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1. INTRODUCTION
Historical museum specimens are the basis of our
knowledge in taxonomy, and for the vast majority
of living things they are fundamental for any bio-
logical work, from systematics to ecology. In most
accredited natural history museums and collections,
specimens are properly curated and well cared for by
skilled and well-trained scientists and technicians.
Nevertheless, with technological advances in mole-
cular biology, coupled with the rapid disappear-
ance of natural landscapes around the world, new
approaches have led the scientic community to
underestimate the morphological characterization
and variation of species. is situation has aected
the practical foundations and taxonomic traditions
for most macroscopic biodiversity, and indirectly
has compromised museum collections and their
associated data.
Most of the Linnean type specimens of rep-
tiles have been placed in the Uppsala University
Museum of Evolution (Uppsala Universitet Evo-
lutionmuseet [UUZM], Uppsala) and the Swed-
ish Museum of Natural History (Naturistoriska
Riksmuseet [NRM], Stockholm). Because addi-
tional specimens may be housed in other European
collections (e.g., Saint Petersburg), this situation
sometimes complicates the task of undertanding
the early taxonomic literature.
Over the years, Swedish museum catalogs and
several works dealing with the Linnean specimens
that were handwritten or published (Hornstedt,
1788; Quensel, 1802; Swartz, 1808; Dalman, 1824;
Lovén, 1887; Lönnberg, 1896; Andersson, 1899,
1900; Löwegren, 1952; Holm, 1957; Madsen, 1959;
Bruno 1968; Wallin, 1977, 1985, 1992, 2001; Fern-
holm and Wheeler, 1983; Wheeler, 1985, 1991; Cro-
chet et al., 2006; Krecsák and Walgren, 2008) initially
may have served as the basis of Linnaeus’ knowledge
and the origin of the specimens avail able to him, and
later the historical context of some European zoolo-
gical collections.
e history of the rst formerly described New
World coralsnake, Coluber lemniscatus (= Micrurus
lemniscatus) (Linnaeus, 1758), is confusing with
regard to the origin and number of specimens avail-
able for its description. More than two centuries
aer its original description, Roze (1989) correctly
interpreted that Linnaeus (1758) never designated
an individual specimen as the type for this Ameri-
can coralsnake, and suggested assigning lectotype
and paralectotype designations for M. lemniscatus.
Roze (1989) stated that, “both syntypes that served
CHAPTER 2 HISTORICAL AND TAXONOMIC RELEVANCE OF
COLUBER LEMNISCATUS LINNAEUS, 1758
Matheus Godoy Pires1, Darlan Tavares Feitosa1,2, Felipe G. Grazziotin3,
Ana Lúcia da Costa Prudente4,5, Nelson Jorge da Silva, Jr. 1,2, and Hussam Zaher3
1 Curso de Biologia, Escola de Ciências Agrárias e Biológicas, Pontifícia Universidade Católica de Goiás, Goiânia, Goiás,
Brazil.
2 Programa de Pós-Graduação em Ciências Ambientais e Saúde, Escola de Ciências Médicas, Farmacêuticas e Biomédi-
cas, Pontifícia Universidade Católica de Goiás, Goiânia, Goiás, Brazil.
3 Museu de Zoologia, Universidade de São Paulo, São Paulo, São Paulo, Brazil.
4 Laboratório de Herpetologia, Museu Paraense Emílio Goeldi, Belém, Pará, Brazil.
5 Programa de Pós-Graduação em Biodiversidade e Evolução, Museu Paraense Emílio Goeldi, Belém, Pará, Brazil.
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38 M. G. Pires, D. T. Feitosa, F. G. Grazziotin, A. L. C. Prudente, N. J. Silva, Jr., and H. Zaher
Linnaeus to describe C. lemniscatus are housed in
the Naturistoriska Riksmuseet, Stockholm” and
noted specimen numbers NRS L-93 and NRS L-94,
followed by a brief description of each specimen
(now NRM 93 and NRM 94). Nonetheless, nine
Linnean specimens of M. lemniscatus were located,
of which three are housed in Stockholm, one in
Uppsala, and ve at the Zoological Museum of the
Zoological Institute (ZISP) (Russian Academy of
Sciences [RAS]), and in this chapter we provide their
redescription and some historical context related to
this fascinating subject, along with related taxono-
mic suggestions (see Chapters 4, 5).
In addition to the biological collections that
house the Linnean specimens (e.g., RAS), other col-
lections are extremely relevant. Together with the rst
publications of Linnaeus, they represent the system
that permeates our basic understanding of taxonomy.
We need to reason on the basis of taxonomy and the
description of new species, which did not change, but
rather is required to adopt a broader combination
of characters and analyses. Here, our intention is to
provide the available information on the specimens
and the historical taxonomic beginnings of the rst
coralsnake described, as well as a review of its closest
taxa (Micrurus l. lemniscatus, M. l. helleri, and M. l.
carvalhoi) and comments on related taxa (M. diutius,
M. frontifasciatus, and M. potyguara).
2. HISTORICAL CONTEXT
2.1. Carolus Linnaeus (1707–1778)
Carolus Linnaeus was a Swedish physician, botanist,
and naturalist who developed the binomial system
of nomenclature for animals and plants, and is refer-
red to as the “father of taxonomy.” e early descrip-
tions of plant and animals were vague and mostly
devoid of real scientic information, and at times
they explored bizarre forms of life to the detriment
of the relevant characteristics of the specimens. e
system developed by Linnaeus, which was based on
the root languages of Latin and Greek, immediately
revolutionized the natural sciences and established
the basis for scientic collections. Linnaeus became
popular in the country in a relatively short period
of time, especially with the nobility and many fellow
naturalists.
Linnaeus was born in Råshult (Småland, Swe-
den) in 1707. As a student, sponsors who recognized
his personal attributes and interest in botany came
to his aid. He began to study medicine at the Univer-
sity of Lund, a practice then common among natu-
ralists, but in 1728 he moved to Uppsala, where he
studied medicine for seven years. While in Uppsala,
his observations and interest promoted his academic
career, and in 1730 he began teaching botany as an
adjunct professor. During this time he made impor-
tant collecting trips to the northern (Lapland, in
1732) and central (Dalarna, in 1734) regions of Swe-
den, which served as the basis for several publica-
tions, including the famous Flora Lapponica (1737)
(Broberg, 2006).
In 1735, Linnaeus moved to the Netherlands
in search of his doctorate in medicine at Hadewijk
University. In Holland he met Albertus Seba and
the botanist Jan Frederik Gronovius, to whom he
showed a manuscript called “Systema Naturae.” With
the sponsorship of Gronovius and Johannes Burman
(another botanist), Linnaeus’ 1st edition of Systema
Naturae was published in 1735 (Broberg, 2006). At
that time, Holland had achieved the status of a world
trade center, due to the supremacy of the Dutch Com-
pany in the transoceanic trade, a fact that inuenced
Linnaeus and enabled a favorable environment for
publications and contacts with navigators, travelers,
important personalities, and scientists.
In 1736, Linnaeus visited the University of
Oxford (United Kingdom) and expanded his contact
with imminent scientists, and then returned to Ams-
terdam, where he remained until 1738. During these
years Linnaeus worked on several important publi-
cations, which can be considered as the base of his
taxonomic system, including Fundamenta botanica
(1736), Bibliotheca botanica (1736), Genera Planta-
rum (1737), and Classes plantarum (1738), as well
as Hortus Cliortianus (1737) and Ichtyologia (1738)
(Broberg, 2006).
In 1738, Linnaeus settled in Stockholm and
practiced medicine, and the following year, along
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with a group of businesspersons and politicians, he
participated in the creation of the Royal Swedish
Academy of Sciences (Kungliga Vetenskapsakade-
min [KVA]), designed in the Royal Society of Lon-
don (United Kingdom) and the Academye Royale
des Sciences of Paris (France). In 1741, he returned
to Uppsala and assumed the position of Professor
of Medicine and Botany, where he remained for the
rest of his life. His dedication earned him the title
of nobility, awarded in 1757 by King Adolf Friderik,
when he signed Carl von Linné or Carl Linné (Bro-
berg, 2006).
Linnaeus’ greatest contribution to zoology was
the establishment of rules for naming living organ-
isms, known as the binomial system, devel oped under
the inuence of preceding ideas and arrangements,
such as the notion of species formulated by John Ray
in his Catalogus plantarum circa Cantabrigiam nas-
centium (1660). In addition, the grouping of plants
according to their morphological characteristics
suggested the notions of gender and species in Cas-
par Bauhin’s work Pinax eatri Botanici (1671), of
which Linnaeus adopted various names, establishing
the hierarchical bases of the classication of living
organisms in kingdoms, classes, orders, genera, and
species (Fig. 1). As a result, Swedish museums still
retain the earliest specimens collected or curated by
Linnaeus or some of his students (sometimes refer-
red as the “Linnean specimens”).
e natural history collections of the Swedish
Museum of Natural History (NRM) and the Uppsala
University Museum of Evolution (UUZM) share
a common history, which is important to address
in order to understand the fate of Linnaeus’ type
specimens.
e Royal Swedish Academy of Sciences (Kung-
liga Vetenskapsakademien [KVA]) was founded in
1739, but did not obtain its physical installations
until 1753. e Naturhistoriska Riksmuseet (NRS
[old acronym]) was established in 1819 through a
merger of the KVA collections and the donations of
the collections of the Baron Gustaf von Paykull, and
later by Charles de Greer (Åhlander et al., 1997).
Linnaeus acted as the curator of the KVA collec-
tions until 1741, when he moved to Uppsala and brou-
ght several specimens that were incorporated into the
collections of the University of Uppsala. At least until
1750, he interfered with the donations to the KVA, by
redirecting biological material of his interest to Upp-
sala (Löwegren, 1952; Åhlander et al., 1997).
Linnaeus maintained excellent contacts and
inuence among the nobility, scientists, and explor-
ers, and his prolic life of publications also came
from providing descriptions for private collections.
e best example came from his major admirers and
sponsors: King Adolf Fredrik I and Queen Lovisa
Ulrika of Sweden. Both maintained natural history
cabinets, following the fashion of the nobility. King
Fredrik had his collection (about 1,100 bottles with
specimens in ethanol) at Ulriksdal Castle, and Queen
Lovisa at Drottningholm Castle (mostly shells,
corals, and insects). e nobles invited Linnaeus to
describe the royal collection, and between 1751 and
1754 he spent nine weeks at Ulriksdal Castle and
prepared a catalog of the king’s collection (Fernholm
and Wheeler, 1983).
While still a crown prince, King Adolf Fredrik
made some donations (duplicate specimens) to
the University of Uppsala collection (Fernholm
and Wheeler, 1983), where Linnaeus was in charge
of the natural history collections. is fact, along
with his personal charisma, served to attract many
Fig. 1. Carolus Linnaeus and the title page of Systema
Naturae (1758).
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40 M. G. Pires, D. T. Feitosa, F. G. Grazziotin, A. L. C. Prudente, N. J. Silva, Jr., and H. Zaher
donations to the university museum and to inspire
students who formed a group that also were known
as “Linnaeus’ apostles.” Linnaeus sent these apostles
in collecting expeditions in which they provided
natural history reports, to such places as southern
Asia, the Pacic islands, New Zealand, Australia,
South Africa, North America, and northern South
America (Suriname). For a long period, dierent
managers were in charge of the KVA collection, until
Anders Sparrman was hired as an eective curator
in 1776. Sparrman, one of Linnaeus’ apostles, exem-
plied Linnaeus’ inuence on the academic and
scientic life in Sweden, and might explain why so
many types were deposited between the NRM and
the UUZM (Åhlander et al., 1997). e importance
of these collections, collectors, and naturalists is
well documented in the fabulous publication e
Linnaeus Apostles: Global Science and Adventure
(Osbeck et al., 2009) (Fig. 2).
Most of these expeditions were carried out by the
Dutch East India Company (Vereenigde Oost-Indis-
che Compagnie [VOC]) and the Dutch West India
Company (Geoctroyeerde West-Indische Compag-
nie [WIC]), which demonstrates the results of the
contacts made by Linnaeus in Holland between 1735
and 1738, and also by the Swedish East India Com-
pany (Svenska Ostindiska Companiet [SOIC]), where
Linnaeus had built excellent relations. At that time,
Suriname was a Dutch colony and an important trad-
ing post of the VOC, and three of Linnaeus’ apostles
collected in this region (Carl Gustaf Dahlberg, Daniel
Rolander, and Pehr Löing). e collections were
analyzed and described by Linnaeus, as academic
dissertations by his students, and later were included
in a serial publication called Amoenitates Academi-
cae, with Linnaeus as the senior author (Löwegren,
1952; Åhlander et al., 1997; Broberg, 2006).
Fig. 2. e collecting travels of Linnaeus’ apostles. Map: Courtesy of Stig Söderlind.
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Advances in Coralsnake Biology: with an Emphasis on South America 41
Lönnberg (1896) noted important collections
donated to the UUZM, and categorized them as
follows: (a) collections described by Linnaeus and
cited in Systema Naturae (1758); and (b) collections
known by Linnaeus but not mentioned. Among the
collections described by Linnaeus, the ones included
in the two editions of Amoenitates Academicae (Lin-
naeus, 1749a, b) are of special interest. is publica-
tion includes four dissertations by Linnaeus’ students
that describe zoological specimens in important col-
lections. Only two, however, are of our interest: (a) the
dissertation by Barthold Rudolph Hast (elaborated
in 1745), Amphibia Gyllenborgiana, which describes
the collection donated by Carl Gyllenborg in 1744
and 1745; and (b) the dissertation by Peter Sundius
(elaborated in 1748), Surinamensia Grilliana, which
describes the collection donated by Claes Grill in
1746. ese dissertations are considered extremely
important, since they mention specimens of Coluber
lemniscatus, and they helped us understand part of
the historical context of these specimens.
Another important source of specimens for the
European collections came from Albertus Seba’s
cabinet of curiosities. Seba was a pharmacist and a
well-established businessperson in Amsterdam, who
used the inuence and power of the Dutch East and
West India companies (VOC and WIC) to obtain
countless specimens for his cabinet. His fame and
contacts led him to sell the rst great natural history
collection to Czar Peter I (e Great), the rst empe-
ror of the Russian Empire, in 1717. is collection,
together with the collection obtained from botanist
Frederik Ruysch, formed the basis of the collections
at the Russian Academy of Sciences (RAS) in Saint
Petersburg (Engel, 1937).
e remainder of Seba’s collection, illustrated in
his bilingual (Latin and Dutch) “esaurus” (Locu-
pletissimi rerum naturalium thesauri accurata des-
criptio – Naaukeurige beschryving van het schatryke
kabinet der voornaamste seldzaamheden der natuur
[Accurate description of the very rich thesaurus
of the principal and rarest natural objects] 1734,
1735, 1761, 1765), was auctioned aer his death in
1752. So, many of Seba’s specimens are housed in
collections in Sweden (Stockholm and Uppsala),
Great Britain, France, e Netherlands, Denmark,
and Russia (omas, 1906; Engel, 1961; Åhlander et
al., 1997) (Fig. 3).
Fig. 3. Albertus Seba in his cabinet of natural curiosities,
and the rst page of his monumental work, esaurus (1734,
1735, 1761, 1765).
2.1. e Linnean description of Coluber
lemniscatus
Although the 1st edition of Systema Naturae was
published in 1735 (while Linnaeus was studying in
Holland), Linnaeus actually did not consecrate the
famous binomial system until the 10th edition (1758).
Nonetheless, it was used in some of his earlier publi-
cations (Figs. 9, 10 [description of King Fredrik’s
collection in 1754]). e 10th and 11th editions of
Systema Naturae (Linnaeus, 1758, 1766) are, in fact,
the lists of species previously described by Linnaeus
or his disciples in pre-Linnean works. In these publi-
cations Linnaeus divided the order Serpentes into
six genera (Crotalus, Boa, Coluber, Anguis, Amphis-
baena, and Caecilia), with Coluber being the richest
species and containing the rst formally recognized
description of an American coralsnake (Coluber
lemniscatus Linnaeus, 1758).
