes St.
Lawrence County, N. Y., 1852, p. 23.—GtJNTHER, Cat. Colbur. Snakes
Brit. Mus., 1858, p. ^l.—Ophibolus eximius Baird and Girard, Cat. N.
Amer. Rept., pt. 1, 1853, p. 87.—Baird, Serp. New York, 1854, p. 21;
Pacif. R. R. Surv., vol. 10, pt. 3, no. 1, 1859, pi. 30, fig. 61.—Miles, Rep.
Geol. Surv. Michigan, 1861, p. 233.—Fogg, Proc. Portland Soc. Nat.
Hist., vol. 1,1862, p. 86.—OpMbolus eximia Hoy, Geol. Surv., Wisconsin,
vol. 1, 1883, p. 424.
—
Coronella eximia Jan, Arch. Zool. Anat., vol. 2,
fasc. 2, 1863, pp. 237, 242; Icon. Gen., livr. 17, 1866, pi. 1, figs. 3, 3A,
3B. 3C.
1842. Pseudoelaps Y Berthold, Abh. d. k. Ges. d. Wiss. Gottingen, vol. 1, p. 23,
pi. 1, figs. 11, 12 (type locality, North America).
1853. Ophibolus clericus Baird and Girard, Cat. N. Amer. Rept., pt. 1, p. 88 (type
locaUty, Clarke County, Virginia; type, U. S. Nat. Mus., no. 2380; 0. B.
Kennerly, collector).—Baird, Pacif. R. R. Surv., vol. 10, pt. 3, no. 1,
1859, pi. 30, fig. 62.
1856. Ablabes triangulum var. clericus Hallowell, Proc. Acad. Nat. Sci. Phila-
delphia, p. 246.
/ REVISION OF THE KING SNAKES. 189
1860. Lampropeltis doliata (part) Cope, Proc. Acad. Nat. Sci. Philadelphia, p. 257
(Del., Washington, D. C).
—
Abbott, Geol. of New Jersey, Append. E,
1868, p. 802.
—
OpMbolus doliatus Nelson, Rep. State Geol. New Jersey,
vol. 2, 1890, p. 647.
—
Osceola doliata Atkinson, Ann. Carnegie Mus., vol.
1, 1901, p. 150.
—
Lampropeltis doliahis Stone, Amer. Nat., vol. 40, no. 471,
1906, p. 167. Evermann and Clark, Proc. Indiana Acad. Sci. for 1914,
1915, p. 345.—Fowler, Copeia, no. 22, 1915, p. 40.
1875. Ophibolus doliatiis triungulus Cope, Bull. U. S. Nat. Mus., no. 1, p. 37.
—
Yarrow, Bull. U. S. Nat. Mus., no. 24, 1882, p. 90.—Davis and Rice,
N. Amer. Rept. and Batr., 1883, p. 34; Batr. Rept. Illinois, 1883, p. 29.—
Butler, Joum. Cincinnati Soc. Nat. Hist., vol. 10, 1887, p. 148.
—
Hay,
36th Annual Rep. State Board Agric. Indiana for 1886, vol. 28, 1887, p.
210; Joum. Cincinnati Soc. Nat. Hist., vol. 10, no. 2, 1887, p. 64.—Cope,
Proc. U. S. Nat. Mus., vol. 11, 1888, p. 383.—Nelson, Rep. State Geol.
New Jersey, vol. 2, 1890, p. 647.
—
Blatchley, Journ. Cincinnati Soc.
Nat. Hist., 1891, p. 32.—Cope, Proc. U. s. Nat. Mus., vol. 14, 1891, p.
609.—Hay, 17th Annual Rep. Indiana Dept. Geol. Nat. Res., 1892,
p. 517.—Cope, Amer. Nat., vol. 27, 1893, p. 1068, pi. 24, fig. 1.—Ditmars,
Proc. Linn. Soc. New York, 1896, p. 13.—Cope, Rep. U. S. Nat. Mus. for
1898, 1900, pi. 32, fig. 1.—Brown, Proc. Acad. Nat. Sci. Philadelphia,
1901, p. 72.—Branson, Kansas Univ. Sci. Bull., vol. 2, no. 3, 1904, p.
399.—Wallace, 56th Annual Rep. New York State Mus. for 1902, 1904,
p. rl40.
—
Brimley, Amer. Nat., vol. 37, 1903, p. 264.
—
Notestein, 7th
Rep. Michigan Acad. Sci., 1905, p. 118.
—
Brimley, Joum. Elieha Mitchell
Sci. Soc, 1907, pp. 146, 148.—Ditmars, Reptile Book, 1907, pp. 341, 342,
pi. 103, figs. 1, 3, pi. 105 (upper fig.); Reptiles of the World, 1910, p.
271.—Thompson, 13th Rep. Michigan Acad. Sci., 1911, p. 107.
—
Somes,
Proc. Iowa Acad. Sci., 1912, p. 150.
—
Brimley, Joum. Elisha Mitchell
Sci. Soc, vol. 30, no. 4, 1915, p. 202. Lampropeltis doliatus triangulus
Ditmars, Proc. Linn. Soc. New York, 1896, p. 13.
—
Mearns, Bull. Amer.
Mus. Nat. Hist., vol. 10, art. 16, 1898, p. 326.—Hay, Proc. Biol. Soc.
Washington, vol. 15, 1902, p. 138.—Clark, 7th Rep. Michigan Acad. Sci.,
1905, p. 110. Ditmars, Amer. Mus. Joum., vol. 5, no. 3, July, 1905, p.
98, figs. 4. 5. Surface, Bull. Div. Zool. Pennsylvania, vol. 4, 1906,
p. 174.
—
Fowler, Annual Rep. New Jersey State Mus. for 1906, 1907, p.
180.—Hahn, Proc. U. S. Nat. Mus., vol 35, 1908, p. 566.—Ruthven,
Michigan Geol. Biol. Suta^, publ. 4, biol. ser. 2, 1911, pp. 2G1, 2(57; 13th
Rep. Michigan Acad. Sci., 1911, p. 115; Michigan Geol. and Biol. Surv.,
publ. 10, ser. 3, 1912, p. 110.
—
^Thompson and Thompson, 14th Rep.
Michigan Acad. Sci., 1912, p. 158.
—
Shufeldt, Guide to Nature, vol. 5,
no. 9, 1913, p. 270, fig. 4.
—
Dunn, Copeia, no. 16, 1915, p. 1.
—
Engelhardt,
etal., Copeia, no. 17, 1915, p. 4.
—
Fowler, Proc. Delaware County Inst.
Sci., vol. 7, no. 2, 1915, p. 43. Thompson, Occ Pap. Mus. Zool. Univ.,
Michigan, no. 18, 1915, p. 5. Ellis, 19th Annual Rep. Michigan Acad.
Sci., 1917, p. 49.
—
Lampropeltis doliata triangula McLain, Notes Coll.
Rept., 1899, p. 3. Osceola doliata triangula Cope, Rep. U. S. Nat. Mus.
for 1898, 1900, p. 885, fig. 210. Atkinson, Ann. Carnegie Mus., vol. 1,
1901, p. 150.—Eckel, Amer. Nat., vol. 35, no. 40, 1901, p. 152.—Morse,
Ohio Nat., vol. 1, 1901, p. 127.—Reed, [Snakes Vic. Ithaca, New York,]
1901.—Eckel, Bull, New York State Mus., no. 51, 1902, p. 374.—
Henshaw, S., Occ. Pap. Boston Soc. Nat. Hist., vol. 7, 1904, p. 8.
—
Morse, Proc. Ohio State Acad. Sci., vol. 4, pt. 3, spec. pap. no. 9, p.
130.
—
Drowne, Men. Roger Williams Park Mus., no. 15, 1905, p. 11.
McAtee, Proc Biol. Soc Washington, vol. 20, 1907, p. 10.
190 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
1888. Ophiholus doliatus collaris Cope, Proc. U. S. Nat. Mus., vol. 11, p. 383 (type
locality, Elmira, Illinois; cotype, U. S. Nat. Mus., no. 2433, E. R. Board-
man, collector); same, vol. 14, 1891, p. 609; Amer. Nat., vol. 27, 1893, p.
1068, pi. 25, fig. 3; Rep. U. S. Nat. Mus. for 1898, 1900, pi. 33, fig. 3.—
Osceola doliata collaris Cope, Rep. U. S. Nat. Mus. for 1898, 1900, p. 887,
fig. 211.
—
Lampropeltis doliatus collaris Hay, Proc. Biol. Soc. Washington,
vol. 15, 1902, p. 138.
1888. Ophibolus doliatus clericus Cope, Proc. U. S. Nat. Mus., vol. 11, p. 383;
vol. 14, 1891, p. 609; Amer. Nat., vol. 27, 1893, p. 1068, pi. 24, fig. 2;
Rep. TJ. S. Nat. Mus. for 1898, 1900, pi. 32, fig. 2.—Brown, Proc. Acad.
Nat. Sci. Philadelphia, 1901, p. 72.
—
Ditmars, Reptile Book, 1907, pp.
341, 346, pi. 103, figs. 2, 4, pi. 105 (middle fig.).—OsceoZa doliata clerica
Cope, Rep. U. S. Nat. Mus. for 1898, 1900, p. 888, fig. 212.—Eckel,
Amer. Nat., vol. 35, no. 40, 1901, p. 152.
—
Lampropeltis doliatus clericus
Fowler, Annual Rep. New Jersey State Mus. for 1906, 1907, p. 181,
text figs., pi. 46.
1892. Ophibolus triangulus triangulvs Garman, H., Bull. Illinois State Lab.
Nat. Hist., vol. 2, art. 13, p. 295.
—
Lampropeltis triangulum triangulum
Stejneger and Barbour, Check List, 1917, p. 89.
—
Noble, Copeia, no.
88, 1920, p. 98.
1893. Ophibolus doliatus temporalis Cope, Amer. Nat., vol. 27, p. 1068, pi. 25,
fig. 4 (type locality, Delaware; type specimen, Acad. Nat. Sci. Phila-
delphia, no. 3597; Mr. Drexler, collector); Rep. U. S. Nat. Mus. for
1898, 1900, pi. 33, fig. 4.
—
Osceola doliata temporalis Cope, Rep. U. S.
Nat. Mus. for 1898, 1900, p. 889, fig. 213.
1900. Osceola doliata triangulum Cope, Rep. U. S. Nat. Mus. for 1898, pi. 18,
fig. 4.
1902. Lampropeltis doliatus doliatus Hay, Proc. Biol. Soc. Washington, vol. 15,
1902, p. 138.
Description.—This is tlie best known form of the genus, and the
one of which the most specimens have been available for study. The
following summary of the scutellation is based upon the examination
of about 400 specimens: Ventrals, 180 to 213; caudals, 29 to 54
(males average about 47 and females about 44); supralabials 7,
rarely 8; infralabials 9, occasionally 8, less often 10, very rarely 11;
1, very rarely 2, preoculars; 2, rarely 1, postoculars; temporals com-
monly 2 + 3 -f 4, often some lower combination, as 2 + 3 + 3, 2 + 2 + 3,
1 + 2 + 3; posterior chin shields usually shorter than anterior, in
contact with each other or separated by 1 or 2 small scaJes; loreal
distinctly longer than high, rarely absent; scale rows on middle of
body usually 21, occasionally 23, rarely only 19 (the formula is most
commonly 21-19-17 or 19-21-19-17, often 21-19, sometimes
21-23-21-19, rarely 19-17).
This snake is of moderate size. An adult is commonly 2^ to 3
feet long. The largest specimen examined was 1,085 mm. in length,
and was taken at Cold Spring, New York. Proportions: Head but
slightly distinct from the neck, tapering from the temples forward,
and truncate at the end; body, rather slender, tapering slightly to-
ward the neck and toward the tail, cylindrical above or very slightly
compressed; belly flat, and meeting the sides at right angles; tail
BEVISION OF THE KING SNAKES. 191
0.100 to 0.155 of the total length (males average about 0.135 and
females about 0.130).
Pattern: From head to tip of tail, 28 to 62 dorsal blotches or sad-
dles of brown, gray, or red (young and some adults), extending down on
the sides as far as to the fifth to second rows of scales. These blotches
are narrowly edged with black, and set on a ground color of whitish,
yellowish or grayish, minutely mottled with darker. On the sides,
alternating with the dorsal blotches, one or two series of roundish
spots, the upper row brown edged with black, the lower mostly
black; belly checked with black and white, sometimes suffused with
red; on back of head (fig. 54) usually a Y-shaped or oval spot of
white; from the side of the neck forward over the eye, a white band,
meeting its fellow across the frontal plate; a black band from the
eye to the angle of the mouth.
Typical individuals from the northern portion of the range may
be described as follows: Body above with 45 to 60 squarish brown
Pig. 54.—Lampropeltis Triangulum Triangulum (Acad. Phila., no. 3621, from Belmont, Phila-
delphia, Pennsylvania). About IJ x nat. size. Showing typical pattern of anterior end ot
BOOT,
blotches, narrowly edged with black, that extend down on the sides
to the fifth row of scales; on each side just below the dorsal blotches
is a series of roundish spots, also edged with black, that alternate
with the dorsal blotches; below these lateral spots and alternating
with them is a second series; these are mostly black, but often in-
close a little brown, and lie partly on the lower rows of scales and
partly on the ends of the ventrals; between these series of spots
or blotches is a ground color of grayish, minutely mottled with
black, the mottling appearing most strongly on the sides; the belly
is white (sometimes tinged with red), variegated with small quach-ate
blotches of black. Prominent on the posterior portion of the head
(fig. 54) is a Y-shaped mark of the ground color, edged with black,
and separated from the white of the neck by a brown band on each
side that joins the first dorsal blotch with the brown of the head.
The ground color of the side of the neck is carried forward above the
eye to meet a transverse band of the same color across the frontal
plate. This cross band is bounded in front by a brown bar, narrowly
edged with black, which crosses the posterior portions of the pre-
192 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
frontal plates and overlaps the anterior edge of the frontal. An-
terior to this the snout is light, mottled with dark, except for a narrow
bar of black across the prefrontal-internasal suture. There are
usually two oval light spots near the parietal suture; a black band
extends diagonally from the eye to the angle of the mouth. The
labials and chin shields are whitish, except for more or less black
near the sutures.
The dentition, as indicated by a dozen specimens from all parte
of the range, is as follows: Maxillaries, more often 12, less often 13,
larger in front, decreasing behind, except that the last two are some-
what longer and stouter than any of the preceding; mandibulars,
11 to 15, most often 13, distinctly larger anteriorly, decreasing rapidly
behind, the third, fourth, and fifth largest, a slightly greater space
between the fourth and fifth or between the fifth and sixth; palatines,
9, 10, or 1], decreasing slightly in size posteriorly; pterygoids, 17
to 23, becoming smaller posteriorly.
Although sufficiently distinct as a rule, there is no characteristic
which will always distinguish triangulum from syspila. These two
forms have been contrasted under the description of the latter.
Specimens of doubtful identity will be found near the common
boundary of their ranges, and in Virginia, Maryland, eastern Penn-
sylvania, New Jersey, Delaware, and perhaps in West Virginia.
Typical triangulum occurs in all these states, but many individuals
show marked degeneracy in pattern and in scutellation. Specimens
wUl be found with as few as 28 dorsal saddles, which extend to the
second or first row of scales, and with head markings typical of
syspila, but such individuals average to have a less number of ven-
trals and caudals, and a lower scale formula than similar exam-
ples of syspila. Specimens from these localities identifiable in the
key as syspila must, for the present at least, be referred on geo-
graphic grounds to triangulum.
Habitat and habits.—From Wisconsin and Illinois, east to New
England, this is a common and well-laiown snake, but in the south-
eastern States, except in the mountains, it appears to be rare. It is of
very general occurrence in all upland situations, woods and fields, and
is not uncommon in the vicinity of dwellings and stables. It does
not appear to haunt the shores of streams or lakes, but may be
expected in most other situations. Its wide choice of habitat is
related to the widespread occurrence of its principal food—mice.
For the vicinity of Saginaw Bay, A-fichigan, Ruthven (1911, 267)
says that ''they were mostly found in the decaying logs on the fossil
beaches and pine ridges, where they fed, in part at least, on the
Michigan mouse (Peromyscus maniculatus bairdii) and the blue-tailed
skink (Eumeces quinquelineatus) , as remains of these animals were
found in the stomachs examined," A specimen taken in a hay field
REVISION OF THE KING SNAKES. 193
in Cheboygan County, jMichigan, disgorged an adult meadow vole
(Microtus pennsylvanicus); another had eaten an adult short-tailed
shrew (Blarina hrevicaudata)
.
Its presence in the vicinity of dwellings may be largely explained
by its fondness for mice. Better than a cat, it can follow a mouse
to its nest and eat the whole brood of young. Ditmars (1907, 344)
mentions finding one that had been killed in a barn, in Sullivan
County, Now York, that had in its stomach five very young rats.
Surface (1906, 178-179) finds that in Pennsylvania three-fourths
of the food of this snake consists of small mammals, chiefly mice,
and that the destructive meadow mouse (Microtus feniisylvanicus)
constitutes about half its food. This fact alone should be sufficient
to assure its protection at the hands of farmers and others; and,
when one considers that it is a perfectly harmless, non-aggressive,
unfearing creature, he may well wonder why it is so persistently
persecuted. Its lack of fear is one cause of its great decrease in
numbers. As a farmer will say, ' ' You have to kick it out of the way."
