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FIELDIANA . ZOOLOGY

Published by
CHICAGO NATURAL HISTORY MUSEUM

Volume 34 DECEMBER 29, 1955 No. 34

CORAL SNAKES OF THE GENUS MICRURUS
IN COLOMBIA

KARL P. SCHMIDT

CHIEF CURATOR EMERITUS, DEPARTMENT OF ZOOLOGY

More than thirty years have passed since the Hermanos Nice'foro
and Apolinar Maria, of the Institute de la Salle in Bogota, Co-
lombia, began their correspondence with various American museums,
enlisting the aid, for the identification of Colombian animals of all
kinds, of zoologists in museums of natural history in both North and
South America. On various occasions Chicago Natural History
Museum has been consulted, especially with regard to bats and
venomous coral snakes of the genus Micrurus from Colombia.
Occasional specimens collected by Hermano Nice'foro Maria have
been retained for our collections, while other identified material has
been returned.

When I engaged in renewed studies on coral snakes in 1951, my
colleague, Philip Hershkovitz, was engaged in zoological field work
in Colombia and was in close contact with the Museo de His-
toria Natural de la Salle, of which Hermano Nice'foro Maria
is director. On the occasion of Mr. Hershkovitz' return to the
Museum from his Colombian mammal survey, the available col-
lections of coral snakes in the Museo de la Salle were packed up and
sent along with him. These specimens nearly double the number of
specimens I have been able to examine from Colombia and this
accumulated material, amounting to 201 specimens, is the occasion
for the present paper. I am indebted to Dr. Robert Mertens, Direc-
tor of the Senckenberg Natur-Museum, for the loan of specimens of
Micrurus dissoleucus for re-examination. I am further indebted to
the John Simon Guggenheim Memorial Foundation and to the
individuals and museums that have so generously entertained me
in the past, and from whom I have borrowed specimens for the
present study. I have examined specimens from the following
institutions and private collections: the American Museum of Nat-
ural History (AMNH), the British Museum (Natural History) (BM),

No. 782 337 THF ll8R*tfY OF THF

338 FIELDIANA: ZOOLOGY, VOLUME 34

the Museum of Comparative Zoology (MCZ), the Academy of
Natural Sciences of Philadelphia (ANSP), Carnegie Museum of
Pittsburgh (CM), and the Emmet R. Dunn collection (ERD).

Much of the material new to me is recorded by Hermano Nice"foro
Maria (1942). There are gratifyingly few differences between his
list and the present one. His map shows the localities named.
H. Daniel (1949) has also listed these species, mainly following
Amaral.

Any brightly colored snake with a pattern of red and black or
of red, yellow, and black, in sharply defined but variously arranged
rings, is commonly referred to as a culebra cor&l or simply as a coral
in Latin American countries. Such snakes in Colombia include
various harmless species of the families Anilidae and Colubridae.
The snakes of the genera Micrurus and Leptomicrurus are the only
types of the venomous Elapidae in South America. Leptomicrurus
narducci barely enters Colombia in the Amazonian headwaters;
Micrurus, however, ranges throughout Colombia with a bewildering
diversity of forms.

The snakes of the genus Micrurus in Colombia are distinguished,
of course, from those harmless snakes that have a similar bicolor
or tricolor ringed pattern, by the presence of the enlarged, venom-
conducting fang on the maxillary bone. This is normally the only
tooth on the maxilla, but when the replacement tooth persists there
may be two fangs side by side. If a living snake is to be determined,
the venomous Micrurus lack the loreal plate on the side of the snout
and never have the black rings in pairs.

The fifteen species and twenty subspecies are the following:

Micrurus mipartitus mipartitus Micrurus ornatissimus

mipartitus semifasciatus ancoralis jani

nigrocinctus mosquitensis dissoleucus dissoleucus

carinicaudus carinicaudus dissoleucus melanogenys

carinicaudus antioquiensis dissoleucus nigrirostris

carinicaudus transandinus lemniscatus

dumerili filiformis

ecuadorianus sangilensis spixi obscurus

nicefori surinamensis surinamensis

psyches hemprichi ortoni

A tentative key for the identification of these species and sub-
species has been drawn up. The difficulties arising in the cases of
subspecies not sharply defined will be discussed under the headings
of the species carinicaudus and dissoleucus.

SCHMIDT: COLOMBIAN CORAL SNAKES 339

1. Black rings in well-defined groups of three, these "triads" separated by
red zones 7

Black rings not in triads 2

2. A broad red band across anterior part of parietals, usually across posterior
tip of frontal shield 3

Top of head black, usually covering most of parietals, with or without a narrow
light band across snout 4

3. Black rings very numerous (41 to 76), the light spaces between them (red in
life, often yellowish or white in preserved specimens) usually narrower than

the black rings Micrurus mipartitus

(See text for distinction of subspecies.)

Black rings few, widely separated by much broader red rings.

Micrurus nigrocinctus mosquitensis

4. Top of head with two narrow light crossbands, one across snout, one across

parietals Micrurus nicefori

No light crossband on snout; parietal light band usually bordering pari-
etals 5

5. Red zones usually much wider than black Micrurus carinicaudus

(See text for distinction of subspecies.)

Red zones, if distinguishable, about equal in length to black 6

6. Yellow rings reduced to spots (i.e. to half-scales adjacent to the black rings) ;
anterior head shields, at least the supraoculars, each with sharply defined

light spot Micrurus ornatissimus

Yellow rings continuous; red rings, if distinguishable, greatly obscured by
invasion of dark pigment Micrurus psyches

7. Anal plate entire Micrurus hemprichi ortoni

Anal plate divided 8

8. Head with a narrow light crossband on snout and a broader one across

parietals 9

No crossband on snout; parietal crossband present or absent 10

9. A moderately slender snake, ventrals 220-263, triads of black rings 8-14

(9-11 in Colombia) Micrurus lemniscatus

Body very slender, ventrals 266-321, triads 11-20 (av. 16) . .Micrurus filiformis

10. Head with black pileus, a narrow light post-parietal band broadened on the

temporals and posterior labials (as in carinicaudus) 11

Head variously colored; if black, the scales sharply outlined with light. . . .12

11. Outer black rings of triad much narrower than middle one. . Micrurus dumerili
Outer black rings of triads subequal to middle one.