In the 10th edition of Systema Naturae (Linnaeus,
1758), he included Coluber lemniscatus (number
285), followed by the specimen’s ventral and subcau-
dal counts (250 and 37, respectively). In the rst two
publications (pre-binomial or PreLinnean) it was
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42 M. G. Pires, D. T. Feitosa, F. G. Grazziotin, A. L. C. Prudente, N. J. Silva, Jr., and H. Zaher
available under the name Coluber in the Amoenita-
tes Academicae, cited in the two rst lines of entry
(Amoen. acad. I. p. 118. n.6. [Linnaeus, 1749a];
and p. 413 n.9. [Linnaeus, 1749b]), followed by the
mention of an illustration of a specimen from King
Adolf Friderik collection (Mus. Ad. Fr. 1. p.34 t.14
f.1. [Linnaeus, 1754]), and two illustrations of Seba’s
esaurus (1734) (Seb. mus. 1. t.10 f. ult. and 2. t.76.
f.3). e type locality is noted as “Asia,” and included
a brief morphological description (Fig. 4).
e description of Linnaeus’ Coluber lemnisca-
tus (1758) is the primary source we used to assess
the early literature on the type specimens involved
in the description of the rst New World coralsnake.
Nonetheless, we need to understand the history
behind the descriptions of the pre-Linnean phase
(any publication printed before 1758) by using the
C. lemniscatus entry as a reference (Fig. 4).
Fig. 4. e Coluber lemniscatus entry in the 10th ed. of Lin-
naeus’ Systema Naturae clearly shows, aer the name of the
species, the reference to the ventral and subcaudal scale
counts of “250—37” obtained from the rst described spec-
imen.
2.1.1. e rst Linnean specimen of Coluber
lemniscatus (in Amoenitates Academicae)
e rst description of a specimen of Coluber lem-
niscatus by Linnaeus was published in 1745, in the
dissertation Amphibia Gyllenborgiana, defended by
Barthold Rudolph Hast and reprinted in Amoenita-
tes Academicae (Amoen. Acad. Vol. I, n° V, p.107)
(Linnaeus, 1749a; Figs. 5, 6). is specimen is men-
tioned in the rst entry of the Systema Naturae (Lin-
naeus, 1758; Fig. 4).
At that time, Linnaeus apparently only knew
of a single specimen of this species. is particular
specimen was part of the donation made by Carl
Gyllenborg to the University of Uppsala in 1744, and
was designated as “COLUBER scutis abdominali-
bus CCL, caudalibus XXXV.” It was followed by two
entries from Seba’s esaurus, and a third from the
Musei Imperialis Petropolitani, Russia (Linnaeus,
1749a). Moreover, the description of the specimen is
very detailed and characterizes the head, trunk, and
tail, as well as the coloration and length (1+ ½ foot
= sesquipedalis); it also reports 43 white body rings
with faded dark rings (Fig. 7).
Fig. 5. e disertation Amphibia Gyllenborgiana, defend-
ed by Barthold Rudolph Hast on 18 June 1745.
Fig. 6. A = the rst page of Amoenitates Academicae Vol -
ume I; and B = the rst page of the reprint of Barthold
Rudolph Hast’s thesis (Amoen. Acad. Vol. I, n° V).
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Fig. 7. e description of Gyllenborg’s specimen in Amoe-
nitates Academicae Vol. I, n° V, pages 118–119, entry 6.
2.1.2. e second Linnean specimen of Coluber
lemniscatus (in Amoenitates Academicae)
e second description of a specimen of Coluber lem-
niscatus was provided to Linnaeus in 1746, through
a donation made to Uppsala University by Claes
Grill. is collection, which originally consisted of
24 specimens collected in Suriname by the farm-
er Gerret and his son, included mammals, birds,
amphibians, reptiles, and invertebrates (Löwegren,
1952). Unfortunately, over the years the collection
suered several losses, and Holm (1957) reported
just nine remaining specimens and provided the last
known report of Grill’s specimen of C. lemniscatus,
from the 1828 Uppsala manuscript catalog. is spe-
cimen is mentioned in the second entry of Systema
Naturae (1758; Fig. 4).
e original description of Grill’s specimen of
Coluber lemniscatus was presented by Linnaeus in
the dissertation Surinamensia Grilliana, defended
by Peter Sundius in 1748 and reprinted in Amoeni-
tates Academicae (Amoen. Acad. Vol. I, n° XVI, p.
483) (Linnaeus, 1749b). is second specimen was
described in a concise manner, and under the nomi-
nation according the previously mentioned work
(Linnaeus, 1749a): “9 COLUBER scutis abdomina-
libus CCL, caudalibus XXXV, Amphib. Gyllenborg.
n.6.” Grill’s specimen had 248 ventral scales (“Mag-
nitudo, color, omnia ut in Gyllenborgiano, sed scuta
abdominalia circiter ducenta & quadraginta octo”)
but no mention of the tail scales, perhaps because
they were assumed to be the same as in Gyllenborg’s
specimen (Fig. 8). Curiously, the reference to this
specimen in the 10th edition of Systema Naturae
(Linnaeus, 1758) is inaccurate, as it indicates page
413, but the correct is page 493 (Fig. 4). Since its last
report (the 1828 Uppsala manuscript catalog), this
specimen is presumed lost, and its whereabouts are
unknown (Holm, 1957).
Fig. 8. e description of Grill’s specimen in Amoenitates
Academicae I. A = title page (483p.); and B = the descrip-
tion of Coluber lemniscatus (here referred only as Coluber)
under number 10 (493 p.).
2.1.3. e specimens in King Adolf Fredrik’s
collection
e specimens in King Adolf Fredrik’s collection
were documented by Linnaeus at Ulriksdal in 1753,
and published in 1754, and included an impressive
array of mammals (Quadrupedia), birds, amphi-
bians (amphibians and reptiles), shes, insects, and
worms, described in Latin and Swedish (Linnaeus,
1754; Fig. 9). It included a brief description of
ve specimens of Coluber lemniscatus, and appar-
ently only the rst specimen was deposited in the
NRM (244-35 scale counts). e second specimen
(250-35 scale counts) may be referred to as the
Gyllenborg’s specimen (Fig. 7), followed by the de-
scription of three specimens referred to as “α”, “β,”
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44 M. G. Pires, D. T. Feitosa, F. G. Grazziotin, A. L. C. Prudente, N. J. Silva, Jr., and H. Zaher
and “γ”. It gives a general statement of the coloration,
followed by an account of the number of ventrals and
subcaudals. According to Linnaeus (1754), three (or
four) specimens of C. lemniscatus were housed in the
royal collection at Ulriksdal (Fig. 10). e use of the
binominal Linnaean system in this publication (i.e.,
Coluber lemniscatus) is noteworthy, as it appeared
ve years before the publication of Systema Naturae.
Fig. 9. e opening pages of Linnaeus’ publication de-
scribing King Fredrik’s collection (1754).
Fig. 10. Linnaeus’ account and description of three spec-
imens of Coluber lemniscatus housed at Ulriksdal Castle).
Based on Linnaeus’ citation of the Amoenitates
Academicae, a publication that in principle dealt only
with the material housed in Uppsala, these addi-
tional specimens (α, β, and γ) were expected to be
in the UUZM collection. When Linnaeus described
the collection of King Adolf Fredrik I (Linnaeus,
1754), the monarch had not donated it the Swed-
ish Royal Academy, but instead had transferred it
from Ulriksdal Castle to Drottningholm Castle. e
collection remained there for many years aer his
death, and thus the three reference specimens were
not in the UUZM collection.
In 1801, only aer the insistence of Quensel,
King Gustav IV (the grandson of King Adolf Fredrik
I) transferred the Drottningholm collection to the
Royal Swedish Academy of Sciences by royal decree,
and also sent a collection to the museum of Uppsala
University (Åhlander et al., 1997). Several impor-
tant specimens were included in the parcel sent to
Uppsala, among which was a specimen of Coluber
lemniscatus (which might be the one we found at the
UUZM collection [see ahead]).
e “α” specimen was described as having 250
and 30 (ventrals and subcaudals, respectively, here
and below), the “β” as having 244 and 35, and the “γ”
specimen as having 241 and 37 (Fig. 8). In our opinion
the “γ” specimen, described by Linnaeus as having
241 and 37, is the same that Andersson (1899) re-
ported as having 240 and 37, and in our analysis as
having 238 and 36 (possibly specimen NRM 93). We
belive that the “β” specimen, described by Linnaeus
as having 244 and 35, is the one Andersson reported
as having 243 and 35, and in our analysis as having
242 and 37 (possibly specimen NRM 94); interest-
ingly, it has the same scale counts as in the rst entry
of this publication (244 and 35). e second entry
represents a reference to Gyllenborg’s specimen
(Figs. 6, 7). We could not nd a match for the “α”
specimen within the available material at NRM.
2.1.4. e specimens in Seba’s esaurus
As pictorial references for this species, Linnaeus
(1749a) cited Seba’s reports of similar snakes, and
specically “Serpentula ex ins. Ceilon” (Seba, 1734,
Tome I, page 19, plate 10, last g.), “Serpens Ceilo-
nica lemniscis latis” (Seba, 1735, Tome II, page 76,
plate 76, g. 3) (Fig. 7), and provided a brief descrip-
tion of specimens from the Kunstkamera, referred
to in Anonymous (1742; see ahead). Linnaeus met
with Albertus Seba twice in 1735, while he was stu-
dying in the Netherlands, but there is no evidence
that he provided a description for Seba’s specimens.
e reference to Seba’s plates makes these two
illustrations iconotypes for Coluber lemniscatus, as
well as another specimen possibly housed in the
Kunstkamera, in Saint Petersburg, at the time. e
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Kunstkamera specimen(s) probably came from the
acquisition of Seba’s specimens by Czar Peter I (e
Great; see ahead).
In the original description of Coluber lemnisca-
tus (Linnaeus, 1758; Fig. 4), both entries in Seba’s
esaurus are based solely on the illustrations in two
dierent volumes: (A) “Serpentula ex ins. Ceilon”
(Seba, 1734; Seb. esaur. I. pag. 19. tab. 10. g. ult.),
which deals with plants and includes two snakes
as “decoration”; both illustrations resemble coral-
snakes, but they could be interpreted as specimens
of Bungarus fasciatus (Fig. 1) and C. lemniscatus
(Figs. 2, 11); and (B) “Serpens Ceilonica lemniscis
latis” (Seba, 1735; Seb. th. T.II. pag. 76. g. 3); simi-
larly, gure 3 might be an excellent representation of
a specimen of C. lemniscatus (Figs. 12, 13).
Fig. 11. A coralsnake depicted in Seba’s esaurus, vol-
ume 1, tabula X, page 19 (1734).
Curiously, in Tomus II of the Seba’s esaurus
(1735), three additional illustrations resemble speci-
mens of Micrurus lemniscatus. In plate VI, gure 2,
the depiction of a coralsnake resembles a specimen
of M. surinamensis. e possibility of an error in the
legend, however, is noticeable. In this plate, gure
1 is described as an “Ibiboboca,” a Brazilian Indian
name for coralsnake, which we believe is mislabeled,
with gure 2 being the correct gure (Figs. 14, 15). It
reads, “Serpens, Ibiboboca; seu Serpens, perelegans,
Brasiliensis,” which clearly indicates a specimen
from Brazil, possibly based on the early publication
of Willen Piso (1648) entitled Historia Naturalis Bra-
siliae, where in two dierent chapters he and Georg
Marcgrave (Georg Marggra) wrote about the coral-
snake Ibiboboca (see Introduction in the book).
Fig. 12. A coralsnake depicted in Seba’s esaurus, vol-
ume 2, tabula LXXVI, gure 3–bottom right (1735).
Fig. 13. e text of tabula LXXVI depicted in Seba’s e-
saurus, volume 2, tabula LXXVI–text (1735). Note: “Num.
3. Serpens, Ceilonica, lemniscis latis” (Pages 79 and 80).
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46 M. G. Pires, D. T. Feitosa, F. G. Grazziotin, A. L. C. Prudente, N. J. Silva, Jr., and H. Zaher
Fig. 14. A coralsnake depicted in Seba’s esaurus, vol-
ume 2, tabula VI, gure 2 (1735). Note the similarity of
the second illustration with Micrurus surinamensis.
Fig. 15. e text related to Seba’s esaurus, volume 2,
tabula VI (1735). We believe this is a labelling error, and
that the correct caption is in Num. 1. Pages 7 and 8.
A second example is plate IX, gure 3, with an
illustration that resembles a coralsnake that in the
text is referred to as “Serpens” (Fig. 16). A third
example (possibly the best) is in text + plate XXVII,
gure 2, (“Serpens anulatus”), which easily could
be interpreted as a specimen of Coluber lemniscatus
(Fig. 17).
Fig. 16. A coralsnake depicted in Seba’s esaurus, volume
2, text + tabula IX, gure 3 (1735). Page 28 and Tabula IX.
Fig. 17. A coralsnake depicted in Seba’s esaurus, vol-
ume 2, text + tabula XXVII, gure 2 (1735).
A nal illustration apparently depicts a speci-
men of Micrurus surinamensis. e caption (Num.
1. Serpens, Aesculapius, Orientalis, maximus) and a
closer look, however, perhaps make it easier to iden-
tify the snake as Erythrolamprus aesculapii (Figs. 18,
19). Linnaeus did not mention these illustrations,
and Schneider (1801) later associated some with this
species. Apparently, Linnaeus had no information
on the collecting site of this specimen.
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Fig. 18. A possible coralsnake depicted in Seba’s esau-
rus, volume 2, tabula LXXXVI, gure 1 (1735). e snake
in this illustration resembles a Micrurus surinamensis, or
most likely, an Erythrolamprus aesculapii.
Fig. 19. e text related to Seba’s esaurus, volume 2,
tabula LXXXVI. Pages 91 and 92.
2.1.5. e specimen from Saint Petersburg
e textual reference “Serpens americanus niger,
annulis latis albis, ac inter ipsos singulos duo annuli
albi tenuiores, ore squamularum albarum nigricante.
Mus. Petropol. p. 464, n. 218” in Linnaeus’ (1758)
description may be related to a specimen that pro-
bably was acquired by Czar Peter I (e Great)
in Amsterdam from Albertus Seba and Frederik
Ruysch (Fig. 20). ese collections constituted the
Kunstkamera (originally a cabinet of curiosities,
established in 1727), or the zoological specimens of
the Saint Petersburg Academy of Sciences (today, the
Russian Academy of Sciences [RAS]).
Fig. 20. Anonymous (1742) Musei Imperialis Petropo-
litani frontispiece (above), and an excerpt from page 464
showing entry 218, which was mentioned in Linnaeus
(1749a).
Nevertheless, ve specimens of Micrurus lem-
niscatus originally were stored in the Kunstkamera,
and later were deposited in the Zoological Museum
(now Zoological Institute of the Russian Academy
of Sciences [RAS]; N. L. Orlov and K. Milto, pers.
comm.), which made it dicult to match the brief
color description of the Nr. 218 entry (Fig. 17) to any
of the specimens housed at the RAS (see ahead).
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48 M. G. Pires, D. T. Feitosa, F. G. Grazziotin, A. L. C. Prudente, N. J. Silva, Jr., and H. Zaher
2.1.4. On the type locality of Coluber lemniscatus
With regard to the type locality for this species, Lin-
naeus evidently committed an error by indicating
“Asia” (Linnaeus, 1758; Fig. 4). A possible origin of
this mistake may be the allusion to Ceylon by Seba
(1734) as the collecting site for some of his speci-
mens (see Section 4).
2.1.5. Color pattern
e original text (Linnaeus, 1758) also contains
minimal information on the color pattern of Coluber
lemniscatus: “Fasciae albae nigraesque, hae annu-
lis duabus saepe interruptae. Corpus glaberrimum.”
Since Linnaeus used his previous publications as
the main references for the 10th Edition of Systema
Naturae, the best description of the color pattern
of C. lemniscatus appears in the description of
Gyllenborg’s specimen (Fig. 7).
