Biological Journal of the Linncon Socie& (1992), 45: 235-254.

With 6 figures

The colouration of the venomous coral snakes

(family Elapidae) and their mimics (families
Aniliidae and Colubridae)
JAY M. SAVAGE, F.L.S. AND JOSEPH B. SLOWINSKI

Department of Biology, University of Miami, P.O. Box 249118, Coral Gables,

FL 33124, U.S.A.

Received 7 August 1990, accepted for publication 30 October 1990

The bright coloured, highly venomous coral snakes, Lcptominurus, Micrurus and Mimroidcs (family
Elapidae) and a series of harmless or mildly toxic mimics form an important component of the snake
fauna of the Americas. Coral snake patterns are defined as any dorsal pattern found in any species of
venomous coral snake and/or any dorsal pattern containing a substantial amount of red, pink or
orange distributed so as to resemble that of some species of venomous coral snake. The components
of coral snake colouration are described and four principal dorsal patterns are recognized: unicolour,
bicolour, tricolour and quadricolour. The tricolour patterns may be further clustered based on the
number of black bands or rings separating the red ones as: monads, dyads, triads, tetrads or pentads.
A detailed classification of all coral snake colour patterns is presented and each pattern is illustrated.
The taxonomic distribution of these patterns is surveyed for mimics and the 56 species of highly
venomous coral snakes. Among the latter, the most frequent encountered patterns are tricolour
monads, tricolour triads and bicolour rings, in that order. No venomous coral snakes have a
tricolour dyad, tricolour tetrad or quadricolour pattern.
As many as I15 species of harmless or mildly toxic species, c. 18% of all American snakes, are
regarded as coral snake mimics. The colouration and behavioural traits of venomous coral snakes
combine to form a significant antipredator defence of an aposematic type. The mimics in turn
rereive protection from predators that innately or through learning avoid coral snake colour
patterns. The precise resemblances in colouration between sympatric non-coral snakes and
venomous coral snakes and the concordant geographic variation between the two strongly support
this view. Batesian mimicry with the highly venomous coral snakes as the models and the other
forms as the mimics is the favoured explanation for this situation. I t is further concluded that a
number of species in the genera Elaphc, Farancia, Nerodia and Thamnophis, although having red in
their colouration, should not be included in the coral snake mimic guild.

KEY WORDS:-Coral snakes ~ colouration ~ mimicry.

CONTENTS

Introduction . . . . . . . . . . . . . . . . . . . 236
Definition of coral snake colouration . . . . . . . . . . . . . 236
Components of the colour pattern . . . . . . . . . . . . . . 237
A classification of coral snake colour patterns . . . . . . . . . . . 240
Taxonomic distribution in American species . . . . . . . . . . . 245
Non-venomous or mildly venomous presumptive coral snake mimics. . . . . 245
Venomous coral snakes . . . . . . . . . . . . . . . 248
Colouration and mimicry . . . . . . . . . . . . . . . . 250
References . . . . . . . . . . . . . . . . . . . 253
235
002+4066/92/030235 + 20 S03.00/0 0 1992 The Linnean Society of London
236 J. M. SAVAGE AND J. B. SLOWINSKI
INTRODUCTION

During a course of studies on the New World venomous coral snakes (family
Elapidae) and their presumed aglyphous and opisthoglyphous mimics we were
struck by the absence of consistent terminology to describe major aspects of
their colour patterns. Since these patterns are often utilized to distinguish
between allied taxa and may be of significance in establishing relationships, this
inconsistency hampers systematic studies of both the highly venomous species
and the guild of coral snake mimics. In addition, the lack of a uniform
terminology inhibits attempts to establish congruence in colouration between the
supposed mimics and their models.
We have previously published a preliminary paper (Savage & Slowinski,
1990) outlining broad categories of coral snake patterns. In the present paper a
much more detailed analysis is developed and we address the problem in depth
by: ( 1 ) identifying and defining the major components of colour pattern in these
groups; (2) establishing a consistent terminology to describe variation in the
patterns and; (3) summarizing the taxonomic occurrence of the various patterns.
In the following sections the term ‘venomous coral snakes’ refers to
proteroglyphous elapids of the genera Leptomicrurus, Micrurus and Micruroides.
The term ‘mimic’ is used to denote aniliid and colubrid snakes that resemble
venomous coral snakes and which are non-venomous or mildly toxic and lack the
‘advanced’ venom delivery system (rigid canaliculate anteriorly placed fangs on
the upper jaw and venom gland compressor muscles) found in elapids.