Micrurus ecuadorianus sangilensis

12. Top of head light, the upper head scales all sharply outlined with black;

frontal shield very narrow Micrurus surinamensis surinamensis

Top of head not light, with dark bordered head shields; frontal normal in
shape 13

13. Head with broad light crossband across posterior ends of supraocular and
frontal shields and anterior ends of parietals; snout black; nuchal ring not

part of first triad Micrurus dissoleucus

Top of head without well-defined light crossband 14

14. Yellow rings very wide, as wide as outer black rings of the triads, top of head

dark, the shields with light outlines Micrurus spixi obscurus

Yellow rings narrower above than outer black rings of triads; head light
anteriorly, with black spots Micrurus ancoralis jani

340 FIELDIANA: ZOOLOGY, VOLUME 34

In this key I have avoided the adult male character of presence
or absence of supra-anal tubercles, feeling that the student or
teacher with a single female or juvenile coral snake at hand would
be hopelessly frustrated if faced with this alternative. Nevertheless,
this character so greatly sharpens the taxonomic distinction of
sangilensis, psyches, and ornatissimus (which do not have it) as
compared with dumerili and carinicaudus, that it must again be
pointed out that the determination of this character for each species
is essential before it can be brought into the system. This feature
is unknown for Micrurus nicefori. The tubercles are present in
adult male mipartitus, but not developed to the extreme of carini-
caudus or nigrocinctus.

LIST OF SPECIES
Micrurus mipartitus Dume'ril and Bibron

Elaps mipartitus Dumeril and Bibron, 1854, Erpet. Gen., 7: 1220 — Rio Sucio.
Micrurus mipartitus Ruthven, 1922, Misc. Publ. Mus. Zool. Univ. Mich.,
8:68.

Diagnosis. — A coral snake with alternating black and red bands,
the black bands wider, a broad red band across the posterior part
of the head; a tendency to very high numbers of ventrals (up to
326); tail short, caudals 20-38; supra-anal tubercles present in adult
males, mainly post-anal.

Discussion. — The fortunate circumstance that Dume'ril and
Bibron named Rio Sucio as type locality of Elaps mipartitus greatly
simplifies the problems of nomenclature for this species, which
exhibits great variability and some obscure geographic partition
into subspecies. The ventrals in this species range from 214 to
326; such a wide variability is to be expected in a character that
tends to the extreme — and is not yet phylogenetically fixed. The
extremes of variation are much less in geographically limited regions;
but the range of variation is nevertheless so great as to prevent
any very accurate analysis into subspecies. A Central American,
a Colombian, and a Colombo- Venezuelan subspecies may ten-
tatively be recognized :

Ventral plates 234-260 in males, 263-274 in females, Nicaragua to Panama.

mipartitus mullifasciatus

Ventral plates 244-289 in males, 251-326 in females, Colombia west and south
of Santa Marta mountains mipartitus mipartitus

Ventral plates 214-239 in males, 233-260 in females, Santa Marta region eastward
to Caracas, Venezuela mipartitus semipartitus

SCHMIDT: COLOMBIAN CORAL SNAKES 341

Micrurus mi par t i t us mipart it us Dume>il and Bibron

Micrurus multiscutatus Rendahl and Vestergren, 1940, Ark. Zool., 33A, no. 1,
pp. 1-16.

The typical subspecies ranges from Darien, Panama, to western
Ecuador, and the populations thus far known from the Cauca and
Magdalena valleys are only feebly distinguished from the trans-
Andean ones. The variation in ventrals is as follows:

No. of
specimens Ventrals Average

Trans-Andean d" 10 259-289 270

9 9 253-326 297

Cauca Valley <? 5 244-273 259

9 8 280-325 296

Magdalena Valley d* 2 246-265 255

9 5 251-282 264

Santa Marta region d* 6 224-234 231

9 8 250-260 256

Venezuela tf 1 214-239 228

9 9 233-258 247

The extent of variation in Micrurus mipartitus was not under-
stood by Rendahl and Vestergren (1940), who established a supposed
new species, M . multiscutatus for specimens from El Tambo in the
Cauca Valley, based on three specimens with ventrals "more numer-
ous than in any known species of Micrurus." The ventrals in their
types and cotypes number 295, 328, and 329. The range of ventrals
in eight female specimens from the Cauca Valley examined by myself
is from 280 to 325, and from Colombia west of the Cordillera Oc-
cidental (nine specimens) from 253 to 326. The low counts within
these ranges are quite uniformly bridged over to the highest by
intermediate ones. The El Tambo specimens do represent the
extreme of high ventrals in mipartitus; but a distinct form for the
upper Cauca Valley is not discernible; if there should be a subspecies
in this area, the name mentalis, Boulenger 1896, would be available
for it.

Micrurus mipartitus semipartitus Jan

Elaps semipartitus Jan, 1858, Rev. Mag. Zool., 1858: 113.

The tabulation above shows that the variation in ventrals in the
Santa Marta mipartitus comes much closer to that of the Venezuelan
form than to that of central and western Colombia. I do not have
at hand the material of this species of the Museo de la Salle, and

342 FIELDIANA: ZOOLOGY, VOLUME 34

without much additional material it is profitless to examine further
into the variation of mipartitus.

Micrurus nigrocinctus mosquitensis Schmidt

Micrurus nigrocinctus mosquitensis Schmidt, 1933, Field Mus. Nat. Hist.,
Zool. Ser., 20: 33 — Limon, Costa Rica.