2.2. Current Linnean specimens of Micrurus
lemniscatus at the NRM
ree Linnean specimens of Micrurus lemniscatus
are housed at the Naturistoriska Riksmuseet (NRM)
in Stockholm, which currently are cataloged as NRM
93, NRM 94, and NRM 95. e specimens labeled
NRM 93 and NRM 94 are the ones designated as the
lectotype and paralectotype by Roze (1983). Previou-
sly, however, Linnaeus (1754) and some later cura-
tors noted a third specimen, but nobody associated
this specimen (NRM 95) as a type of M. lemniscatus.
e curators of the KVA, and later the NRM,
compiled several catalogs. Clas Fredrik Hornstedt
followed Anders Sparrman as curator of the KVA,
and in 1788 he compiled the oldest known catalog of
the collection (“Museum Regiae Academiae Scientia-
rum Svecicae. Pars Prima. Quae Spectat Regnum Ani-
male”) and listed two specimens of Coluber lemnis-
catus without museum voucher numbers. Initially,
Hornstedt did not report the number of specimens
of C. lemniscatus, but aer his initial eort someone
with distinct calligraphy added the information
(Fig. 21). Whether this information was added to
the catalog before or aer the 1801 donation of the
Drottningholm specimens to the KVA is unknown
(Hornstedt, 1788; Löwegren, 1952; Åhlander et al.,
1997). Whether these specimens correspond to the
Linnaeus denition of C. lemniscatus is another
mystery, since Hornstedt (or someone else) appa-
rently raised some doubt about its identication
by placing a question mark aer its denomination
in the catalog (Fig. 21), and at that time specimens
of dierent ringed snakes could have been confused
with C. lemniscatus (e.g., C. aesculapii and Hydrops
triangularis).
In 1802, Conrad Quensel prepared a catalog in
which the snakes were listed in a separate volume
(“Catalogus Amphibiorum Musei R:iae Ac:miae
Scientiarum Holmiensis. Catalogus Musei Zoolo-
gici Pars 2da Quensel. Holmiae Anno 1802”). e
snakes were included in a separate volume (Pars 2da,
Amphibia). Quensel listed three specimens of Colu-
ber lemniscatus under number 51 of this catalog (Fig.
22), and associated at least one of the specimens with
the material from Drottningholm (Quensel, 1802;
Åhlander et al., 1997).
Fig. 21. e Coluber lemniscatus entry in the manuscript
catalog of the KVA, written by Hornstedt in 1788.
Fig. 22. e Coluber lemniscatus entry in the manuscript
catalogue written by Quensel in 1802, the rst report of
this species at the NRM. At least one specimen is linked
to Musei Adolphi Friderici Regis, which arrived from
Drottningholm aer 1801.
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Olof Peter Swartz succeeded Quensel as curator
of the NRM, and compiled a catalog in 1808 (“Cata-
log öfver Kongl. Svenska Wettensk. Academiens
Museum, författad 1808, af O. Individual: 1268 Spec.
Distincta = 905”). Under Coluber, he noted three
specimens of C. lemniscatus (Fig. 23) with the same
comments made by Quensel about one specimen
having arrived from the Drottningholm collection
(Swartz, 1808; Åhlander et al., 1997).
Fig. 23. e Coluber lemniscatus entry in the manuscript
catalogue written by Swartz in 1808, repeating the infor-
mation provided by Quensel in 1802.
Johan Wilhelm Dalman compiled another cata-
log in 1824 “Amphibia Musei Holmiensis Secundum
Systemata Cuvierii et Merremii disposita. (Enume-
rantur tantun Species in Catalogo Svartzii [ab anno
1808]),” in which he indicated three specimens of
Coluber lemniscatus, aer adopting the name Elaps
lemniscatus (Dalman, 1824; Åhlander et al., 1997;
Fig. 24).
Fig. 24. e Elaps lemniscatus entry in the manuscript cat-
alogue written by Dalman in 1824.
Lars Gabriel Andersson wrote two separate
catalogs that were published, one on the type spe-
cies of snakes (1899) and the other on the reptiles
(1900) housed at the NRM. In his catalogs, Ander-
sson commented on each species and on individual
specimens, and made comparisons with Linnaeus’
papers. He indicated two bottles with two specimens
in each, both labeled as Coluber lemniscatus. One
of the bottles, however, contained two specimens of
Hydrops triangularis, which suggests a labeling error
in the collection (Fig. 25). e two specimens of C.
lemniscatus noted by Andersson (1899) are the same
ones labeled NRM 93 and NRM 94 (see sections
2.1.3. and 3.1.).
Fig. 25. e specimens of Hydrops triangularis (NRM 58a
and NRM 58b) maintained in a bottle labeled as Coluber
lemniscatus. Photos: Site of the NRM as the Adolphi Frid-
erici Collection.
Considering the early works of Linnaeus
(1749a,b, 1754) and Seba (1734), we can speculate
that Linnaeus’ knowledge of Coluber lemniscatus
was not based only on the two specimens designated
by Roze (1989) as the lectotype and paralectotype
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50 M. G. Pires, D. T. Feitosa, F. G. Grazziotin, A. L. C. Prudente, N. J. Silva, Jr., and H. Zaher
(NRM specimens), but a third specimen in the
UMMZ collection perhaps is another type that we
found, which can be traced back in the cloudy his-
tory of these specimens. Later, we obtained the infor-
mation for the Russian specimens that also were part
of Linnaeus’ entries in Systema Naturae.
As stated by Åhlander (2009) in the remarks of
the NRM database, the two specimens (NRM 93 and
NRM 94) are labeled as having arrived from Drott-
ningholm, thus from Ulriksdal Castle, and should
have been two of the three specimens described by
Linnaeus in 1754.
e third Linnean specimen of Coluber lemnis-
catus at the NRM (NRM 95) is the one Andersson
(1899) noted as having been found mixed among
specimens of C. aesculapii. Andersson determined
this specimen to be Elaps lemniscatus, and provided
the following description: (total length = 385 mm;
tail length = 55 mm; middorsal scales = 15; ventrals
= 261; and subcaudals = 35). e ventral and sub-
caudal counts, as well as the tail length, agree with
our data, but the total length obviously is incorrect.
When we visited the NRM, in addition to the
type specimens designated by Roze (NRM 93 and
94), we found specimen NRM 95 marked as a type
assigned to the Linnean series. None of the counts
on the NRM specimens are compatible with those
noted by Linnaeus (1754), and none of the speci-
mens contained a black round mark on the nuchal
region, such as in the Uppsala specimen (see ahead).
Our measurements and counts for specimen
NRM 95 are as follows: total length = 706 mm; tail
length = 56 mm; ventrals = 261; and subcaudals =
35. Oddly, this particular specimen also has a Drott-
ningholm label, as well as a Dalman’s label, and cur-
rently is regarded as a syntype of Coluber aesculapii,
as it was stored among specimens of this species;
however, we consider it unlikely that Linnaeus did
not recognize the dissimilarity of this specimen. In
the past, perhaps this specimen was mixed in with
specimens of C. aesculapii, especially because none
of the descriptions provided for this species by Lin-
naeus apply to this specimen (scale counts and a pat-
tern of paired rings). Further, this specimen actually
is a Micrurus lemniscatus, and may be a replacement
for a lost specimen (Figs. 26, 27).
Fig. 26. A specimen of Coluber lemniscatus (NRM 95)
found mixed in with specimens of C. aesculapii. Photo:
Site of the NRM as the Adolphi Friderici Collection.
Fig. 27. Specimens of Coluber aesculapii (NRM 84 and
NRM 85) mixed in with the specimen of Micrurus lemnis-
catus (NRM 95). Photos: Site of the NRM as the Adolphi
Friderici Collection.
2.3. e Linnean specimen of Micrurus
lemniscatus at the UUZM
When we analyzed Linnaeus’ work describing the
collection of King Adolf Fredrik I, we noted that
Table XIV shows a Coluber lemniscatus swallowing
another vertebrate, possibly a Typhlonectes (Fig.
28). is specimen would have been unmistakable
among the specimens examined by Linnaeus, since
it is the only one with a black, round spot in the
region between the parietals and the anterior ring of
the rst triad.
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At the UUZM museum in Uppsala Univer-
sity, we found a specimen of Micrurus lemniscatus
(UUZM 06) that in the collection was identied as
assiged to the type series of M. lemniscatus (Lin-
naeus, 1758). Although the specimen had faded, the
scale counts show 250 ventrals and 34 subcaudals,
which are compatible with those in the original de-
scription of the species (Fig. 4), and it also contains
a marking identical to the specimen in King Adolf
Fredrik’s collection that was illustrated and de-
scribed by Linnaeus (1754; Fig. 24), which corrobo-
rates the history of the partial transfer of the King’s
collection to Uppsala (see 2.1.3.).
Lönnberg (1896) described the type-specimens
of vertebrates housed at the UUZM. Under entry
number 6 he described a specimen of Elaps lemnis-
catus with a reference of Amoenitates Academicae
I, number V, page 118, entry 6 (Fig. 7). In his own
words: “is species of snake has been recognized
by all later authors from Linnaeus’ description and
gure of the same, given in Museum Regis Adolphi
Friderici tab. XIV g. 1. e black spot on the occi-
put of this Gyllenborg’s specimen is slightly larger
than that on the gure mentioned, but rather smaller
than on Jan’s gure 1 plate V of Livraison 42” (Figs.
28, 29).
e comment on Jan and Sordelli’s specimen by
Lönnberg might lead to dierent interpretations. Jan
and Sordelli (1872) noted that two specimens used
in volume 42 (Atlas) of their iconography came from
Pernambuco, Brazil (Musée de Hamburg), and Suri-
name (Musée de Munich), respectively. Lönnberg’s
comment has to be interpreted as a comparison of a
similar black marking on the occiput of both speci-
mens. One may interpret that Jan and Sordelli’s spe-
cimen is the same one used in Linnaeus’ description
of King Adolf Fredrik’s collection (1754). Jan and
Sordelli’s gure is quite dierent from the one depic-
ted in Figure 24 of Linnaeus (1754); see Fig. 28, in
which the black marking is much smaller and loca-
ted slightly on the le side of the head (see Fig. 29).
e same entry number (6), also listed by
Holm (1957), is the current collection number at
the UUMZ (UUMZ 06), which makes it the rst
specimen of Micrurus lemniscatus described in a
pre-Linnean publication, and originally linked to the
collection of King Adolf Fredrik I (Fig. 30).
Fig. 28. Table XIV of the Museum Regis Adolphi Frideri-
ci, gure 1 (Linnaeus,). A specimen of Coluber lemniscatus
swallowing what appears to be a typhlonectid.
Fig. 29. Elapiens. Iconographie Genérale des Ophidiens
(Jan and Sordelli, 1864). Tome Troisème, 42º Livraison,
plate V, gure 1.
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52 M. G. Pires, D. T. Feitosa, F. G. Grazziotin, A. L. C. Prudente, N. J. Silva, Jr., and H. Zaher
3. THE TYPE SPECIMENS
In this section we divide the specimens we accessed
in loco, and some that later were revealed during the
course of our studies, but did not have a chance to
examine properly. Despite the signicance of the
UUZM 06 specimen as the rst described Linnean
specimen of Micrurus lemniscatus, we did not con-
sider it necessary to change the designation of the
actual lectotype (Roze, 1983) and oer it as a new
lectotype, since all of the specimens somehow are
linked to Linnaeus.
3.1. e specimen at the Uppsala University
Museum of Evolution (UUZM)
UUZM 06—new paralectotype (Fig. 30): An adult
female, with 4 preventrals, 250 ventrals, and 34
subcaudals; the eye diameter is shorter than ½ of
its distance from the border of the mouth; the ros-
tral is higher than wide; the internasals are ⅔ the
length of the prefrontals; the frontal is hexagonal,
1⁄ times longer than wide, as long as its distance
from the snout, and wider anteriorly; the parietals
are as long as their distance from the snout; the
preoculars = 1/1; the postoculars = 2/2; the tem-
porals = 1+1/1+1; and the 3rd and 4th supralabials are
in contact with the eye. Dorsally, the snout (rostral,
anterior nasals, internasals, anterior ⁄of prefront-
als, and the 1st and anteriormost part of the 2nd pair
of supralabials) is black, without any visible white
markings; a white preocular band behind the black
snout includes the posterior ⁄of the prefrontals, the
anterior part of the preoculars, the posterior nasals,
and the posterior portions of the 2nd and 3rd pairs
of supralabials. A black head cap follows the white
preocular band over the frontal and supraoculars,
the posterior portion of preoculars, the anterior
⅓ of the parietals (until the level of the posterior
extreme of the frontal), the postoculars, the anterior
temporals, and the 4th, 5th and anterior portion of
the 6th supralabials. A round black marking covers
the posterior ⅓ of the parietals, the 1st and anterior
portion of the 2nd vertebral scales, the anterior ⅔ of
the 1st adjacent paravertebral scales, and the medial
margins of the anteriormost scales of the adjacent
parvertebral scale rows. e remaining surface
of the head is red. Ventrally, the head appears to
be entirely red, including the four preventral and
the 1st ventral scales, except for the upper margins
of the 1st–4th le infralabials. e body triads = 13
+ 1⅔ on the tail. e anterior black ring of the 1st
triad is separated from the parietals by ve ver-
tebral scales. e middle black ring is longer than
the external ones. e white rings are much shorter
than the black external rings. e total length (ToL)
= 480.8 mm; the head length (HL) = 14.35 mm; the
snout–vent length (SVL) = 445 mm; and the tail
length (TL) = 35.8 mm. e TL/SVL ratio = 0.080.
Fig. 30. A Linnaean specimen (UUZM 06) of Coluber
lemniscatus. Photo and Drawings by D. T. Feitosa.
Remarks: is specimen probably was part of the
King Adolf Fredrik I collection, which was incorpo-
rated into the UUZM collection in 1801, when King
Gustav IV (grandson of King Adolf Fredrik I) deci-
ded to split the original collection between the KVA
and UUMZ.
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3.2. e specimens at Swedish Museum of
Natural History (NRM)
NRM 93—lectotype (Fig. 31): An adult male, with 4
preventrals, 238 ventrals, and 36 subcaudals; the eye
diameter is slightly longer than ½ of its distance from
the border of the mouth; the rostral is higher than
wide; the internasals are ⅔ the length of the prefron-
tals; the frontal is hexagonal, ⁄ times longer than
wide, as long as its distance from snout, and wider
anteriorly; the parietals are as long as their distance
from the snout; the preoculars = 1/1; the postoculars
= 2/2; the temporals = 1+1/1+1; and the 3rd and 4th
supralabials are in contact with the eye. Dorsally, the
snout (the rostral, the anterior nasals, the internasals,
and the 1st and the anterior margin of the 2nd pair of
supralabials) is black; a white preocular band behind
the black snout includes the anterior ¼ of the fron-
tal, the prefrontals (except for their posterior mar-
gin, they are lined with black, but this does not reach
the frontal-prefrontal-supraocular suture on the le,
and from the medial margin of the right prefrontal
to the right margin of the frontal), the anterior ⅓
of the preoculars, the posterior nasals, the 2nd and
antero-superior border of the 3rd le supralabials,
and the 2nd and 3rd supralabials on the right. A black
head cap extends over the posterior ¾ of the frontal,
the supraoculars, the posterior ¾ of the preoculars,
the anterior ⅓ of the parietals, the postoculars, the
anterior temporals, and most of the 3rd, followed by
4th, 5th, and the anterior part of the 6th pair of supra-
labials. A black blotch is present on the posterior tip
of the le parietal. e remaining surface of the head
is red. Ventrally, the head is red, with the 1st–4th infra-
labials completely black, and four irregular black
blotches are present on the chin shields. e body
triads are 12 + 1⅔ on the tail. e anterior black ring
of the 1st triad is separated from the parietals by ve
vertebral scales, and the majority of the red nuchal
scales are tipped with black. e middle and the
external black rings are similar in length. e white
rings are very short, about ¼ the length of the black
ones. e ToL = 684 mm; the HL = 16.57 mm; the
SVL = 625 mm; and the TL = 59 mm. e TL/SVL
ratio = 0.094.
Fig. 31. A Linnaean specimen (NRM 93) of Coluber lem-
niscatus. Photo and drawings by D. T. Feitosa.
