Noteworthy dietary records of the Variable Coral Snake Micrurus diastema (Serpentes: Elapidae) in America

Abstract

Diet is an essential component of the natural history and ecology of organisms. Many snake species' trophic ecology is poorly characterized, mainly because predation events are rarely documented in nature. The diet of Micrurus diastema has been reported so far to include only other snakes. To obtain further knowledge on the species diet, we performed intermittent field excursions in Veracruz, Mexico, and surveyed published information for this snake in Mexico. Thereby, we report four items of prey in the diet of M. diastema, three of which are previously unreported, including a lizard (Sauria: Holcosus amphigrammus, Serpentes: Amerotyphlops tenuis, and Drymarchon melanurus); the fourth prey previously reported is another snake (Serpentes: Coniophanes imperialis). Our results extend the knowledge on the Variable Coral Snake's trophic ecology in the northern extreme of its geographical distribution in Mexico.

Full text

NORTH-WESTERN JOURNAL OF ZOOLOGY 17 (1): 100-105 ©NWJZ, Oradea, Romania, 2021
Article No.: e211507 http://biozoojournals.ro/nwjz/index.html
Noteworthy dietary records of the Variable Coral Snake
Micrurus diastema (Serpentes: Elapidae) in America
Edgar Ahmed BELLO-SÁNCHEZ1,2, Christian Alejandro DELFÍN-ALFONSO2,*,
Carlos PÉREZ-ALVARADO3 and Felipe LARA-HERNÁNDEZ3
1. Laboratorio de Biología de la Conducta, Instituto de Neuroetología, Universidad Veracruzana,
Av. Luis Castelazo Ayala S/N, Col. Industrial Ánimas, Xalapa, Veracruz 91190, México.
2. Laboratorio de Zoología, Instituto de Investigaciones Biológicas, Universidad Veracruzana,
Av. Luis Castelazo Ayala S/N, Col. Industrial Ánimas, Xalapa, Veracruz 91190, México.
3. PIMVS, Herpetario Palancoatl. Avenida 19, 5225, Córdoba, Veracruz 94540, México.
* Corresponding autor, C.A. Delfín-Alfonso, E-mail: cada7305@gmail.com
Received: 09. April 2020 / Accepted: 23. March 2021 / Available online: 25. March 2021 / Printed: June 2021
Abstract. Diet is an essential component of the natural history and ecology of organisms. Many snake species' trophic ecology is
poorly characterized, mainly because predation events are rarely documented in nature. The diet of Micrurus diastema has been
reported so far to include only other snakes. To obtain further knowledge on the species diet, we performed intermittent field
excursions in Veracruz, Mexico, and surveyed published information for this snake in Mexico. Thereby, we report four items of prey
in the diet of M. diastema, three of which are previously unreported, including a lizard (Sauria: Holcosus amphigrammus, Serpentes:
Amerotyphlops tenuis, and Drymarchon melanurus); the fourth prey previously reported is another snake (Serpentes: Coniophanes
imperialis). Our results extend the knowledge on the Variable Coral Snake's trophic ecology in the northern extreme of its
geographical distribution in Mexico.
Key words: Amerotyphlops, diet, Drymarchon, Holcosus, predator, prey.
Introduction
The New World coral snakes are represented by two genera
(Micruroides and Micrurus) and 89 species that inhabit tropi-
cal and subtropical regions in the Americas (Castoe et al.
2007, Uetz 2020). The species of this genus are venomous
and consume vertebrate and invertebrate prey (Roze 1996,
Campbell & Lamar 2004). The diet of some Micrurus spp.,
such as the Eastern Coral Snake M. fulvius (Linnaeus, 1766),
Texas Coral Snake M. tener Baird & Girard, 1853 and Dou-
ble-Banded Coral-Snake M. altirostris (Cope, 1860), is well-
documented (Jackson & Franz 1981, Greene 1984, Roze 1996,
Reams et al. 1999, da Silva & Aird 2001, Campbell & Lamar
2004). However, little is known about the feeding ecology of
many other coral snakes due to their elusive behavior and
fossorial habits (Cavalcanti et al. 2010).
The Variable Coral Snake, M. diastema (Duméril, Bibron
& Duméril, 1854), is a medium-sized snake (average total
length: 600–750 mm but can exceed 900 mm); it occurs along
the Atlantic Coast from the center of the state of Veracruz in
Mexico to northwestern Honduras (Campbell & Lamar
2004). However, a recent molecular study of the species
changed its taxonomy, suggesting a separation of species
and reducing the number of taxa as well as their geographic
distribution for the M. diastema group (Reyes-Velasco et al.
2020), restricts its distribution only to the low and moderate
elevations of the Atlantic drainage from central Veracruz,
Tabasco, and northern Oaxaca, which was formerly the dis-
tribution of M. d. diastema, M. d. affinis, and M. d. macdougalli
(Campbell & Lamar 2004). The Variable Coral Snake M. dia-
stema occurs in the tropical rain forest and deciduous forest
at lowlands and cloud forest and mixed forest at highlands
(Campbell & Lamar 2004, Reyes-Velasco et al. 2020). The
records in the literature about the diet of Micrurus diastema
are scarce and only include snakes (Rodríguez-García et al.
1998, West et al. 2019). Therefore, the present contribution is
relevant because we point out the records of items unreport-
ed in the literature about the diet of M. diastema, three of
which are previously unreported in this coral snake's diet.
Material and Methods
Study area
We performed intermittent field sampling, specifically for M. diaste-
ma from 2011 to 2016, in the municipalities of Atoyac and Amatlán
de Los Reyes in the region known as "the High Mountains" and in
the Actopan municipality in the Coastal Plain of the Gulf, both re-
gions in central Veracruz state, Mexico (Fig. 1). These municipalities
are dominated by tropical, subtropical, and semi-temperate vegeta-
tion, including cloud mountain forest and tropical deciduous forest;
mono- and polycultures are also present, mostly sugar cane and cit-
rus trees (INEGI 2016). The study area encompasses elevations from
sea level (e.g., Actopan) to 1600 meters above sea level (m a.s.l.) (e.g.,
Atoyac). The average rainfall varies between 800 and 1500 mm year-
ly; the average temperature ranges between 20 °C in the upper part
of Atoyac and 28 °C in Actopan (García 2004).
Diet data review
Considering the new reclassification of the species (Reyes-Velasco et
al. 2020), we carried out a literature review for the diet of M. diaste-
ma, excluding cases for taxa previously included in the species. We
focused on the search terms "Micrurus diastema + diet + feeding +
Variable Coral Snake + diet + serpiente coralillo del sureste + dieta +
alimentación"; we used three online sources: ResearchGate
(www.researchgate.net), BioOne (https://bioone.org/), and Google
Scholar (https://scholar.google.com/). The information was com-
piled in a table and projected on a map to spatially locate the studies
in a geographic context. For visualization purposes, we used specific
coordinates when available or we estimated a centroid using the spa-
tial references of the studies, through the software Google EarthTM
platform. We used the M. diastema distributions maps generated by
Campbell & Lamar (2004) and Acevedo et al. (2020) to geographical-
ly locate contributions, which allowed us to identify geographical
gaps in the diet study of the species. All information was processed
and projected in geographic coordinates in ArcMap 10.3 (ESRI 2014),
to help us identify geographical gaps in the diet study of the species.