Diagnosis. — A subspecies of nigrocinctus characterized by much
lower numbers of ventrals in both sexes than are to be found in
nigrocinctus nigrocinctus. The species nigrocinctus as a whole is
allied to the Colombian carinicaudus in its pattern of widely spaced
black rings with narrow yellow rings bordering the black, in having
the sexes sharply distinct in tail length, and the adult males with
well-defined supra-anal tubercles.

Range. — The Caribbean slopes of Central America from Costa
Rica to northwestern Colombia.

Colombian specimens. — A male specimen from Turbo, Antioquia,
MLS no. 602, collected in May, 1943, forms the sole record for the
nigrocinctus group in Colombia. This specimen agrees excellently
with the original description of mosquitensis; it has 195 ventrals;
52 caudals, of which 2 are entire; and 18 black rings on the body,
with 5 on the tail.

Micrurus carinicaudus Schmidt

Micrurus carinicauda Schmidt, 1936, Field Mus. Nat. Hist., Zool. Ser., 20:
194 — Orope, Zulia, Venezuela; Niceforo Marfa, 1942, Rev. Acad. Colomb.
Ciencias, 5: 98.

Diagnosis. — A coral snake with black rings not in triads, with
narrow yellow borders often not discernible in preserved specimens,
the red zones always longer than the black; top of head black, no
red crossband on the anterior portion of the parietal shields; sexes
sharply different in tail length, adult males with strongly developed
supra-anal tubercles.

Range. — From the Maracaibo Basin eastward as far as Caracas,
westward to the Magdalena Basin, and southward along the central
Andes to the latitude of Bogota.

Remarks. — The species carinicaudus is the common coral snake
of northern and central Colombia. The three subspecies, c. carini-
caudus, c. antioquiensis, and c. transandinus, are not sharply definable,
and though their intergradation has not yet been traced, their
ranges are expected to meet in the northern lowlands of Colombia.

SCHMIDT: COLOMBIAN CORAL SNAKES 343

Micrurus carinicaudus carinicaudus Schmidt

Diagnosis. — Distinguished from the two western subspecies by
its higher number of black rings and in having the nuchal ring
present and complete. In Magdalena Valley intergradient spec-
imens the number of rings may be high, with much irregularity
of the nuchal ring.

Range. — Northwestern Venezuela to Santander Norte, Colombia,
with intergradient specimens from Olanche, Landazuri, Sasaima,
and Charala in Cundinamarca.

Colombian specimens. — Specimens from Bucaramanga, Gram-
alote, Catatumbo, Chimacote, and Cucuta are unequivocally
assignable to the typical subspecies. The 17 specimens from Cundi-
namarca agree with c. carinicaudus in their high average number
of black rings, but tend strongly to reduction of the nuchal ring,
which is a half -ring or is otherwise irregular in seven specimens. In
10 male specimens the black rings vary from 14 to 24; in 14 female
specimens the rings range from 17 to 27.

Micrurus carinicaudus antioquiensis Schmidt

Micrurus antioquiensis Schmidt, 1936, Field Mus. Nat. Hist., Zool. Ser.,
20: 195 — Santa Rita, north of Medellin; Niceforo Maria, 1942, Rev.
Acad. Colomb. Ciencias, 5: 98.

Diagnosis. — Distinguished from the coral snakes of the non-
triad type west of the Andes (Micrurus carinicaudus transandinus)
by having the nuchal ring reduced or irregular in the majority of
specimens; and from carinicaudus carinicaudus by a much smaller
number of black rings, average 14.

Range. — Restricted to the Cauca Valley, but intergrading over
a broad area in the Magdalena Valley with c. carinicaudus.

Colombian specimens. — In the six male and five female specimens
from the Cauca drainage (Medellin and vicinity and Segovia) five
have the nuchal ring wanting and four have it reduced to a half
ring; it is very irregular in one and normally developed as a complete
ring in one. The number of black rings on the body in this series
ranges from 10 to 17, averaging 14.

Micrurus carinicaudus transandinus Schmidt

Micrurus transandinus Schmidt, 1936, Field Mus. Nat. Hist., Zool. Ser., 20:
195 — Andagoya, Choco, Colombia; Niceforo Maria, 1942, Rev. Acad.
Colomb. Ciencias, 5: 100.

344 FIELDIANA: ZOOLOGY, VOLUME 34

Diagnosis. — A coral snake of the species carinicaudus with normal
nuchal ring and with relatively few black rings on the body (11-21,
average 15).

Range. — The Choco region west of the Andes from the Gulf of
Uraba to Ecuador.

Colombian specimens. — The large series of transandinus now
available is relatively uniform in both coloration and scutellation.
In fifteen male specimens the range of ventrals and caudals is 194
to 204 and 50 to 58, with 11 to 16 black rings on the body. In
fifteen females the ventral range is 208 to 217, caudals 36 to 42,
black rings 14 to 21. The localities represented are Pueblo Rico,
Condoto, Andagoya, Barbacoas, and Acandi.

Micrurus dumerili Jan

Elaps dumerilii Jan, 1858, Rev. Mag. Zool., 1858: 522 — Cartagena.
Micrurus dumerilii Ruthven, 1922, Misc. Publ. Mus. Zool. Univ. Mich.,
8: 68; Niceforo Maria, 1942, Rev. Acad. Colomb. Ciencias, 5: 98.

Diagnosis. — A species of Micrurus with a black pileus, with the
middle black band of the first triad as the nuchal band; accessory
black bands in the triads narrow; strongly developed supra-anal
keels in adult males.

Range. — From Cartagena to the Santa Marta region, southward
to Santander do Norte.

Colombian specimens. — This species is confined to Colombia;
a specimen in the Museo de la Salle, no. 545, was collected south
of the Santa Marta region — the first known extension of its range.