NRM 94—paralectotype (Fig. 32): An adult male,
with 4 preventrals, 242 ventrals, and 37 subcaudals;
the eye diameter is slightly longer than ½ of its dis-
tance from the border of the mouth; the rostral is
higher than wide; the internasals are ⅔ the length of
the prefrontals; the frontal is hexagonal, 1⁄ times
longer than wide, as long as its distance from the
snout, and wider anteriorly; the parietals are shorter
than their distance from the snout; the preoculars =
1/1; the postoculars = 2/2; the temporals = 1+1/1+1;
and the 3rd and 4th supralabials are in contact with
the eye. Dorsally, the snout (the rostral, the anterior
nasals, the border of the nostrils with the posterior
nasals, the anterior ¾ of the internasals, and the 1st
and anterior part of the 2nd pair of supralabials) is
black, without any visible white markings; a white
preocular band behind the black snout includes the
posterior margin of the internasals, the prefrontals,
the anterior ½ of the preoculars, the posterior nasals
behind the nostril, the anterior margin of the fron-
tal, and the posterior ½ of the 2nd and anterior ⅓ of
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54 M. G. Pires, D. T. Feitosa, F. G. Grazziotin, A. L. C. Prudente, N. J. Silva, Jr., and H. Zaher
the 3rd pair of supralabials. A black head cap covers
most of the frontal, the supraoculars, the posterior
½ of the preoculars, the anterior ⅓ of the parietals,
the postoculars, ⅔ of the anterior pair of temporals,
and ⅔ of the 3rd, 4th, 5th, and anterior part of the 6th
supralabials. e remaining surface of the head is
red. Ventrally, the head is red, with a black, irregular
marking covering the inferior border of the 2nd, the
posterior ¾ of the 3rd, and the anterior ¼ of the 4th
infralabials on the le side. e remaining ventral
head scales are immaculate. e body triads are 15
+ 1⅔ on the tail. e anterior black ring of the 1st
triad is separated from the parietals by four vertebral
scales. e middle black rings on the triads are 1½–2
times longer than the external ones, and as long or
slightly longer than the red rings. e white rings are
very short, about ½ the length of the external black
rings, and about ⁄ or ¼ the length of the middle
black rings. e ToL = 545 mm; the HL = 13.47 mm;
the SVL = 504 mm; and the TL = 41 mm. e TL/
SVL ratio = 0.081.
Fig. 32. A Linnaean specimen (NRM 94) of Coluber lem-
niscatus. Photo and drawings by D. T. Feitosa.
NRM 95—new paralectotype (Fig. 33): An adult
female with 4 preventrals, 261 ventrals, and 35 sub-
caudals; the eye diameter is slightly longer than ½
of its distance from the border of the mouth; the
rostral is higher than wide; the internasals are ⅔
the
length of the prefrontals; the frontal is hexagonal,
1⁄ times longer than wide, as long as its distance
from the snout, and broader anteriorly; the parie-
tals are as long as their distance from the snout; the
preoculars = 1/1; the postoculars = 2/2; the tem-
porals = 1+1/1+1; and the 3rd and 4th supralabials
are in contact with the eye. Dorsally, the snout (the
rostral, the anterior nasals, the border of the nostrils
with the posterior nasals, the internasals, the ante-
rior margins of the prefrontals, and the 1st and ante-
rior portion of the 2nd supralabials) is black, without
any visible white markings; a white preocular band
behind the black snout includes the anterior margin
of the frontal, the prefrontals, the posterior margin
of the internasals, the anterior ⅓ of the preoculars,
and the posterior ⅔ of the 2nd and the anterior ⅓ of
the 3rd supralabials. A black head cap covers the fron-
tal (except for a small portion of the anterior margin
on the le), the supraoculars, the posterior ½ of the
preoculars, the anterior ⅓ of parietals, the post-
oculars, the anterior le and anterior ⅔ of the right
temporal, and the posterior ⅔ of the 3rd, 4th, 5th and
the anterior ⅓ of the 6th supralabials. e remaining
surface of the head is red, except for a black blotch
covering the posterior extremity of the le parietal
and some irregular black tipping on a few scales on
the occipital region. Ventrally, the head appears to
be entirely red, except for a few irregular markings
on the anterior infralabials and the anterior margin
of the right anterior chin shields. e body triads =
12 + 1⅔ on the tail. e anterior black ring of the
1st triad is separated from the parietals by 3½ verte-
bral scale rows. e middle black rings of the triads
are slightly longer than the external ones. e white
rings are short, about ¼ of the length of the external
black rings and ⁄ the length of the middle one. e
red rings are slightly longer than the black external
rings, but shorter than the middle black triad rings.
e ToL = 706 mm; the HL = 15.09 mm; the SVL =
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Advances in Coralsnake Biology: with an Emphasis on South America 55
650 mm; and the TL = 56 mm. e TL/SVL ratio =
0.086.
Fig. 33. A Linnaean specimen (NRM 95) of Coluber lem-
niscatus. Photo and drawings by D. T. Feitosa.
Remarks: Specimen NRM 95 likely is a replacement for a
lost specimen bearing the Drottningholm label.
3.3. e specimens at the Russian Academy of
Sciences—new paralectotypes
e reference of possible specimen(s) at the Musei
Imperialis Petropolitani in Russia by Linnaeus
(1749a, p. 464, n. 218) led us in a dicult quest that
revealed interesting and important information. e
entry number 218 either can be related to a specimen,
or just to the species. If it followed the publications
of the time, in all Linnean publications the number
clearly is the entry of a species name. All of the spe-
cimens in the Kunstkamera (originally conceived
as a cabinet of curiosities, established in 1727) were
recorded as the zoological specimens of the Saint
Petersburg Academy of Sciences (today the Zoolo-
gical Institute [ZISP]; Russian Academy of Sciences
[RAS]). With the relevant information provided by
N. L. Orlov and K. Milto (pers. comm.), the RAS still
holds ve specimens of Micrurus lemniscatus (ZISP
1258, ZISP 1261, ZISP 1262, ZISP 1263, and ZISP
1264), but their roles still have not been determined
in the Linnean descriptions and type designations.
Despite having been cited by Linnaeus (1749a) as
specimen(s) of Coluber lemniscatus in Saint Peters-
burg, he never acknowledged them in Systema Natu-
rae (1758), which created a dubious interpretation.
Nevertheless, since historically Czar Peter I might
have linked these specimens to the partial acquisi-
tion of Seba’s esaurus, in our opinion these speci-
mens also should be considered as paralectotypes of
M. lemniscatus. We did not have an opportunity to
personally examine these specimens, but N. L. Orlov
and K. Milto kindly prepared and sent us a large
series of their photographs (Figs. 34–38).
Fig. 34. Micrurus lemniscatus (ZISP 1258). A = ventral
view; and B = dorsal view. Photos: K. Milto and N.L. Orlov.
With all of these facts, the best course of action
would be to allocate specimen UUZM 06 as the
third paralectotype of Micrurus lemniscatus (even
knowing that this was the rst specimen of M. lem-
niscatus described), together with NRM 94 and
NRM 95, and maintaining specimen NRM 93 as
the lectotype. Undoubtedly, the Russian specimens
somehow are involved as comparative specimens
to the type series, even though none were cited in
later publications, but listed by Linnaeus (1749a).
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56 M. G. Pires, D. T. Feitosa, F. G. Grazziotin, A. L. C. Prudente, N. J. Silva, Jr., and H. Zaher
Accordingly, we suggest including these ve speci-
mens (ZISP 1258, ZISP 1261, ZISP 1262, ZISP 1263,
and ZISP 1264) in the type series of M. lemniscatus,
as additional paralectotypes.
4. THE TYPE LOCALITY
Schmidt and Walker (1943) examined specimens
of snakes housed at the Uniniversity of Arequipa
(Peru), and with respect to a specimen of Micrurus
lemniscatus, which lacked collecting data, suggested
restricting the type locality of this species to Belém,
Pará, Brazil. e authors, however, did not provide
justication for assigning this as the type locality.
Previously, we indicated that most of the Linnean
collection from South America came from Suri-
name, and we do not believe that any material from
Belém (Brazil) ever reached Linnaeus. Members
of the Dutch West India Company (WIC) were in
Brazil from 1624 to 1654, and their travels primarily
were restricted to the northeastern region. us, the
geographic area of trade and inuence of the WIC
never included the mouth of the Amazon River,
and its activities ended at least 90 years before the
Fig. 35. Micrurus lemniscatus (ZISP 1261). A = ventral
view; and B = dorsal view. Photos: K. Milto and N.L. Orlov.
Fig. 36. Coluber lemniscatus (ZISP 1262). A = ventral view;
and B = dorsal view. Photos: K. Milto and N.L. Orlov.
Fig. 37. Coluber lemniscatus (ZISP 1263). A = ventral view;
and B = dorsal view. Photos: K. Milto and N.L. Orlov.
Fig. 38. Coluber lemniscatus (ZISP 1264). A = ventral view;
and B = dorsal view. Photos: K. Milto and N.L. Orlov.
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Advances in Coralsnake Biology: with an Emphasis on South America 57
publication of Hast’s dissertation (1745) and 80 years
before Seba’s esaurus (1734).
e geographic distribution of Micrurus lem-
niscatus is extensive (Campbell and Lamar, 1989,
2004; Silva Jr. et al., 2016; see Chapter 4). When Carl
Gyllenborg’s original collection was donated to the
UUZM, M. Beronius, J. Ihre and D. Solander conduc-
ted an inventory that listed nine snakes (9 Indianiske
ormar af dierente sorter) (Holm, 1957). e term
Indianiske might have a general translation of tro-
pical, southern, exotic (de A. Mattisson, Svenska
Akademiens ordboksredaktion, pers. comm.) which
makes sense if we note that from the nine snakes list-
ed, seven are typical South American taxa (Coluber
Ahaetulla = Leptophis ahaetulla; Coluber Cobella =
Liophis cobella; Coluber Domicella = Liophis poecilo-
gyrus; Coluber lemniscatus = Micrurus lemniscatus;
Coluber angulatus = Helicops angulatus; Coluber
Petola = Oxyrhopus petola; Coluber annulatus =
Leptodeira annulata). is only can be explained by
the inuence of the Dutch East and West India com-
panies (VOC and WIC) in Suriname, a known trade
post and Dutch colony in northern South America.
Roze (1967) agreed that the northern Guiana
populations of Micrurus l. lemniscatus resemble the
general description of Linnaeus’ syntypes, but oddly
he suggested considering Belém as the type locality
invalid (even though the geographic distribution
of this species was well known in this region), and
maintaining it as “Asia.”
We also rely on Carl Gyllenborg’s well-docu-
mented collection, which was donated to Uppsala
University, as the source of the rst described spe-
cimen of Micrurus lemniscatus (Linnaeus, 1749a).
Based on these assumptions, the type locality of M.
lemniscatus could not be Belém, Brazil (de Schmidt
and Walker, 1943) or unknown (Roze, 1967), so we
suggest restricting the type locality of M. lemnisca-
tus to Paramaribo (Suriname) in order to settle this
matter.
5. NEW TAXONOMIC PROPOSAL
Micrurus lemniscatus has undergone an intricate
taxonomic history, on account of the denition of its
generic status (Laurenti, 1768; Schneider, 1801; Dau-
din, 1803; Beebe, 1919) and several contradictory
attempts to lump or split this taxon into dierent
species or subspecies (Schmidt and Schmidt, 1925;
Werner, 1927; Schmidt, 1936; Amaral, 1944; Burger,
1955; Roze, 1967, 1989; Giraudo and Scrocchi, 2002;
Harvey et al., 2003; Starace, 2013; Pires et al., 2014).
Even with the recent specic relocation of M. diutius
(Starace, 2013) and the description of M. potyguara
(Pires et al., 2014), M. lemniscatus apparently is com-
posed of more than one species.
Concerning the species groups, Silva, Jr. and
Sites, Jr. (1999, 2001), in two studies dealing with the
systematics of the triadal species based on morph-
ology, DNA, and isoenzyme evidence, suggest-
ed that the Micrurus frontalis (Duméril, Bibron
and Duméril, 1854) and M. lemniscatus (Linnaeus,
1758) groups were complexes of non-monophyletic
species. eir hypothesis, in part, was corroborated
by additional studies (Giraudo and Scrocchi, 2002;
Castoe et al., 2007; Pyron et al., 2011, 2013; Renjifo
et al., 2012; Lee et al., 2016; Zaher et al., 2016, 2019),
thereby justifying the need for a taxonomic revision
of both groups.
Silva, Jr. and Sites, Jr. (1999) resolved some of the
taxonomic enigma of the Micrurus frontalis species
complex by recognizing seven subspecies of M. fron-
talis as full species—M. frontalis (Duméril, Bibron
and Duméril, 1854), M. altirostris (Cope, 1860), M.
baliocoryphus (Cope, 1862), M. brasiliensis Roze,
1967, M. diana Roze, 1983, M. pyrrhocryptus (Cope,
1862), and M. tricolor Hoge, 1956. e M. lemnisca-
tus species complex (Pires, 2011; Pires et al., 2014)
remains unresolved, as it consists of three subspe-
cies, M. l. lemniscatus (Linnaeus, 1758), M. l. helleri
Schmidt and Schmidt, 1925, M. l. carvalhoi Roze,
1967, and three associated species (M. diutius Bur-
ger, 1955; M. frontifasciatus Werner, 1927; and M.
potyguara Pires, Silva, Jr., Feitosa, Prudente, Pereira
Filho, and Zaher, 2014).
Herein, we provide a taxonomic revision of
the Micrurus lemniscatus complex based on mor-
phological (biometric, meristic, hemipenial, cra-
nial, and color pattern) and molecular data (two
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58 M. G. Pires, D. T. Feitosa, F. G. Grazziotin, A. L. C. Prudente, N. J. Silva, Jr., and H. Zaher
mitochondrial genes). We compare all the species
with each other and with other sympatric triadal
coralsnakes species, using univariate and multivar-
iate statistics. We review, revisit, and discuss the
type-series of taxa in a historical context, and de-
scribe a previously neglected lectotype of M. lemnis-
catus, in an eort to stabilize the nomenclature and
raise awareness of important specimens that have
not been examined in previous literature. Our results
suggest the obsolescence of the existing taxonomic
arrangement for these taxa, and reveal problems in
the typication of M. l. lemniscatus and M. l. car-
valhoi. We include one additional specimen of M.
lemniscatus with characteristics that conform with
the original description of the species more pre-
cisely, which was examined and used by Linnaeus as
a reference in the type series, as a paralectotype of M.
lemniscatus. Our results also highlight the unlike-
lihood that Linnaean specimens of M. lemniscatus
originated in Brazil, and thus we suggest changing
the type locality of this species. We propose a new
taxonomic arrangement that considers M. lemnisca-
tus and M. carvalhoi as full species, and M. l. helleri
as a synonym of M. lemniscatus, and characterize M.
diutius and M. frontifasciatus.
We examined 1,088 specimens of the Micrurus
lemniscatus complex (Pires, 2011; Pires et al., 2014)
with associated taxa that remain or formerly have
been recognized as its subspecies, in an attempt to
provide a congruent diagnosis for these forms, and
with an aim at stabilizing their taxonomic status
(167 M. diutius; 388 M. l. carvalhoi; 310 M. l. lem-
niscatus; 200 M. l. helleri; 16 M. frontifasciatus; and
seven M. potyguara; Appendices I, II). For com-
parative analyses, we also used specimens of 29
species of South American coralsnakes that occur
sympatrically or within the distributional limits of
the M. lemniscatus complex, as follows: Leptomi-
crurus collaris; L. narduccii; L. scutiventris; Micrurus
annellatus; M. albicinctus; M. averyi; M. corallinus;
M. langsdor; M. ornatissimus; M. pacaraimae; M.
paraensis; M. putumayensis; M. psyches; M. remo-
tus; M. tikuna; Micrurus mipartitus; M. altirostris;
M. baliocoryphus; M. brasiliensis; M. decoratus; M.
liformis; M. hemprichii; M. ibiboboca; M. isozonus;
M. nattereri; M. serranus; M. spixii; M. surinamensis;
and M. tricolor.
Our proposal is based on quantitative (meris-
tic and morphometric characters) and qualitative
analyses of morphological characters (hemipenial
and cranial characters, and color pattern), as well as
a limited molecular analysis (Appendices III, IV).