DEFINITION OF CORAL SNAKE COLOURATION

An initial consideration in undertaking this review was the need to delimit

those elements of pattern that constitute coral snake colouration since the
venomous coral snakes (the nominal genera Leptomicrurus, Micruroides and
Micrurus) and their harmless to mildly toxic mimics display a substantial range of
variation in colouration. Throughout the Americas snakes having a considerable
amount of red, pink or orange in their colouration are called coral snakes, corales
(Spanish) or cobras-coraes (Portuguese) and almost all venomous coral snakes and
their mimics are known by these names. Consequently we have taken our cue
from the vernacular names and define as a coral snake pattern any one having a
substantial amount of red, pink or orange in the dorsal colouration and/or found
in some species of coral snake.
The patterns usually consist of alternating bands or rings of black (or dark
brown) and red, pink or orange, but some coral snakes have a uniform red dorsal
(and ventral) colouration and a few others may only be spotted or blotched with
black. In addition, while most venomous coral snakes have the body encircled by
black and red rings and usually by yellow or white rings as well, a few species
have bicolour black and yellow or black and white ringed patterns. For this
reason we have included in this survey all kinds of bicolour snakes having
alternating black and light rings around the body. Essentially bicoloured
harmless and mildly toxic snakes having dark brown or black blotches or bands
restricted to the dorsal and lateral areas of the body (e.g. Leptodeira nigrofasciata)
are not considered to be mimics of bicolour coral snakes and are not included in
this review. Also excluded from our definition of coral snake patterns, since they
 THE COLOURATION OF THE VENOMOUS CORAL SNAKES 237

occur in no venomous coral snakes, are patterns of snakes having narrow red,
pink or orange dorsal and lateral stripes (e.g. some lhamnophis and Liophis),
others with uniform or striped dorsums that have red, pink or orange venters
(e.g. some species of Coniophanes, Rhadinaea, Trimetopon and Urotheca) or red, pink
or orange venters and nuchal collars (e.g. Diadophis) or the dorsum suffused with
one of these bright colours (e.g. Coluberjagellum).
In view of the above discussion any definition of coral snake colouration must
include two components, one emphasizing the presence of red, pink or orange
colours and the second including all patterns found in the venomous coral
snakes. Consequently our definition is inclusive and takes into account both
colour and taxonomic parameters. Simply stated, coral snake colouration is
defined as: any dorsal pattern found in a species of venomous coral snake andlor any dorsal
pattern containing a substantial amount of red, pink or orange distributed in a fashion that
resembles a pattern found in some species of venomous coral snake.

COMPONENTS OF THE COLOUR PATTERN

The following paragraphs are designed to identify and define major

components of the colour pattern in coral snakes (venomous and otherwise).
Although emphasis in the remaining sections of this paper will be on the dorsal
pattern, the characteristics of other pattern components are often useful in
discriminating among taxa and in evaluating mimicry. The principal elements of
the colour pattern in these snakes are described below:
1. snout patern-if different from or discontinuous from the pattern on more
posterior head shields, the snout region is most often uniform black (Fig. lA, B)
in the venomous coral snakes but it may be red, white or yellow (Fig. 1C) or the
shields may be light and edged with dark or the reverse (Fig. 1D);
2. black head cap-the uniform black area variously covering the enlarged
head shields (rostral, internasals, prefrontals, parietals, nasals, loreal,
supralabials and circumocular series) on the top and sides of the head; the head
cap may be continuous from the snout posteriorly (Fig. 1A) but in other cases
the snout may be light in colour with the head cap restricted to the more
posterior shields or it may be entirely lacking; in some cases the black snout
colouration is separated from the head cap by a light bar across the prefrontal
and/or interorbital area (Fig. 1B);
3. light prefrontal band-the light band running transversely across the
prefrontal region in some forms that separates the dark (usually black) snout
from the black head cap (Fig. 1B);
4. light prefrontal-interocular band-a light band running transversely
across the head in some mimics that separates the dark snout colour from the
dark head cap;
5 . light interocular band-a light band running transversely across the head
in some mimics that separates the dark snout from the dark head cap;
6. light head band-a red, orange, yellow or white area crossing the posterior
head shields on the upper surface of the head and continuing a short distance
onto the neck in most cases (Fig. 1A-C);
7. light nuchal collar-a light coloured collar around the neck that may cross
the posterior portions of the parietals (Fig. 1D);
8. dark nuchal collar-a dark coloured band or ring across neck, posterior to
238 J. M. SAVAGE AND J . B. SLOWINSKI

A C

B D

Figure I . Characteristic patterns of the head and nuchal regions in coral snakes. Stippled areas
represent red; white areas yellow or white; black areas black. See text for explanation of
components.

the light head band or light nuchal collar when one of these is present (Fig.
1A-D) ;
9. postnuchal light collar-a contrasting light collar located on the body
posterior to the dark nuchal collar (Fig. 1A-D);
10. dorsal pattern-uniform or variously marked with dark or light;
1 1. tail pattern-uniform or variously marked with dark or light; similar to or
different from dorsal pattern;
 THE COLOURATION OF THE VENOMOUS CORAL SNAKES 239

12. ventral pattern-uniform or variously marked with dark or light;