In this form, as in Micrurus ecuadorianus, I am certain that the
triad arrangement of the black ring is developed independently of
that of the great number of triad-ringed species of South America.
Black pigment, originally distributed throughout the red zones,
becomes concentrated at the borders of the yellow zones, and this
produces the primitive triad arrangement. I regard dumerili as
closely allied to carinicaudus.

Micrurus ecuadorianus sangilensis Niceforo Maria

Micrurus ecuadorianus sangilensis Niceforo Maria, 1942, Rev. Acad. Colomb.
Ciencias, 5: 98, pi. 3, fig. 10 — San Gil, Santander, Colombia.

Diagnosis. — A coral snake with black pileus, with the black
bands in triads, with the nuchal band the middle band of the first
triad, and without supra-anal tubercles in the adult male.

SCHMIDT: COLOMBIAN CORAL SNAKES 345

Range. — Confined to the region between the Cordillera Central
and the Cordillera Oriental, northern Colombia.

Colombian specimens. — The range of variation of the specimens
at hand (8) falls within that of the larger series of paratypes de-
scribed by Nice"foro Maria.

Micrurus ornatissimus Jan

Elaps ornatissimus Jan, 1858, Rev. Mag. Zool., 1858: 521 — Mexico (corrected

to Colombia, and restricted to Rio Putumayo).
Micrurus ornatissimus Schmidt, 1936, Field Mus. Nat. Hist., Zool. Ser., 20:

191.
Micrurus mimosus Amaral, 1935, Mem. Inst. Butantan, 9: 221, fig. 6 — Rio

Putumayo; Niceforo Marfa, 1942, Rev. Acad. Colomb. Ciencias, 5: 99,

pi. 3, fig. 11.

Diagnosis. — A coral snake with a light spot on each supraocular
shield in most specimens, black rings not in triads, the black rings
edged with very narrow yellow rings that are reduced to yellow spots
on half scales, so that the yellow rings are discontinuous. The red
rings are subject to darkening with black pigment and the black
rings to fading, producing an almost uniform coloration from which
the yellow half-scales stand out as strikingly distinct spots; the
venter fades to complete loss of the dark rings.

Range. — Region of headwaters of the Amazon, from the Rio
Caqueta in Colombia to southern Peru. Occasional specimens from
the lower Amazon are regarded as having been transported, acci-
dentally or by human agency.

Colombian specimens. — The three specimens at hand are from
La Pedrera, Rio Caqueta, from Putumayo, and from Puerto Asis,
Putumayo. These three specimens agree in having the diagnostic
character of ornatissimus, in the presence of a light spot on each
of the supraocular scales; their dark bands are alternately lighter
and darker, corresponding evidently to alternating red and black
zones; the venter in all three is at first glance without markings.

When the coloration is attentively examined, it is found that
the black rings can plainly be seen on the smallest specimen; that
in another specimen (the largest), they cannot be seen at all; and
that in the other, the red rings can be detected, with no trace what-
ever of black or yellow rings ventrally. These seem to be stages
in fading with exposure to light after preservation, rather than
ontogenetic changes in the color pattern. There is much about
ornatissimus that is not understood, and description of the pattern

346 FIELDIANA: ZOOLOGY, VOLUME 34

from life is especially required. Differential fading of the several
colors is well known in coral snakes. In Micrurus mipartitus, for
example, the light rings of the body are red in life but may be
yellowish or white in preserved specimens, while the bands across
the head and tail are still distinctly red. I hope to return to the
ornatissimus-langsdorffi problem in discussing the larger series of
specimens now available from Ecuador and Peru.

Micrurus psyches Daudin

Vipera psyches Daudin, 1803, Hist. Nat. Kept., 8: 320, pi. 100, fig. 1— Suri-
nam.

Micrurus psyches Beebe, 1919, Zoologica, 2: 216; Niceforo Maria, 1942, Rev.
Acad. Colomb. Ciencias, 5: 100, pi. 3, fig. 9.

Diagnosis. — A coral snake with numerous black rings alternating
with narrow yellow (or white) ones, with the alternate black rings
distinguishably different in width; these alternate rings sometimes
much lighter in color, evidently red rings with the scales darkened
by black pigment; top of head black, without crossbands, with
a narrow light band separating the black pileus from the nuchal
ring; no supra-anal tubercles in adult males.

Range. — The Guiana region to the Orinoco Basin, and thus in
the Meta drainage at Villavicencio, Colombia.

Colombian specimens. — Three specimens from Villavicencio have
very slightly higher scale counts than Guianan specimens. MLS
no. 576, a male, has ventrals 201, caudals 50, with 37 + 8 black
rings (all the rings dead black). MLS 573 and 574 have ventrals
217 and 214, caudals 33 and 30, and alternating black and darkened
red rings, 26 + 25 and 23 + 22 on the body, black rings on tail
5 in both.

Micrurus nicefori sp. nov.

Micrurus mipartitus subsp. Niceforo Maria, 1942, Rev. Acad. Colomb.
Ciencias, 5: 99.

Type. — A juvenile male specimen from Villavicencio, Cundi-
namarca, collected in 1937 by Niceforo Maria; Museo de la Salle
no. 571.

Diagnosis. — A species of Micrurus with a high ventral count;
numerous alternating black and light rings, the black rings not
arranged in triads, not alternately wider and narrower; the head
black, with a narrow light crossband across the prefrontals and
a second one across the posterior part of the parietals.

SCHMIDT: COLOMBIAN CORAL SNAKES

347

Description of type. — Body slender, head not distinctly wider
than body, tail short; eyes of moderate size, their distance from thf
labial border about one and a half times their diameter; head
shields in normal arrangement; upper and lower labials 7 on each
side; a single preocular and 2 postoculars on each side; temporals

a b

FIG. 65. Micrurus nicefori sp. nov. a, pattern of head and neck; b, pattern
of body.