Of course it is considered poisonous by the majority of people, and is
killed for that reason, or just because it is a snake. Many snakes
are, indeed, of slight, if any, economic value; some, as the rattle-
snakes that destroy pests, have the disadvantage of being poisonous,
but the spotted adder, or milk snake, has perhaps the distinction
of combining the greatest number of valuable traits with the mini-
mum of undesirable or neutral qualities. It is true that he must
be charged with the eating of young birds when the nests are located
in accessible places, but Surface (1906, 178-179) has shown that
birds constitute but a small proportion of its diet, and doubtless
some of the birds it is found to have eaten were taken dead. Snakes
and lizards constitute a regular, although small, proportion of its diet.
It is known to eat the Butler garter snake (ThamnopMs hutleri), the
smooth green snake (Liopeltis vemalis), the water snake (Natrix
sipedon), the ring-necked snake (DiadopMs punctatus, McAtee, 1907,
10); De Kay's snake, the queen snake or striped water snake, the
red-bellied snake, robin's eggs, jumping mice, the white-footed mouse,
and slugs are charged to it by Surface (1906, 178-179); and young
individuals have been observed, in captivity, to eat small dead
minnows, and pieces of the same.
"In captivity [Ditmars, 1907, 344] this snake is indifferent in
feeding and seldom lives long. It prefers mice, which are quickly
constricted to death in the reptile's strong coils. Young specimens
can seldom be induced to take food of any character. Although
rather a quiet reptile, the milk snake will sometimes resent handling
in a curious and rather treacherous manner. Without a pretense
of striking it will swing the head about suddenly and grasp the hand,
when it deliberately chews in such a manner that the fine, recurved
194 BULLETIN" 114, UNITED STATES NATIONAL MUSEUM.
teeth lacerate the flesh sufficiently to bring the blood, although the
minute punctures are but very superficial wounds and heal at once,
like a scratch from a fine point."
It is likely that this snake does much of its foraging at night, as then
it would have the best chance to find mice and the small snakes
and lizards that hide under boards, logs, and bark. Its occasional
presence in dwellings may be explained, as Surface has indicated
(1906, 177), by the attraction of a warm atmosphere when the air
outside is cold, or its prowling along crevices, into cracks and holes
m.Siy lead it into buildings. Without doubt it is often brought into
cellars and sheds under the bark or in the cracks of firewood.
This snake is oviparous, depositing from 8 to 13 oval eggs (Ditmars,
1907, 344-345) with a leathery shell, and presenting the same soft
and white appearance as the surface of a toadstool. Ditmars records
batches of 8, 11, and 9, laid on July 10, 12, and 28, and hatching on
September 5, 6 to 8, and October 1 to 3, respectively. Mr. E. B.
Williamson took a batch of 13 eggs in Wells County, Indiana, in
August, 1917, 7 of which hatched September 15 to 17. O. P. Hay
(1892, 518) reports finding eggs of this snake in Indiana, in a pile of
manure, more or less glued together. Young snakes are about 6f to
8|- inches in length. They molt very soon after hatching.
Nothing definite seems to be known about its hibernation. It
probably burrows in the ground, as it is often turned out by the plow
in the spring. It appears to be rather late entering winter quarters
and early.emerging, although probably less so than the garter snake.
Hay (1892, 518) reports taking it in Indiana on March 30, and of
having seen a dead one in the road still earlier.
Range.—From Minneapolis south through northern Illinois,
Indiana (except the southwestern section), eastern Tennessee,
Georgia, and Florida, and north through all the eastern States to the
forty-sixth parallel.
On the west from Tennessee to Minnesota it meets and inter-
grades with its only close relative, syspila. In the southeastern
States, except in the mountains, it appears to be a rare snake. The
most northern record is for a specimen taken in 1917 at Cecil Bay,
Emmett County, Michigan, by Dr. J. H. Ehlers; in New England
it has not been reported north of Vassalboro, Maine; it is common
in Cheboygan and Emmett Counties, Michigan, but is not reported
for the Upper Peninsula; it should however be looked for on the
south shore of the northern peninsula of Michigan, as it has been
taken at Newport, Door County, Wisconsin. In Florida it has been
taken as far south as Candler, in Marion County.
The most northern localities for Canada are Spence's Lake, Brace-
bridge, Ontario, and Aylmer, near Quebec.
REVISION OF THE KING SNAKES. 195
Specimens have been examined from the following localities, in
addition to those represented by specimens in the United States
National Museum: Portland, and MiddJetown, in Connecticut;
Delaware; Candler (llarion County), and Gotha (Orange County), in
Florida; Athens, Beverly Hills, Danville (Vermillion County), Glen
EUyn and Hinsdale (Du Page County), Joliet (Will County), Kendall,
FlQ. 55.—Map SHOVIKQ LOCAUTT records for LAMPROPELTIS TRIANGULUU TRIANGULUM.
Kenilworth (Cook County), in Illinois; Bloomington CMunroe
County), Bluffton (Wells County), La Fayette, Lebanon (Boone
County), Mitchell (I^awrence County), North Vernon (Jennings
County), Sims (Grant County), Vigo County, and Winona Lake, in
Indiana; Lewiston, and Vassalboro, Maine; Brookline, Cambridge,
Deerfield, Feltonville, Hudson, Lancaster, Lynn, Magnolia, Maiden,
196 BULLETIIT 114, UNITED STATES NATIOITAL MUSEUM.
Medford, Naushon Island, near Northampton, Waltham, Ware,
Warwick, Wenham, and Williamstown, in Massachusetts; Ann Arbor,
Bay View (Emmett County), Berrien County, Brighton (Livingston
County), Cass County, Cecil Bay (Emmett County), Charity Islands
(Lake Huron), Douglas Lake (Cheboygan County), East Lake
(Manistee County), East Lansing (Ingram County), Jackson County,
Kalamazoo, Le Roy (Osceola County), Northport (Leelanau County),
Olivet, Orion and Pontiac (Oakland County), Port Austin (Huron
County), Portage Lake (Washtenaw County), Rollins (Lenawee
County), Shelby (Oceana County), Third Sister Lake (Washtenaw
County), Van Buren County, Walnut Lake (Oakland County),
Wayne County, and Ypsilanti, in Michigan; Fort Snelling (near
Minneapolis), Minnesota; North Conway, New Hampshire; Cald-
well (Essex County), Delair, Haddonfield (Camden County), Morris-
town, Short HiUs (Essex County), Snow Hill (Camden County),
Trenton, Woodstown (Salem County), in New Jersey; Albany,
Ashoken Survey (Ulster County), west side Cayuga Lake, Cold
Spring (Putnam County), Cornell Heights, Cousook, Fallsburg (Sul-
livan County), Forest Hills (Long Island), Hornellsville, Hyde
Park (Dutchess County), Irvington-on-Hudson (Westchester County),
New Lots (Long Island) Perry City (Wyoming County), Purdy Creek
(Steuben County), Rye (Westchester County), Six Mile Creek (near
Ithaca), Staten Island (Grosmere), Taughannock Falls (Thompkins
County), Wascabne Mountain (Westchester County) in New YorTc;
Brevard, Pineola (Avery County), Pisgah Forest Reserve, Roan
Mountain (6,000 feet, Mitchell County), in Nortli Carolina; Columbus,
East Rockport, Franklin County, Hughes (Butler County), Kettlers-
ville (Shelby County), Logan County, Sandusky, Youngstown (Ma-
honing County), in Ohio; Bakerstown (Allegheny County), Beaver
(Beaver County), Belmont (Philadelphia County), Carlisle (Cumber-
land County), Chambersburg (Franklin County), Cherry Run (In-
diana County), Concord (Delaware County), Conneaut Lake (Crawford
County), Cumberland Valley, Diamond Valley (Huntington County),
Fairmont Park and Germantown and Holmesburg (Philadelphia
County), Indiana (Indiana County), Jackson Valley, Jennerstown,
Johnstown, LaGrange (Philadelphia County), Lewisburg, Linsvillo
(Crawford County), Lopez (SuUivan County), Ludlow (McKean
County), Mill Creek (Montgomery County), Mount Washington (Alle-
gheny County), Northumberland, Overbrook (Philadelphia County),
Pike County, Pittsburgh, Presque Isle, Raymonds (Potter County),
Sewickley (Allegheny County), Summit, Tarentum (AlleghenyCounty),
Two Lick Creek (Indiana County), Venango County, Walnut Hill
(Montgomery County), Wama (Delaware County), Water Street (Hunt-
ington County), Waynesburg (Green County), Westchester (Delaware
County), Williamsport (Lycoming County), Wilmerding (Allegheny
County, in Pennsylvania; Hampton County, South Carolina; Franklin
REVISION OF THE KING SNAKES. 197
County), Knoxville, and Nashville, Tennessee; Bethel, Vermont; Lou-
doun County, and Staunton (Augusta County), Fw'^rmia; Cheat River,
"^est Virginia; Beaver Lake (Dodge County), Cedarburg, Cedar
Lake (Washington County), Madison, Mauston, Milwaukee, Newport
(Door County), and Waukesha, in Wisconsin; Aylmer (Quebec),
Hull (Ottawa), near Ottawa, Point Pelee (Ontario), Spence's Lake
(Bracebfidge, Ontario), and near Toronto, in Canada.
Published records, that appear to be reliable, for other localities,
are as follows: Huntington, Connecticut, (Linsley, 1843, 43); Nor-
way, Maine, (Verrill, 1863, 197); Springfield, Massachusetts, (Allen,
1868, 180); near Waveland, Montgomery County, (Butler, 1887,
148), and Greencastle C^fcLain, 1899, 3), Indiana; London, Ohio
(Morse, 1901, 127), Hamilton County, Akron, and Nelsonville, Ohio
(Morse, 1904, 130); Alma, Michigan (Ruthven, 1912, 113); Antrim,
Montcalm, Kent, Ottawa, and Barry Counties, Michigan, (Clark
1905, 110); Rockland (Wallace, 1904, rl40), Rossic, St. Lawrence
County, New York (Hough, 1852, 23), and Cold Spring Harbor,
Yaphank, Orient, Greenport, and Southold, on Long Island, New
York (Engelhardt, 1915, 4); Stony Run, Cecil County, Maryland
(Fowler, 1915, 40); St Clair township (Allegheny County), Pennsyl-
vania (Atkinson, 1901, 150); Shower Hill, New Jersey (Fowler, 1907,
181); Sunburst, Hay^vood County (Brimley, 1915, 202), and Chest-
nut Knob Mountain, North Carolina (Dunn, 1917, 630).
Variation.—In spite of the apparent constancy and distinctness of
triangulum in the northern tier of States, the form as a whole is one
of the most variable. The diagram showing geographic differences
in number of ventrals (fig. 56) brings out some notable variations with
locality. The dependability of the figures varies somewha't with the
number of specimens upon which they are based. It is evident, how-
ever, that in the northern and western portion of its range the ventrals
average higher, more than 200, and the extremes are higher than for
the central and northeastern parts. The few specimens from the
southeastern States, except in the mountains, show the high num-
bers of the northwest. From New England to Virginia and in the
Alleghenies the averages and the extremes are low, and distinctly
lower in New Jersey and Delaware, and near the District of Columbia.
This variation in ventrals should be compared with the table of
scale formulae. It will be noted that in the states close to the range
of syspila the scale rows average about the same as for that form,
that is, a formula lower than 21-19 is very common in males and a
little less so in females. In New York and Pemisylvania the only
change is a slightly lower average, but in New England, the most dis-
tant region, only one male is recorded with a formula of 21-19, more
than half the females have a lower formula than this, and the smallest
proportion of specimens reaches 23 rows. This compares closely with
the southern Alleghenies. The formulae for New Jersey and the
198 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
District of Columbia average still lower than in New England and the
Alleghenies, and it is here that the lowest point of all, 19-17, is
attained.
Of other features of the scutellation the temporals are perhaps the
most variable. The normal number for the group and for the genus
is 2 + 3+4, but in this group in particular a reduced number is very
Ventral
plates.
216
210
204
198
192
186
'
180
In- North- Wis- Mich- Can- New South- East- NJ. D.C. Ohio. West- Alle- N. C. Ten- Fla.
diana. ern con- igan. ada. Eng- ern ern Del. 23 7 em ghen- S.C. nes- 3
19 HI. sin. 61 6 land. N.Y. Pa. 14 Pa. ies. 3 see.
10 16 63 25 19 52 19 6
Localities, and ntjmbees of specimens.
Fig. 56.—Diagram sho-wing geographic variation in number of ventral plates in Lampropeltis
triangulum triangulum.
frequent—the last row is oftenest reduced by one, the second row
less often so reduced, and the first row is the most constant. The
specimens from the District of Columbia show the greatest reduction
in temporals. The accompanying table gives at a glance the aver-
ages to two places of decimals, of the temporals in the first and
third rows for Missouri, Massachusetts, and the District of Columbia.
Temporals
of the
first rov
.
Temporals
of the
third row.
Syspila in Missouri L88
1.94
L63
3.65
3.83
3.26
Triangulum in Massachusetts.
Triangulum in District of
Columbia
REVISION OF THE KING SNAKES. 199
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200 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
This indicates that the District of Columbia specimens have suf-
fered an actual reduction in the number of temporal scutes. In the
extent of this reduction they can be compared only with micwpholis,
elapsoides, and virginiana, reduced forms belonging to the same group,
but only distantly related to triangulum.
Bearing in mind the peculiarities of the geographic variation in
ventrals, scale rows, and temporals we have now to consider the color
pattern. As will be evident from the descriptions, triangulum is
separated from syspila almost entirely on the basis of color pattern
differences. Typical specimens of the two forms are very different
appearing animals, and, although intergradation certainly takes place
along the common boundary of their ranges in Illinois, Indiana, and
Ohio, each form seems to retain its distinctive features very close to
the border of its range. Just how true this statement is for Kentucky
and Tennessee can not be told now for lack of specimens, but the few
we do possess indicate that syspila extends over about the western
third of these States. Throughout the northern tier of States the
typical pattern, as given in the description, is seldom departed from.
The dorsal blotches average about 47, are almost always more than 35,
and are usually accompanied by two more or less well-developed
series of lateral alternating spots on each side. But in the southern
AUeghenies the dorsal blotches average somewhat fewer, the higher
numbers of the north are rarely, if ever, attained, the blotches often
extend lower down the sides, the two rows of lateral alternating
blotches also often extend down lower, and are rather frequently
fused, either partially or completely, into a single series. This does
not mean that typical examples do not occur, but that they are less
frequent than in the north. This reduction in number of dorsal
blotches, accompanied by increase in width and consequent loss of
the upper series of alternating spots, reaches its extreme, as the dia-
gram shows (fig. 57), in New Jersey, Delaware, Maryland, and
northern Virginia.
The complicated and characteristic pattern of the head follows, in
a general way, the pattern of the body. It is most perfectly developed
in the northern parts of the range. Even here the chevron on the
back of the head may appear only as a transversely oval spot, and the
white line between, over, and behind the eyes may be partly bridged
by extension of the brown pattern. But in practically all of these
northern specimens the half collar is completely replaced by the
chevron or its representative. Close to the range of syspila the half
collar is often partly or completely developed. Many specimens from
western Pennsylvania and northern West Virginia, while typically
triangulum in other respects, possess the half collar instead of the chev-
ron. South in the AUeghenies aU conditions of chevrons and half
collars may be found. Often when the half collar is present there
will be two notches in its anterior border or one in its posterior, rep-
REVISION OF THE KING SNAKES. 201
resenting the arms of the chevron. The District of Columbia speci-
mens often have the half collar complete and the other head mark-
ings much reduced. Some specimens cannot be distinguished from
examples of syspila. Those from New Jersey, and Delaware and
some from eastern Pennsylvania and parts of Maryland are reduced
in the same way.
Some of these specimens have received varietal names. Thus
Baird and Girard described an individual from Clarke County,
Virginia, as clcricus, and the name has been frequently used for speci-
mens from New Jersey and other states, and the name has been
supposed to apply to a close relative of triangulum that replaced it
on the south. It can be stated, however, that the material at hand
Dorsal
blotches.
61
49
I 1 1 I ,:m« '
New District Southern
Jersey. of Columbia. Alleghenies
14 22 14
25
Wis- Michi- New New
consin. gan. England. York.
16 61 63 45
LOCAUTIES, AND NUMBERS Ot SPECIMENS.
FlQ. 57.—DUGRAM SHOWING GEOGRAPHIC VABUTION IN NUMBER OF DORSAL BLOTCHES IN LAMPROPELTB
TRLANGULUM TRLiNGULUM.
indicates that the supposably distinguishing features of clericus have
no constancy and no geographic basis. However, it is not improbable
that when sufficient specimens are available for study, a local race
wiU be found having its center in New Jersey and Delaware, and
perhaps southern Virginia. Stone (1906, 167) and others have noted
that typical triangulum appears to be replaced in the coastal plain
regions of these States by individuals with fewer and broader dorsal
blotches and reduced head markings, but as ah'eady stated the
present material is insufiicient for identifying a separate race. The
collaris * of Cope was based upon selected individuals from Elmira,
Illinois, St. Louis, Mssouri, and Washington, District of Columbia,
that exhibited the half collar, regardless of whatever other char-
acters they possessed.