A. Morphological characteristics
We observed a large overlap of variation among
the species, as revealed by the maximum and mini-
mum values, but generally the use of standard errors
shows statistical dierences among species and sex.
e number of ventral and subcaudal scales varies
among Micrurus l. carvalhoi, M. diutius, M. fronti-
fasciatus, M. l. lemniscatus, and M. potyguara, with
an overlap owing to extreme values.
e loadings obtained from a Principal Compo-
nent Analysis (PCA) showed in their PC1 that, for
both sexes, morphometric characters contributed
the most to group separation (> 68%), as this com-
ponent accumulated most of the data variance (54%
in males, 52.5% in females). In PC2, the characters
that contributed the most to group separation (>
58%) were the number of body triads (TriB) for both
sexes, and subcaudals (SC) for males, and ventrals
(VE) for females.
Wilk’s statistic was equal to 0.129 (approx. F
(65.2433) = 20.21442; P < 0.0000) for males, and
0.116 (approx. F (65.1223) = 10.86184; P < 0.0000)
for females, indicating signicant dierences among
species in the multivariate space of the characters
analyzed. For males, the number of the body triads
(TriB) and the length of the middle black ring (MB)
composed the rst canonical axis, and VE, SC, and
the posterior black ring dened the second axis. For
females, the number of the body triads (TriB) com-
posed the rst canonical axis, and VE, SC, and the
anterior black ring dened the second axis.
In males, the values for TriB generally were posi-
tively related with axis 1, whereas the values for MB
were negatively related with this axis. ese cha-
racters clearly dierentiated Micrurus lemniscatus
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Advances in Coralsnake Biology: with an Emphasis on South America 59
from M. l. carvalhoi and M. diutius. We observed the
highest values for TriB in M. l. carvalhoi. Similarly,
axis 2 demonstrated that M. diutius has lower num-
bers for VE compared to M. l. lemniscatus and M. l.
helleri. Males of M. l. carvalhoi showed higher num-
bers for TriB than females. Females of M. potyguara
also showed higher values for VE than males. Most of
the data (males and females) of M. l. helleri fall within
the range of M. l. lemniscatus. We found a remark-
able overlap of data between M. frontifasciatus and M.
diutius (both sexes). e data for male M. potyguara
overlap those of M. l. carvalhoi, M. l. helleri, M. l. lem-
niscatus, M. diutius, and females with M. diutius, M. l.
helleri, and M. l. lemniscatus. Micrurus frontifasciatus
(males and females) fall within the range of M. diu-
tius (males and females) (Fig. 39).
Fig. 39. Bivariate plot for the rst two axes derived from scores of DFA analysis for males (A) and females (B) of Micru-
rus lemniscatus carvalhoi, M. diutius, M. frontifasciatus, M. potyguara, M. l. lemniscatus, and M. l. helleri.
(A)
(B)
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60 M. G. Pires, D. T. Feitosa, F. G. Grazziotin, A. L. C. Prudente, N. J. Silva, Jr., and H. Zaher
B. Molecular phylogeny
Zaher et al. (see Chapter 3) conducted a molecu-
lar phylogeny for 197 species of elapids, in which
Micrurus lemniscatus is nested with other Amazo-
nian taxa; however, several observations involved
limited sampling, the representation of valid taxa,
and the quality of the taxonomic identication of
samples and museum vouchers. Consequently, we
used the available samples from the M. lemnisca-
tus species complex, and by using mitochondrial
sequences of cytochrome b (cytb) and subunit 4 from
NADH dehydrogenase (nd4) for Micrurus (Appen-
dix IV) we recovered a topology that supports our
taxonomic decision (Fig. 40).
Among the monadal species (Monadal clade)
we retrieved three highly supported subclades, as
follows: M01 (100/1.0)—M. alleni; M02 (100/0.99)—
six species clustered in pairs into three strongly
supported small clades: (1) M. browni + M. diastema
(100/1.0); (2) M. fulvius + M. tener (95/1.0); and (3)
M. nigrocinctus + M. mosquitensis (64/0.99); and
M03—(93/0.99), composed of a highly support-
ed clade formed by M. albicinctus + M. corallinus
(92/1.0), which includes M. ornatissimus, M. psy-
ches, M. dumerilii, and a clade of M. mertensi + M.
bocourti (97/99), as successive sister clades. In both
trees, the position of M. elegans showed low support.
Subclade M01 is the sister group of M02, and M03
is the most basal subclade among them. Micrurus
laticollaris is a sister group of all monadal subclades
(detailed topology not shown in Fig. 42).
Within the triadal clade we recovered three
Fig. 40. A summary ML tree highlighting the relationship among the species of Micrurus in the triadal clade. e num-
bers near nodes represent the bootstrap values (above the branches) and the posterior probabilities (below the branches).
Bootstrap values less than 70% and posterior probabilities less than 0.8 are not shown. e branches and tip labels colored
in red indicate the M. lemniscatus species complex. T01, T02 and T03 represent the three main subclades found within
the triadal clade.
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Advances in Coralsnake Biology: with an Emphasis on South America 61
well-supported subclades, as follows: T01 (92/1.0)—
M. hemprichii, M. surinamensis, M. diutius, M. l.
lemniscatus, and M. l. helleri; T02 (96/1.0)—M.
mipartitus and M. dissoleucus; and T03 (< 70/0.99)—
M. altirostris, M. obscurus, M. pyrrhocryptus, M.
baliocoryphus, M. frontalis, M. brasiliensis, and M. cf.
ibiboboca. Subclade T01 includes the three sampled
subspecies of M. lemniscatus (M. l. lemniscatus, M.
l. helleri and M. l. carvalhoi) in a highly supported
clade (89/1.0), which as successive sister groups of
M. diutius (< 70/0.99) contains M. surinamensis (<
70/0.9) and M. hemprichii (92/1.0) (detailed topo-
logy not shown in Fig. 42). In the tree, L. narduccii is
a sister group of subclade T01. In both analyses, we
recovered M. decoratus as the sister group of a weakly
supported clade formed by subclades T02 and T03.
e latter species is a sister group in both analyses
(Appendix 2) of a weakly supported clade formed
by subclades T02 and T03. e odd situation of M.
decoratus already was evident from an early study
(Silva, Jr. and Sites, Jr., 2001), but this situation is a
direct result of the type of data, the number of taxa
included, and the analyses performed (see Chapter
3). Within subclade T03, all of the samples identied
as M. cf. ibiboboca cluster together in a well-support-
ed clade (96/1.0) that is a sister group (87/1.0) of
a clade formed by M. frontalis and M. brasiliensis
(99/1.0). Our analysis recovered, with low support,
a clade composed of M. pyrrhocryptus and M. balio-
coryphus (100/1.0) as a sister group of the clade ibi-
boboca / frontalis / brasiliensis. We found the species
M. obscurus (84/1.0) and M. altirostris (< 70/0.99)
positioned as the successive sister groups of the
entire clade composed of ve previous species (M.
pyrrhocryptus / M. baliocoryphus / M. ibiboboca / M.
frontalis / M. brasiliensis).
e average of the p-distances among all the
species of Micrurus is 0.13, which range from 0.018
(between M. frontalis and M. brasiliensis) to 0.203
(between M. browni and M. mipartitus). Within the
monadal clade the average distance is 0.078, ranging
from 0.02 (between M. corallinus, and M. albicinctus)
to 0.175 (between M. dumerilii and M. laticollaris),
whereas within the triadal clade the average distance
is 0.113, ranging from 0.018 (between M. frontalis
and M. brasiliensis) to 0.186 (between M. dissoleucus
and L. narduccii).
Between the three subspecies sampled in the
Micrurus lemniscatus complex, the p-distance is
0.067, whereas the distance between M. l. carvalhoi
and M. diutius is 0.074, and between M. l. lemnisca-
tus and M. diutius the distance is 0.077. Between M.
l. lemniscatus and M. l. helleri the p-distance is only
0.036, and the average p-distance for the M. lemnis-
catus complex is 0.072 (Subclade T01). In our opi-
nion, this is a strong indication of the synonymy of
M. l. helleri with M. l. lemniscatus, as well the recog-
nition of M. l. carvalhoi as full species (M. carvalhoi).
C. Taxonomic decision
e comparative morphological analysis between
specimens of Micrurus l. lemniscatus and M. l.
helleri, as well as understanding variation in the
color pattern of both forms, does not support the
separation of these taxa as subspecies, which justies
the synonymization of M. l. helleri under M. lemnis-
catus. e characteristics of the triads and the length
of the triad rings also are highly variable, as are the
black markings on the gular region of M. lemnisca-
tus. e variation in M. l. helleri falls within that of
the nominal species. e general characters of the
holotype of M. helleri are as follows: male specimen;
ventrals = 255; subcaudals = 35; triads = 11 + 1⅓;
ToL = 635.3 mm; SVL = 586.0 mm; HL = 14.9 mm;
and TL = 49.3 mm.
We recognize the specic status of Micrurus lem-
niscatus and M. carvalhoi, and provide a better char-
acterization of M. diutius, M. frontifasciatus, and M.
potyguara. Consequently, herein we propose the place-
ment of M. lemniscatus helleri in the synonymy of
M. lemniscatus and assign new types, and contrary
to the arguments of Harvey et al. (2003) recognize
M. frontifasciatus as separate from M. lemniscatus.
e general morphological characters of the
head and body color, the triadal pattern, and the
scale counts are diagnosable. All of the taxa have
the same general head color pattern, which con-
sists of a black snout, followed by a narrow white
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62 M. G. Pires, D. T. Feitosa, F. G. Grazziotin, A. L. C. Prudente, N. J. Silva, Jr., and H. Zaher
preocular band and a longer black transversal band.
is character varies among the species, along with
the gular color pattern. e same triadal pattern of
a middle black ring, longer (or the same length) as
the external ones, and white rings that are shorter
(or the same length) as the external black rings, is
common to all taxa, but varies in the length of the
white rings among the species. Although there is an
overlap in the number of body triads and the scale
counts, a combination of other characters (compa-
rative length of the body rings, head color pattern,
hemipenial, and cranial morphology) can be used to
separate the forms related to Micrurus lemniscatus.
D. Hemipenial morphology of members of the
Micrurus lemniscatus complex
e species in the Micrurus lemniscatus complex
are characterized by the presence of a short and
slightly bilobed hemipenis, which is capitate and
covered with irregularly-distributed spines that
are more densely arranged on the sulcate surface,
and decrease in the proximal to distal direction.
In M. lemniscatus and M. carvalhoi, the capitulum
comprises about ⅔ of the total length of the hemi-
penis, but it is only about ¾ of the total length in
M. diutius and M. potyguara. In all the species, the
capitulum is ornamented with irregular and dense-
ly arranged spines, similar in size throughout the
circumference of the organ, and in most species it
extends the length of the capitulum. In M. lemnisca-
tus, M. diutius, and M. potyguara the lobes represent
approximately ⁄ of the total length of the organ, in
M. carvalhoi ¼, and in M. lemniscatus about ½, and
the lobes are ornamented with spines that decrease
in size and number from the apexes toward the base
of the lobes, especially on the asulcate side. e
sulcus spermaticus is deep, with smooth borders
bifurcating at a level more proximal than the bases
of the lobes (sometimes almost at the same level on
the base of the lobes). In M. carvalhoi and M. poty-
guara, the sulcus spermaticus bifurcates on the distal
⅓ of the hemipenis, and in M. lemniscatus and M.
diutius in the distal ¼. In all the taxa, the interlo-
bar region contains semi-equal spines that are more
densely arranged than the ones on the asulcate side.
e capitular sulcus is deep, and crosses the organ
perpendicularly at the distal limit of the proximal ⅓
of the organ, and reaches the sulcate surface on both
sides on a decreasing curve, with its limits reach-
ing the margins of the sulcus spermaticus. Aer
bifurcating, the sulcus spermaticus extends centripe-
tally to the distal tip of the lobes. e body of the
hemipenis is covered with smaller spines than on the
capitularis, which are larger on the distal region and
decrease in size toward the base of the organ, which
is ormamented with tiny and irregularly distributed
spines. e basal pocket is small and naked (Fig. 41).
Fig. 41. e hemipenes of the Micrurus lemniscatus spe-
cies complex. A = M. carvalhoi (MZUSP 15534); B = M.
diutius (MZUSP 4792); C = M. lemniscatus (CEPB 9056);
and D = M. potyguara (UFPB 4359). Scale bars = 5 mm.
Photos: Matheus G. Pires.
E. Cranial morphology of the Micrurus
lemniscatus complex
e skull is slender in dorsal aspect, and convex
laterally. An oblong posterior extremity (curved or
rounded) is present on the parietal, with the pro-
otic bones visible and on which lie the slender and
slightly curved supratemporal bones. e parietal is
wider on the posterior ½ and narrower toward the
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Advances in Coralsnake Biology: with an Emphasis on South America 63
frontals. e posterior end of the parietal is roun d-
ed in Micrurus lemniscatus, but pointed in M. car-
valhoi and M. diutius. e sagittal crest is well
developed in M. carvalhoi, but subtle in M. diutius
and absent in M. lemniscatus, and extends from the
posterior extremity of the parietal to the proximity
of the frontoparietal suture, where it bifurcates and
follows the contour of this articulation. e frontal
bones are triangular, laminar, lack processes or pro-
jections, and their posterior extremities are round-
ed in M. lemniscatus and pointed in M. carvalhoi and
M. diutius, and are approximately ½ the length of the
parietal. e prefrontals are short, wide, and contact
each other through the elongated dorsomedial pro-
cesses of the frontals, and also form a concave arch
with the nasal complex in the articulation region.
e dorsomedial processes of the prefrontals are well
developed and connected widely at the midline. e
nasals are small and narrow and allow a view of the
lateral margins of the septomaxillas and the posterior
vomer, as well as the contour of the premaxilla. e
anterior extremity of the nasals does not reach the
anterior border of the premaxilla in M. lemniscatus,
is of the same length in M. diutius, and sometimes is
slightly longer in M. carvalhoi. e maxillary bones
are stout and longer than the frontals. Owing to the
rounded posterior extremity of the parietal bone,
the antero-superior border of the prootic bones are
slightly projected anteriorly, almost in line with the
prootic-sphenoid suture. e quadrate bones are
triangular and contain a wide articular contact with
the supratemporals, which represents up to 70.6% of
the total length of the bones. In M. lemniscatus, the
posterior extremities of the supratemporals do not
exceed the posterior limit of the exoccipital bones,
whereas in M. carvalhoi and M. diutius they are lon-
ger and clearly reach beyond the posterior limit of
exoccipitals. In M. diutius, the pseudocoronoid pro-
cess of the compound bone is well developed, but it
is discrete in M. lemniscatus and M. carvalhoi, and
extends between 10.2% and 16.9% (height) of the
entire bone. ere are 7–8 palatine, 3–4 pterygoid,
and 8–12 dentary teeth (Fig. 42).
Fig. 42. Selected skulls of the Micrurus lemniscatus species
complex: A–E = M. carvalhoi (IBSP 67825); F–J = M. diu-
tius (MPEG 21169); and K–O = M. lemniscatus (MPEG
24536). Scale bars = 5 mm. Photos: Matheus G. Pires.
F. Species accounts
To better characterize the taxa herein designated as
the Micrurus lemniscatus complex, below we use a
combination of cranial and hemipenial morpholo-
gical features, as well as the described dierences in
color pattern. As previously noted, extreme varia-
tions in color pattern and the length of the body
triad rings are evident in M. lemniscatus and M.
carvalhoi, and also might involve other forms, as the
geographic distributions of these taxa is large and the
variation is not well described and understood. Pires
(2011) provided additional data and references.
Micrurus lemniscatus (Linnaeus, 1758)—new
status
Coluber lemniscatus Linnaeus 1758: 224.
Natrix lemniscata—Laurenti 1768: 76.
Elaps lemniscatus—Schneider 1801: 291.
Vipera lemniscata—Daudin 1803: 13.
Micrurus lemniscatus—Beebe 1919: 216.
Micrurus helleri—Schmidt and Schmidt 1925: 129.
Micrurus lemniscatus lemniscatus—Amaral 1944: 89.
Micrurus lemniscatus lemniscatus—Roze 1967: 32.