13. subcaudal pattern-uniform or variously marked with dark or light;
similar to or different from ventral pattern.
Components 3, 4 and 5 are essentially the same feature differing only in how
far forward the dark head cap extends. Similarly, components 6 and 7, differ
only in the degree to which the black nuchal collar extends anteriorly; if far
anteriorly a light head band is produced (Fig. 1A-C), if not a light nuchal collar
is present (Fig. 1D).
Much of the confusion regarding the concordance or lack thereof between
presumed mimics and venomous coral snake models derives from the absence of
a standardized system for categorizing the variation in dorsal colour patterns. In
the system adopted in this paper emphasis is placed upon the nature of the black
(sometimes dark brown) portion of the pattern for establishing a consistent
terminology. The kinds of black markings found in snakes included in this review
are defined as follows:
spots-small regular or irregular markings covering a portion of a single scale,
a single scale or several scales;
blotches-regular or irregular figures covering many scales;
saddles-saddle-shaped blotches extending only a short distance onto flanks;
bands-continuous transverse markings that extend across the dorsum well
onto the flanks or to the margin of the ventrals;
rings-regular transverse markings that completely encircle the body;
stripes-continuous longitudinal lines.
A common cause of difficulty in studying colour pattern is the failure of many
authors to discriminate between bands and rings. Because this difference is
crucial for identification in many cases, the term band should only be used for
markings restricted to and crossing the upper and lateral body surfaces.
When the body pattern consists of bands or rings the several distinctive
patterns may be defined based on the number of black components per light
interspace or ring. For example in the simplest case (bicolour) the pattern is one
of alternating black bands or rings and light (red, orange, yellow or white) bands
or rings (one dark marking, one light). In tricolour and quadricolour banded or
ringed snakes there are a variety of patterns. A classification of these elements for
banded and ringed snakes follows:
(I) Bicolour-alternating bands or rings of black and light (red, pink, orange,
yellow or white; frequently red in the venomous coral snakes); one black
component for each light component ( 1 black: 1 light).
(11) Tricolour-alternating bands or rings of black, red and light (usually
yellow, sometimes white):
(a) tricolour monads-red bands or rings separated from one another by a
sequence of alternating bands or rings of light-black-light; one black component
between two red components ( 1 black: 1 red);
(b) tricolour dyads-red bands or rings separated from one another by a
sequence of alternating bands or rings of black-light-black; two black
components between two red components (2 black: 1 red);
(c) tricolour triads-red bands or rings separated from one another by a
sequence of alternating bands or rings of black-light-black-light-black; three
black components between two red components (3 black: 1 red);
(d) tricolour tetrads-red bands or rings separated from one another by a
240 J. M. SAVAGE AND J. B. SLOWINSKI

sequence of alternating bands or rings of black-light-black-light-black-light-

black; four black components between two red components (4 black: 1 red);
(e) tricolour pentads-red bands or rings separated from one another by a
sequence of alternating bands or rings of black-light-black-light-black-light-
black-light-black; five black components between two red components (5 black:
1 red).
(111) Tricolour lacking black bands or rings-red rings separated from one
another by an alternating sequence of white, yellow, white rings (0 black: 1
yellow: 1 red);
(IV) Quadricolour-red or orange centred dyads (black-red-black or black-
orange-black) outlined by white on a grey ground colour (2 black: 1 grey
in terspace).
Some authors (Taylor & Smith, 1943; Smith & Fitzgerald, 1986, for
Scaphiodontophis) have called condition 11(b) above a ‘triad’ because it consists of
three elements (2 black and 1 light). Others (Klauber, 1943; Tanner, 1953,
1983; Zweifel, 1952, 1974, for some Lampropeltis) have taken the opposite tack for
this pattern, calling the red ring plus the bordering black ring on each side a
‘triad’ and assuming that the light ring separating the black rings is an
interspace. We believe that these usages are inappropriate and confusing since
the term triad has long been used for condition II(c) above (at least since
Schmidt & Schmidt, 1925), especially to describe the pattern of many venomous
coral snakes. Consequently, the terms dyad (for pairs of black bands or rings:
I I ( b ) ) and triad (for trios of black bands or rings: II(c)) are adopted here to
provide a consistent and logical terminology for these aspects of colour pattern.
It should be noted that a few taxa with coral snake patterns (particularly some
Lampropeltis) have the black body bands continuing irregularly onto the ventrals
but these usually do not form complete rings. In these cases both white (or
yellow) and/or red rings may be present. In the system adopted below these
patterns are considered to be ringed and the light ring colour(s) will be
mentioned.

A CLASSIFICA‘TION OF CORAL SNAKE COLOUR PATTERNS

The following outline indicates the principal colour patterns found in coral
snakes. Each pattern type is denoted by a letter code that will be used in the
taxonomic survey to follow (Figs 2-6)
I. UNICOLOUR (U )
A. Dorsum and venter red
1. unmarked ( U )
2. nuchal black collar (Uc)
B. Only dorsum red
1. unmarked (Ud)
2. nuchal black collar (Ucd)
11. BICOLOUR (BI)
A. Spots or blotches
1. black
a. spots (BIS)
b. blotches (BIL)
 THE COLOURATION OF THE VENOMOUS CORAL SNAKES 24 1

Figure 2. Diagrammatic representations of spotted, blotched and striped coral snake patterns.

2. red blotches on light ground colour (BIr)

3. red blotches on black ground colour (Brb)
B. Black bands
1. alternating (BIB)
2. fused dorsally (BIFb)
C. Rings
1. black rings alternating with light (BIR)
2. black rings fused dorsally (BIF)
3. red bands on light (RB)
D. Secondarily bicolour-black rings
1. suffused tricolour monads (BIM)
2. suffused tricolour dyads (BID)

Figure 3. Diagrammatic representations of bicolour coral snake patterns. Colours as indicated in

Fig. 2.
242 J. M.SAVAGE AND J. B. SLOWINSKI

Figure 4. Diagrammatic representations of non-ringed tricoiour coral snake patterns. Colours as

indicated in Fig. 2.