« Although no type was specifically designated, Cope twice figured United States National Museum
specimen No. 2433 from Elmira, Illinois, in coimection \vith descriptions of collaris, and this may therefore
properly be considered the type. As this specimen is a triangulum, the name collaris becomes a synonym.
202 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
Similarly an aberrant individual from Delaware (Acad. Nat. Sci.
Philadelphia, no. 3597) became the type of temporalis Cope. There
may be others like this in Delaware (Stone, 1906, 167) but so far
this appears to be the only example of temporalis. The head pattern
is obsolete; the dorsal blotches are wide, few, and somewhat fused
with each other; and the scale formula is 19-17.
Affinities.—Under variation it was shown that triangulum attains
its most perfect development in the northern States, that in the
Alleghenies and on the Atlantic coast it has suffered more or less
reduction in scutellation and pattern, and that the greatest reduction
occurs from southern New York to Virginia. The meaning of this
extreme condition is not at all clear, and, in attempting an explana-
tion, it must be remembered that from the whole peninsula of Dela-
ware and eastern Maryland we have but a single specimen, and that
all of those from Virginia are from close to the District of Columbia
or from the Alleghenies. Some of those from the District have been
known under the name of "doliatus," and have been supposed to be
the same thing as what we are here calling syspila. Admittedly some
young individuals can not be told from the latter. Others, however,
are typically triangulum, and still more are mixtures. Perhaps this
is a region of intergradation between a typical triangulum and a
coastal variety extending from New Jersey to North Carohna. There
seems to be nothing constant. In this connection it may be remarked
that more than half of these specimens are very small, and perhaps
some of the aberrant ones would have died a natural death very
soon if they had not been found and preserved. It is a fact that
some of the most aberrant individuals, described and not described,
are juveniles. Of the former multistrata Kennicott is an example.
If these Washington specimens were fairly constant in pattern and
scalation and possessed a definite geographic range, they could be
recognized with a name, but such does not appear to be the case.
The resemblance to syspila must be regarded as secondary because
the scutellation is definitely reduced, and the pattern as weU bears
every evidence of being capable of derivation by reduction from typi-
cal triangulum. Whatever may be the reason it seems evident that
in the parts of its range where triangulum is farthest removed from
syspila it has a reduced scutellation or pattern or both; for example,
New England east of the Berkshires, New Jersey and Virginia east
of the Alleghenies.
The fact that the pattern of triangulum seems to be developed in
perfection only north of the southern limit of glaciers argues against
the assumption of this pattern as primitive. As brought out in the
discussion of syspila, triangulum must be considered as at one end
of the chain, amaura-syspila-triangulum. If we are to favor evolu-
tion or differentiation accompanying migration, we can not do other-
wise than regard triangulum as the latest of this chain.
REVISION OF THE KING SNAKES. 203
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204 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
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REVISION OF THE KING SNAKES. 205
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206 BULLETIN 114-, UNITED STATES ISTATIOlSrAL MUSEUM.
LAMPROPELTIS ELAPSOIDES ELAPSOIDES (Holbrook),
SCARLET KING SNAKE, BED KING SNAKE, THUNDER SNAKE, "COEAL SNAKE."
Figs. 58, 73.
1789. Coluber doliatus Lacepedb, Hist. Nat. Serp., vol. 2, tabl. meth., pp. 104,
294.
—
Daudin, Hist. Nat., vol. 7, 1800, p. 74.
—
Harlan, Joum. Acad.
Nat. Sci. Philadelphia, vol. 5, pt. 2, 1827, p. 362; Med. Phys. Researches
1835, p. 125.
—
INatrix doliatus Merrem, Versuch Syst. Amphibien, 1820
p. 129.
—
Coronella doliata Holbrook, N. Amer. Herp., ed. 2, vol. 3,
1842, p. 105, pi. 24.
—
Ofliiholus doliatus Garman, S., Mem. Mas. Comp,
Zool., vol. 8, no. 3, pt. 1, 1883, pp. 64, 154, pi. 5, fig. 2.
—
Coronella doliatus,
Bote, Isis, 1827, p. 539.
—
Coronella doliata Boulenger, Cat. Snakes
Brit. Mus., 1894, p. 205.
1837. Coronella cocdnea Schleqel, Essai Phys. Seri>., p. 57, pi. 2, fig. 11.
—
GiJNTHER, Cat. Colubr. Snakes Brit. Mus., 1858, p. 42.
—
Jan, Arch. Zool.
Anat., vol. 2, fasc. 2, 1863, pp. 237, 239; Icon. Gen. Ophid., livr 17,
1866, pi. 1, fig. 1.—BocouRT, Miss. Sci. Max., pt. 3, vol. 2, 1886, p. 608,
vol. 3, pi. 39, figs, la to le.—Lampropeltis cocdnea Cope, Proc. Acad.
Nat. Sci. Philadelphia, 1860, p. 257.
—
Ophibolv^ cocdneus Brimley,
Joum. Elisha Mitchell Sci. Soc, vol. 21, no. 4, 1905, p. 152.
1838. Coluber elapsoides Holbrook, N. Amer. Herp., ed. 1, vol. 2, p. 123, pi. 28
(type locality. South Carolina and Georgia).
—
Calamaria elapsoidea Hol-
brook, N. Amer. Herp., ed. 2, vol. 3, 1842, p. 119, pi. 28.—DeKay,
New York Fauna, pt. 3, 1842, p. 49. Osceola elapsoidea Baird and
GiRARD, Cat. N. Amer. Rept., pt. 1, 1853, p. 133.
—
Lichtenstein, Mus.
Zool. Berolinenses, 1856, p. 24.
—
Cope, Proc. Amer. Philos. Soc, vol. 17,
1878, p. 65.—Yarrow, Bull. U. S. Nat. Mus., no. 24, 1882, p. 89.—Davis
and Rice, N. Amer. Batr. Rept., 1883, p. 33.—Cope, Proc. U. S. Nat.
Mus., vol. 11, 1888, p. 381; Proc. U. S. Nat. Mus., vol. 14, 1891, p. 606.—
Loennberg, Proc. U. S. Nat. Mus., vol. 17, 1894, p. 325.
—
Cory, Hunting
and Fishing in Florida, 1896, p. 130.—Cope, Rep. U. S. Nat. Mus. for
1898, 1900, p. 900, fig. 221, pi. 18, fig. 3.—Wright, Proc. Acad. Nat. Sci.
Philadelph;ia, 1915, pp. 139, 140, 148.
—
Ophibolus elapsoideus Garman,
H., Bull. Illinois State Lab. Nat. Hist., vol. 3, art. 13, 1892, p. 229 (Anna,
III.).
—
Lampropeltis elapsoides Stejneger and Barbour, Check List,
1917, p. 88.—SrEJNEGER, Proc. Biol. Soc. Washington, vol. 31, 1918, p. 99.
1883. Ophibolus doliatus elapsoideus Gabman, S., Mem. Mus. Comp. Zool., vol. 8,
no. 3, pt. 1, pp. 65, 155.
1901. Ophibolus doliatus cocdneus Brown, Proc. Acad. Nat. Sci. Philadelphia,
p. 74.
—
Brimley, Joum. Elisha Mitchell Sci. Soc, vol. 21, no. 4, 1905,
p. 145; 1907, pp. 145, 148.—Ditmars, Reptile Book, 1907, pp. 341, 350,
pi. 105 (lower figure).
—
Brimley, Joum. Elisha Mitchell Sci. Soc, vol.
30, no. 4, 1915, p. 202.
—
Lampropeltis doliatus cocdneus Wright, Proc.
Acad. Nat. Sci. Philadelphia, 1915, pp. 142 and following, also p. 165,
fig. 5.
1915. Osceola doliata doliata Wright, Proc. Acad. Nat. Sci. Philadelphia, p. 139.
The name Coluber doliatus of Linnaeus must, as Stejneger has
pomted out (1918, 99), be rejected as unidentifiable; it is unlikely
that it was intended for any form that we now know as Lampropeltis,
The next name to be considered, therefore, is the Coronella cocdnea
of Schlegel. He, without doubt, had the present species before him
REVISION OF THE KING SNAKES. 207
but he referred it to the foPin described as Coluber cocdneus by
Latreiile (Sonini and Latreille, 1799, 138); in the belief that the latter
had had the same thing that he (Schlegel) had but erroneously thought
it to be the Coluber coccinca of Blumenbach (1788, 11). As a matter
of fact, there can be no doubt that Latreiile had only Blumenbach's
coccinea, and this, as is well known, is the Cemphora cocdnea that
we know to-day, and which is a very different snake from our L.
elapso'ides, but one that presents an unusually strong superficial resem-
blance to it. Therefore, since Schlegel refers his coccinea to Latreille's,
and since the latter was plainly a very different thing, the name coc-
cinea Sclilegel is a synonym of Cemophora coccinea (Blumenbach).
Description.—This form is the smallest of the genus and the most
removed structurally from the ancestral type. The scutellation, based
upon 83 specimens, is as follows: Ventral plates, 152 to 193, average
about 175; caudals, 32 to 48 (males, 38 to 48, average 42; females
32 to 42, average 37.5); supralabials, 7; infralabials, 8, sometimes 9,
rarely 7; 1 preocular; 2, very rarely 1 or 3, postoculars; temporals,
1+2 + 3, occasionally an additional scute in one of the rows; pos-
terior chin shields usually in contact, shorter than or about as long
as the anterior; loreal decidedly longer than high, or entirely replaced
by a downward extension of the prefrontal; scale rows iisually
17-19-17, som.etimes 19-17, or 17-19-17-15 or 17-15, very rarely
reaching 21, always having at least as few as 19 rows o:i the neck
and never ending with more than 17.
Head but slightly distinct from the neck; body slender, somewhat
compressed, varying but little in diameter, sides meeting belly in a
well-defined angle; tail slender, tapering, 0.118 to 0.169 of the total
length (males, 0.129 to 0.169, average 0.149; females, 0.118 to 0.147,
average 0.135). The largest specimen examined measured 599 mm.
and came from the Okefinokee Swamp in southern Georgia.
The pattern is made up of 15 to 25 paire of black rings encircling
the body from head to tip of tail. These border narrower rings of
yellow (Brown, 1901, 74, says "white in the young, je]lo\v in adults")
and are separated by broader ones of red. The black rings are a
little Av4der on the dorsal line and sometimes meet here across the
red; they become narrower on the first row of scales and usually
extend across the belly, but if interrupted here, they extend at least
onto the ends of the ventrals. The yellow rings are about 1^ to 2
scales wide on the middorsal line and 2 to 4 on the firet row of scales;
they are complete on the belly or partially interrupted by a black
spot; on the sides they are often more or less mottled with darker.
The red bands average to be about as wide as the groups of black and
yellow rings that separate them; they are complete on the belly,
although som.etimes sliglitly mottled or spotted there with black.
208 BULLETIN" 114, UNITED STATES NATIONAL MUSEUM.
The occiput is crossed by a black band (fig. 58) that may extend
forward between the eyes. The snout is reddish and immaculate.
Behind the occipital black band is a yellow collar, and behind this
is another black band which is generally interrupted on the throat.
The chin is a mingled red and j^ellow passing into yellow on the throat.
The actual colors of an adult specimen from Florida are as follows
(color names from Ridgway, 1912) : Morocco red above, changing
to a dragon's blood red below; encircled by narrow bands of a baryta
yeUow on the sides, slightly deeper chrome above and paler below,
which are bordered with black, the latter becoming blackish brown
below. On the occiput a sooty black band; snout a claret brown;
labials, morocco red mingled with a little yellow, anterior and pos-
terior edges black; chin and throat, mingled Morocco red and pale
martin's yellow. First yellow ring conspicuously mottled with red,
succeeding ones slightly so down to the second or third row of scales
Fig. 5S.—Lampkopeltis elapsoides elapsoides (U.S.N.M. no. 55903, St. John County, Florida).
About 2| X nat. size. Showing typical patteen of anterior end of body.
(the red in minute flecks on the yellow). Iris, Morocco red; pupil
black; tongue Morocco red, or browner.
The dentition, as based upon examination of eight specimens, is as
follows: Maxillaries, 13, 12, or 14, subequal, the last two slightly
larger; mandibulars, 12 to 14, the third, fourth, and fifth the largest,
decreasing decidedly behind, the greatest space between the fourth
and fifth; palatines, 11, 10, or 12, subequal; pterygoids, 17 to 23
(except that a specimen from Raleigh, North Carolina, has 26 and
27), smaller than the palatines, and decreasing posteriorly.
The copulatory organ may be described as follows: Bilobed (but
not forked) ; sulcus single, and extending over the side of the larger
lobe. A small space at the distal end of the organ may be practically
free of calyces, or exhibit only ridges of the latter extending across
it. Calyces not numerous ; fringes few, extending distinctly, although
only a little, below the end of the organ before changing to spines.
The latter numerous, closely set together, increasing gradually in
size to about the middle of the organ, then getting smaller again,
but not abruptly so. Thus the spines extend more than half way
to the base, sometimes nearly all of the way. Unless the organ was
fully everted when the specimen was fresh, but little more will be
made out than that it has a few calyces at the extreme distal end,
REVISION OF THE KING SNAKES. 209
bearing short fringes, followedTay slender, closely set spines, extending
distinctly more than half way to the base of the orgsiii.
Habitat and habits.—The best account of its habits is given by
Ditmars (1907, 352):
The Scarlet King Snake is a burrowing species, thus demonstrating in habits, as
well as in form, its degenerate character as compared with the other snakes of the
genus Ophibolus. Specimens may be most commonly foimd under the loose bark
of fallen and decaying trees. Here they prey upon the smaller species of snakes,
lizards, or upon very young mice that are yet in the nest. It is probable that this
little snake issues at night from its lair and searches in crevices in the bark for
various lizards that crawl into such places to sleep. This theory appears logical
after an examination of a series of preserved specimens, in wHch the stomachs of
several contained the remains of swifts (Sceloporus) and "blue-tail" lizards (Eumeces).
In captivity this snake evinces a very gentle disposition, seldom offering to bite.
However, it invariably burrows into the soil of its cage or hides under loose objects,
such as pieces of bark or fiat stones. Thus it constitutes a very indifferent object
for observation and study. While displaying a very indifferent appetite as a captive,
it may be occasionally induced to take verj' young mice. Although diminutive in
make-up, the reptile constricts its prey in exactly the same fashion as its larger and
more powerful allies.
Wright says (from observations in the Okelinokee Swamp, 1915,
167):
In food habits this species is more or less of a constrictor. It feeds on gi'ound
lizards, skinks, swifts, and other snakes and insects. In the stomach of No. 6242
we found an angle worm and the remains of two killifishes, suggesting more of an
aquatic nature tlian usually ascribed, but after every rain Billy's Island is covered
with little water pools containing fish which as evaporation goes on become stranded.
Such would be easy of capture. Our specimens jdelded no clue to the oviparity or
breeding of this species.
Suggesting its probable climbing abihty, Wright records the cap-
ture of a specimen "on one of the frames of an old building, the
snake being 3^ to 4 feet above the ground."
Range.—Tliis form is known to occur from New Orleans east to
Mobile, thence north as far as central Kentucky, Blnoxville, Ten-
nessee, and Raleigh, North Carolina, and south throughout Florida.
Besides the localities represented by the list of specimens examined,
this species is recorded by Loennburg (1894, 325) from Toronto,
Orange County, and Key West, Florida, and by Brimley (1905, 146)
from Tarpon Springs, Florida, and (1920, 108) from Rutherfordton,
North Carolina.
Variation and affinities.—It takes no great familiarity with the
snakes of the triangulum group to become aware that in elapsoides
we have a form that occupies a distinctly isolated position, struc-
turally, from all the others. Specimens from the most diverse
localities—New Orleans, Kentucky, Florida, North Carolina—pre-
serve the distinctive characters of the species with the greatest
fidelity. It apparently does not intergrade with any other member
of the group (except perhaps with its own derivative, virginianxi, in
210 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
the north.). In the vicinity of New Orleans its range appears to
overlap that of its close relative amaura, yet each form preserves its
identity.^
With the exception of Florida and the vicmity of Raleigh, North
Carolina, specimens of elapsoides are not numerous in collections.
This must be attributed to lack of collecting in other parts of its
range. Thus, although questions of geographic variation can not
&S-
Fig. 59.—Map showing locauty eecoeds for Lampbopeltis Kuapsoidms elapsoides.
be treated as they should, somethhig may be learned from a com-
parison of selected localities.
6 We may at this point, perhaps, best record a most puzzling specimen (Cat. no. 56196, U.S.N.M.)
beiongin? originaay to the Hurter collection and labeled "Florida, 1902." In size it resembles an adult
trianguJum; ventrals, 200, temporals, 2+3+4, and scale rows 19-21-19-17 also suggest triangulum (the
only other nearly related Lampropeltis m this region). Perhaps the only structural feature indicating
elapsoides is the reduced number of lower labials, eight, on each side. The pattern is most similar to that
of virginiana, but the intervals on the belly opposite the dorsal yellow bands are filled with black. There
are 18 red areas on the body and tail; the head is mostly red except for a black band across the first dorsal
scales and the posterior ends of the parietals. This specimen quite evidently fits no recognized form in
the genus nor can it be regarded as the first specimen of a form hitherto unknown, for its characters are
apparently not such as could belong to any form theoretically possible in Florida. It is not, however,
inconceivable that a young adult of triangulum might mate with a large adult of elapsoides. Such a coinci-
dence could easily explain the make-up of this unusual specimen, and to regard it as such ahybrid is per.
haps the best disposal that can be made of It at present.