Micrurus lemniscatus helleri—Roze 1967: 35.
Micrurus lemniscatus—Pires 2011: 88.
Lectotype: NRM 93(M), unknown locality.
Paralectotypes: NRM 94(M) and NRM 95(F), unknown
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64 M. G. Pires, D. T. Feitosa, F. G. Grazziotin, A. L. C. Prudente, N. J. Silva, Jr., and H. Zaher
locality; UUMZ 06, unknown locality; ZISP 1258,
ZISP 1261, ZISP 1262, ZISP 1263, and ZISP 1264
(not sexed), unknown locality.
Type locality: Paramaribo, Suriname.
Diagnosis. Micrurus lemniscatus can be distin-
guished from all other triadal species of Micrurus
by the following combination of morphological
characters: black snout (rostral, internasals, anterior
border of prefrontals and nasals, and 1–2 supralabi-
als); white preocular transverse band covers most of
prefrontals, anterior border of supraoculars, poste-
rior nasals, preoculars, and 2–3 supralabials; black
cephalic cap includes frontal, supraoculars, post-
oculars, anterior ⅓ of parietals, temporals, and 2–3
supralabials; remainder of head red, with or with-
out black-tipped scales; gulars red with mental and
some or most of infralabials black; 1st triad separated
from parietals by 1–6 scales; triadal pattern: middle
black ring longer or same length as outer ones, white
rings usually shorter than outer black rings (1½–2
dorsal scales in length), red rings shorter than entire
triad or even shorter than middle black ring; red
and white rings usually black-tipped, but might be
immaculate on rst ¼ of body; and 7–16 body triads
and ⅓–2 tail triads. Ventral scales 230–250 in males,
and 245–270 in females. Subcaudals 26–48 in males,
and 24–48 in females (Fig. 43A).
Comparison with sympatric species. Micrurus
lemniscatus occurs sympatrically with three spe-
cies of Leptomicrurus, and 11 species of monadal,
one species of bicolored, and nine triadal species of
Micrurus. It diers from Leptomicrurus by the con-
tact of the 1st infralabials behind the mental scale
(vs. the 1st infralabials are separated by an extended
mental that contacts the chin shields), a shorter and
stouter body (vs. a long and slender body with more
ventrals than in any other species of Micrurus), and
the presence of a triadal pattern (vs. the absence of
complete rings on the body). Micrurus lenniscatus
diers from all of the monadal and bicolored species
in its genus by the presence of a triadal color pattern.
It diers from M. carvalhoi by a higher mean number
of ventrals (245.8 vs. 242.6) and subcaudals (36.4 vs.
30.8), a lower number of body triads (10.8 vs. 12.6),
and a shorter cephalic cap (vs. a long cephalic cap);
from M. diutius by a higher mean number of ven-
trals (245.8 vs. 227.6), a lower number of vertebral
scales between the parietals and the 1st triad (6–8 vs.
2–4), and the presence of shorter white rings (1.5–2
vs. 2.5–4 dorsal scales in length); from M. liformis
by a lower number of ventrals (221–277 vs. 259–
335), and a lower number of triads (7–16 vs. 12–23);
and from M. hemprichii, M. isozonus, M. spixii, M.
obscurus, M. nattereri, and M. surinamensis by the
presence of a white preocular transverse band (vs.
the absence of a white preocular transverse band)
(Figs. 43, 44).
Fig. 43. Comparative morphological pattern of the Mi-
crurus lemniscatus species complex: A = M. lemniscatus
(UUZM 06); B = M. carvalhoi (USNM 76341); C = M.
diutius (FMNH 34472); D = M. frontifasciatus (NMW
18298); and E = M. potyguara (UFPB 4359). Drawings:
Marcus A. Buononato.
Variation. e head is red with irregular black
markings, black-tipped scales, or both. A black
cephalic cap generally reaches the anterior ⅓
(75.5%) or ½ of the parietals (12.8%), but rarely
reaches ⅔ (9.8%) or the entire parietals (1.5%). A
varying degree of irregular black markings are pres-
ent on the chin, including on the mental and at
A B C D E
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Advances in Coralsnake Biology: with an Emphasis on South America 65
least the rst four infralabials. e markings on
the chin can include the mental and the rst 3–4
infralabials (64.8%) with the inclusion of the 1
st
pair of gulars as in a semicircle (17%), and can be
immaculate (4.2%), with irregular black markings
(6.5%), with irregular lateral markings (5%), or
reduced to the mental (2.5%) (Fig. 44). In males
(N = 298), the ventrals = 221–268, the subcaudals
= 26–48, and the body triads = 7–16; the HL =
7.9–47.3 mm, the SVL = 197–1,280 mm, and the
TL = 14.3–126.8 mm. In females (N = 212), the
ventrals = 230–277, the subcaudals = 24–48, and
the body triads = 7–14; the HL = 7.3–30.9 mm,
the SVL = 200–1,220 mm, and the TL 10.3–119.7
mm. e 1
st
triad is separated from the parietals
by 1–6 vertebral scales: 3 (43.5%), 4 (29.7%),
2 (10.5%), 5 (9.9%), 1 (2.5%), and 6 (2.5%). e
black middle triad rings are longer that the outer
black rings (78.7%), or are semi-equal in length
(21.3%). e white rings vary in length (0.5–4.5
dorsal scales), with the majority (83.7%) consist-
ing of 1½ (33.1%), 2 (37.6), and 2½ (13%) scales.
Fig. 44. Variation in the head coloration of Micrurus lem-
niscatus: A = MUSM 3256; B = MPEG 13517; C = NRM
6308; D = MPEG 16164; E = MPEG 17513; and F = FMT
2425. Drawings: Matheus G. Pires.
e tail triads are 1–2, with a predominance of the
1⅔ pattern (1⅔ = 87.7%; 1 ⅓ = 9.6%; 2 = 1.8%; and 1
= 0.9%). e degree of black pigmentation on the red
rings is variable, and can be irregular (65%), with all
of the scales tipped with black (19%), or immaculate
(16%). e white rings can be irregularly marked
with black (17%), immaculate (40%), or the scales
might contain ne black tipping (43%) (Fig. 44).
Hemipenis. CEPB 9056 (Municipality of Altamira,
State of Pará, Brazil; le organ). e hemipenis is
short, slightly bilobed, capitate, and covered with
irregularly distributed spines that are arranged more
densely on the sulcate surface, and decrease slightly
in number from a proximal to distal direction. e
lobes are approximately ¼ of the total length of the
organ, or ½ of the capitulum length. Semi-equal
spines are present in the interlobar region, which
are arranged more densely than the spines on the
asulcate side. e capitular groove is deep, and
crosses the organ perpendicularly at the distal limit
of the proximal ⅓ of the organ, and on both sides
reaches the sulcate surface on a proximally orient-
ed curve, until reaching the margins of the sulcus
spermaticus. e capitulum is ⅓ of the total length
of the hemipenis, and contains irregular and dense-
ly arranged spines that are subequal in size along
the perimeter and extend almost to the capitulum,
and decrease slightly in size and number toward the
base of the lobes, especially on the asulcate surface.
e sulcus spermaticus is deep and contains smooth
borders, and bifurcates more proximally than the
interlobar crotch (in some cases, at the interlobar
crotch). Aer bifurcating, the sulcus spermaticus
extends centripetally to the distal tip of the lobes.
e stem is covered with smaller spines than in
the capitulum, which are unevenly distributed and
decrease in size proximally. e basal pocket is
small and naked (Fig. 40C).
Geographic distribution. Micrurus lemniscatus
inhabits the valleys of the Araguaia, Branco, Japurá/
Caquetá, Juruá, Madeira, Negro, Purus, Solimões,
Tapajós, Tocantins, Trombetas, Uatumã, Ucayali,
and Xingu rivers. It also occurs in the oodplains
of Marajó Island (Brazil), the mountainous forests
of the Cordillera Oriental (Colombia) and the
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66 M. G. Pires, D. T. Feitosa, F. G. Grazziotin, A. L. C. Prudente, N. J. Silva, Jr., and H. Zaher
Cordillera Real (Ecuador), the cloud forests of Napo
(Ecuador), the Babaçu forests (in the states of Mar-
anhão and eastern Pará, Brazil), the tropical dryland
forests of the state of Mato Grosso (Brazil) and in
Apure/Villavicencio (Colombia), and the mangrove
areas of Pará (Brazil), toward the Guianan region.
is species also might occur in southern Venezuela,
Roraima (Brazil), Guyana (French Guyana), and in
northern Mato Grosso (Brazil). e distributional
gap in the middle of the Brazilian Amazon may reect
the poor representation of specimens in museums.
e area of sympatry with M. diutius is well dened
in the north-central area of Amazonas and in west-
ern Pará (Brazil). Sympatry with M. carvalhoi occurs
in the northern region of Tocantins, and perhaps in
northern Mato Grosso (Brazil) (Fig. 45).
Micrurus carvalhoi Roze, 1967—new status
Micrurus lemniscatus carvalhoi Roze, 1967: 33.
Micrurus carvalhoi—Pires, 2011: 139.
Holotype: USNM 76341 (M) (Fig. 46).
Paratypes: AMNH 27344 (F), AMNH 96999(M),
FMNH 37201(F), FMNH 37739(M), FMNH
37740(F), FMNH 37741(M), FMNH 42193(F),
FMNH 42194(F), MCZ 16683(M), MCZ 17762(M),
MCZ 17852(M), MNRJ 1317(M), MNRJ 1318(M),
MNRJ 957(M), NHMW 13384:2(M), NHMW
18296(M), NHMW 18640(F), USNM 100718(F),
USNM 76342(F), CAS 49297(F), USNM 39074(M),
USNM 76340(M), USNM 76343(M), SMF
20767(M).
Type locality: Catanduva, State of São Paulo, Brazil.
Fig. 45. e geographic distribution of species in the Micrurus lemniscatus complex. Map: Sérgio H. M. Nogueira.
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Advances in Coralsnake Biology: with an Emphasis on South America 67
Fig. 46. Holotype of Micrurus carvalhoi (USNM 76341).
Photo and drawings: Drawings: D. T. Feitosa
Diagnosis. Micrurus carvalhoi can be distinguished
from all other triadal species of Micrurus by the
following combination of morphological characters:
black snout (rostral, internasals, anterior border of
prefrontals and nasals, and 1–2 supralabials); white
preocular transverse band covers most of prefrontals,
anterior border of supraoculars, posterior nasals,
preoculars, and 2–3 supralabials; black cephalic
cap includes frontal, supraoculars, and postoculars,
anterior ⅔ of parietals, temporals, and 4–5 suprala-
bials; head red, with or without black tipped scales;
gulars red, with mental and some or most of infra-
labials black, in a horseshoe-shaped pattern; 1st triad
separated from parietals by 3–4 scales; middle black
ring longer or same length as outer ones; white rings
shorter than outer black rings; red rings shorter that
entire triad or even middle black ring; red and white
rings usually with black-tipped scales (Fig. 43); and
7–18 body triads and ⅓–2 tail triads.
Comparison with sympatric species. In northeast-
ern Brazil, Micrurus carvalhoi is sympatric with M.
ibiboboca, M. brasiliensis, M. potyguara, and M. lem-
niscatus (in the western limit of its distribution). In
the open areas of central Brazil, M. carvalhoi is sympa-
tric with M. frontalis, and in the southeast ern region
with M. decoratus and the monadal M. corallinus. In
the southern region, it is sympatric with M. altirostris
and M. silviae, and in the southwestern region, with
M. baliocoryphus and M. tricolor, but less probable
with M. pyrrhocryptus based on the actual distribu-
tional information (Silva Jr., 2016b; see Chapter 5).
Micrurus carvalhoi diers from M. lemniscatus by a
mean lower number of ventrals (242.6 vs. 245.8) and
subcaudals (30.8 vs. 36.4), a higher number of body
triads (12.6 vs. 10.8), and a longer black cephalic cap;
from M. potyguara by a shorter black cephalic cap
that does not cover the entire surface of the parietals,
and the head is not distinct from the neck (vs. long
cephalic cap entirely covering the parietals, and the
head is broader than the neck); from M. decoratus
by the presence of a complete set of black rings on
the rst triad (vs. an incomplete rst triad), 20–41
SC (vs. 14–35), and 9–16 body triads (vs. 12–23);
from M. corallinus by the presence of a triadal pat-
tern (vs. a monadal pattern); from M. ibiboboca by
the presence of an immaculate black snout (vs. the
snout motled with white); and from M. altirostris, M.
baliocoryphus, M. brasiliensis, M. frontalis, M. pyr-
rhocryptus, M. silviae, M. tricolor by the presence of
a white prefrontal transverse band (vs. the absence
of a white prefrontal transverse band).
Variation. e head is red and might contain irre-
gular black markings, black-tipped scales, or both.
e length of the white transverse band and the
black cephalic cap is variable. e black cephalic
cap generally reaches the anterior ⅔ (57.6%), ½
of parietals (26.4%), or the anterior ⅓ (6.9%), but
rarely reaches the entire parietals (9%). Ventrally, the
chin contains irregular black markings that include
the mental and most or all of the infralabials, in a
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68 M. G. Pires, D. T. Feitosa, F. G. Grazziotin, A. L. C. Prudente, N. J. Silva, Jr., and H. Zaher
typical horseshoe-shaped pattern (86.4%). Another
pattern might be an irregular semicircle (8.5%),
black posterior infralabials (2.2%), black anterior
gulars (1.4%), and other aberrant forms (1.5%) (Fig.
47). In males (N = 301), the ventrals = 186–270, the
subcaudals = 20–41, and the body triads = 7–11; the
HL = 7.9–38 mm, the SVL = 210–1,550 mm, and
the TL = 12.8–100.9 mm. In females (N = 132), the
ventrals = 210–276, the subcaudals = 21–40, and the
body triads = 9–16; the HL = 8.0–34.3 mm, the SVL
= 96–1,274 mm, and the TL = 13.1–85.7 mm. e
1st triad is separated from the parietals by 2–5 ver-
tebral scales: 3 (54%), 4 (31%), 2 (11%), and 5 (4%).
e black middle triad rings might be longer that
the outer rings (84.8%), or all three are semi-equal
in length (15.2%). e white rings vary in length
(0.5–3 dorsal scales) with the majority (84.5%)
consisting of 1½ (33.8%), 2 (30.4), and 2½ (20.3%)
scales. e tail triads are 1–2, with a predominance
of the 1⅔ pattern (1⅔ = 50%; 1⅓ = 30.6%; 2 = 16.2%;
1 = 3.2%). e degree of black pigmentation on the
red rings is variable and can be irregular (86%), with
all of the scales containing black tipping (7.3%) or
immaculate (6.7%). Similarly, the white rings can
be irregular and marked with black (31.3%), imma-
culate (22%), or the scales might contain ne black
tipping (46.7%).
Hemipenis. IBSP 49507 (Batatais, São Paulo, Brazil;
le organ). e hemipenis is short, slightly bilobed,
capitate, and covered with irregularly distributed
spines on both sides. e size of the spines increas-
es from the proximal to the distal extremities. e
lobes represent approximately ⁄ of the total length
of the hemipenis, and ⅓ of the length of capitulum.
Smaller, semi-equal spines are present in the inter-
lobar region, which are more densely arranged on
the asulcate surface. e capitular sulcus is deep,
and crosses the organ perpendicularly at the distal
limit of the proximal ¼ of the organ. e capitulum
is about ⅔ of the total length of the hemipenis, and
contains a large number of irregular and densely
arranged and equally sized spines along the circum-
ference of the organ, which extend to the capitulum
and then slightly decrease in size and number at the
base of the lobes, especially on the asulcate surface.
e sulcus spermaticus is deep and contains smooth
borders, and bifurcates on the distal ½ of the capi-
tulum below the base of the lobes, and on the dis-
tal ⅓ of the hemipenis. Aer bifurcating, the sulcus
spermaticus extends centripetally to the distal tip of
the lobes. e body is covered with smaller spines
(compared to the capitulum) that are larger on the
distal region and decrease in size toward the base
of the organ, which contains unevenly distributed
smaller spines. e basal pocket is small and naked
(Fig. 40A).