E. Longitudinal pattern
1. dorsal area red, dark stripe(s) (LS)
2. dorsal area red, lateroventral area light (RL)
3. dorsum dark, lateral red stripe (S)
111. TRICOLOUR (T)
A. Non-ringed patterns
1. black dorsal blotches and/or bands
a. interspaces red dorsally (Tb)
b. interspaces red ventrally (Tbx)
c. interspaces yellow, red, yellow (Tbi)
2. black dorsal and lateroventral bands with red saddles on light
ground colour (TYv)
3. black bands and lateroventral spots, with red bands above
a. solid black bands (TXa)
b. central red area in black bands (TX)
4. red bands (outlined by black) on light (TSR)
5. alternating black and red blotches (TBr)
6. saddles
a. yellow or white (outlined in black) on red (Tsy)
b. black (outlined in yellow) on red (Tsb)
c. red (outlined in black) on yellow (Tsr)
7. tricolour monads: one black band per red band separated from one
another by yellow (TMb)
 THE COLOURATION OF THE VENOMOUS CORAL SNAKES 243

Figure 5. Diagrammatic representations of tricolour banded and ringed coral snake patterns.
Colours as indicated in Fig. 2.

8. tricolour dyads: two black bands per red band, black bands in
contact with red separated from one another by yellow or white
(TDb)
9. tricolour triads: three black bands per red band, outer black bands

Figure 6. Diagrammatic representations of tricolour and quadricolour banded and ringed coral
snake patterns. Colours as indicated in Fig. 2.
244 J. M. SAVAGE AND J. B. SLOWINSKI
in contact with red, separated from inner black band by yellow or
white (TTb)
10. tricolour striped (TS)
B. Ringed in black, red and usually yellow (or white)
1. black rings, interspaces red above yellow or white below
a. interspaces with red saddles (TZs)
b. interspaces uniform red dorsally (TZ)
2. tricolour monads: one black ring per red ring, separated from one
another by yellow rings (TM)
3. tricolour monads with accessory black elements (spots, bands,
saddles or rings): one black ring per red ring with each red ring
split by an accessory black element (TMa)
4. tricolour monads: black and/or yellow rings incomplete dorsally
(TMi)
5. tricolour monads with incomplete white (or yellow) rings partially
splitting black rings dorsally; red rings completely separated from
black rings by complete yellow (or white) rings (TMs)
6. tricolour dyads: two black rings per red ring, black rings in contact
with red rings and separated from one another by yellow (or white)
rings (TD)
7. tricolour dyads with incomplete yellow (or white) rings partially
splitting black rings dorsally
a. red rings contact black rings on both sides (Td)
b. red ring contacts black ring on one side, a yellow (or white) ring
contacts black ring on other side (tdte)
8. tricolour dyads with red rings incomplete ventrally and venter
mostly black (tdv)
9. tricolour dyads with incomplete red rings partially splitting black
rings lateroventrally; black rings separated by yellow or white rings
(td)
10. tricolour dyads with accessory yellow rings separating black from
red rings (Tda)
11. tricolour triads: three black rings for each red ring; outer black
rings in contact with red rings, separated from inner black ring by
yellow or white rings (TT)
12. tricolour tetrads: four black rings for each red ring; outer black
rings in contact with red rings, other black rings separated from one
another by yellow (or white) rings (TTE)
13. tricolour pentads: five black rings per red ring; outer black rings in
contact with red rings, other black rings separated from one
another by yellow (or white) rings (TP)
C. Ringed in white, yellow and red (NBR)
IV. QUADRICOLOUR
A. Saddles of red or orange outlined by black with a light margin on a
grey ground colour (Qs)
B. Banded with light-outlined red or orange centred dyads on a grey
ground colour (QDb)
C. Ringed by light-outlined red or orange centred dyads on a grey
ground colour (QD)
 THE COLOURATION OF THE VENOMOUS CORAL SNAKES 245

It should be noted that in many species of coral snakes and their mimics the
scales in the red areas have black tips. In certain cases the amount of black
pigment increases ontogenetically and suffuses the red areas, sometimes
obliterating the red entirely (e.g. bicolour Micrurus alleni). We have referred to
these patterns as secondarily bicolour patterns (IID above).