1-1, with a large post- temporal along the parietal shield, on each
side; the sixth upper labials and the parietals widened so that they
nearly separate the anterior and posterior temporals; ventrals 244
(counted from the chin shields) ; anal divided; caudals 37, all divided.

Head black above; a narrow light band on the snout extending
from the third labial on each side across the pref rentals; an equally
narrow band from the seventh upper labials crossing the posterior
temporal and the post-temporal and the parietals, with a forward
projection on the temporal suture, the tips of the parietals being
black; labial border black beneath, middle of chin light with obscure
black spots. Nuchal ring open below, extending from the posterior
ends of the parietals to the sixth scale behind them.

Black rings, including the nuchal, 52 (two divided so that on
one side they may be counted as 54) ; black rings two to three ventrals

348 FIELDIANA: ZOOLOGY, VOLUME 34

in length; light rings (presumably red in life) usually two ventrals
in length, rarely three; a few of the scales in the light rings with
small dark spots; tail with seven dark rings and the intervening
light rings dark-spotted like those on the body.

Measurements. — Total length 270 mm., tail 23 mm.; diameter
of body 3.5 mm.

Remarks. — As Hermano Nice"foro Maria relates, this specimen
has passed through my hands before (probably about 1938) and
through those of Dr. Alcides Prado of the Instituto Butantan. We
both reported it to him as undescribed and returned his specimen
with the hope that additional ones might turn up in order to provide
a better basis for a description. I still find no possible allocation
for this specimen among the known species. I am now much more
impressed than formerly with the validity of the characters of the
color pattern in Micrurus, and I do not regard the new form as
subspecifically related to mipartitus, of which I have examined
large numbers of specimens. Lest the type be worn out in further
travels, I have given this little coral snake waif a name.

Micrurus hemprichi ortoni Schmidt

Micrurus hemprichi ortoni Schmidt, 1953, Fieldiana, Zool., 34: 166 — Pebas,

Peru.
Micrurus hemprichii Niceforo Maria, 1942, Rev. Acad. Colomb. Ciencias,

5:99.

Diagnosis. — A species of coral snake of the genus Micrurus with
the anal plate entire; with the black rings in groups of three (triads) ;
and with 5 to 6 triads on the body.

Range. — The region of the upper Amazon.

Colombian specimens. — Only a single specimen from Colombia
is at hand, MLS no. 568, from Florencia, Caqueta. The specimen
recorded by Niceforo Maria from Putumayo appears to represent
another record. In the present specimen, a female, the ventrals
number 187; the caudals 30, with four undivided; and the black
triads 5, with only two- thirds of a triad on the tail. The total
length is 760 mm., tail 72 mm.

The type locality of hemprichi was "Colombie," but the descrip-
tion of the type places it unequivocally with the Guianan series,
and as the species has not been recorded from Venezuela, and as
I believe Jan's statement of the origin of his specimen to be in error,
I have suggested as amended type locality "Vicinity of Bartica,

SCHMIDT: COLOMBIAN CORAL SNAKES 349

British Guiana." The Guianan subspecies has 7 to 10 triads on the
body instead of 5 to 6 as in ortoni.

Micrurus surinamensis surinamensis Cuvier

Micrurus surinamensis surinamensis Schmidt, 1952, Fieldiana, Zool., 34: 29.
Micrurus surinamensis Niceioro Maria, 1942, Rev. Acad. Colomb. Cien-
cias, 5: 100.

Diagnosis. — A coral snake with the black rings in triads; the
head broad, with a distinctively narrow frontal shield; the shields
of the top of the head all red, narrowly outlined with black; ventrals
162-174 in males, 173-187 in females.

Range. — The headwaters of the Amazon tributaries, northward
to Villavicencio (Orinoco drainage) ; the Guiana region.

Colombian specimens. — The Museo de la Salle has two specimens,
MLS no. 592, a male, from Villavicencio, and no. 593, a female,
from Tres Esquinas, Rio Orteguaza, Caqueta. In the male specimen
the ventrals number 172^, the caudals 34 (with 12 undivided);
the temporals are 1-2 on each side, and the triad formula is 6% +
^1 (body + tail). In the female specimen the ventrals number
169, the caudals 31 (two undivided), the temporals are 1-2, and the
total length is 495 mm., tail 54 mm. In this specimen the triads,
about 8 + ^1, are somewhat irregular, obscured dorsally by
a general darkening of the red zones, and still more by the irregular
fusion of black rings posteriorly, where they may be arranged
in "quadrads."

I have examined also a specimen in the British Museum labeled
Villeta, and one from Puerto Boy, Caqueta, sent by Hermano
NiceToro Maria for my examination on an earlier occasion; and
Amaral records a second specimen from Villavicencio, likewise from
the Museo de la Salle.

Micrurus lemniscatus Linnaeus

Coluber lemniscatus Linnaeus, 1758, Syst. Nat., ed. 10, p. 224 — Asia (in errore;

type locality restricted to lower Amazon region).
Micrurus lemniscatus Beebe, 1919, Zoologica, 2: 216; Nic6foro Marfa, 1942,

Rev. Acad. Colomb. Ciencias, 5: 99.

Diagnosis. — A coral snake of the genus Micrurus of slender
habitus; head with a distinctively small eye; black rings in triads,
of which the first is invariably a complete group of three; always
with a light crossband across the head anterior to the eyes.

350 FIELDIANA: ZOOLOGY, VOLUME 34

Range. — The Amazon and Orinoco basins, extending to Trinidad
and the Guianas; the species reaches Colombia along the Putumayo
and Caquetd rivers and occurs at Villavicencio, in the drainage
of the Meta.

Colombian specimens. — The Colombian specimens available for
examination, all collected by Nice"foro Maria, agree closely with the
Amazonian population. In seven males the ventrals range from
236 to 252, and the caudals from 34 to 41; in four females these
ranges are respectively 243 to 261 and 40 to 43. The number of
triads of black rings in the whole series varies from 9 to 11 on the
body and from 1^ to 2 on the tail. The largest specimen, a skin,
may be estimated as a little more than a meter in length, the length
of the tail being 93 mm.