REVISION OF THE KING SNAKES. 211
Reference to the accompanying table of geographic variation
shov.'s that the averages for ventral jilates are highest in the region
from New Orleans to Florida and near Kaleigh, lower at the Oke-
finokee Swam.]), and lowest in southern Florida. Tiie maxmia and
minima are correlated with the averages—the lowest number occurs
at Lemon City and the highest at Mobile. Precisely the same situ-
ation holds with regard to the infralabials, and this is closely paral-
leled by the presence or absence of the loreal. The scale rows, too,
show the same relation to locality. While the formula is corajnonly
17-19-17, it is very noticeable that 19-17 or 19-20-19-17 appears
in the extreme western portion of the range, and in the extreme
north, while in penhisular Florida nothhig higher than 17-19-17 has
been noted (except one hidividual, said to be from Marion County,
which has the unusual formula 18-21-19-17); at Lemon City 17 rows
throughout have been noted m 3 specimens, 17-15 in 2, and 15-17-15
in 1.
Table showing geographic variation in elapsoides.
Num-
ber of
speci-
mens.
Ventral plates. Loreal. Infralabials.
Locality.
Extremes. Aver-
age.
Pres-
ent.
Ab-
sent.
Extremes. Aver-
age.
New Orleans to western Flor-
ida 8
6
10
11
168-193
170-185
173-186
152-180
180
176
5 3
1
2
6
8-9
8-9
8-9
7-9
8.63
Okefinokee Swamp 8.15
Raleigh, North Carolina
Lemon City, Florida
181 1 8
172 5
8.38
7.95
It seems quite evident that the tendency to reduction in size and
scutellation has been able to proceed faster in the most isolated
portion of the range, that is, Florida. We can not doubt that the
form as it occurs here is a direct derivative of the same as it occurs
in Georgia and Alabama, for the reason that Florida is geologically
too recent to be a region of preservation of animal types. For this
reason and because the only difference between the Florida ela'psoides
and the same from the region north of Florida is that the former
attauis lower structural limits and averages than the latter, we must
conclude that evolution in this form is here proceedmg hi the direc-
tion of reduction or degeneration. When therefore we see elapsoides
in Alabama presenting such an approach and close resemblance to
a nearly related form, overlappmg in individual instances every
difference between the two, we can not but conclude that the same
processes by which elapsoides is bccomhig changed in Florida have
operated to evolve elapsoides from amaura. A study of all the
material available shows that the structural differences between them
are only such as would result from hitensification of degenerative
212 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
tendencies expressed as individual variations in amaura and other
forms of the triangulum group. Foliowmg is a summary of the most
evident differences:
Decrease in bodily size has resulted in (1) a reduction in number
of ventral plates—a change that is paralleled by several other forms
in the group and genus, the only difference being that here the de-
crease is more pronounced; (2) a slight decrease in number of caudal
scutes, resulting in a slightly greater proportionate tail length; (3)
a decrease in number of dorsal scale rows to a point that averages
even lower than the lowest individual variation in other members
of the group and genus. Accompanying decrease in bodily size the
head has become smaller and slightly different in shape; (4) the lower
labials are reduced by one in the majority of individuals; (5) the
temporals are reduced by one in each row; (6) the loreal has become
narrow and elongate, and in man}^ mdividuals it has been replaced
entirely by a downward extension of the prefrontal plate; (7) the
supraoculars have become small; (8) the rostral has become more
pointed and slightly extended beyond the lower jaw. The pattern
has suffered less marked change; (9) the paired black annuli have
been reduced by about two in correlation with the change in number
of ventrals and caudals; (10) the ringed pattern has become well
fixed and there is less widening of the yellov/ annuli on the sides;
(11) the black pigment has receded entirely from the snout, and in
most cases has become restricted to the posterior portion of the head.
That two forms, directly related, can be so distinct at the common
boundary of their ranges calls for an explanation. Fhst, it must be
recognized that elapsoides is more profoundly modified than any
other member of the group. This undoubtedly means that it is
older than the others, or that it has been isolated for a considerable
time, or both. If we assume a northeastward migration from
northern Mexico antedating that by which triangulum, sysinla, and
the others may be supposed to have reached the United States, we
have an explanation for elapsoides. If, after this form or its fore-
runner reached the southeast, an embayment from the GuK of
Mexico extended up the lower Mississippi Valley (and it is probable
that such actually occurred during the Quaternary), it might have
divided the ancestral stock into an east and a west division. The
section in the southeast might then very conceivably have undergone
a modification, due to its isolation, into what we now know as elap-
soides, whUe the main stock, still connected with its Mexican rela-
tives, was but slightly changed. When the gulf waters receded the
ranges of the isolated sections would have been joined again. It
then appeared that differentiation had been so great that intergrada-
tion seldom if ever occurred, while it was not sufficiently great to
allow either form to encroach much upon the range of the other.
REVISION OF THE KING SNAKES. 213
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REVISION OF THE KING SNAKES. 215
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216 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
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REVISION OF THE KING SNAKES. 217
LAMPROPELTIS ELAPSOIDES VIRGINIANA Blanchard.
1920. Lampropeltis elapsoides virginianaBhAyicnA-TiT), Occ. Pap., Mua. Zool., Univ.
Michigan, no. 81, p. 2 (type locality, Raleigh, North Carolina; type
specimen, U.S.N.M. no. 21163; collected June 16, 1893).—Brimley, C. S.,
Copeia, no. 89, 1920, pp. 106-109.
Since no new specimens of this form have been examined since the
publication of the original description, the latter, with the remarks
following it, is here quoted in full
:
Diagnosis.—Similar in sciitellation and proportions to L. elapsoides elapsoides (Hol-
brook), but the red areas instead of completely encircling the body are restricted to
black-bordered dorsal saddles which extend upon the ends of the ventral plates.
Fig. 60.—Map showinq locality records for Lampropeltis klapsoides vntQiNiANA.
Range.—^Northern North Carolina and Virginia, east of the Allegheny Mountains.
Type specimen.—United States National Miiseum No. 21163; Raleigh, North Caro-
lina, June 16, 1893.
Description of type specimen.—Ventral plates 183; anal single and entire; caudals
44 pairs; dorsal scale rows 19 on middle of body, 17 anteriorly and posteriorly (formula
19-17-19-17); upper labials 7; lower labials 7 on the right side and 8 on the left; 1
preocular, 2 postoculars; 1 temporal of the fu-st row, and 2 of the second; no loreal
plates, the prefrontal in contact with the second upper labial on each side; nasal
divided; anterior chin shields in contact with each other, the posterior nearly as
large, in contact with each other anteriorly, diverging somewhat posteriorly; rostral
protruding, aa in L. elapsoides; other head shields normal for the genua.
218 BULLETIN 114, U^TITED STATES I^TATIOlsrAL. MUSEUM.
Total length 473 mm.; tail length 73 mm.; tail therefore about 13 per cent of total
length. Sex, male.
The dentition is as follows: Maxillary teeth 13 on each side; mandibular teeth 14
on the left side, 15 on the right; palatine teeth 10 on the left side; pterygoid teeth 22
on the left, and 19 on the right.
Pattern of body composed of 18 dorsal saddles of red, bordered with black, separated
by whitish areas, and extending upon the ends of the ventral plates; 4 additional
saddles of red on the tail. The dorsal saddles are from 5 to 8 scales in length above,
narrowing to 2 to 4 scales on the first row. The black borders are li to 2 scales wide
on the middorsal line, and ^ to 1 scale on the first row. The whitish cross bands are
about 1| scales wide above, widening suddenly on the lower rows to 4 or 5 scales on
the first row. On the belly, opposite each dorsal whitish cross band is a large squarish
blotch of black.
On the head a black band 2 mm. in width crosses the posterior portions of the parietal
plates, leaving their tips whitish, and ending on the seventh upper labials. The
frontal and temporals are mostly black, the rest of the head mostly light, probably red
in life. The chin and throat are immaculate whitish.
Remarks.—So few specimens of this form are known that no general description can
be drawn up. It is by no means certain even that all the specimens here referred to
virginiana are conspecific with the type.
The other specimen from Raleigh (cat. no. 56197, U.S.N.M.) is almost identical with
the type in structural features, but the red saddles extend farther upon the belly.
Anteriorly their black borders are separated by only a narrow midventral strip of
whitish, while posteriorly they meet below. The specimen from Cuscowilla, Vir-
ginia (cat. no. 26181, U.S.N.M.) is a juvenile closely similar in scutellation to the
two preceding and nearly like the last in pattern, but there is a tendency to develop
black pigment on the fore part of the belly between the ends of the red saddles. This
tendency becomes more and more pronounced posteriorly, developing also opposite
the whitish dorsal cross bands, so that the latter half of the belly is nearly all black.
The specimen from Appomattox County, Virginia (Cat. no. 4466, U.S.N.M.) lacks
the head but otherwise shows itself to be almost identical with the type. The ventral
borders of the red saddles are, however, less well defined, and black pigment on the
belly is less regularly distributed.
The specimen from Alberene, Virginia (Cat. no. 25321, U.S.N.M.) shows a few
differences from the others. The twenty-first row of scales is represented by four
scales on one side of the body, the lower labials are 9 on each side, the loreals are pres-
ent, the upper anterior temporals are present, although small, and the number of red
saddles reaches 25. Furthermore, the black borders of these saddles show very little
of that widening in the middorsal region that is so characteristic of elapsoides. The
red saddles extend well upon the ventral plates and are sharply delimited by their
black borders. Otherwise there is very little black pigment on the belly. The
whitish crossbands are rather strongly mottled with darker on the sides, and all the
dorsal scales in the red areas are less strongly but very distinctly mottled with dark.
In scutellation this specimen is perhaps nearer to L. triangulum triangulum, but it
can certainly never be regarded as identical with that form, and, all things considered,
it seems much better to identify it provisionally as virginiana.
The specimen from the District of Columbia (Museum of Zoology, University of
Michigan, no. 52203) is much more puzzling. The scutellation is closely like elap-
soides, but the whole snake is larger and stouter, measming 581 mm., even with the
tip of the tail missing. Furthermore the red saddles number 27 and overlap the ends
of the ventral plates only a little. The black borders of the saddles show scant if any
tendency to widen in the middorsal region, and the head shows faint but recognizable
vestiges of the common parietal and supraocular spots of triangulum and some speci-
REVISION OF THE KING SNAKES. 219
mens of L. trianguluvi syspila (Cope). There are no lateral spots alternating with the
dorsal saddles, but anteriorly on the belly there is, as in the type, a single large black
blotch opposite each of the dorsal whitish bands. Posteriorly, however, these blotches
become irregular and the black pigment is much increased in amount. Whatever
this specimen is, it is not a Iriangulum. It may represent a derivative of triangulum
as yet unrecognized, but in view of the great variability exhibited by specimens
from this middle Atlantic region and the lack of representatives from large areas of
Virginia, Delaware, and Maryland it is not possible at present to define the characters
of this unknown form or even to be certain that such a form exists. The present plan
of identifying this aberrant specimen as virginiana is admittedly an expedient for
delaying the settlement of the difficulty until more specimens shall be available for
study. The dentition of this individual, except for the pterj'goid teeth is as follows:
maxillaries, 13; mandibulars, 13; palatines, 11.
The specimen in the Academy of Natural Sciences of Philadelphia (no. 3601) is of
uncertain origin. It may be from Delaware. It is closely similar to the Cuscowilla
specimen, and is with little doubt a virginiana.
The form virginiana, as defined by the type and represented liy all the others listed
except the two doubtful ones (those from Alberene and the District of Columbia),
is without doubt a derivative of ela-psoides by an alteration of the pattern closely
similar to that which took place when the su])species of triangulum changed from the
ringed to the saddled type of pattern. Whether this change toward the triangulum
type went so far that individuals of virginiana were able to hybridize with the degen-
erating section of triangulum of the mid-Atlantic States seems improbable. It is
more likely, as indicated above, that there exists in this region a degenerate derivative
of triangulum. This question can bo settled only when specimens shall be available
from the Delaware-Maryland peninsula, and from eastern Virginia.
Mr. C. S. Brimley has recently published notes on numerous exam-
ples of red king snakes from the vicinity of Raleigh, North Carolina
(1920, 106-109), which confirm our expectation that this is the
boundary region between the ranges of elapsoides and virginiana and
that the two forms here intergrade.
186550—21—Bull. 114 15
220 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
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REVISION OF THE KING SNAKES. 221
I.AMPROPELTIS RUTHVENI Blanrhard.
Fig. 74.
1920. Lampropdtis rulhveni Blanchard, Occ. Pap., Univ. Mich., no. 81, p. 8,
pi. 1, fig. 2 (type locality, Patzcuaro. Michoacan, Mexico; tj'pe specimen,
U. S. Nat. Mus., no. 46558; collected by E. W. Nelson, Aug. 2, 1802),
Since the t^^pe specimen is the only one known, the original descrip-
tion is quoted in full:
Diafjnosi.i.—A ringed form similar in coloration to L. polyzona Cope, L. triangnlum
nelsoni Blanchard, and L. midticincia (Yanow). From L. polyzona it (\iiierb in ih.e
complete absence of black lips on the red and •whitish scales, in the completely black
head, and in the low number of ventral plates, 189. From nelsoni it differs princi-
pally in the higher number of aunuli, 30, and the much narrower red rings, 2 to 3
scales in width. From L. multicincta it is distinguished by the low numbers of ven-
trals and annuli, by the fact that the black rings show scant, if any, tendency to
overspread the red areas dorsally, and by the mottling of the yellow rings with darker,
particularly on the sides.
Range.—Patzcuaro, Michoacan, Mexico.
Type Specimen.—United States National Museum No. 46558 (skin and head only);
Patzcuaro, ^Michoacan, Mexico; collected by E. W. Nelson, August 2, 1892.
Description of type specimen.—Ventrals, 189; anal single and entire; caudal scutes
50 plus, divided (tip of tail missing); dorsal scale rows, 21 anteriorly, 23 on middle of
of body, and 19 posteriorly (formula therefore 21-23-21-19); upper labials 8 on left
side, 7 on right; lower labials 9; 1 preocular, 2 postoculars; temporals somewhat irreg-
ular, about 2+3+4, the upper left anterior temporals much reduced; loreal longer
than high; nasals injured on each side; anterior chin shields in contact with each
other and with the first four lower laljials; posterior chin shields shorter than the
anterior and separated from each other by 2 or 3 small scales; other head sluelds normal
for the genus.
Total length (lip of tail missing), about 745 mm., tail length 112 mm.
The dentition is as follows: Maxillary teeth, 14 on the left side, 15 on the right,
the last two distinctly enlarged; mandibular teeth, 14 on the left side 13 on the right,
decreasing slightly in size; palatine teeth, 13 on the left, 11 on the right; pterygoid
teeth, 21 on the right side.
The color pattern is composed of 24 whitish rings on the body and 6 on the tail.
These rings are about 1^ scales wide on the middorsal line and 2 scales wide on the
first row of scales; on the sides and on the belly, posteriorly, they are mottled with
darker. Bordering the whitish rings are black annuli about 2 scales vide dorsally
and 1 scale vide on the belly. The black rings are separated by red rings, 2 to 3
scales in vidth. The actual colors can not be determined from so old a specimen,
but there are indications that the whitish rings may have been suffused \\ith pink.
The head ia black nearly to the tips of the parietal shields, except for flecks of whitish
on the lower portions of some of the upper labials. The chin is whitish except for
some black on the first 5 or 6 of the lower labials. The ftrst black ring begins about
2 scales behind the parietals and is continuous across the tlu:oat.
Remarks.—The status and significance of this form, represented as it is by only a
single specimen, must remain in doubt for the present. It appears, however, to be
more closely allied to L. vtuliidnda than to any other form in the genus.
222 BTJIiLETIJSr 114, UNITED STATES NATIONAL MUSEUM.
LAMPROPELTIS MULTICINCTA (Yarrow).
CORAL KING SNAKE; EING SNAKE; RED MILK SNAKE; HARLEQUIN SNAKE.
Fig. 75.
1876. Bellophis zonatus Lockington, Proc. California Acad. Sci., vol. 7, May 1,
p. 52 (type locality, northern California^).
—
Coronella zonata Boulengeu,
Cat. Snakes Brit. Mus., yol. 2, 1894, p. 202.
—
Lampropeltis zonata Van
Denbtjrgh, Occ. Papers California Acad. Sci., vol. 5, 1897, p. 167, fig.
p. 168.—McLain, Coll. Kept. W. Coast TJ. S., 1899, p. 11.—Meek, Field
Columb. MuB. Pub., 104, vol. 7, no. 1, 1906, p. 15.
—
Ophiholus zonatus
DiTMARS, Reptile Book, p. 357, pi. 103, figs. 7, 10, pi. 107 (middle fig.),
1882. Ophibolus getulus multidnctus Yarrow, Proc. XJ. S. Nat. Mus., vol. 5, p.
440 (type locality, Fresno, California; type specimen, U. S. Nat. Mus.,
no. 11753; G. Eisen, collector); Bull. U. S. Nat. Mus., no. 24, 1882,
p. 94.