TAXONOMIC DISTRIBUTION IN AMERICAN SPECIES

Non-venomous or mildly venomous presumptive coral snake mimics

The following list is based on published reports and photographs and our own
observations on living or recently killed specimens. Principal literature sources
include, but are not restricted to: Campbell & Lamar (1989); Cei (1986);
Conant (1975); Dixon & Soini (1986); do Amaral (1976); Duellman (1978);
Freiberg (1982); Hecht & Marien (1956); Lancini (1979); Mehrtens (1987);
Mertens (1956); Moonen, Eriks & Deursen (1979); Perez-Santos & Moreno
( 1988); Pough ( 1988); Roze ( 1966); and Stebbins ( 1966).
Brattstrom (1955) and Pough (1988) provided lists of snakes having coral
snake patterns. We have found that their surveys are rather incomplete and
Pough's ( 1988; table 1 ) regional comparisons are consequently misleading. The
number of American species (1 15; exclusive of the venomous coral snakes)
potentially involved in this complex is really quite astonishing and approaches
18% of the total number of snake species found in the Americas. Of these about
90% have red in the colour pattern. We believe that we have included in this list
all authentic records of New World snakes having a coral snake pattern as
defined above. However, a number of species within the genera Apostolepis,
Atractus, and Elapomorphus that have not been described or illustrated from life
may prove to have substantial amounts of red in their colouration and might be
added to this listing.
An asterisk ( * ) indicates no red, orange or pink in the dorsal colouration.
Light areas other than the red ones in the tricolour patterns are usually yellow or
white. If some other light colour is involved it is noted. For species in which the
dorsal pattern does not continue onto the venter, the ventral colouration is
indicated. Codes separated by a comma indicate patterns found as variants on
different individuals of the same species (e.g. T M , TT). Codes separated by a
slash indicate variant patterns found on different parts of the body of one
individual (e.g. TM/Ttb).
Anilius
scytale-BIR
Apostolepis (black nuchal collar variably present in some striped forms)
ambiniger-Ucd; venter cream
ussimilis-Ucd; venter white
bum'oi-LS; venter cream
cearensis-Ucd; venter cream
dorbignyi-Ucd; venter white
erythronota--1s; venter cream with paired lateral black spots
fluvitorquata-LS, Ucd (rarely); yellow venter
ventrimaculatus--1s; venter cream with black crossbands
villaricae-LS; venter cream
Atructus
elaps-BIR, TD, TT, TTE
246 J. M. SAVAGE AND J. B. SLOWINSKI
etyfhromelas-BIB, BIS; venter red with black spots
guenfheri-BIB; venter cream
latifrons-TD, TT, T T E
obesus-Td
poeppingi-BIF
reticulatus-U; venter cream
Cemophora
coccinea-TDb, Tsr; venter white
Chilomeniscus
cinch-Tb; venter cream
Chionactis
occipitalis-TBr, TZs; often no red; venter cream or white
palarostris-TBr, TZs; venter cream
Clelia
juveniles of all species-Ucd; venter white
Dipsas
articulata-TM
bicolor-TZs
breuifacies-BIR
gaigeae-Tb
gracilis-BIR
pauonina BIB* anteriorly; venter tan
femporalis-BIR
fenuissima-B I R
uiguieri-BIR
Drepanoides
anomalus-Ucd; venter cream
Elaphe
guffafa-BIr, Tsr (rarely); venter blotched
Javirufa-Tsr
Elapomorphus (black nuchal collar variably present in striped forms)
lemniscafus-LS; ventrals black with light edges
lefiidur-Ucd, LS; venter white
rnerfmi-Ucd; venter white
puncfatw-LS, Ucd; venter white
tticolor-ucd; venter white
Erythrolamprus
aesculapii-BIR, TD, Tsy, TTE, Tda
buupnfhuisiz-TDltdte, TTE
bizona-TD
gumtheri-BID above/BIR below, T M
mimus-TM, TMs
pseudocorallw-TM
Farancia
abacura-BIF, BIFb adults
BIB, BIF, BIFb juveniles
cryfrogramma-TS
Geophis
brachycephalw-Brb, S; venter white to grey, usually banded with dark
cancellafur-BIB; venter cream
duellman-BIB (red or white); venter red or white
luficincfw-BIB; venter blackish
russafus-BIB; venter cream
scmidoliafus-BIB (white)/BIB (red); venter immaculate (whitelred) or mottled
zcledoni--S; venter uniform black
Gyalopion
quadranguluris-Tbi; venter yellowish
Helicofis
angulata-Tbx (tan)
 THE COLOURATION OF THE VENOMOUS CORAL SNAKES 247
HydrodFtes
bicinctu-TX, T X a
Hydrops
martii-TM
triangulum-BIR; venter red
Lampropeltis
getulw-BIB*, BIR* (some)
mexicanur-Tsr, QS, QDb, QD, BID*
pyromclunu-TD; red and sometimes white rings
triangulum-Tsr, TSR, TDb, tdv, td, TD, BIR, BID (red rings only)
zonutu-BID*, TD, td
Lcptodciru
rubric&-BIB; venter red
Liophis
unomulur-Tb (some individuals), othen with lateral black blotches as well; venter yellow
cpincphalur-BIB; venter with dark markings
frenata-BIF
poccilogyrus-TDb; venter with dark markings
Lystrophis
dorbigny-Tsb, TYv, BIF, TbR; most of venter red
histricus-TMb; venter red
smicinctus-TDb; venter mostly black
Nerodia
fyciah-U, RB, TSR (some individuals); venter blotched
Nma
sebue-TMb, BIS, Ucd, BIS/UcD; venter cream
Oryrhopus
formosus-U (adults), BIB; venter cream
leucomela-TZ; venter cream
mclunogcnys-TTb; venter cream
occipitulis-Ucd; venter cream
pctolurius-BIB (most individuals), TZs; venter cream
rhombifcr-Tbx, Tsb; venter cream
trigeminur-TTb; venter cream
vmezuclanur-TMb; venter cream
Pseudoboa
coronuta-Ucd in juveniles; venter cream
huai-RL, sometimes with a mid-dorsal stripe; venter cream
ncuwiedii-Ucd in juveniles and some adults; venter cream
Rhinobothryum
bovullii-TM
lmtiginosum-TM
Rhinochcilus
lecontci-BIB, TMb, Tsb; venter cream or white
Scuphiodontophis
annulutur-TDb; venter cream
vmwtissimur-TMb; venter cream
Scolecophis
atrocinctur-TZ; venter yellow
Sibon
anthracops-TM, TZ, TZs (no red on venter)
unnulutur-BIR, TD/TSR
Sirnophis
rhinostom-TT, TP, T P / T T
rohdei-TT
Siphlophis
ccrvinur-Tb, blotches often separated on mid-line; venter yellow
Sonoru
ucmulu-Uc, TZs, TM/TD/TT
248 J. M. SAVAGE AND J. B. SLOWINSKI