The Colombian localities from which lemniscatus has been
received are Florencia, Rio Orteguaza, and Rio Putumayo, Inten-
dencia del Caqueta, and Villavicencio, Cundinamarca.

Micrurus filiformis Giinther

Elaps filiformis Giinther, 1859, Proc. Zool. Soc. London, 1859: 86, pi. 17

— Para, Brazil.
Micrurus filiformis Amaral, 1925, Proc. U. S. Nat. Mus., 67, Art. 24, p. 19;

Niceforo Maria, 1942, Rev. Acad. Colomb. Ciencias, 5: 99.

Diagnosis. — An elongate and excessively thin-bodied species in
which the elongate character of lemniscatus is exaggerated; with the
lemniscatus head and body pattern, but with more numerous triads
and a higher ventral count in both sexes.

Range. — Known throughout the northern Amazon Basin, absent
in the Guianas, reaching Colombia along the Putumayo and Caqueta
rivers, northward to Villavicencio at the headwaters of the Meta.

Colombian specimens. — The three available specimens, all male,
are from Villavicencio, Cundinamarca; and from Puerto Boy and
La Pedrera, Intendencia del Caqueta. Ventrals range from 266 to
281, caudals 39 to 41, and triads of black rings 12 to 16. The longest
specimen measures 540 mm., tail 44 mm.

Micrurus spixi obscurus Jan

Elaps corallinus var. obscura Jan, 1872, Icon. Gen. Ophidiens, 41: pi. 6,

fig. 3 — Lima (in errore; corrected to eastern Peru).
Micrurus spixii obscura Schmidt and Walker, 1943, Field Mus. Nat. Hist.,

Zool. Ser., 24: 294.
Micrurus spixii spixii Niceforo Maria, 1942, Rev. Acad. Colomb. Ciencias,

5: 100.

SCHMIDT: COLOMBIAN CORAL SNAKES 351

Diagnosis. — A large coral snake with stocky body and very
short tail, the black rings arranged in triad groups in which the
yellow rings are as broad as or broader than the outer black rings;
the nuchal black marking (the middle ring of a triad) has a forward
projection extending to the parietals; there are usually a number of
entire subcaudals.

Range. — Micrurus spixi ranges throughout the Amazon Basin,
but is absent from the Guianas and most of Venezuela. The sub-
species obscurus ranges from northern Bolivia through Amazonian
Peru and Ecuador to Colombia; it is replaced in Amazonas by the
typical spixi spixi.

Colombian specimens. — The seven Colombian specimens are
from Florencia, Puerto Boy, and Rio Putumayo, Intendencia del
Caqueta. In the six males the ventrals range from 211 to 215, the
caudals from 17 to 22 (0 to 10 entire) . In the single female specimen
the ventrals number 219 and caudals 19 (6 entire). The number
of triads of black rings on the body (as seen from above) is 5 in five
specimens, with the posterior two-thirds of a triad on the neck and
an anterior two-thirds of a triad on the tail. In two specimens
the nuchal triad is complete, and the triad formula is 6 + %. The
anterior member of the first triad is present on the parietal shield,
usually broken into spots or otherwise obscure; on the under side
of the head it is present as a narrow transverse half-ring in the five
specimens listed as having two-thirds of a triad anteriorly. In other
subspecies of spixi this anterior ring is much more obscure; and the
two specimens, MLS 580 and 589 (both from Florencia) are the
only ones known in obscura in which the anterior ring is fully com-
plete on the upper side.

The largest specimen, a male from Puerto Boy, measures 884
mm., tail 46.

Micrurus ancoralis jani Schmidt

Micrurus ancoralis jani Schmidt, 1936, Field Mus. Nat. Hist., Zool. Ser.,
20: 197 — Andagoya, Choco, Colombia; Nic^foro Maria, 1942, Rev. Acad.
Colomb. Ciencias, 5: 98.

Diagnosis. — An elongate but stout-bodied coral snake with the
black rings in triads, and with an "anchor-shaped" nuchal marking;
ventrals 248-266 in males, 271-290 in females; triads on body 12-15
in males, 14-16 in females.

Range. — The Choco region of Colombia, northward to Panama.

352 FIELDIANA: ZOOLOGY, VOLUME 34

Colombian specimens. — For my original description I had eighteen
specimens, and only four additional specimens have appeared in the
intervening years, two in Chicago Natural History Museum and
two in the Museo de la Salle. One of these latter exceeds a meter
in length, and this specimen has temporals 1-2 on each side (1-1 in
all others). In range of ventral count and in color pattern these
specimens agree fully with the original series. Micrurus ancoralis
jani is known from Tado, Condoto, Pena Lisa, Novita, Pueblo Rico,
and Andagoya, all in the Rio San Juan drainage, and from Quibdo
(MLS 500 and 501) on the upper Atrato.

Micrurus dissoleucus Cope

Elaps dissoleucus Cope, 1859, Proc. Acad. Nat. Sci. Phila., 11 : 345 — Venezuela.

MATERIAL EXAMINED

PANAMA: Panama City, MCZ 38244; Juan Diaz, MCZ 49943; Aguadulce,
AMNH 67058; no locality, ERD Coll., no number (1), Munich, no number
(1), ANSP 7173, 22602-35. CANAL ZONE: Ancon, MCZ 16304 (type of dunni);
Golf Club, MCZ 45405; Corozal, MCZ 18813.