1886. Coronella multifasdata Bocourt, Miss. Sci. Mex., pt. 3, p. 616, pi. 40, figs. 2,
2a-2c (type locality, California; type specimen in Paris Museum; M. de
Cessac, collector).
1900. Ophibolus getulus hoylii Cope (part). Rep. U. S. Nat. Mus. for 1898, p. 921.
1902. Lampropeltis pyrrhomelaena multicincta Stejneger, Proc. U. S. Nat. Mus.,
vol. 25, p. 153.
—
Grinnell and Grinnell, Throop Inst. Bull., no. 35,
1907, p. 39, fig. 16.
—
Grinnell, Univ. California Publ. Zool., vol. 5,
no. 1, 1908, p. 165.—Atsatt, Univ. California Publ . Zool., vol. 12, no. 3,
p. 41.
—
Grinnell and Camp, Univ. California Publ. Zool., vol. 17, no.
10, 1917, p. 184. Stejneger and Barbour, Check List, 1917, p. 89.
—
Lampropeltis pyromelana multicincta Hall and Grinnell, Proc. Cali-
fornia Acad. Sci., ser. 4, vol. 9, no. 2, 1919, p. 60.
As Stejneger has already shown (1902, 153) the Coluber (Zacholus)
zonatus of Blainville (1835, 293) is of too uncertain identity for its
name to be used for the present form. Therefore, although no
specific rule applies in this case, it is in the interest of stability to
ignore the name as revived by Lockington.
2>6Scnpiion.—Forty-nine specimens of this form have been exam-
ined. Their scutellation is as follows: Ventrals, 202 to 222; caudals,
45 to 61 (males, 45 to 61, average, 55; females, 46 to 56, average 52);
supralabials, 7, very rarely 8; infralabials, 9, rarely 10 or 8; preocu-
lars, 1, very rarely 2; postoculars, 2; temporals, 2-F3+4, infre-
quently 1 less in any row; posterior chin shields as long as anterior,
or but little shorter, in contact or separated by one or two small
scales; loreal longer than high, rarely absent; scale rows on middle of
body usually 23, sometimes only 21, rarely 25.
In proportions this form differs somewhat from the varieties of
triangulum in having the head a little more distinct from the neck,
but less so than in pyrrTiomelaena, in having a proportionately some-
what longer and blunter snout, and in having a longer tail. The latter
» Dr. .John Van Denburgh, of the California Academy of Sciences writes: " Regarding tlie types of Lock-
Ington's Bellophis zonatus I can state that they were two in number and were labeled Santa Barbara, but
probably were collected in the mountains near there, and that they were in the Academy's collection until
destroyed by the great fire in April, 1906."
REVISION OF THE KING SNAKES. 223
divided by the total length varies from 0.131 to 0.161, averaging about
0,146 for each sex. The largest specimen examined came from Cal-
lahan, California, and measm-ed 973 mm.
The pattern is formed of about 45 whitish rings separated by black
ones which are more or less completely split with red. The white
rings, 23 to 57 in number, are from 1 to 3 scales in width, not widened
on the sides nor mottled there with darker, and are complete on the
belly. The black rings are from two to several times as wide as the
white, and are usually more or less completely split with red, which is
wider on the lower rows of scales. The first black ring is usually
but not always, complete on the throat. The head is black, only
occasionally lightened in the frontal, prefrontal, and loreal region.
The dentition is indicated by a few specimens to be as follows:
Maxillary teeth, 11 to 13, commonly 12, second to fifth longest,
rest smaller, except the last two, which are blade-like and distinctly
enlarged; mandibular teeth, 12 to 14, usually 12, second to fifth
longest, becoming much smaller posteriorly; palatines, 9 to 11,
commonly 9, becoming smaller posteriorly; pterygoids, 15 to 18,
small, diminishing in size posteriorly.
This form may be distinguished from gentilis, as it occurs west of
the Rocky Mountains, by the greater number of ventral plates,
more than 200, and by the fact that the white rings do not widen
on the sides; the snout of gentilis may be red at the tip, but in
multicincta it is only black. From pyrrliomelaena it may be known
by the black instead of whitish snout, by the number of caudal
plates, not more than 61 in multicincta nor less than that number in
pyrrhamelaena, by the lower numbers of ventral plates and white
annuli, and by the infrequency of 10 infralabials. Occasional speci-
mens in which the red is reduced to practical disappearance may
resemble hoylii, but may be immediately distinguished by the totally
black snout, and the fact that the wliite rings are not particularly
mder below than above. From annulata it may be known by the
greater number of white rings, nearly always more than 30.
Habitat and Tiabits.—^According to Van Denburgh (1897, 169)
"this brilliant snake seems to prefer the moister, cooler portions of
the state [of California], such as are occupied by coniferous forests."
According to Grinnell and Camp (1917, 186) it "inhabits forest floors
and chaparral-covered hillsides." For Los Angeles County Grinnell
and Grirmell (1907, 40) say it is of "very general distribution in the
large canyons all through our mountains. In most of its range it
appears to be the commonest snake; at least it is the one most often
to attract attention. The coral king snake is a relatively small
snake, of slow movements, and perfectly harmless. Yet when
roughly handled it bites to the best of its strength. The senior
224 BULLETIN 114, UJflTED STATES NATIONAL MUSEUM.
author has allowed himself to be bitten on two different occasions
by king snakes, and although the needle-sharp teeth sank deeply
enough into liis hand to draw blood, the resulting wounds healed
promptly with no unpleasant complications whatever." ''Very
Fig. 61.—Map showing LOCAUTt eecoeds foe LAMPEOPELTia multincixcta.
little is known of the habits of this snake (Van Denburgh, 1893, 169).
Old hunters say that it destroys many rattlers and other snakes.
One of my specimens had eaten two blue-bellied Lizards (Sceloporus
occidentalis) . It is popularly supposed to be very poisonous."
REVISION OF THE KING SNAKES. 225
A specimen kept in captivity by the writer ate an adult ThamTwphis
hutleri, coiling about it in true constrictor style.
Range.—Occurs throughout the State of California except in the
deserts. A fine large specimen from Callahan, Siskiyou County
(California Academy of Sciences, no. 36062), constitutes the most
northern record; San Diego is the most southern locality. A speci-
men in the collection of the Field Museum of Chicago (no. 1426)
from Mount Whitney in Inyo County proves its occurrence east of
the Sierra Nevada. It is found at greater elevations than any other
form of the genus (Hot Springs, Long Canyon, Mount Whitney,
8,000 feet; Strawberry Valley, San Jacinto Mountains, 6,000 feet;
Lyth Crick, San Bernardino Mountains, 6,000 feet; Santa Ana
River, San Bernardino Mountains, 5,600 feet; San Jacinto Mountains,
5,400 feet; King River Canyon, Fresno County, 5,000 feet; Yosemite
Valley, 5,000 and 4,000 feet; San Bernardino Mountains, 4,000 feet).
In addition to the localities included in the list of specimens
examined. Van Denburgh (1893, 169) records Santa Barbara; Soquel,
Santa Cruz, and Glenwood, in Santa Cruz County; Hodgdon's,
Tuolumne County; Heaven's Gate, near Little Kern Lake, Tulare
County. Grinnell (1908, 165) records a specimen from Cedar Cabin,
at about 5,500 feet in the San Bernardino Mountains.
Variation.—^The different scale formulae, as shown by the table,
are numerous. Little can be said about their geographic variation,
except to note indications of higher averages in the Sierra Nevada
and lower averages in western California. But between the sexes
a decided difference is observable. Reference to the table of scale
formulae wall show that the highest formulae are not reached by
the males, and that, in this sex, there are usually only 17 rows at
the posterior end of the body, while in females there are 19; the
few of the latter that do have 17 at the end fail to reach 23 at the
middle of the body. Therefore from the scale formulae alone one
may make a close guess as to the sex.
The numbers of ventrals and annuli are likcAvise unsatisfactory
from a geographic standpoint, but the indications are in much the
same sense as the scale rows. While but little difference is noticeable
between eastern California and Los Angeles County, there is sugges-
tion of decrease in both in the northwest. And in this connection
it may be remarked that the most heavily pigmented individuals
come from the Sierra Nevada, and the least from western and north-
western California. One specimen from Santa Clara County has but
23 white rings, and the pairs of black ones are separated by wide
intervals of red. In fact individuals from this region seem generally
to have the red in complete rings, and there is sometimes a develop-
ment of red in the prefrontal region.
226 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
Scaleformulae of multidncta.
Formula.
El Dorado
to Kern
County.
Los Angeles
and vicin-
ity.
Vicinity of
San Fran-
cisco.
Other locali-
ties.
Total.
Male. Fe-
male. Male.
Fe-
male. Male.
Fe-
male. Male. JTate. Male.
Fe-
male.
23-25-23-21-19
23-21-19
21-23-21-19
21-19
23-21-19-17
21-23-21-19-17
21-19-17
19-21-19-17
Total
2
1
3
2
1
12
3
"3"
1 4
1
1 1
"i"
1
4
2
2
1
3
5
11
1
2
5
1 1 3 1 3 4
1
1
8 7 4 6 4 1 7 7 23 21
Ventrals and annuli of multidncta.
Localities.
Num-
ber of
speci-
mens.
Ventrals. White annuU.
Extremes. Aver-
age.
Extremes. Aver-
age.
El Dorado County to Kern County
Los Angeles County
15
16
5
202-222
207-218
205-210
215
213
208
37-57
44-51
23-40
46
48
35Vicinity of San Francisco Bay
Affinities.—The affinities of this form are much in doubt. It was
originally described as a subspecies of getulus. More recently
Stejneger (1902, 153) rated it as a subspecies of pyrrliomelaena. Tliis
is undoubtedly much nearer to its true relation. In favor of this
it may be noted that (1) the two forms present a striking superficial
resemblance to each other; (2) they occupy adjacent regions; (3)
a considerable proportion of the specimens of each possess the scale
formula 23-21-19-17, a formula very rare in other forms of the
genus. It is generally accepted that two forms to be rated as sub-
species must intergrade in the region of the common boundary of
their ranges. Multidncta is certainly not now known to intergrade
with pyrrhomelaena, and the differences between them are so great
as to render it unlikely that they do intergrade. Pyrrhomelaena is
much further removed in structural characters from the mean of the
group than multidncta, so that we could hardly derive the latter from
it; to derive the former from multidncta we must assume the west
coast as the center of origin of this group, but, even assuming this,
we have far too much specialization in pyrrhomelaena for the short
geographical distance that it is removed from its ancestor, and further-
more we have great differentiation in structural features with but
REVISION OF THE KING SNAKES. 227
little in pattern—a reverse of the normal situation. It is difficult
for the writer to conceive these forms as directly related.
Besides pyrrhomelaena we have to consider gentilis. The range
of the latter broadly overlaps that of the former. We do not know
how far west it extends, as it is a secretive species, and very little
careful collecting has been done in Utah and Nevada. In the western
portion of its range gentilis certainly takes on a striking resemblance
to multicincta. It must be noted, however, that it appears to undergo
a decided reduction in number of ventrals. It is perhaps not impos-
sible that on reaching California it could become modified into
multicincta, but it seems unlikely.
This would leave multicincta quite isolated, were it not for the
single specimen found by E. W. Nelson at Patzcuaro, Michoacan,
Mexico, and named by the writer, ruthveni. This specimen appears
to be very closely allied to multicincta, yet distinct from it. The
discovery of confirmatory specimens would be gratifying.
228 BULLETIN" 114, UNITED STATES NATIONAL MUSEUM.
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REVISION OF THE KING SNAKES. 229
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230 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
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REVISION OF THE KING SNAKES. 231
LAMPROPELTIS PYRRHOMELAENA (Cope).
ARIZONA RINGED SXVKE.
Figs. 5, 71.
1866. Ophibolus pyroinelanns Cope, Proc. Acad. Nat. Sci. Philadelphia, p. 305
(type locality. Fort Whipple, Arizona).
1875. Ophibolus pyrrhomclas Yarrow, Geog. Geol. Explor. Surv. W. 100th Mer.,
vol. 5, chap. 4, p. 537, pi. 19.—Cope, Bull. U. S. Nat. Mus., no. 1, 1875,
p. 37.—Yarrow, same, no. 24, 1882, p. 91.—Cope, Proc. U. S. Nat. Mus.,
vol. 14, 1891, p. 610; Rep. U. S. Nat. Miis. for 1898, 1900, p. 907, fig.
224.
—
Lampropeltis pyrrhomelas Van Denuurgh, Proc. California Acad.
Sci., ser. 2, vol. 6, 1896, p. 347.
1875. Ophibolus pyromelas Coues, Geog. Geol. Explor. Surv. W. 100th Mer.,
vol. 5, chap. 5, p. 619.
1883. Ophibolus gelulus pyromelanus Garman, S., Mem. Mus. Comp., Zool., vol.
8, no. 3, pt. 1, pp. 67, 157.
1901. Ophibolus zonatiis Brown, Proc. Acad. Nat. Sci. Philadelphia, p. 79.
—
DiTMARs, Reptile Book, 1907, pi. 103, fig. 6, pi. 107 (upper fig.).—Tucker,
Dang. Pois. Snakes U. S., 1912, p. 1, pi. 1, fig. F.
1902. LampropeUis pyrrhomelaena Stejneger, Proc. U. S. Nat. Mus., vol. 25,
p. 152.
—
Van Denburgh and Slevin, Proc. California Acad. Sci., ser.
4, vol. 3, 1913, p. 415.
1917. Lampropeltis pyrrhomelaena pyrrhomelaena Stejneger and Barbour,
Check List, p. 88.
Description.—The scutellation, based upon 32 specimens, is as
follows: Ventrals, 216 to 235; caudals, 61 to 79; supralabials, 7,
occasionally 8; infralabials, 10, sometimes 9, rarely 11 or 12; a single
preocular; 2, rarely 3, postoculars; temporals, 2 + 3 + 4, occasionally
an extra scute in any row; posterior chin shields usually shorter and
narrower than the anterior, and generally separated from each other
by 1 or 2 small scales; loreal distinctly longer than high; scale rows
23 or 25 on middle of body, formulae varying from 23-25-23-21-19 to
21-23-21-19-17.
Body long, slender, varying but little in diameter; sides meeting
belly in a fairly well defined angle; head distinct from neck, widest
at the temples; snout rather long, tapering, and almost truncate; eye
proportionately larger than in other forms of the genus; tail long and
slender, from 0.153 to 0.182 of the total length (males, 0.162 to 0.182;
average, 0.172; females, 0.153 to 0.176, average, 0.167). The longest
specimen examined came from the Huachuca Mountains in southern
Arizona and measured 1067 mm.
The pattern (fig. 71) is of 35 to 71 whitish annuli on body and tail,
separated by black which is more or less completely split with red.
Some specimens are ringed throughout with white, red, and black, in
others the red is continuous across the dorsal line only at the anterior
end of the body and near the anal region. The red widens at the
expense of the black on the lower rows of scales and usually crosses
the belly. The black becomes much narrower on the sides, and
traverses the bellv or sometimes fails to reach it. The whitish annuli
232 BULLETIN ]14, UZsTlTED STATES NATIONAL MUSEUM.
are from ^ to 2|, usually 1 to 2, scales in width, not wider on the
sides, and usually more or less obstructed on the belly with black.
The head is black from about the anterior portion of the frontal
to the posterior part of the parietals and laterally to the eyes or
middle upper labials. The whole snout is whitish, and there is some-
times a light spot above each eye; the chin and gular region is white
and quite or nearly immaculate. A white band crosses the tips of
-o
ss
Fig. 62.
—
Map showing locality records for Lampropeltis pyeehomelaena.
the parietals, widening anteriorly on the sides; behind this is a black
band, which narrows on the sides and is rarely, if ever, complete on
the throat.
The dentition is as foliovvs: Maxillary teeth, 13 to 20, commonly
14 to 16, somewhat longer anteriorly, the last two a little stouter and
more flattened laterally; mandibular teeth, 14 to 18, second to sixth
largest, becoming much smaller posteriorly; palatine teeth, 11, less
REVISION OF THE KING SNAKES. 233
often 12 or 13, rarely 14, becoming slightly smaller posteriorly;
pterygoids, 16 to 22, smaller than the palatines, likewise diminishing
in size posteriorly.
The penial characters are approximately as follows: Organ small
and slender in comparison with size of animal; calyces few, with
fringes few and short; latter succeeded by slender spines, which
increase in size only slightly; about halfway from the base the latter
decrease rapidly in size and disappear entirely, or remain over most
of basal portion of organ as minute spines more or less imbedded.
This form may be distinguished from gcntilifi, multicincia, and
annulata by the prominently whitish snout, the greater number of
ventrals and caudals, the more numerous annuli, the head distinct
from the neck, and the frequent occurrence of 10 infralabials. All
but the first two of these points will serve to distinguish it from
nchoni.
Habitat and Jiahits.—From a few brief notes accompanying speci-
mens and from the localities it seems that this snake prefers the
canyons and mountains, and is not found at §,11 in the deserts. Van
Denburgh and Slevin (1913, 415) record three specimens from pine
woods m Bear Canyon in the Catalma Mountains, in Ramsey Canyon,
Huachuca Mountains, and Oak Creek Canyon in Coconimo County,
Arizona. One from the Hualapai Mountains was also taken in a pine
forest.