michoacanensis-Uc, BIR, TD, TD/td

semiannulata-Tb, T Z
Stenorrhim
fminvillii-Ud, LS (rarely); dark ventral markings
Tantilla
rubra-Ucd
supracinch-TDb; venter red
Thamnophis
Couchi-TbR
sirtalis-TbR
Tripanurgos
compressus-BIL, BIB
Tropidodipsas
annulifera-BIR
fasciata-BIR*, BIB*
Jlscheri-BIR*, BIB*
philippii-BIR*
sartorii-BIR, BIB
zweifcli-BIR*
Urotheca
elapoides-Uc, BIR, BIS, BIL, TM, TT, TTb, TMb, TMi
euryrom-TZ (no red on venter), BIR*, BIR
Xenopholis
scalari-BIB; venter cream
Among these genera Apostolepis, Clelia, Drepanoides, Elapomorphus, Oxyrhopus,
Pseudoboa, Rhinobothpm, Siphlophis, Sonora, Stenorrhina, Tantilla, Tripanurgos and
Xenopholis have grooved rear-fangs and a venom gland (Duvernoy’s gland).
Urotheca contains one striped species with grooved fangs, but the forms listed
above lack them although their bites are known to cause definite toxic effects in
human beings (Dunn, 1954; Seib, 1980). The putative coral snake mimics listed
for these genera are regarded as mildly toxic and not dangerous to human
beings, but their venoms are doubtless used to subdue prey. The listed species of
the genera Clelia and Erythrolamprus, like the venomous coral snakes, are
principally ophiophagous and Lampropeltis often consumes other snakes.

Venomous coral snakes

This review includes all species and putative subspecies listed by Roze (1983,
1987, 1989) and Campbell & Lamar (1989). When all subspecies have the same
pattern only the inclusive species name is listed. A double asterisk (**) indicates
that some representatives of that taxon have a tricolour monad pattern; a triple
asterisk (***) indicates that some have a bicolour ringed (BIR, BIM) or banded
(BIMb) pattern. Unless otherwise noted bicolour species have a black and red
pattern.

UNIFORM (U)
Micrurus diastcma alicnus (U, Uc)**
BICOLOUR (BI)

BLOTCHES RINGS
BIL Micrurus b d i * * * BIF Leptomirmrus collaris
BIL Micruw limbatus (some)*** BIF Lcptomicrurus narduccii
BIF Lcptomicrurus scutiuentfir
 THE COLOURATION OF THE VENOMOUS CORAL SNAKES 249