COLOMBIA: No other data, or questioned data, MCZ 29602; "Trinidad,"
AMNH 3912; "Cartago," AMNH 20407. Bolivar (by inference): Magdalena
Valley, ANSP 6806, BM 1902-5-15-20. Atlantico: Barranquilla, MLS 542, Senck-
enberg Mus. 20734-5, 43586, Berlin 3853 (2); mouth of the Magdalena, Vienna,
no number (3). Magdalena: Salamanca Island (east of Barranquilla), Vienna,
no number (4). Santa Marta, CNHM 2050, MCZ 15967; Bonda, CM 206
(type of hollandi), 207; no data, ANSP 6807 (type of melanogenys), AMNH
45095-96. Norte de Santander: Cucuta, CNHM 5685, 27009, MLS 537, 539.

VENEZUELA: No other data, ANSP 6781 (type of dissoleucus). Federal
District: Caracas, Munich, no number (1); El Valle, AMNH 59393; Puerto La
Cruz, CM 7346. Aragua: Maracay, Munich, no number (5); Fijirias, Munich, no
number (1); Pie del Cerro, CM 7397. Zulia: Maracaibo, Basel 2309-10; Cabinas,
BM 1932-12-5-1.

Diagnosis. — A small species of coral snake of the genus Micrurus,
rarely exceeding 400 mm. in length (maximum known 620 mm.);
tail relatively short but with distinct sex dimorphism in length; the
black rings in triads; the nuchal ring representing the posterior ring
of a triad; triad formula ^6 to ^11 for the body, and % to 1%
for the tail; no light crossband on the snout; head crossed posteriorly
by a broad light band. Four subspecies may be recognized.

Synopsis of subspecies of Micrurus dissoleucus:

1. Caudals in males 23-28 2

Caudals in males 21-22 dissoleucus melanogenys

(Santa Marta region)

FIG. 66. Pattern of head of type of Micrurus dissoleucus Cope, ANSP no.
6781; X 2.

FIG. 67. A triad of black rings at mid-body in type of Micrurus dissoleucus
Cope, ANSP no. 6781.

353

354 FIELDIANA: ZOOLOGY, VOLUME 34

2. Ventrals in females fewer than 212 dissoleucus dissoleucus

(northeastern Colombia and northwestern Venezuela)
Ventrals in females more than 212 3

3. Triads of black rings (in both sexes) 6-8 dissoleucus nigrirostris

(lower Magdalena region)

Triads of black rings 8-11 dissoleucus dunni

(Panama and presumably adjacent Colombia)

Comparisons.— By way of amplification of the comparison of the
subspecies of dissoleucus in the key above, the range of variation
in numerical characters may be tabulated as follows:

No. of Average

specimens Ventrals Caudals Triads of triads

dissoleucus rf1 8 175-201 24-28 8-10 9

9 10 201-212 21-23 6-11 9

melanogenys d" 3 185-199 21-22 6-9 7

9 4 197-210 17-19 7-8 7

nigrirostris d" 11 190-208 23-27 6-8 7

9 7 212-228 17-22 7-8 8

dunni d1 8 185-200 24-26 8-10 9

9 3 208-217 19 10-11 10

Distribution. — The vicariating subspecies extend along the
Caribbean versant of South America from as far east as Caracas
in Venezuela to the Canal Zone and to the Pacific coast in Panama.
They occur inland in Venezuela to moderate altitudes but are other-
wise characteristically lowland forms. The subspecies melanogenys
is narrowly separated from nigrirostris of the Magdalena delta by
the Cienega de Santa Marta. The typical subspecies extends into
Colombia in the region of Cucuta, rather well separated from the
range of the Santa Marta subspecies by the barrier of the Sierra de
Perija. There is no presumable barrier between the subspecies
nigrirostris and the Panamanian dunni, and it is somewhat to be
inferred that the latter form may reach Colombia. There is a gap
of some 500 km. between the known ranges of dunni and nigri-
rostris.

M icrurus dissoleucus dissoleucus Cope

Micrurus dissoleucus dissoleucus Schmidt, 1936, Field Mus. Nat. Hist., Zool.
Ser., 20: 202; Niceforo Maria, 1942, Rev. Acad. Colomb. Ciencias, 5: 98.

Diagnosis. — A small species of coral snake in which the nuchal
black ring is not part of the first triad on the body; head crossed
posteriorly by a broad light band; no light crossband on snout;
tail relatively short; caudals 21-23 in females, 24-28 in males;

SCHMIDT: COLOMBIAN CORAL SNAKES 355

ventrals 201-212 in females, 175-201 in males; and triads of black
rings on the body averaging 9 in both sexes.

Range. — From northeastern Colombia (North Santander) east-
ward through northern Venezuela to Caracas.

Colombian specimens. — This species occurs at Cucuta; Nice'foro
Maria gives the additional localities El Rosario, Rio Tulia, Puerto
Santander, and Petrolea. I do not have the Museo de la Salle
series at hand.

Micrurus dissoleucus melanogenys Cope

Elaps melanogenys Cope, 1860, Proc. Acad. Nat. Sci. Phila., 12: 72 — South

America (restricted to Santa Marta region, Colombia).
Elaps hollandi Griffin, 1916, Mem. Carnegie Mus., 7: 218, pi. 18, figs. 10-12

— Bonda, Colombia.
Micrurus dissoleucus melanogenys Schmidt, 1936, Field Mus. Nat. Hist.,

Zool. Ser., 20: 203; Niceforo Maria, 1942, Rev. Acad. Colomb. Ciencias,

5: 98 (part only).

Diagnosis. — A small coral snake with the black ring in triads,
the nuchal ring not part of the first triad, and with an extremely
short tail; caudals 17-19 in female specimens, 21-22 in male speci-
mens. The number of black rings on the tail is correspondingly
reduced, with no more than one complete triad present in the
specimens known.

Range. — Confined to the Santa Marta region in northeastern
Colombia.

Colombian specimens. — I have available only four specimens, two
males and two females (including the type and paratype of hollandi},
with data. In addition to the type of melanogenys (which I have
examined) I associate with this series two specimens without data,
in the collection of the American Museum of Natural History.

Micrurus dissoleucus nigrirostris subsp. nov.