Range.—Pyrrhomelaena is known to occm* from the Great Salt
Lake south through Utah, Arizona, and western New Mexico to
Sonora and Chihuahua. Careful collecting may be expected to
extend its range in all directions. The vertical range of this form is
comparable, so far as we now know, onh^ with that of multicincia. The
following altitudes are recorded with specimens: 7,000 feet—Bear
Canyon on Mount Lemon in the Catalina Mountahis, Arizona, and
Beaver Canyon, Beaver County, Utah; 6,300 and 5,800 feet
—
Hualapai Mountains; 5,410 feet—Carr's llanch. Sierra Ancha, Gila
County, Ai'izona.
No records have been noted for localities not included in the list
of specimens examined.
Varlalion and affvniiics.—The present material indicates that this
form is remarkably constant throughout its range, but a fuller series
of specimens may be expected to reveal geographic differences that
can not now be detected.
234 BULLETIN" 114, UNITED STATES NATIONAL MUSEUM.
The distribution of the scale formulae between the sexes is of
interest. It is as follows:
Formula. Male. Female. Total.
23-25-23-21-19
23-21-19
21-23-21-19
23-21-19-17
21-23-21-19-17
Totals
2
7
4
5
2
3
3
3
1
5
10
7
6
2
20 10 30
Only 1 female out of 10 ends with 17 rows. A striking feature in
which this form resembles multicincta and in which both apparently
differ from all other forms in the genus is the possession by many of
the same individuals of a maximum of 23 rows and a minimum of
17. Three changes from the middle to the posterior end of the body
is common when the maximum is 25 rows, but is elsewhere very rare
when the maximum is 23, and is quite unknoAvn when the maximum
is 21 or 19. In this feature there is certainly support to the propo-
sition that pyrrliomelaena and multicincta are more closely related to
each other than to any other form of the genus. In other respects,
however, the gap between the two seems to be rather wide. Pyrrlio-
melaena is in every way a more specialized form. Its scale rows
average higher—the maximum in no individual yet examined being
lower than 23; there are more ventrals; the tail is the longest in the
genus; the lower labials are commonly 10, the upper labials are
oftener 8 than in any other form in the genus; the head compares in
shape and in distinctness from the neck only with alterna; the denti-
tion is the highest for the genus.
All these features proclaim this form an old and specialized deriva-
tive of some more normal representative of the genus. While we do
not say that it can not have been derived from multicincta, it is never-
theless difficult to conceive of such great structural differentiation
where the geographical separation is so little and where the habitat
preferences of the two seem to be so similar. It is not impossible
that the relationships of multicincta and gentilis may yet prove to be
so close as to exclude the former from the possibility of direct rela-
tionship with pyrrliomelaena. More material from the Great Basin
region is the first essential. We would therefore prefer at present to
regard pyrrliomelaena as a relatively ancient and isolated type,
because of (1) its high specialization, (2) its great constancy through-
out its wide range, and (3) the lack of any form with which it may
safely be considered as closely related.
REVISION OF THE KING SNAKEG. 235
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KEVISIOF OF THE KING SNAKES. 237
SUMMARY.
Included in this group, on the basis of intergradation along the
common boundaries of their ranges, are, nelsoni, annulota, gentilis,
amaura, syspila, and triangulum; by reason of close similarity wdth
the lii"st two, polyzona and micro2)holis; elapsoides and virginiana on
account of close resemblance to amaura; because of apparently
closer relationsliip to this than to any other group in the genus;
rt'tliveni, multicincta, and pijrrliomelaoM.
As already shown, micropliolis can be derived only from polyzona.
The most evident reasons for this are (1) it is closely allied to tliis
form and to tliis one only; (2) it is decidedly isolated geographically
from all other forms of the genus; (3) it is inadmissable as a primi-
tive type on account of specialization in important structural and
color pattern characters, particularly (a) in reduction of anterior
temporals to one, (&) in the frequency of entire caudal plates, and
(c) in the white rings of the pattern being much widened and suffused
with red. Polyzona must be regarded as closely allied to annvlata
and particularly, perhaps, to nelsoni, but its exact relationship is
uncertain, due to insufficient material.
The evolution of elapsoides from amaura has already been explained
imder the discussion of the former, and we need not do more here
than repeat our belief that it represents an earlier and less extensive
migration than that which gave rise to the subspecies of triangulum;
that it underwent differentiation and specialization through an
isolation which may have been affected by a not very extensive
embayment of the Gulf of Mexico up the lower ^lississippi Valley.
Its close ally, virginiana; was differentiated from elapsoides by a
joining of the adjacent black rings on the belly across the red spaces,
and their lateral migration, thus restricting the red to wide dorsal
saddles. Tliis, as vdW be shown below, is almost exactly the same
method as that by which syspila was derived from amaura.
If now we consider the subspecies of triangulum, we fuid tliat the
structural differences, separating any two adjacent forms are only
average and so quite insufficient to distinguish them, and even ex-
treme forms are but moderately distinct in structure. The differ-
ences are thus too slight to be a main reliance in determining the
center of origin of the series. This structural similarity is, however,
of much value in assuring us of the close genetic relationships exist-
ing between all these forms, and, if we are able by other means to
determine direction of relationship, structural tendencies will then
be of value in either strengthening or weakening those conclusions.
Aside from the geographic demands which must be met, our main
reliance must be upon color pattern. This, as well as the scutella-
tion, demonstrates the close relationship of the whole series and in par-
238 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
ticular of any two adjacent forms, but, as the differences registered
in the pattern are more evident and more distinctive than any others,
these have been made the main basis of distinction between differ-
ent forms. However, near the bomidaries between any two ranges
Fig. 63.—Map showing distribution of the various forms of the Triangulum group.
even these distinctions break down and we find individuals which
can not be definitely assigned to either form. This completes the
proof furnished by the structural features of the close relationship
existing between all these forms.
REVISION OF THE KING SXAKI5S. 2B9
The races of triangulum form a continuous chain from southern
Mexico to southeastern Canada. Some form in the series must be
closer to the ancestral type than all the others. Let us consider
the northern end of this chain.
If we start with typical triangulum, as it occurs in western New
York, and examine a series of representative specimens from localities
between here and Central Florida, we will notice a very gradual
change in pattern in Pennsylvania, West Virginia, and southward;
the half collar behind the neck very frequently replaces the chevron
mark; the latter, when present, is more imperfectly shaped than in
the north, and the other characteristic head markings are often
very badly formed. Throughout this wide extent of latitude we
will fuid no distinctive change in triangulum and no closely allied
form—nothing closer than elapsoides, getulus, and rhomhomaculata.
If we now examine a representative series of specimens from northern
Indiana and Illinois southward into Louisiana, we will find a very
different situation. In southern Illinois and southern Indiana
triangulum intergrades perfectly with syspila. The latter is spotted
like triangulum but the spots are typically longer, wider nnd fevvcr;
normally they are red, but occasional examples have them brown
or gray as in triangulum. The head pattern of tlie latter (fig. 54)
rarely appears in full in syspila (fig. 52) but portions of it are usuidly
present, as, for instance, the anterior black border of the half collar
behind the neck is sometimes indented, indic;;ting the position of
the arms of the chevron mark of triangulum; the half collar may
be more or less interrupted by the approximations of its black borders
on one or both sides of the median line; light superciliary spots witli
black borders are often present over the eyes, and these are some-
times joined, as in triangulum; the light postocular band is often
represented by lateral indentations of the anterior black border of
the half collar and sometimes it is present in entirety. As we go south
and west from southern Illinois the frequency and perfection of these
markings decreases. To one who has examined considerable num-
bers of these specimens the geographic change is most striking.
Typical triangulum (figs. 54 and 69) occurs about as far south as
Vigo County, Indiana; at this point there is a relatively rapid change
through intergrades to syspila (figs. 52 and 68); then south into
Arkansas we find a gradual, progressive change which leads directly
into amaura (fig. 67), in the southern part of the state. That syspila
is the only close relative of trianyulum, and that the former passes
by perfect intergradation into amaura, can not be denied. One of
these forms must then be ancestral to the others.
It has been said that the color pattern of trian^gulum must be re-
garded as primitive, because its distinctive features are repeated in
other forms in widely separated groups, as, for instance, the calli-
240 BULLETIN lU, UinTED STATES NATIONAL MUSEUM.
gaster group of the present genus, the genus Elai^lie, in particular, and
numerous other genera. This, however, is an assumption that must
be proved. Simplicity and specialization in color pattern can not be
decided on a priori grounds. It is not safe to say that because a
certain type of pattern is primitive or specialized in one or several
genera that it is so in any given case. Close examination may bring
out reasons why the rule does not hold in a particular instance.
Thus we have a spotted pattern in Drymolius margaritiferus and in
the three forms of the getulus group, Jloridano,, lioTbrooTci, and spleiv-
dida. In Drymolius there is good reason for considering this pat-
tern speciahzed, namely, as belonging to an end form. As these
are all closely allied and all different, they are probably not all
specialized, or, if so, some are less specialized than others, jn the
case oi jloridana geographic probabilities alone are enough to place
its pattern as recent, and as derived from something else, even if we
failed to note that it still bears the vestiges of an earlier pattern;
and careful study of geographic probabilities and structural and pat-
tern differences have shown that the pattern of hoTbrooJci is a deriva-
tive of that of splendida, and that the pattern of the latter is the only
one from which aU the others in the group may be derived. Thus
we would have been in error had we said in the first place that because
the pattern of splendida looked similar to that of Drymolius that it
was specialized and not a suitable starting point for the derivation
of other patterns. A type of pattern therefore that is specialized in
one form may prove, for reasons quite evident upon examination and
comparison, to be primitive in another.
In the case of triangulum there are numerous real difficulties in
the way of considering this form as primitive. It should be noticed
that (1) the pattern is best and most perfectly developed only in a
region that can not have been a center of preservation of reptilian
life, the glaciated portion of the northeast; (2) in the southeast, from
West Virginia and Maryland to Florida, it is erratic and not typical;
(3) it is much less stable and widespread than the ringed type of pat-
tern, and in the central Atlantic States it is undergoing degeneration
in company with structural reduction; (4) no other form of the
triangulum group presents any approach to this pattern except syspih,
the nearest relative of triangulum; (5) if triangulum be considered the
stem form of the group the explanation of elapjsoides becomes ex-
ceedingly difficult. It is well known that the pattern and color of
some snakes change diuring development from young to adult, and
the young stages in some cases are supposed to indicate ancestral
conditions. This is the case with the racer, Coluler constrictor,
which changes from a spotted to a plain black or bluish snake
during growth from young to adult. In the young of conjuncta the
body pattern can not be told from that of loylii, but that of the
REVISION OF THE KING SNAKES. 241
adult is quite distinct. Conjuncta is in all probability a derivative
of hoylii. A similar situation probably holds with respect to hoi.
broolci, and probably also with several other forms of this genus-
The probability is therefore strong that the fact that the young of
triangulum have red instead of brown blotches indicates that the
ancestor of tnangvlurn was a red instead of brown spotted snake.
(6) Jf we have any regard for the evidence derived from the getulus
and calligaster groups that marked reduction in structural features
is evidence of specialization, we can not ignore the fact that triangu-
lum is the most reduced form in this whole series. This reduction is
expressed particularly in the great frequency of the scale formulae
21-19-17 and 19-21-19-17 and the large number of specimens that
possess a temporal formula lower than 2 + 3 + 4, i. e., 2 + 3 + 3, 2 + 2 + 3,
1+2 + 3.
if we accept these reasons for not regarding triangulum as ancestral
we are led next to syspila. But this form does not possess the pat-
tern that was alleged to be primitive in triangulum. In scale rows
and temporals it shows the same degenerative tendencies that are
more strongly expressed in triangulum. Tn fact, there appear to be
no good reasons for considering sysjnl/i- as primitive. This leads us
directly to amaura. This form is restricted to the lower Mississippi
Valley—a region geologically new and therefore totally unsuited to
be a center of origin of a group of animals, Gentilis could scarcely
be regarded as an ancestral type as its color pattern is very evidently
a derived one, it is a reduced form in number of ventrals, and it
inhabits a region entirely unsuited for preservation of old types.
The only other possibilities are the Mexican forms nelsoni and
annuhta, and for present purposes it is immaterial which of these
may be the oldest. Nelsoni is less likely to be primitive because it
inhabits chiefly the coastal area, and it presents a derived appear-
ance in the extreme width of the red interspaces. There are for
annulata, however, several positive reasons for regarding it as the
ancestral type of the triangulum series. (1) In structural features
it presents the closest approach to the primitive t3^pes of the other
large groups: The loreal is oblong with no peculiarity in shape;
the scales rows are 23 or 21 (no specimens drop to 17 at the
end); the temporals are 2 + 3 + 4; the labials are 7 and 9; the chin
shields are normal; the proportions of head, body, and tail are
average; the ventral plates average but slightly less than for splen-
dida, (2) It occupies a geographically favorable region, the Mexi-
can plateau. (3) It is in a region inhabited b}^ the ancestors of the
getulus and calligaster groups. (4) It is in a region sho^Mi to have
been the center of dispersal for another widespread and recent group
of North American snakes, the garter snakes. (5) The color pattern
of all the forms of triangulum may be most simply derived from that
242 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
of annulata. (6) Stillmoreimportant is the fact that annvlata lies at the
point of convergence of three separate genetic series of the triangulum
group : The triangulum-syspila-amaura-annulata series, the virginiana-
elapsoides-amaura-annulata series, and the gentilis-annulata series.
This situation fulfills therefore the most convincing of the criteria
formulated by Adams (1902, 122) for determination of centers of dis-
persal, namely, "continuity and convergence of lines of dispersal."
If we accept annulata as the stem form of the triangulum series we
may describe the derivation of the color patterns as follows: From
the plateau region of southern Mexico north to extreme southern
Texas we have a form, annulata (fig. 66) , ringed with black and white
or yellow, the black more or less split with red dorsally but continuous
between the white rings on the belly, except at the extreme anterior
end of the body, where the red may cross the belly. The white rings
are nearly uniform in width on the dorsal scales and only slightly, if
any, widened on the ventrals; they are not encroached upon by the
black, but may be more or less mottled with black, particularly on
the sides. Figure 66 shows this pattern as exhibited by a typical
specimen from Puebla, Mexico. Passing to amaura (fig. 67), we see
that the white rings have become distinctly widened on the lower
rows of scales, that the black on the belly has become separated on
the midventral line, and that the red has increased in extent at the
expense of the black. The black has here ceased to encircle the body,
and has become restricted to a border for wide dorsal saddles of red.
Figure 68 is a typical example of syspila from- Arkansas. This pat-
tern is derived from the last by a lateral contraction of the dorsal
red saddles, a narrowing of the black borders, the development of a
ventro-lateral series of black spots in alternation with the dorsal
blotches, a checking of the belly with small quadrate spots of black,
and an increase in the mottling of the white dorsal scales (no mottling
was shown on the white scales of figure 67 because the specimen was
much faded). The alternating spots and perhaps some of the small
quadrate spots may be the result of a pinching off, as suggested by
figures 67 and 68. Figure 69 shows the pattern of a typical triangulum
from northern Michigan. Whether the upper series of alternating
spots is the result of an upward migration of the first series (fig. 69)
is uncertain, but it appears much more likely that it is due to a pinch-
ing off of segments of the dorsal blotches in the course of their con-
traction. This is the end of the series in this direction.
To derive gentilis we must return to annulata. The course of evo-
lution has here been different. Instead of a lateral shrinking of the
black and red, there has been a longitudinal shrinking. The begin-
ning was the same as for the other series. The white rings widened
on the lower rows of scales and on the ventrals, thus greatly increasing
the width of the white rings in proportion to the black on the belly.
^7.—Lampropeltis Triangulum amaxhia (M.C.Z.,
I, Dallas, Texas). About IJ x nat. size. A
OR PATTERN OF THIS FORM (THE BLACK AND RED,
;R, OFTEN COMPLETELY ENCIRCLE THE BODY).
lIRIANGULUM TRIANGULUM (U.S.N.M.,
coMMfY. Michigan). About ij xnat.size.
HCt^-
(G. 73.-LAMPROPELT1S ELAPSOIDES ELAPSOIDES 'ICINCTA (STAN . UnIV., NO. 4309, FYrFE, E
L-
(U.S.N.M., NO. 23807, Raleigh, NORTH Caro- a). About li xnat.size. Typical pa t-
una). About ij x nat. size. Typical color
pattern.
186550—21. (To face page 242.)

70
-LamPROI hLTIS MKltOl llOIJb (AtAD NAT
^ci PmuDLLPuu NO 18211 Ecuador)
Abocijxhat. size. Typical cotOR pattern
w KcuADOBjiN Colombia th:
1NTERVA13 ABE NARROWER.
i'lG 71 —LAMPROPELTIS PlRRHOiIELAE^A (U fa N M ,
51-126, Graham Mountains, Arizona). About :
SIZE, Ttpicalco
Ki\\\>}{S'
^^\%m^
FIO. OO.-I-AJ
NO. 01700, KUUKTCquNTV, MiailUAN).
Typical c<
Fig. 73.—I-4«:
(U.S.N.M., so. 2
UNA). ABOUT 1)
fATTERN.
Flo. 74.—LAHPBOPELIU BUTUVENI (U.SJJ.U,, MO. 4«US, PaTZCUAKO,
North Caeo- MicHOACAN»UExtco). About IJXNat.mw. OOLOBPATTEBWorTYPt
Typical colos specdieii.