BANDS RINGS
BIB Micrurus bernadi BIM Micrurus alleni yatesi adult (black and
BIR Micrurus ephippijier ephippijier (black and yellow above, T M below) **
yellow above, T M below) BIM Micrurus annellatus annellatus adult (black
and white or red)**
BIM Micrurus annellatus balzani adult (black
and white or red)**
BIR Micrurus diastema apiatus
BIR Micrurus langsdorfi langsdorfi** (black
and white above, T M below; some
TM/BIR)
BIR Micrurus langsdorfi omtissimus (black and
white above, T M below)**
BIR Micrurus limbatus (some)
BIR Micrurus margaritijierus (black and white;
white discontinuous; venter black and
white or black, white and brown)
BIR Micrurus mipartitus (black and white, or
ye11ow )
BIR Micrurus multijiasciatus (black and white,
red or yellow)
BIR Micrurus multkcutatus
BIR Micrurus nigrocinctus divaricatus* *
BIR Micrurus nigrocinctus zunilmis* *
BIR Micrurus peruvianus (black and yellow) * *
BIR Micrurus petersi (black and yellow) *
BIR Micrurus psyches psyches (black and white
or yellow)
BIF Micrurus psyches donosoi
BIR Micrurus psyches medemi (black and
yellow) * *
BIR Micrurus putumaynrris (black and yellow)
BIR Micrurus rwtanus
BIR Micrurus spurrelli (black and white)
BIR Micrurus stewarti
TRICOLOUR MONADS (TM)
RINGS
Micruroides euryxanthus
Micrurus alleni alleni
Micrurus alleni yatesi juvenile* * *
Micrurus annellatus annellatus juvenile* **
Micrurus annellatus balzani juvenile* **
Micrurus annellatus bolivianus
Micrurus averyi
Micrurus bogcrti
Micrurus browni
Micrurus clarki
Micrurus corallinus
Micrurus diastema @nis
Micrurus diastema aglaeope (TMa)
Micrurus diastema alienus
Micrurus diastema diastema (some TMb)
Micrurus diastema macdougalli
Micrurus diastema sappm'
Micrurus d i s t m
Micrurus dumm'lii antioguiensis
Micrurus dumm'lii carinicauda
Micrurus dumm'lii transandinus
Micrurus dumm'lii venezuelmu
 250 J. M. SAVAGE AND J. B. SLOWINSKI
RINGS RINGS
M i m r u s ephippifcr zapotecus (TMa, black M ‘imcrus catamayenu
spots, bands or saddles in red rings) M ‘icrurus decoratus
Micrurus fulvius M ‘icrurus dissolnccus
Micrurus hippocrepis TM/TMi M‘icrurus dumm’lii dumcrilii
Micrurus langsdo@ langsdorfz (some M ‘icrurus dumm’lii colombianus
TM/BIR)*** M‘icrurusjlifomis
Micrurus langsdorfz omatissimus* * M‘ i m r u sfronhJkwiatus
Micrurus 1atiJiiciatus M‘icrurusfronhlis
M i m r u s mcrtm‘ M‘icrurus hcmprichii
Micrurus ncbularis M ‘icrurus ibiboboca
Micrurus nigrocinctw nigrocinctus M‘ i m r u s isozonus
Micrurus nigrocinctus babaspul M~ i m r u laticollaris
s
Micrurus nicrocinctus coibmris M i m r u s lcmniscatus
Micrurus nigrocinctus divaricatuss * * Micrurus sangilmis
Micrurus nigrocinctus mosguilmFis Micrurus spixii
Micrurus nigrocinctus zunilm’s* * Micrurus surinammis
M i m r u s pmvianus*** Micrurus tschudii
Micrurus petcrsi*** TRICOLOUR TETRADS (TTE)
M i m r u s proximans (None)
Micrurus psychs circinalis TRICOLOUR PENTADS (TP)
Micrurus psyches mcdcmi* * RINGS
Micrurus psyches paraensis Micrurus elcgans (black blotches in some
Micrurus psyches remotus orange rings)
Micrurus stdndachncri TRICOLOUR WHITE, YELLOW
Micrurus stuarti AND RED (NBR)
TRICOLOUR DYADS RINGS
(None) Micrurus langsdorfi langsdorfz**. * * *
TRICOLOUR TRIADS (TT)
RINGS
Micrurus bocourti

, Significantly, the head pattern and colouration in tricolour venomous coral

snakes generally corresponds to that on the body. In most species having the
tricolour monad pattern, the white or yellow head band or nuchal collar, black
nuchal collar and postnuchal light collar represent the first monad. Micruroides
euryxanthus is an exception for although it has a white nuchal collar it lacks a
black one and the first monad is located several head lengths posterior to the
nuchal region.
In typical tricolour triad forms the black nuchal collar is the anterior black
element in the first triad and the light head band or nuchal collar is usually red.
Micrurus dissoleucus, Micrurus laticollaris and some Micrurus spixii are unusual in
lacking red on the dorsum of the head.

COLOURATION AND MIMICRY

The 56 species of venomous coral snakes are essentially restricted to the

Neotropics in distribution, although one species (Micrurus fuluius) occurs in the
south-eastern United States and another (Micruroides euryxanthus) is found in the
southern arid regions of New Mexico and Arizona and in adjacent Mexico.
Members of this lineage are active at night and during the day under conditions
of low illumination (i.e. inside forests). They tend to be secretive and feed
primarily on other snakes (including their own species), although several South
American forms (Micrurus lemniscatus and Micrurus surinamensis) also eat
 THE COLOURATION OF THE VENOMOUS CORAL SNAKES 25 I

freshwater fishes and one (Micrurus hemprichii ortoni) prefers onychophorans

(genus Periputus). Known predators on coral snakes are primarily birds (see
Brugger, 1989) and snake-eating snakes. Available evidence indicates that
envenomation by coral snakes may kill large raptors but there are recorded cases
of successful avian predation on these snakes (Smith, 1969).
Many species of coral snakes exhibit defensive behaviours that are keyed to
reducing predation. Some or all of these behaviours may be used by a single
individual and include hiding the head under the body coils, flattening the body,
coiling the tail to expose the bright colours on the underside and waving the tail
about. In addition the head may be withdrawn from hiding and swung from side
to side with the mouth open and ready to bite any object that is contacted, the
body may be snapped back and forth almost spasmodically, the tail may be used
to strike aggressively or waved to divert attention from the head and the contents
of the cloaca may be discharged with or without a series of loud popping sounds.
The idea that the bright colouration typical of venomous coral snakes is also
an antipredator defense that performs an aposematic (warning) function and
that other similarly coloured snakes are mimics has been long and hotly debated
(Dunn, 1954; Brattstrom, 1955; Hecht & Marien, 1956; Mertens, 1956; Wickler,
1968; Grobman, 1978), although A. R. Wallace (1867, 1870) long ago
concluded that mimicry was involved. Contrasting patterns of alternating black
and light bands or rings are found in many species of snakes (Brattstrom, 1955).
In these forms, including the venomous coral snakes, the pattern serves a cryptic
function serving to hide them from predators (Brattstrom, 1955; Jackson,
Ingram & Campbell, 1976; Pough, 1976). However, there now seems to be little
question that the bright coral snake colour patterns combined with the unusual
behaviours described above are important antipredator defences that act
through the vision of potential predators on venomous coral snakes to warn them
that they may be subject to a serious venomous bite (Greene, 1988; Pough, 1988;
Campbell & Lamar, 1989).
In some cases, predator avoidance of coral snake colour patterns probably
involves learning based upon an unpleasant experience with a coral snake bite.
Because of their short fangs and the need to chew to ensure envenomation, coral
snakes frequently inject only small and not necessarily deadly amounts of venom
so that predators probably often survive bites (e.g. Smith, 1969). The carefully
designed experiments of S. Smith (1975, 1977), however, demonstrate innate
avoidance of a coral snake pattern by several potential avian predators,
including motmots (Eumomota) and kiskadees (Pitunow). Smith’s studies
convincingly established the reality of aposematic colouration in coral snakes
and their mimics.
Both Batesian and Miillerian mimicry have been proposed as explanations for
the coral snake mimicry guild. Additionally, Wickler (1968) popularized the
concept that the highly venomous coral snakes and the harmless mimics were
mimicking the mildly toxic, rear-fanged forms, a type of mimicry he called
“Mertensian”. However, we agree with Greene & McDiarmid (1981) that the
term is unnecessary since it represents a form of Batesian mimicry. That some
kind of mimicry is involved is attested to by the precise resemblances in
colouration between non-coral snakes and sympatric venomous coral snakes and
the concordant geographic variation between the two (Savage & Vial, 1974;
Greene & McDiarmid, 1981; Roze, 1983). We support the concept that Batesian
252 J. M. SAVAGE AND J. B. SLOWINSKI