Micrurus dissoleucus dunni Schmidt, 1936, Field Mus. Nat. Hist., Zool. Ser.,
20: 203 (part).

Type. — Senckenberg Museum no. 20734, a male, from Barran-
quilla, Colombia, collected by Fr. Regel in 1897.

Diagnosis. — A coral snake of small size, with the black rings in
triads, the nuchal ring not a part of the first triad; snout entirely
black; ventrals 190-208 in males, 212-228 in females; caudals 23-27
in males, 17-22 in females; number of triads, in both sexes, six
to eight.

FIG. 68. Pattern of head of type of Micrurus dissoleucus melanogenys,
ANSP no. 6807; X 4.

FIG. 69. Pattern of body in type of Micrurus dissoleucus melanogenys,
ANSP no. 6807.

356

SCHMIDT: COLOMBIAN CORAL SNAKES 357

Description of type. — A coral snake of the Micrurus dissoleucus
group, with small head and rounded snout; rostral narrowly visible
from above; internasals half as long as the pref rentals; frontal
obtusely angulate in front, relatively wide; one preocular and 2
postoculars; the temporals on each side (i.e. the normally single
anterior and posterior temporals) fused into a single elongate shield,
followed by two shields posteriorly; 7 upper and 7 lower labials;
scale rows 15 at the first widened ventral; ventrals 208, caudals 27,
two of which are undivided.

Head black, with a yellow band from the fifth and sixth labials
across the head, occupying the posterior two-thirds of the supra-
oculars and frontal, the postoculars, the anterior half of the elon-
gate temporal, and the anterior two-fifths of the parietals; lower
labials and chin entirely dark except for the yellow posterior chin
shields opposite the light band on the upper side of the head; black
bands in triads, nuchal band not included in the first, seven com-
plete triads on the body plus one and one-third on the tail (counting
the black tail tip) ; black rings subequal above, the outer rings of the
triads narrowed on the ventrals; the third triad in number of ventrals,
as follows:

Red Black Yellow Black Yellow Black Red
94 35 349

Measurements of type. — Total length 355 mm., tail 27 mm.

Notes on paratypes. — I have examined seventeen specimens, in
addition to the type, that are referred to this form. Of these one
has no data; the specimen labeled as from Cartago may be explain-
able by a lapsus as coming from Cartagena; a specimen labeled
"Trinidad" may be entirely mislabeled, or may be from some
obscure Trinidad in Colombia unknown to me. The remaining
specimens with positive and plausible data are from the region of
the Magdalena delta or the lower Magdalena Valley.

The type is exceeded in length only by MCZ no. 29602, a female
367 mm. long, with tail 20 mm. There is a frequent tendency (as in
Senck. no. 20735) for the reduction of the contact between the
anterior and posterior temporals, so that the upper angle of the
sixth labial may meet, or nearly meet, a lateral angle of the parietal.
It is gratifying to note that the material now at hand confirms my
prediction of 1936 that the Magdalena Basin specimens might be
distinguishable from the Panamanian dunni.

358 FIELDIANA: ZOOLOGY, VOLUME 34

Micrurus dissoleucus dunni Barbour

Micrurus dunni Barbour, 1923, Qcc. Papers Mus. Zool. Univ. Mich., 129: 35

— Ancon, Canal Zone, Panama.
Micrurus dissoleucus dunni Schmidt, 1936, Field Mus. Nat. Hist., Zool. Ser.,

20: 203 (part).

Diagnosis.— A small coral snake with the nuchal black ring not
a part of the first triad of black rings on the body; caudals in males
23-28; ventrals in female specimens more than 212; and triads of
black rings on the body 8-11.

Range. — Panama from the Canal Zone eastward, the eastern
boundary unknown, perhaps extending into Colombia.

Remarks. — The formal reference to dunni is included in order
to complete the treatment of the species dissoleucus. The subspecies
dunni probably occurs in northwestern Colombia. I am indebted
to Mr. Arthur Loveridge for corrections of the original description
of the type, which has 217 ventrals instead of 224, and measures
313 mm., tail 18 mm., instead of 130 mm. and 20 mm. Barbour
stated that the light rings within the triads are red, which, on the
analogy of the great majority of coral snakes, is highly improbable.
He correctly discerned the close relation to dissoleucus, but his
suggestion of a relation to tschudi of coastal Peru is rejected.

REFERENCES

AMARAL, AFRANIO DO

1935. Estudos sobre ophidios neotropicos., XXXIII. Novas especies de
ophidios da Colombia. Mem. Inst. Butantan, 9: 219-223, 1 pi.

BARBOUR, THOMAS

1923. Notes on reptiles and amphibians from Panama. Occ. Papers Mus.
Zool. Univ. Mich., 129: 1-16.

DANIEL, H.

1949. Las serpientes en Colombia. Rev. Fac. Nac. Agronomia Medellin, 9:
301-333.

MAR!A, HERMANO NICEFORO

1942. Los ofidios de Colombia. Rev. Acad. Colomb. Ciencias, 5: 84-103,
3 pis., 1 map.

RENDAHL, HIALMAR, and VESTERGREN, GRETA

1940. Notes on Colombian snakes. Ark. Zool., 33A, no. 1, pp. 1-16, 4 figs.

SCHMIDT, KARL P.

1936. Preliminary account of coral snakes of South America. Field Mus.
Nat. Hist., Zool. Ser., 20: 189-203.

1952. The Surinam coral snake Micrurus surinamensis. Fieldiana, Zool., 34:
25-34, figs. 4-6.

SCHMIDT: COLOMBIAN CORAL SNAKES 359

SCHMIDT, KARL P., and WALKER, WARREN F., JR.

1943. Peruvian snakes from the University of Arequipa. Field Mus. Nat.
Hist., Zool. Ser., 24: 279-296.

UNIVERSITY OF ILLINOIS-URBANA