Fio. 76.—LAMmoi'Ei.TiBiiULTiaKori(fti*i*. Univ.,
OOEADOCOUIirY.CALlPOBKU). ABOUT li X NAT.
TEEN.
I8A&SO-31. (To raco page 313.)
•
REVISION OF THE KING SNAKES. 243
Here the black failed to split on the midline, or, if it did split, it failed
to recede. The red failed to increase in area at the expense of the
black, so that in this form the black is usually wider on the middorsal
line than on the lower rows of scales and often excludes the red en-
tirely from here, as shown in figure 72 (a specimen from Provo,
Utah). Mottling of the white rings is variable in gentilis, but usually
present to some degree at least.
Examination of the available material indicates that the split in
these two courses of evolution is not a clearly defined one, but that
the tendency to develop the pattern shown in figure 68 is gradually
intensified to the east. A fairly well-marked division is, howevei',
noticeable in the region of the ninety-seventh meridian. Here there
is a distinction in head pattern as well as in body pattern. To the
east the head is mostly red, and alternating spots are developed on
the lower rows of scales; to the west these are not developed and the
head is mostly black. The situation would be met if the ancestral
form spread from northeastern Mexico at the same time northeast,
north, and northv/est.
The relationships of fyrrliomelaena and midticincta to each other
and to the other forms of the genus are still in doubt. There is little
if anything in the penial characters to ally them closely with the
getuluR group. It is true that the calyces are but few and slightly
fringed, but the spines are much more numerous and more slender,
and extend more than half way to the base of the organ, as is char-
acteristic of the triangulum series. Muliicmcta was described by
Yarrow as a subspecies of getulus and more recently has been regarded
by Stejneger and others as a subspecies of vyrrhomelaena.
Cope confused it with both of these forms. It can hardly be a
close ally of hoylii, since their ranges overlap and their structural
differences are too great, particularly in respect to scale rows, penial
characters, and color pattern. Its relation to gentilis may be closer
than has been supposed. Typical patterns of gentilis and multicincta
(figs, 72 and 75) show that the white rings in gentilis are but little
widened in the most western specimens, perhaps an approach to
multicincta in which they are practically uniform in diameter; and
it is noticeable that in the latter form, as in gentilis, the black on the
belly is mostly concentrated o])posite the dorsal red areas. The
color pattern of multicincta could without violence be regarded as an
intensification of that of gentilis; that is, the black has still further
encroached on the red and has completely excluded the latter color
from the snout. The scutellation would be considered as a wide, but
perhaps not impossible, step. Thorough collecting in Nevada, eastern
California, and Arizona should be of much assistance in deciding this
matter. Scale rows and ventrals average at least as high as may be
expected for annulata in the plateau portion of its range, and higher
244 BULLETIISr 114, UNITED STATES NATIONAL MUSEUM.
than for any of the subspecies of triangulum, except nelsoni, of the
west coast of Mexico. In proportions, dentition, and scalation it is
so close to the average for the genus that it is difficult to conceive of
it as much specialized or as a derivative of pyrrliomelaena. In fact,
for regarding it as a subspecies of the latter there is no evidence.
We do not know whether their ranges meet, but the specimens on
hand show no approach of one form to the other in pattern or scala-
tion. It is even possible that muUicincta may be only distantly
related to pyrrliomelaena. Its color pattern is similar, to be sure, and
their ranges are practically adjacent, but if pyrrliomelaena did not
exist we would probably not be unwilling to regard muUicincta as the
west coast representative of the subspecies of triangulum. The
discovery of the Patzcuaro specimen, indicating, as it does, the occur-
rence in southern Mexico of a close relative of muUicincta, puts a new
face on the situation. Nothing definite can be said, however, until
more specimens of the new form, ruthveni, shall have been found.
Pyrrliomelaena gives much evidence of specialization in (1) its
high number of labials, (2) its aty])ical pattern, (3) long tail, (4) high
dentition, and (5) wide head. If we examine its color pattern
(fig. 71) we notice that while there is the same marked tendency for
the black to overspread the red dorsally, the former narrows decidedly
on the sides and often fails to reach the ventrals—a decided difference
from gentilis and muUicincta; and the black of the belly, contrary to the
situation in the latter two forms, is concentrated chiefly opposite the
dorsal white rings, and it is the red that most regularly crosses the
belly. It is difficult to say how important this cUstinction may be.
The prominent specializations in structural features indicate old age
in the form and show that there may have been time for the pattern
to develop these peculiarities starting from the annulata condition.
The suspicion can not be avoided, however, that the present simi-
larity in the patterns of muUicincta and pyrrliomelaena are due to
convergent evolution.
It is noticeable that muUicincta is less removed structurally from
the average of the genus than pyrrliomelaena, as it is less removed in
style of color pattern from gentilis and annulata.
On the whole it would seem best to place these two forms with the
triangulum group on the basis of similarity in general style of color
pattern—that is, rings of red, yellow, and black—and to regard ^jyr-
rliomelaena tentatively as an isolated form, at least as old as elapsoides,
and muUicincta as the isolated representative on the Pacific coast of
the rest of the triangulum group.
The preceding discussion may be summarized as follows:
1. Micropliolis must be regarded as a specialized derivative of
polyzona; the relationship of the latter to nelsoni and annulata is
close but not clear.
KEVISTON OF THE KING SNAKES. 245
2. Elapsoides is a derivative of amaura, or of the form that repre-
sented it in the Texas region before the appearance of syspila and
triangulum. In the northeastern portion of its range it has given rise
to the color pattern variety, virginiana.
3. The subspecies of triangulum may all be traced to a center of
dispersal in the plateau region of Mexico.
4. The relationships of pyrrhomelaena and multicincta are doubtful,
but there seems to be the least difficulty in regarding fyrrliomelaena
multicincta
gentilis /"
triangulum
jn/rrhomelaena
nelsoni
\
syspila
gentilis—^amaura
annulata
mrgimana
elapsoides
ruihveni polyzona
\
micropholis
Fig. 76.—a Diagrammatic Presentation of the Rel.*.tionships of the Forms of the Trungclum
Group.
as a specialized and isolated form, much older than the subspecies
of triangulum, and multicincta as the west coast representative of the
rest of the triangulum group.
ISOLATED FORMS.
LAMPROPELTIS MEXICANA (Garman).
Fig. 77.
1883. Ophibolus trinngulus, war. mexicanv^ Garman, S., Mem. Mus. Comp. Zool.,
vol. 7, no. 3, p. 66 (type locality. Mexico, near San Luis Potosi; cotypoa,
probably numbers 4652 and 4653 of the collection of the Museum of
Comparative Zoology at Cambridge, Massachusetts).
1902. Coronclla mericana GtJNTHER, Biol. Cent.-Amer., p. 110.
No type was designated in the original description of mexicana,
and no specimens were listed. The description shows, however,
that there were at least two specimens, and it fits remarkably well
for two from that locality in the collection of the Museum of Com-
parative Zoology. The scales of both are in 23 rows; the ventral
plates are exactly 193 and 199, as in the origmal description; the
head is "much swollen at the temples;" and the pattern and colora-
tion answer well. According to the description, the caudals are
56-58 pairs, and the dorsal blotches are 38-40; these specimens
seem to have 55 pairs each of caudals, and about 39 blotches each.
These differences when compared with the coincidences are too
246 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
slight to render it improbable that these are the specimens from
which the original description of mexicana was prepared, and then*
numbers have therefore been given above as the cotypes of this
form.
Description.—Since the original description is somewhat incomplete,
the following is offered in its stead : Ventrals, 193 and 199 ; caudals, 55,
in two rows (both specimens females); supralabials, 7; infralabials
10, fifth largest, 5 under the last 3 upper labials; preocular single
except that it is divided in number 4652 on the left side; postoculars,
2; temporals, 2 in the first row, 3 in the second, and 4 or 5 in the
FiQ. 77.—Lampropeltis mexicana (M.C.Z., scale pattern from no. 4653, color pattern from no.
652, San Luis Potosi, Mexico). About 1| x nat. size. Showing color pattern.
third; posterior chin shields shorter than the anterior, and separated
from each other by 2 small scales; loreal about twice as long as high;
anal plate entire; scales smooth, with 2 apical pits; dorsal scale
rows, 23-21-19.
Body moderately slender, tapering slightly toward the tail; belly
flat, meeting the sides in a rather distinct angle; head flat, distinct
from the neck, swollen at the temples, tapering anteriorly, snout
truncate; tail, 0.16 to 0.17 of the total length; eye large, its diameter
nearly twice the height of the third upper labials. One of the speci-
mens measures 803 millimeters, the other 452.
The pattern of the body (fig. 77) recalls that of triangulum. There
are about 39 transversely oblong red blotches on head and tail
REVISION OF THE KING SNAKES. 247
bordered with black, and separated by narrower grayish mottled
spaces. The red blotches are about 2 to 3^ scales long, and they
maintain this length through a width of about 8 scales across the
back, then narrowing, they extend down on the sides to about the
second or third row of scales. These downward extensions are some-
times nearly or quite isolated from the dorsal blotches. The black
borders are i- to 1 scale in width doreally, becoming narrower on
the sides. The spaces between the blotches are whitish, and strongly
but minutely mottled with dark, except close to the black borders.
The belly is heavily blotched with black; the tendency is for narrow
transverse bands to cross the belly and end on the first row of scales
in alternation with the dorsal blotches, but these are often broken
in the middle and obscured by large black patches between them
and opposite to the dorsal blotches. On the head of the small
specimen are two red V-shaped marks black-bordered, opening
forward, the larger chiefly on the parietals, the smaller chiefly on
the frontal. Behind the eye is a large black blotch. Rest of head,
chin, and throat whitish, minutely mottled with darker.
The dentition is as follows: Maxillar^^ teeth, 13, the last two
enlarged, not grooved nor separated from the rest by an interspace;
mandibular teeth, 14 in one specimen (number 4652) and 16 and 17
in the other, the third, fourth, and fifth large, the last small; pala-
tines, 13 in each instance; ptery^goids, 22 (only one set counted).
Remarks.—If, as is entirely probable, the two specimens described
above are the originals, then they are the only specimens of the form
known. That they belong to the genus Lampropeltis as at present
defined, must be conceded. They can not, however, be closely related
to any other form in the genus; the head pattern is unique, and,
although the body pattern bears a superficial resemblance to trian-
gulum and to the members of the caUigaster group, it yet bears a
distinct stamp of originality; the 10 lower labials, long tail, swollen
temples, and high number of palatine teeth are all features that
mark it as specialized. It is doubtless more closely allied to the
tnangulum group than to either of the other two.
LAMPROPELTIS ALTERNA (Brown).
Fig. 78.
1902. Ophibolus altemus Brown, Proc. Acad. Nat. Sci. Philadelphia for 1901
(Feb. 6, 1902), p. 612, pi. 34 (type locality, Davis Mountains, Jeff Davis
County, Texas; type specimen, number 14977, Acad. Nat. Sci., Phila-
delphia, E. Meyenberg, collector); same, 1903, p. 550.—Ditmars,
Reptile Book, 1907, p. 356.—Strecker, Baylor Bull., vol. 18, no. 4,
1915, p. 39.
—
Lampropeltis altema Stejneger and Barbour, Check
List, 1917, p, 87.
This name rests upon a single specimen, received alive at the
Zoological Gardens in Philadelphia, and said to have been found in
248 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
the Davis Mountains, Texas. It answers so closely in structural
features to the genus Lampropeltis that, in spite of its unique type
of pattern, and lesser structural peculiarities, it must be included
here until further specimens make possible a determination of its
true status.
Description.—Since the original description is very accurate it is
quoted in entirety
:
Maxillary teeth 13; mandibular 1-^15. Body moderately slender; head distinct,
muzzle contracted; eye rather large. Rostral low and broad, barely visible from
Fig. 78. -Lampropeltis alterna (Acad. Nat. Sa., Philadelphia, no. 14977, type, Davis Mounta i.> s»
Texas). 2 X nat. size. Showing sttle of color pattern.
above; intemasals about half the length of prefrontals; frontal a little longer than the
suture between parietals, longer than the snout; pariotals large, wide in front, narrow
behind; nasals 2, the nostril between them; loreal small, longer than high; preocular
1; postoculars, 2 on one side, 3 on the other; temporals, 2+3 on one side, 3+4 on the
other; upper labials 7, third and fourth in orbit; lower labials 11. Posterior chin
shields a little shorter than the anterior, not separated by scales. Scales smooth,
with two inconspicuous pits, in 25 rows. Ventrals 217; anal entire; subcaudals 60
pairs. Total length 710 mm. (tail 115).
The ground color is slate gray, crossed on the back, at intervals of 3 to 5 scales, by
bands of black which are alternately wider and narrower, the wide ones covering from
2 to 3 scales on the middle of the back, and more or less divided transversely on their
centers with scarlet. The narrow bands are about 1 scale wide and wholly black,
occasionally broken through by the ground color. On the neck the bands are narrower
REVISION OF THE KING SNAKES. 249
and less defined, while the red is more pronounced on the posterior part of the body.
There are 19 red and black bands on the body, and an equal numlicr of the inter-
mediate black ones. On the tail* there are 5 bands, which form quite distinct rings,
on the last two of which the red is absent. The head, including the labials, is dark
gray with small dark mottlings, not well defined, and a narrow black streak from the
postoculars to the angle of the mouth. Ventral surface grayish white, heavily blotched
with black, into which the black portion of the cross bands runs.
By way of correction and addition to the above it may be noted
that the temporals are 3+4 + 5 on each side; the left lower labials
are 11, the right, 10; the third postocular is on the left side and is a
derivative of the fourth supralabial ; the last two maxillary teeth are
slightly enlarged, and the palatines are 12 on each side; the dorsal
scale formula is 25-23-25-23-21-19-20, and the changes in the
number of rows take place in the manner usual for this genus ; the
tail is about 0.17 of the total length.
Remarks.—It will be evident from the above description that this
specimen is about as far removed in its structural features from the
normal forms of the genus as is pyrrhomelaena. Like the latter it
has a long tail, wide head, high numbers of ventrals, caudals, scale
rows, temporals, infralabials, and a long, narrow loreal. Its style
of coloration is, however, quite different from anything else in the
genus, but knowing how easily one pattern may be changed into
another radically different in appearance, we can not assign great
importance to that fact alone. The writer would agree with Brown
(1901, 613) in placing it nearer to pyrrJiomelaena in structural features
than to any other form of the genus, but it does not appear to lie at
all near to leonis, as that author suggested. It would seem host to
await the finding of more specimens before making any definite
statement as to its status.
CONCLUSION.
The preceding descriptions and discussions have brought out the
fact that the genus Lampropeltis is naturally divided into three m.ain
groups, of closely related forms (exclusive of two forms of doubtful
relationships, mexicana and altema). Two (the GETTJIUS and
CALLIGASTER groups) are more closely allied to each other than
either to the third, and the latter (the TRIANGULUM group) is
composed of at least three minor groups, representing different
degrees and kinds of differentiation, and different periods of dispersal
.
On account of these facts and because the most primitive forms of
the groups are apparently very far from being directly related, each
group has been treated very nearly independently in searching for
its center of dispersal, for it is conceivable that the groups as now
known may have started from different centers without affecting the
propriety of uniting them all in a single genus. This treatment has
resulted in showing that in all probability each of these groups origi-
250 BUULETIN" 114, UNITED STATES NATIONAL MUSEUM.
nated in some portion of the region between Texas and Nicaragua.
This, together with the fact that the only forms of doubtful relation-
ships are located in this region, indicates that the center of dispersal
of the entire genus is in the Southwest.
Let us now consider how the genus as a whole answers to the criteria
for determination of centers of dispersal, as formulated by Adams
(1902, 122). There are 10 of these criteria, as follows:
1. Location of greatest differentiation of a type.
2. Location of dominance or great abundance of individuals.
3. Location of synthetic or closely related forms (AUen).
4. Location of maximum size of individuals (Kidgway-Allen).
5. Location of greatest productiveness and its relative stability,
in crops (Hyde).
6. Continuity and convergence of lines of dispersal.
7. Location of least dependence upon a restricted habitat.
8. Continuity and directness of individual variations or modifica-
tions radiating from the center of origin along the highways of
dispersal.
9. Direction indicated by biogeographical affinities.
10. Dii'ection indicated by the annual migration routes, in birds
(Palmen).
Some of these criteria are of only limited value, and the fifth,
ninth, and tenth can not be used at all in the present instance. The
others wiU be discussed in order.
1. Location of greatest differentiation of a type.—In the Southwest
we have polyzona, nelsoni, annulata, leonis, calligaster, splendida,
pyrrhomelaena, ruthveni, and alterna. The only region at aU com-
parable with this in diversity of type is the Southeast. Here we
have four forms, all specialized, and two of them {elapsoides and
rJiombomaculata) obvious derivatives of western types; but there is
no representative of pyrrhomelaena in the Southeast, nor of mexicana,
nor of alterna. The greatest differentiation is therefore imquestion-
ably in the Southwest.
2
.
Location of dominance or great abmndance of individuals.—^This
criterion is of only minor value; exceptions may be readily called to
mind. It is valueless in this case, however, since, in the present
unsatisfactory state of our knowledge, there is as much to be said
on one side as on the other.