mimicry is involved with the venomous coral snakes being the models and the
other forms the mimics. Nevertheless, it seems unlikely that all of the diverse
colour patterns described in this paper actually evoke an avoidance response in
predators. Consequently, we have attempted to identify those colour patterns
that are most likely to be involved in coral snake mimicry and the taxa which
may be regarded with some confidence to be members of the coral snake mimic
guild.
There can be little question that snakes having ringed tricolour patterns
(IIIB) are mimics since these patterns are identical to, or closely resemble the
most common colourations in venomous coral snakes. Similarly, tricolour forms
having the black body bands arranged in monads, dyads or triads (IIA 8-10)
resemble both tricolour ringed coral snakes and ringed mimics in their dorsal
patterns and are almost certainly mimics as well.
Most snakes exhibiting other non-ringed tricolour patterns are probably
mimics. However, we are sceptical regarding the role of the colour patterns in
Elaphe (Tsr, BIr) and Nerodia (U, TSR, RB) and do not consider the patterns in
Thamnophis and Farancia erytrogramma (TS) to be involved in mimicry since they
do not even superficially resemble the pattern of any venomous coral snake. The
several quadricolour patterns (IVA-C) are essentially modifications of tricolour
ones and snakes having them are considered mimics.
Snakes having bicolour ringed or banded patterns (IIB-C) of black and red
are doubtless mimics of similarly marked venomous coral snakes. The same may
be said for bicolour black and white or yellow ringed forms since they have
patterns typical of some bicolour venomous coral snakes. We are dubious as to
whether all black and white or yellow banded snakes (BIB*) are mimics since
this pattern has a cryptic function and is frequently found in arboreal forms or in
species that do not co-occur geographically with venomous coral snakes. The
mimetic role of all other bicolour black and red patterns is questionable although
those with a substantial amount of red may sufficiently resemble banded or
unicolour species to gain some protection.
Although only one species (Micrurus diastema) of venomous coral snakes
includes individuals that have a uniform red body, we believe that this
colouration serves an aposematic function in these individuals and in a series of
mildly toxic and harmless snakes. Consequently, we regard the latter as members
of the mimic guild although the suspected role of the uniform, bright red
colouration remains to be confirmed.
According to this analysis the following genera contain species that may be
regarded as coral snake mimics; an asterisk (*) means that all members of the
genus may be so considered: Anilius* , Apostolepis, Atractus, Cemophora* ,
Elapomorphus, Erythrolamprus*, Farancia, Geophis, Gyalopion, Hydrops*, Lampropeltis,
Leptodeira, Liophis, Lystrophis, Ninia, Oxyrhopus, Pseudoboa, Rhinobothryum*,
Rhinocheilus*, Scaphiodontophis*, Scolecophis* , Sibon, Simophis*, Siphlophis, Sonora,
Stenorrhina, Stilosoma* , Tantilla, Tripanurgos* , Tropidodipsas, Urotheca and
Xenopholis* .
Those individuals interested in further investigation of coral snake mimicry
should concentrate their attention on these genera and co-occurring venomous
coral snakes. More studies are certainly in order to determine how widespread
innate avoidance of coral snakes by predators may be. In addition, although
difficult to test, the role of empathetic learning in the origin and maintenance of
 T H E COLOURATION O F T H E VENOMOUS CORAL SNAKES 253

coral snake mimicry is an area needing further exploration. We suspect that both
innate and learned behaviours, each specific to different groups of predators, are
involved in predator avoidance of the coral snake mimic guild. Hopefully the
detailed description of colouration presented above will provide a basis for such
studies and precise evaluations of aposematic behaviour involving specific
models and their sympatric look-alike mimic(s).

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