Distribution. Micrurus carvalhoi occurs from the
coastal areas of Brazil, including most of the Caa-
tinga, Cerrado, and the Atlantic Forest, to south-
western Brazil (southern Mato Grosso and Mato
Grosso do Sul), eastern Paraguay, and the Argenti-
nian Mesopotamia (Fig. 45).
Remarks: Originally described in 1967 based on
a type series of 25 specimens, Micrurus carvalhoi
represents a species of the M. lemniscatus com-
plex with a predominant distribution in the Cer-
rado and dryland areas of northern Argentina
and east ern Paraguay. At that time, Roze (1967) of-
fered a diagnosis based of the presence of irregular
black markings on the red and white rings, which
Fig. 47. Variation in the head coloration of Micrurus car-
valhoi: A = IBSP 29909; B = IBSP 40317; C = IBSP 49507;
D = IBSP 26966; E = IBSP 51599; and F = IBSP 28272.
Drawings: Matheus G. Pires.
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Advances in Coralsnake Biology: with an Emphasis on South America 69
some times formed irregular black transverse lines
on the white rings, in addition to lower subcaudal
counts in both sexes.
e large area of sympatry between Micrurus car-
valhoi and other triadal species, especially M. ibibo-
boca, whose pattern also includes a white preocular
transverse band, along with its tendency for longer
white rings and a varying degree of black markings
on the red and white rings, were contributing factors
for eliciting taxonomic confusion. Consequently,
aberrant individuals were described as new taxa (e.g.,
Elaps marcgravii, E. gravenhorstii) or synonymized
with other taxa (Wied-Neuwied, 1820; Boulenger,
1896; Schmidt, 1936; Amaral, 1944). In addition to
M. carvalhoi, the extensive type series also is a poly-
typical assemblage comprised of three other taxa,
which contributed to this problem. A specimen (CAS
49279) from Lagoa de Papary (in the municipality of
Nísia Floresta, Paraíba, Brazil) actually is identied
as M. potyguara. Another specimen (SMF 20767)
from Ilhéus, Bahia, Brazil, indicated by Roze (1967)
as SMF 9428b (Boettger’s collection number, which
currently is not used), is identied as a specimen of
M. ibiboboca, which contributed some skepticism
as to the variation data for M. carvalhoi in the ori-
ginal description. In addition, a specimen collected
by Natterer in 1824 (NHMW 13384:2) supposedly
from Cuiabá, in the state of Mato Grosso, Brazil, is
a specimen of M. lemnis catus. Since Cuiabá lies out-
side of the currently known geographic distribution
for M. lemniscatus, this place ment may have resulted
from the vandalization of Natterer’s collection before
it was shipped to Europe (Schrockinger, in Vanzo-
lini, 1993). Natterer spent about 18 years traveling
throughout Brazil, and particularly in the Brazilian
Amazon, so this specimen might have been collec-
ted elsewhere.
Micrurus diutius Burger, 1955
Micrurus lemniscatus diutius Burger, 1955.
Micrurus diutius—Starace 2013: 502.
Micrurus diutius—Pires, 2011: 192.
Holotype: FMNH 34472 (M) (Fig. 48).
Paratypes: CM 7925 (M), UIMNH 22626 (F).
Type locality: Tunapuna, Trinidad.
Fig. 48. e holotype of Micrurus diutius (FMNH 34472).
Photo and drawings: D. T. Feitosa
Diagnosis. Micrurus diutius can be distinguished
from all other triadal species of Micrurus by the
following combination of morphological characters:
black snout (rostral, internasals, anterior border of
prefrontals, most of nasals, and 1–2 supralabials);
white preocular transverse band covers most of
prefrontals, anterior border of supraoculars, most
of preoculars, and 1–2 supralabials; black cephalic
cap includes most of frontal, supraoculars, postocu-
lars, anterior ⅓ of parietals, ⅓–½ of temporals, and
2–3 supralabials; remainder of the head red, with
or without black-tipped scales; venter of head red;
irregular black markings on gulars, and infralabials
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70 M. G. Pires, D. T. Feitosa, F. G. Grazziotin, A. L. C. Prudente, N. J. Silva, Jr., and H. Zaher
highly variable; 1st triad separated from parietals by
2–7 scales; middle black ring slightly longer or same
length as outer ones; white rings usually shorter
than outer black rings; red rings shorter than entire
triad or even middle black ring; red and white rings
usually black-tipped, but with tendency to a less
melanic pattern (Fig. 43C); and 6–16 body triads
and 1 ⅓ –1⅔ tail triads.
Comparison with sympatric species. Micrurus diu-
tius occurs sympatrically with Leptomicrurus colla-
ris, four species of monadal Micrurus (M. averyi, M.
pacaraimae, M. paraensis, and M. psyches) and eight
triadal species (M. liformis, M. hemprichii, M. iso-
zonus, M. lemniscatus, M. obscurus, M. nattereri, M.
spixii, and M. surinamensis). Micrurus diutius can be
distinguished from L. collaris by the contact of the
anteriormost infralabials behind the mental scale,
and a triadal pattern (vs. the 1st infralabials separa-
ted by contact between the mental and anterior chin
shields, and the bicolored pattern with incomplete
rings). It diers from M. averyi, M. pacaraimae, M.
paraensis, and M. psyches by the presence of a triadal
pattern (vs. a monadal pattern); from M. liformis
by the number of ventrals (214–242 vs. 259–335),
the number of body triads (6–15 vs. 12–23), and the
number of white rings on the triads (2–5 vs. 1–2);
from M. lemniscatus by a lower mean number of
ventrals (227.6 vs. 245.8), a higher number of ver-
tebral scales between the parietals and the rst triad
(2–4 vs. 6–8), and longer white rings on the triads
(2.5–5 vs. 1.5–2); and from M. hemprichii, M. iso-
zonus, M. spixii, M. obscurus, M. nattereri, and M.
surinamensis by the presence of a white preocular
transverse band (vs. the absence of a white preocular
transverse band).
Variation. e head is red and might contain irre-
gular black markings, black-tipped scales, or both.
A white transverse band and a black cephalic cap
are variable in length. e black cephalic cap usually
reaches the anterior ⅔ (57.6%), ½ of the parietals
(26.4%), or the anterior ⅓ (6.9%), but rarely reach-
es the entire parietals (9%). Ventrally, the chin
contains irregular black markings that include
the mental and most or all of the infralabials, in a
typical horseshoe-shaped pattern (86.4%). Other
patterns may include irregular semicircles (8.5%),
black posterior infralabials (2.2%), black anterior
gulars (1.4%), and other aberrant forms (1.5%) (Fig.
49). In males (N = 301), the ventrals = 186–270, the
subcaudals = 20–41, and the body triads = 7–18; the
HL = 7.9–38.0 mm, the SVL = 210–1,550 mm, and
the TL = 12.8–100.9 mm. In females (N = 132), the
ventrals = 210–276, the subcaudals = 21–40, and
the body triads = 9–16; the HL = 8.0–34.3 mm, the
SVL = 96–1,274 mm, and the TL = 13.1–85.7 mm.
e 1st triad is separated from the parietals by 2–5
vertebral scales: 3 (54%), 4 (31%), 2 (11%), and 5
(4%). e black middle triad rings might be longer
than the outer rings (84.8%), or all three might be
semi-equal in length (15.2%). e white rings vary in
length (½–3 dorsal scales) with the majority (84.5%)
consisting of 1½ (33.8%), 2 (30.4), and 2½ (20.3%)
scales. e tail triads are 1–2, with a predominance
of the 1⅔ pattern (1⅔ = 50%; 1 ⅓ = 30.6%; 2 = 16.2%;
1= 3.2%). e degree of black pigmentation on the
red rings is variable and can be irregular (86%),
with all the scales containing black tipping (7.3%)
or immaculate (6.7%). Similarly, the white rings can
be irregular and marked with black (31.3%), imma-
culate (22%), or the scales might contain ne black
tipping (46.7%).
Fig. 49. Variation in the head coloration of Micrurus diutius:
A = MNHN 1989.3151; B = MPEG 21169; C = FMT 1820;
D = FMNH 34472; E = MPEG 22281; and F = INPA 11122.
Drawings: Matheus G. Pires.
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Advances in Coralsnake Biology: with an Emphasis on South America 71
Hemipenis. MZUSP 5465 (Barcelos, Amazonas,
Brazil; right organ). e hemipenis is short, bi-
lobed, capitate, and covered with large and irregularly
distributed spines on both the sulcate and asulcate
surfaces, which decrease in size from a proximal to
distal direction. e lobes are approximately ¼ of the
total length of the organ, or ⅓ the length of the capi-
tulum. e interlobar region contains a few smaller
spines, which are more visible on the asulcate side.
e capitular groove is deep, and crosses the organ
perpendicularly at the distal limit of the proximal ¼
of the organ, and on both sides reaches the sulcate sur-
face on a decreasing curve until reaching the margins
of the sulcus spermaticus. e capitulum is ¾ of total
length of the hemipenis and contains a large number
of
spines, which are large and irregular along the cir-
cumference of the organ, and almost extend to the
capitulum. e spines decrease slightly in size and
number along the base of the lobes, especially on the
asulcate surface. e sulcus spermaticus is deep and
contains smooth borders, and bifurcates at a more
proximal level than the base of the lobes and on
the distal ⅓ of the hemipenis. Aer bifurcating, the
sulcus spermaticus extends centripetally to the dis-
tal tip of the lobes. e body is covered with much
smaller spines that are larger on the distal region,
and decrease in size toward the base of the organ,
which contains unevenly distributed smaller spines.
e basal pocket is small and naked (Fig. 40B).
Distribution. Micrurus diutius occurs in Trinidad
and Tobago and in the Guianan region, including
eastern Venezuela, to northern Amazonia in Bra-
zil, in the states of Amazonas, Pará, Amapá, and
Roraima (Fig. 45).
Remarks: Burger (1955) described this taxon as a
subspecies of Micrurus lemniscatus based on three
specimens (all males), with the holotype (FMNH
34472) from Tunapuna, Trinidad, and two paratypes,
one (CM 7925) from Cumanacoa, Sucre, Venezuela,
and the other (UIMNH 22626) from Hoso Rora,
near Morawhanna, in northeastern Guyana. e
diagnosis of M. diutius was based on the pres ence
of heavily melanic white rings, moderately melanic
red rings, a fewer number of ventrals (219–226 in
males, 226–239 in females), and an overall longer
body when compared to M. lemniscatus.
Gasc and Rodrigues (1980) and Roze (1986) also
considered Micrurus diutius as a subspecies of M.
lemniscatus. Cunha and Nascimento (1980) report-
ed the rst record of this taxon from Brazil, and
Chippaux (1986) and Welch (1994) cited other spe-
cimens. Starace (1998) noticed dierences between
M. l. lemniscatus and M. l. diutius, and reported
the sympatric occurrence of both subspecies in
French Guiana and questioned the cataloguing of
these two forms as subspecies. Gorzula and Seña-
ris (1999) commented on the apparently disjunct
populations along coastal southeastern Brazil, in an
apparent equivocal inversion of Micrurus ibiboboca/
Elaps marcgravii reports for the Guianan region.
Later, Frota et al. (2005) rectied the identication
of a specimen considered by Guedes (2000) as M.
frontifasciatus, and determined it as M. l. diutius,
followed by Natera-Mumaw et al. (2007), who re-
corded the westernmost known locality for this
taxon at Parcelamiento El Cují, near Tocuyito, Cara-
bobo, Venezuela.
As with other taxa in the Micrurus lemniscatus
complex, the degree of melanism on the red and
white rings varies widely in M. diutius. e rings of
the specimens herein assigned to this taxon range
from immaculate to extremely melanic, which also
applies to the chin. is problem originated with
Boulenger (1896), who combined the NHMUK spe-
cimens from Trinidad and Tobago (with fewer ven-
tral scales and longer white rings) with specimens
from eastern Brazil (M. ibiboboca) having these
characteristics (previously identied by Günther
[1858] as Elaps lemniscatus) as Elaps marcgravii.
Further, Werner (1901) also reported a specimen of
E. marcgravii from Trinidad and repeated the same
determination for specimens from Peru and Bolivia
(Werner, 1901) with fewer ventral scales, as had Boett-
ger (1894), even though the latter noted numerous
similarities with M. lemniscatus.
Beebe (1919) reported the occurrence of
Micrurus marcgravii and M. lemniscatus in Guyana,
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72 M. G. Pires, D. T. Feitosa, F. G. Grazziotin, A. L. C. Prudente, N. J. Silva, Jr., and H. Zaher
and adopted the generic name Micrurus Wagler,
1824 in place of Elaps Schneider, 1801, as used by
Stejneger and Barbour (1917). Amaral (1926) agreed
with this decision, despite his insistence on consider-
ing Elaps lemniscatus (sensu Schneider) as a com-
posite of M. lemniscatus and M. ibiboboca, but of M.
lemniscatus and other forms that were not indicated.
Despite advancements in nomenclature, the
taxonomic problem of the distinction between
Micrurus ibiboboca and M. lemniscatus by including
M. diutius as a subspecies of the latter remained,
either because of the scarcity of museum specimens,
or the lack of knowledge of their morphological
variation and geographic distribution. Further, this
situation was aggravated when Moonen et al. (1979)
included a photograph of M. l. diutius as M. ibibo-
boca, with long white rings, but did not acknowl-
edge the diagnostic characters between these spe-
cies, such as ventral scale counts. Campbell and
Lamar (1989), Golay et al. (1993), Roze (1996), Sta-
race (1998), David and Ineich (1999), Abuys (2003),
and Kok et al. (2003) followed the same reasoning.
ese problems began to be addressed when
Campbell and Lamar (2004) suggested the possi-
bility that these specimens were in fact Micrurus l.
diutius, and nally, MacCulloch and Lathrop (2004)
treated the subject and rejected the presence of M.
ibiboboca in the Guianan region. Curiously, Cole et
al. (2013) did not mention the occurrence of M. diu-
tius in Guyana.
Micrurus frontifasciatus (Werner, 1927) —new
status
Elaps frontifasciatus Werner 1927: 250.
Micrurus lemniscatus—Amaral, 1929: 55
(partim).
Micrurus lemniscatus frontifasciatus—Roze,
1967: 34.
Micrurus frontifasciatus—Campbell and Lamar,
1989: 116.
Micrurus lemniscatus frontifasciatus—Roze,
1996: 175.
Micrurus lemniscatus—Campbell and Lamar,
2004: 222.
Micrurus frontifasciatus—Roze, 1996: 175.
Micrurus lemniscatus—Harvey, Aparicio-E, and
González-A, 2003: 13.
Micrurus frontifasciatus—Pires, 2011: 213.
Holotype: NMW 18298 (M) (Fig. 50).
Type locality: Bolivia.
Fig. 50. Holotype of Micrurus frontifasciatus (NMW
18298). Photo and drawings: D. T. Feitosa
Diagnosis. Micrurus frontifasciatus can be distin-
guished from all other triadal species of Micrurus by
the following combination of morphological charac-
ters: black snout (rostral, internasals, anterior border
of prefrontals, most of nasals, and 1–2 supralabials);
white preocular transverse band covers most of pre-
frontals, anterior border of supraoculars, most of
preoculars, and 1–2 supralabials; black cephalic cap
includes most of frontal, supraoculars, postoculars,
anterior ¼– ⅓ of parietals, ⅓–½ of temporals, and
2–3 supralabials; remainder of head red, with black-
tipped scales; venter of head red; irregular black
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Advances in Coralsnake Biology: with an Emphasis on South America 73
markings on gulars, and infralabials usually small; 1st
triad separated from parietals by 4–6 scales; middle
black ring longer than outer ones; white rings shorter
than outer black rings; red rings shorter than entire
triad; red and white rings usually black-tipped; and
8–11 body triads and 1⅓–1⅔ tail triads. Due to the
rarity of this species, it was not possible to examine
the hemipenis (Fig. 51).
Fig. 51. A live individual of Micrurus frontifasciatus from
San Sebastián, Santa Cruz, Bolivia. Photo: Michael Jensen.