3. Location of synthetic or closely related forms.—It has been noted
frequently that those forms of this genus to which the groups trace
their origin are more closely allied in structure with each other than
with any of the other forms in the genus. For example, the south-
western types, calligaster and leonis may, much more readily than
the specialized rhomhomaculata of the Southeast, be associated with
the GETTJLTJS group, and it is with the western representatives of
REVISION OF THE KING SNAKES. 251
the latter rather than with the eastern that the closest relationship
lies. Of the TRIANGULUM group the outlying types. micropJiolis,
elapsoides, virginiana, and tnanguhim, are much more differentiated
from the other groups than its southwestern representatives, poly-
zona and annulata.. This criterion is therefore admirably fulfilled
by the Southwest, and by this region only, and its value as a criterion
may be regarded as second only to the sixth,
4. Location ofmaximum size of individuals.—Apparently no greater
size is obtained by any forms than that reached by the soutliwostern
representatives of all the groups, with the exception of the purely
peninsular and derived forms, jioridana and hrooTcsi; and it will be
remembered that the forms which are most reduced in size and
scutellation occupy regions most distant from the southwest, namely
conjuncta of the Cape region of Lower California, and elapsoides,
and rhombomaculata of the southeastern States. Of ndsoni and
annulata we have too few specimens to know what the normal or the
maximum size may be but polyzona of southern Mexico may properly
be compared with triangulum of the northeastern States. Of 300
specimens of the latter the largest measured 1,085 mm., and of 61 of
the former the largest measured 1,610 and the next largest 1,580
mm., a decided difference in favor of the Southwest. The forms of
the GETULUS group are of approximately equal size; of the two
well-known forms of the CALLIGASTER group, the western repre-
sentative is decidedly the larger. This criterion therefore points
definitely to the Southwest.
6. Continuity and convergence of lines of dispersal,—Both the
GETULUS and CALLIGASTER groups are represented in the
Southwest, and the isolated types, mexicana, alterna, and pyrrhome-
laena, are not represented outside of the Southwest. But it is the
TRIANGULUM group that illustrates this most admirably. In this
we see the convergence toward Mexico of the polyzonamicropholis
line from the south, the annulatorgentilis line from the north, the
annvlata-elapsoides line from the east, and the annvlata-triangvlum
line from the northeast.
All genetic lines in the genus, therefore, either converge to the
Southwest, or are represented in the Southwest as well as elsewhere,
or ai'e not represented outside of the Southwest, and no genetic
lines converge to any region other than the Southwest. Thus this
most dependable of all criteria is satisfied only by the Southwest.
7. Location of least deperulence upon a restricted habitat.—Too little
is known of the habitat relations of these snakes to discuss this cri-
terion profitably, but it may be noted that the only burrowing forms
(elapsoides, virginiana, and rhombomaculata) are located in the South-
east.
186560—21—BuU. 114 17
252 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
8. Continuity and directness of individual variations or modifications
radiating from the center of origin along the highways of dispersal.—
Lengthy demonstration of this criterion is not necessary here, since
the descriptions and summaries demonstrate conclusively its direct
applicability to the Southwest.
It will be evident that although these criteria are not of equal
value, all point to the Southwest, and that the most dependable of
these, the third, sixth, and eighth, are especially definite in this
respect. The argument for the Southwest is not expected to rest
upon any single piece or line of evidence, but it is believed that the
weight of positive and the lack of negative evidence, as brought out
in the summaries of the separate groups and in the discussion of the
preceding criteria, amount to a demonstration that the center of
dispersal of the genus Lampropeltis is in the southwestern portion of
the North American continent.
Before concluding, a few remarks on the relation between the
environment and the distribution of species and subspecies, as
exhibited by the genus Lampropeltis, may not be out of place. It is
noticeable that each form inhabits a region of rather definite environ-
mental conditions, and that, within any such region, a group of
directly related forms has but a single representative. Thus, re-
stricted to the southeastern States, we have three forms, each repre-
senting a different line of descent. Each of these forms is replaced
west of the Alabama-Mississippi region by a closely allied form, and
the different environmental conditions of southern Florida are re-
flected in these forms as follows: One apparently does not extend
south of the northern portion of the State, one is replaced by a closely
allied form derived from it, and the third expresses an extreme re-
duction that may yet lead to subspecific differentiation. The prairie
region is characterized by two forms (calligaster and syspila), the
lower Mississippi Valley by one (amaura), and another (holhrooJci)
ranges over both of these areas, but does not extend to any adjacent
region. Two characterize the northeastern deciduous forest province ;
one of these (niger) is confined to the southern section, west of the
Alleghenies, the other (triangulum) inhabits the whole province, but
has developed most successfully in the north. At the western limit
of forests in Texas, both hoTbrooTci and amaura are replaced by their
close relatives, splendida and gentilis, respectively. In Mexico the
major environments are represented by polyzona in the lowlands of
the east and south, nelsoni on the west coast, and annulata in the
plateau region. Whatever may be the cause of speciation, or the
breaking up into subspecies, it is a fact that when a form has mi-
grated into a region of decidedly different environment it has become
altered, and the alteration has remained constant in its main features
REVISION OF THE KING SNAKES. 253
throughout that environment. Thus it has come about that the
direct relative of any form is found in an adjacent environment.
The peculiar exception offered by California e (p. 94), where specia-
tion by mutation is indicated, affords opportunity for a most inter-
esting investigation. But in general, the oft emphasized rule holds
for this genus; namely, the occurrence of two different forms of the
same genus in the same locality is prima facie evidence that they
belong to different lines of descent, and, conversely, directly related
forms are not found in the same locality.
The preceding might be detailed more fully, and more complete
information on the little known forms would doubtless result in
added illustrations of the fact that the environment seems to exert
a circumscribing influence on the species, that is, the range of the
species is determined by the environment, and, that when the latter
changes, the species is altered or dies out.
The obvious differences in the relative ages of certain forms and
groups of forms in the genus Lampropeltis, as shown by specializa-
tion, structural isolation and diversification, geographic position, etc.,
leads naturally to speculation as to the time relations of their spread
from the Southwest. Since triangulum is typically developed only
north of the southern limit of glaciers, we may assign it to a post-
Pleistocene migi'ation. The range of syspila and the northern por-
tion of the range of gentilis were doubtless likewise largely uninhabi-
table during the glacial epoch, and tliis together with the perfect
intergradation between these forms indicates that they belong to the
same "wave" of dispersal. The high specialization of elapsoides
and its distinctness from its nearest relative argues for isolation, and
greater age than that of triangulum and syspila. These demands
are satisfied by assuming it to have reached the Southeast at some
favorable time during or before the ice age, and to have been isolated
by an embayment of the lower Mississippi Valley during the Pleisto-
cene (see map in Willis and Salisbury, 1910, 277). The high degree
of specialization and structural isolation of pyrrJiomelaena suggests
that it is as old as any form in the genus. As it certainly has no
relatives outside of the Southwest, it may be regarded as indigenous
to this region, and its structural isolation is doubtless due to an early
differentiation from the primitive stock. In the CALLIQ-ASTER
group the apparently sharp separation and the specialization of its
eastern representative, make natural an assignment to a period
similar to that when elapsoides reached the Southeast. Consequent
separation during a greater or less portion of the Glacial Epoch
induced divergent differentiation. The GETXTLUS group may be
considered nearly or quite as recent as the varieties of triangulum,
with, perhaps, greater age indicated for the western representatives
254 BULLETIN 114, UNITED STATES NATIONAL MUSEUM.
to allow them to reach Cape San Lucas and Santa Catalina Island.
The Central American forms are less easy to account for. Nelsoni
assumed its specific identity before the Tres Marias Islands were
separated from the mainland. This is likely to have been pre-
Pleistocene. Polyzona shows too great variability to be easily
regarded as an ancient type, but it is old enough to have produced
the well-marked micropTiolis in Colombia and Ecuador, and is there-
fore without doubt older than the northern varieties of triangulum.
The preceding paragraph may be summarized as follows: The
genus Lam'pro'peltis underwent a differentiation in the southwestern
portion of the North American continent in the late Tertiary, pro-
ducing a type that favored the mountains and uplands, the ancestor
of the TRIANGUIUM group, and another type that favored the low-
lands, ancestor of the GETULUS and CALIIGASTER groups. The
latter early differentiated into the progenitors of the two last groups,
and of these it was the CAIIIGASTER section that first spread north-
east and east. This must have occurred previous to the Pleistocene,
and at about the same time that the migration took place which
brought the progenitor of elapsoides to the southeast. During the
Pleistocene, the latter and the ancestral type of rhombomaculata
suffered an isolation from their direct relatives west of the present
position of the Mississippi River and were thus free to undergo con-
siderable differentiation. Following the Pleistocene, elapsoides ex-
tended its range south into Florida, there undergoing still further
specialization during the Recent Epoch, and west to Louisiana,
here meeting the form amaura, the post-Pleistocene representative
of its ancestral stock. If rliombomaculata extended its range into
southern Florida, it is yet to be found there. Calligaster, after the
retreat of the glaciers, was able to extend its range throughout the
prairie region northeastward, and north to the limit of its endurance
of cold. The western section of the GETTJITJS group may have
begun its evolution during or perhaps preceding the Pleistocene, but
it was only following that period that the spread east and northeast
occurred. And it was following the Ice Age, also, that the latest
forms of the TRIANGULUM group reached their present ranges
{amaura, syspila, triamgulum, and at least the northern section of
gentilis)
,
This historical outline corresponds with the present distribution
and structural relations of the king snakes, and with probable Pleisto-
cene geography. The genus as a whole seems to be relatively recent,
but the question of its origin is beyond the limits of the present study.
In this connection, however, we may say that there are apparently
no nearly related genera in South America, in the West Indies, nor
in Asia, and that it is entirely likely that its nearest relatives a.re to
be found in the genus Coronella of Europe and northern Africa.
REVISION OF THE KING SNAKES. 255
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INDEX.
[The black-faced numbers indicate specific headings.)
Adder, spotted, checkered 188, 1S3
alterna 5,6,7,17,234,247,249,250,251
aitcrnus 247
amaura 162,169,172,179,181,
184,185,202,210,211,212,237 et seq., 252,25-i
amaurus 172
annulata 142,
140, 155,156, 157,159, 169, 172, 175,
176, 223,233, 237 et seq., 250, 251, 2.i2
annulatus 159, IfiS
anomala 6, 139
arciCera 140
Arizona ringed snake 231
balteata 7j
Bastard bom snake 49
Bellophis 4
zonatus 4
Black and white king snake 75
Black moccasin 49
boylei 75
Boyle's king snake 75
boylii 12,66,68,69,73,75, 89 et seq., 103,
104 et seq., 163,222,223,240,241,250 et seq.
brooksi 4,57,58,66,105,113,115,251
Brown king snake 128
califomiae 6, 58, 91, 103, 105, 115, 163, 253
califomica 04
calligastcr 5, 7, 10, 15, 115, 128,
132, 133, 134, 138, 139, 250, 252, 254
calligastcr group 6,
17, 107, 108, 138, 241, 247, 249 et seq.
Caudals, variation of, in genus 15, 24
celacnops 165
Cemophora coccinca 207
Centers of dispersal, criteria for 250
Chain snake 49
Checkered adder 188
clerica 190
clericus 188, 190, 201
coccinea 172, 206
coccinea, Coronella 4
coccineus 140, 155, 172, 179,206, 207
collaris 179, 201
Color pattern (see ofso description of eachform) . 16, 24
Coluber constrictor 240
Colubridae 5
conjuncta 5, 12, 15, 66, 73, 76, 77, 89,95, 103
,
114, 115, 159, 163, 240 et seq., 251
conjunctus 89, 159
constrictor, Coluber 240
Coral king snako 222, 223
Corallilo 139,142
Coral snake. 206
Coronella 5, 254
coccinea 4
186550—21—Bull. 114- -18
Page.
Cowsucker 49
Dental characters, value of 16
Dentition {see also description of each form) ... 7, 25
doliata 139,165,179,189,206
doliatus 172, 173, 179, 189, 190, 202, 206
Drymobius margaritiferus 240
eiseni 94
Elaphe:
guttata 10
laeta 10, 1 16, 117, 122
obsoleta 10
clapsoidea 173, 206
elapsoides 5, G, 8, 10, 11, 12, 15, 10, 163, 172, 173,
176, 200, 206,219, 237, 250, 251, 253, 254
elapsoidcus 206
evansii 116, 117
Evans king snake 115
Evolution in scutellat ion 11
eximia 188
exiinius 116, 188
floridana 5, 11. 15, 51, 55, 57, 58, C2,
60,105,113,114,115,240,251
formosa 139, 140, 149
Formulae of scale rows 10-12
gentilis. 7, 159, 161, 1G2, 163, 105,172, 174, 175, 176, 181,
184, 185, 223, 227, 233, 234, 237 ct seq., 251, 252, 253
getula 33,49,75,94
getulus 12, 33, 36, 38, 43, 44, 45, 46,49,
62, 64, 80, 104 et seq
.
, 226, 243
getulus group 6, 11, 13, 16, 17,26, 119,
241,243,219 et seq.
map of 104
relationships of 115,132
summary of 104-115
getulus, Herpetodryas 4
C roups of king snakes 17
Guinea snake 33
guttata, Elaphe 10
Harlequin snake 222
Herpetodryas getulus 4
holbrooki 7,12,28,29,30,88,43,44,46,51,
57, 80, 104 et seq., 163, 240, 241 , 252
Horse racer 49
House moccasin 188
House snake 188
Key to forms of Lampropeltis 18
King snake 26, 33, 36, 49, 66
Labials, variation of, in genus 13-15, 24
laeta, Elaphe 116,117
Lampropeltis 4
leonis 138,249,250
Loreal, variation of, in genus 15
figure of 5
Maps. See under description of each form,
margaritiferus, Drsrmobius 240
259
260 INDEX.
Page.
Master snake 49
mexicana 6,7,17,245,249,250,251
mexicanus 245
microphoUs 5, 6, 11, 15, 139, 140, 142, 143,
145, 146, 149, 155, 163, 200, 237 et seq., 251, 254
Milk- snake 75,188,193
Moccasin, House 188
Molecatcher 128
multicincta 7, 11, 106, 168, 221, 222,
226, 227, 233, 234, 237, 243, 244, 215, 249 et seq.
multicinctus 222
multifasciata 222
multistrata 165, 202
multistratus 155, 165
multistriata 165
Mutation 99, 100
Natrix doliatus 206
nelsoni 142,143,145,146,155,
162, 163, 172, 221, 233, 237 et seq., 250, 251, 252
niger 12, 38, 39, 43, 51, 58, 104 et seq., 252
nitida 92,96,100,103,105,115
obsoleta, Elaphe 10
occipitalis 159
oligozona 140, 155
Ophibolus 4,
5
Osceola 4
parallelus 172, 173
Penial characters (see also description of each
form) 7
Penis:
in caUigaster group 17
in getulus group 17, 108-109
in Lampropeltis 7
in triangulum group 17
Pituophis sayi 34
polyzona 11, 139, 149, 151, 152, 155, 156,
161, 162, 163, 221, 237 et seq., 250, 251, 252, 2.54
polyzouus 139,140,149
Pseudoelaps Y 188
pseudogetulus 75
pyromelanus 231
pyromelas 231
pyrrhomelaena 5, 6, 7, 11, 15, 163, 166, 223,
226, 231, 237, 243, 244, 245, 249, 250, 251, 253
pyrrhomelas 231
Rattlesnake pilot 49
Red king snake 206
Red milk snake 179, 222
Red snake 179
Ring snake 222
rhomboniaculata 5,8,
12, 15, 117, 119, 120, 124, 128^
138, 139, 163, 250, 251, 254
rhombomacalatus 116, 128
Page.
rathveni 221, 227,237,245,250
sayi 25,33,34,43
sayi, Pituophis 34
Say's king snake 33
Scale rows:
formulae of 10-12,25
variation of, in genus 9
Scarlet kind snake 206
Scotophis 116
Sex, to distinguish 6
Speckled king snake 33
Sphenophis 4
splendida 7, 26, 35, 38, 39, 44, 66
67, 68, 69, 73, 83, 104 et seq., 240, 250, 252
spendidus 26
Spotted adder 188
Spotted king snake 33
Summary:
calUgaster group 138
characteristics of genus 24, 25
general summary 249
getulus group 104
triangulum group 237
syspila 168,
169, 172, 173, 175, 176, 1 79, 184, 192, 194, 197,
198,200,201,202,212,237 et seq., 252,253
syspilus 179
sysputus 165
Tail length, proportionate 24
temporalis 190,202
Temporals, variation of, in genus 15, 24
Thamnophis 8, 9, 10
Thunder and lightning snake 49
Thunder snake 49,206
tigrina 116
triangula 188, 189
triangulum 6, 7,
11, 15, 163, 168, 175, 181, 183, 184, 185, 188,
210, 212, 219, 237 et seq., 251, 252, 253, 254
triangulum group 1,6,
13, 16, 17, 108, 139, 247, 249-254
map of 238
summary of 237
triangulus 188,189,190
Ventrals, variation of in genus 15, 24
virginiana 5, 8, 10, 15, 172, 200, 209, 217, 237, 251
Wamper 49
Wampum snake 49
Yellow-bellied king snake 115
Y, Pseudoelaps 188
yumensis 7, 28,
29, 30, 66, 76, 77, 80 et seq., 99, 103, 104 et seq.
zonata 222
zonatus 139, 222, 231
zonatus, Bellophis 4
o