Variation. e head is red and sometimes contains
irregular black markings (especially on the frontals),
black-tipped scales, or both. A black cephalic cap
reaches the anterior ⅓, and rarely ½ of the parietals
(5%). Ventrally, the chin is mostly red with irregular
black markings that may include the mental and the
1st three infralabials (96.5%). Sometimes it includes
the anterior 1 ⅓–½ of the anterior gulars (2.3%), or
the markings are irregularly distributed (1.2%) (Fig.
52). In males (N = 11), the ventrals = 212–229, the
subcaudals = 29–40, and the body triads = 8–11.
e HL = 10.5–26.9 mm, the SVL = 223–927 mm,
and the TL = 21.3–70.6 mm. In females (N = 5), the
ventrals = 227–245, the subcaudals = 29–41, and the
body triads = 9–11. e HL = 10.4–19.5 mm, the
SVL = 230–600 mm, and the TL = 19.0–51.3 mm.
e 1st triad is separated from the parietals by 2–4
vertebral scales: 2 (63.7%), 3 (31%), and 4 (19%).
e black middle triad rings are longer that the outer
rings. e white rings vary in length (1½–3 dorsal
scales), and consist of 1½ (29.4%), 2 (56.6%), and 3
(14%) scales. e tail triads are 1–1⅓, with a predo-
minance of the 1⅓ pattern (97.6%). e degree of
black pigmentation on the red rings is variable and
can be irregular (30%), with all the scales containing
black tipping (62%) or immaculate (8%). e white
rings can be immaculate (10.3%), or contain scales
with black tipping (89.7%).
Fig. 52. Variation in the head coloration of Micrurus
frontifasciatus: A = NMW 18298; B = ZMH-R 2702; C =
ZMH-R 2703; D = ZSM 43-1029a; E = ZMH-R 2705; and
F = SMF MJ 637 (eld number). Drawings: Matheus G.
Pires.
Comparison with sympatric species. Micrurus
frontifasciatus occurs sympatrically with one spe-
cies of Leptomicrurus (L. narduccii), one species of
monadal (M. annellatus), and ve triadal species
of Micrurus (M. hemprichii, M. lemniscatus, M.
obscurus, M. spixii, and M. surinamensis). Because
of the limited amount of information available for
this species, it also could be sympatric with another
species of Leptomicrurus (L. scutiventris), and three
monadal (M. langsdor, M. ornatissimus, and M.
putumayensis) and ve triadal species of Micrurus
(M. diana, M. liformis, M. mipartitus, M. pyrrho-
cryptus, and M. serranus). Micrurus frontifasciatus
can be distinguished from L. narduccii by contact
of the anteriormost infralabials behind the mental
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74 M. G. Pires, D. T. Feitosa, F. G. Grazziotin, A. L. C. Prudente, N. J. Silva, Jr., and H. Zaher
scale, and a triadal pattern (vs. the 1st infralabials
separated by contact between the mental and ante-
rior chin shields, and a bicolored pattern consisting
of incomplete rings). It diers from M. annellatus by
the presence of a triadal color pattern (vs. a monadal
pattern); from M. lemniscatus by a lower mean num-
ber of ventrals (224.5 vs. 245.8), a stout body (vs. a
slender body), and reduced chin ornamentation (vs.
a predominantly black horseshoe-shaped pattern on
the chin) (see Remarks); and from M. hemprichii, M.
spixii, M. obscurus, M. nattereri, and M. surinamen-
sis by the presence of a white prefrontal transverse
band (vs. the absence of a white prefrontal trans-
verse band).
Distribution. Micrurus frontifasciatus occurs along
the eastern Andean slopes of Peru and Bolivia, in
places such as Yungas, Puna, and the Chiquitania
forests, at elevations from 550 to 2,840 m (Fig. 45).
Remarks: Micrurus frontifasciatus originally was
described by Werner (1927) based on a single speci-
men collected by Staudinger in Bolivia, and this taxon
remains poorly represented in museum collections.
Campbell and Lamar (1989) reported the type speci-
men as unnumbered, based on a translation of Wer-
ner’s description, which did not include a museum
number for the holotype, and they also examined a
photograph of a specimen from near the type loca-
lity; consequently, they decided to treat this taxon as
a species; in their 2004 book, they listed it under M.
lemniscatus (W. Lamar, pers. comm.). While visit-
ing the NMW collection (in Austria) in 2012 and
2015, we were able to examine the specimen. is
species is similar to M. lemniscatus, which genera-
ted remarkable taxonomic instability, as it some-
times was placed as a subspecies of M. lemniscatus
or included in the synonymy of the latter (Amaral,
1929; Campbell and Lamar, 1989, 2004; Roze, 1983,
1996). Despite this fact, our sample exhibits a com-
bination of characters that characterizes and die-
rentiates it from the taxa within the M. lemniscatus
species complex. Due to the lack of specimens, we
were unable to include morphological data on the
hemipenis or the skull.
Micrurus potyguara Pires, Silva Jr., Feitosa,
Prudente, Alves Filho, and Zaher, 2014
Micrurus potyguara Pires, Silva Jr., Feitosa, Prudente,
Alves Filho, and Zaher, 2014: 571.
Holotype: UFPB 4359 (M).
Paratypes: UFPB 4355 (F), UFPB 4358 (M), UFPB 4361
(F), NRM 1685 (M), and CAS 49297 (M).
Type-locality: Mata do Buraquinho, Municipality of
João Pessoa, Paraíba, Brazil.
Diagnosis. Micrurus potyguara can be distinguished
from all other triadal coralsnake species by the
following combination of morphological characters:
black snout (rostral, most of internasals, anterior
border of prefrontals, most of nasals, and 1–2 supra-
labials); white preocular transverse band covers
prefrontals (sometimes reaches anterior border of
frontal), preoculars, and posterior border of the 2nd,
3rd, and anterior border of 4th supralabials; behind
white band, a black cephalic cap includes frontal
(or most of it), supraoculars, postoculars, parietals,
⅓ of temporals, and 1–2 supralabials (usually part
of 4th, 5th, and part of 6th); remainder of head red
with heavily black-tipped scales; venter of head red;
infralabials mostly black, including part of gulars; 1st
triad separated from parietals by three scales; middle
black ring slightly longer or same length as external
ones; white rings shorter than external black rings;
red rings shorter than entire triad or even middle
black ring; red and white rings tipped with black,
with tendency to melanic pattern; and triads 9–12
plus 1⅓–1⅔ on tail (Fig. 53).
Comparison with sympatric species. Micrurus
potyguara occurs sympatrically with one species of
monadal Micrurus (M. corallinus) and two triadal
species (M. carvalhoi and M. ibiboboca). Micrurus
diutius can be dierentiated from M. corallinus (a
monadal species) by its distinctive pattern of black
rings. Micrurus potyguara diers from M. carva-
lhoi by the presence of a higher number of ventrals
in females (253–263 vs. 221–268), a lower number
of triads (9–12 vs. 9–18), longer white rings, the
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Advances in Coralsnake Biology: with an Emphasis on South America 75
parietals are entirely black (vs. the anterior ⅓ to ½
of the parietals black), and a semicapitate hemipe-
nis (vs. capitate); it diers from M. ibiboboca by the
pres ence of a higher number of ventrals (231–263 vs.
188–254), a higher number of subcaudals (29–38 vs.
16–34), and the parietals are completely black (vs.
the anterior ⅓ to ½ of the parietals black). Micrurus
ibiboboca also has a tendency for containing a white
snout, and longer white rings that usually are imma-
culate on the anterior ⅓ of the body.
Fig. 53. A live individual of Micrurus potyguara. Photo:
Gentil Alves Pereira Filho.
Distribution. Micrurus potyguara is known from
remnants of the Atlantic Forest along the coast of
northeastern Brazil, which today includes the states
of Paraíba, Pernambuco, and Rio Grande do Norte.
Remarks: is taxon recently was described from
the northeastern coastal areas of Brazil, derived
from Micrurus carvalhoi (Pires et al., 2014), and
because of its geographical range might be confused
with populations of M. ibiboboca.
6. CONSIDERATIONS
One of the most important tasks in biologic al research
is the ability to recognize the relevance of all aspects
of taxonomic research. e basis of taxonomy is old
and remains formally organized on the contribution
of Linnaeus (1758). e relevance of the binomial
system of taxonomy is accepted worldwide, but its
regulations constantly are being adapt ed to cope
with advancing descriptive techniques (biochemical
and molecular). Nevertheless, maintaining a precise
history of the type series undoubtedly is one of the
pillars of taxonomy and always will be the basis of
our knowledge of biodiversity, and one of the ways
of valuing museum specimens (and museums them-
selves) as a depository for this information.
Even so, it remains dicult to diagnose the spe-
cies in the Micrurus lemniscatus complex based only
on meristic and morphometric data. is problem
frequently occurs with many species of the genus
Micrurus, and has been well documented in dierent
studies (Silva, Jr. and Sites, Jr., 1999, 2001; Di-Ber-
nardo et al., 2007; Feitosa et al., 2007, 2015; Pires et
al., 2014;). We based our taxonomic decisions on
a combined analysis of these data with other diag-
nostic morphological characters, including the head
and body color pattern, triad pattern, cranial and
hemipenial morphology, and the eye diameter as it
relates to the distance between the lower margin of
the orbit and mouth (de Silva, Jr. and Sites, Jr., 1999;
Di-Bernardo et al., 2007), and also from previously
generated molecular data (Zaher et al., 2016) and
from this work.
We redescribed Micrurus lemniscatus based on
morphological characters, using an ample sample
size that covers most of the known geographic range
of this taxon. We were able to analyze all the types
related to this species. In fact, Linnaeus’ original des-
cription in 1758 did not include a museum voucher
numbers, but only oered general information on
the specimens identied as Coluber lemniscatus.
Originally, the species was described based on
two syntypes (NRM 93, NRM 94) but a third (NRM
95) and a fourth (UUZM 06) still exist, but never
were included in any description or discussion about
this species complex. Historical information and
museum records claried this situation, and herein
we proposed a more stable taxonomic arrange ment
with NRM 93 as the lectotype, and NRM 94, NRM
95, UUZM 06, ZISP 1258, ZISP 1261, ZISP 1262,
ZISP 1263, and ZISP 1264 as paralectotypes.
Advances in Coralsnake Biology Ch. 2.indd 75Advances in Coralsnake Biology Ch. 2.indd 75 12/18/20 2:51 PM12/18/20 2:51 PM
76 M. G. Pires, D. T. Feitosa, F. G. Grazziotin, A. L. C. Prudente, N. J. Silva, Jr., and H. Zaher
We believe that any suggestion to synonymize
Micrurus frontifasciatus with M. lemniscatus is ina-
dequate, because morphologically M. frontifasciatus
is more similar to M. diutius. In doing so, M. fronti-
fasciatus would be composed of two disjunct popu-
lations, an Andean and a Guianan, which conicts
with the known theories of speciation. Recently,
the mistake of using disjunct populations for spe-
cies assumptions has been made for other species
of coralsnakes. Despite the reasonable sample sizes
for M. lemniscatus and M. diutius, the problem with
M. frontifasciatus is that morphological data remains
unavailable for the skull and hemipenis. Because of
the lack of conclusive data that could be used to shed
light on the relationship between M. diutius and M.
frontifasciatus, as a conservative measure we suggest
maintaining both taxa as full species.
Roze (1967) reallocated Micrurus helleri as a
subspecies of M. lemniscatus (Roze, 1967), and its
status has remained unchanged (Campbell and
Lamar, 1989, 2004; Roze, 1996) although no one has
examined and compared specimens of this taxon
with its congeners or recognized their taxonomic sta-
tus. By using a combination of characters (meristic,
morphological, and molecular), we did not nd any
statistical or morphological evidence to support this
arrangement. Consequently, we propose to include
M. l. helleri as a junior synonym of M. lemniscatus.
Converesely, many characteristics support the
elevation of Micrurus lemniscatus carvalhoi to full
species status (M. carvalhoi), and we suggest recog-
nizing these taxa as distinct entities.
Further, the complex taxonomy of the Micrurus
lemniscatus is highlighted with the clear distinction
of the Atlantic coast and interior populations of
South America, as well as strong support for validat-
ing M. altirostris, M. baliocoryphus, and M. pyrrho-
cryptus (Silva, Jr. and Sites, Jr., 2001; Castoe et al.,
2007; Renjifo et al., 2012; Pyron et al., 2011, 2013;
Lee et al., 2016; Zaher et al., 2016, 2019).
Within Micrurus lemniscatus species complex,
we were able to separate and better characterize the
taxonomic units, as follows: (A) Micrurus lemnis-
catus—Amazonia and north of South America; (B)
Micrurus carvalhoi—a wide corridor on the Atlantic
coast and part of central Brazil and northeastern Para-
guay and Argentina; (C) Micrurus diutius—north-
ern South America; and (D) Micrurus potyguara—
eastern Brazil.
From the previously recognized taxa, Micruus
lemniscatus still has the broadest geographical distri-
bution, and pending further studies with com bined
morphological and molecular evaluations, still may
contain other forms that might be recog nized as
separate species.
ACKNOWLEDGMENTS
We are indebted to Eric Åhlander and Sven Kullan-
der for the most precious help and hospitality of-
fered at the Naturistoriska Riksmuseet (NRM) in
Stockholm, and Karolina Larsson, Anders Nilsson,
and Hans Mejlon from the Uppsala Universitet Evo-
lutionmuseet (UUZM), in Uppsala. We also acknow-
ledge Konstantin Milto and Nikolai L. Orlov (RAS)
for their help with the Russian Linnean specimens.
We also thank the following curators and respective
curatorial stas for providing access to specimens
under their care: Darrel Frost and David Kizirian
(AMNH); Ned Gilmore (ANSP); Patrick Campbell,
David Gower, Colin McCarthy, and Mark Wilkinson
(NHMUK); Jens Vindum (CAS); Carlos Grisolía
(CCG); Hélder Lúcio Rodrigues Silva and Wilian
Vaz Silva (CEPB); José M. Gallardo (CHINM); Guar-
ino R. Colli (CHUNB); Stephen Rogers (CMNH);
Pablo Venegas and Lurdes (CORBIDI); Antônio
Jorge Suzart Argôlo (CZGB and MZUESC); Harold
Voris and Alan Resetar (FMNH); Jean Samonek and
Carlos Jatobá (FMT); Claudia Medina and Carlos
Montana (IaVH); Giuseppe Puorto and Francisco L.
Franco (IBSP); Martha Calderón and Bessy (ICN);
Fernanda Werneck (INPA); Anibal Rafael Melgarejo
Gimenez (IVB); Rafe Brown (KU); Jonathan Losos
and José Rosado (MCZ); Juan Carlos Chaparro-Auza
(MHNC); Claudia Torres, Jesús Santiago Córdova,
Betty Milán, and Rina Ramírez (MHNJP – Univer-
sidad Nacional Mayor de San Marcos); María del
Pilar Rivas (MHNUC); Juan Daza (MHUA); Hno.
José Edilson Espitia (MLS); Júlio César Moura Leite
Advances in Coralsnake Biology Ch. 2.indd 76Advances in Coralsnake Biology Ch. 2.indd 76 12/18/20 2:51 PM12/18/20 2:51 PM
Advances in Coralsnake Biology: with an Emphasis on South America 77
(MNHCI); Ivan Ineich (MNHN); Martha Motte
Paredes (MNHNP); Paulo Passos (MNRJ); Roberto
Gutierrez and Roy Santa Cruz Farfán (MUSA);
Carol L. Spencer, and Jimmy A. McGuire (MVZ);
Flora Acuna Juncá (MZUEFS); Heinz Grillitsch
(NHMW); Robert H. Murphy and Pedro Henrique
Bernardo (ROM); Gunther Köhler (SMF); Daniel
Oliveira Mesquita (UFPB); Christopher A. Phillips
(UIMNH), Ronald Nussbaum and Gregory Schnei-
der (UMMZ); W. Ronald Heyer, Roy McDiarmid,
Kenneth Tigue, Robert Wilson, and Steve Gotte
(USNM); Karolina Larsson, Mats Eriksson, Anders
Nilsson, and Hans Mejlon (UUZM); Mark-Oliver
Rödel and Christoph Kucharzewski (ZMB); Alexan-
der Haas and Jakob Hallermann (ZMH); Frank Glaw
and Michael Franzen (ZSM); and Paulo Roberto
Manzani (ZUEC).
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