Dietary records of the Variable Coral Snake
101
Sampling methods
In October 2011, were carried out surveys in the municipality of
Amatlán de Los Reyes; in October 2013, other surveys were carried
out in the municipality of Atoyac; finally, in March 2016, field sur-
veys were carried out in the municipality of Actopan, very close to
the coast of the Gulf of Mexico.
We carried out in each of the three municipalities two days ran-
dom tracks snake visual encounter surveys. The same tracks were
surveys during the day (10:00 to 16:00 h) and then at night (19:00 to
22:00 h), for a total of six days and 12 km sampled in the three sites,
by five persons to document visual encounters. The tracks frequently
crossed various plant associations of tropical and subtropical forests,
rural roads, secondary vegetation, grasslands and pastures, crops,
and even without vegetation areas.
During the surveys, we captured the coral snakes that we locat-
ed alive; each was weighed (using a digital scale, to the nearest g),
photographed with a delineated ruler (for subsequent measure mm),
and palpated to identify if it contained food in the stomach, and fi-
nally released at the capture site. Nevertheless, when dead snakes
(e.g., roadkill) were located, they were collected and processed for
deposit in collections (see below). We also recorded the coordinates
with a GPS Garmin eTrex 30®, the type of vegetation, and the sub-
strate (i.e., rocky, sandbanks, shrub, asphalt, fallen trees) of each in-
dividual. The photographs were used to perform morphometric
measurements on the individuals using ImageJ 1.46 software (Broeke
et al. 2015), widely used in the field of herpetology (Astley et al.
2017).
Furthermore, where possible, we recorded the sex of the indi-
vidual and identified their developmental stage (juvenile, adult),
based on aspects of its natural history (Lee 2000; Pérez-Higareda et
al. 2007). In rural areas of Latin America, farmers frequently end up
killing encountered snakes (Hernández & Bravo 2009, Gutiérrez
2011); therefore, we searched for such dead snakes (i.e., on roads and
around human settlements). Located dead specimens were fixed in
70% alcohol and deposited in the Colección Herpetológica del Insti-
tuto de Investigaciones Biológicas (CHIIB) (Xalapa) and of the Fac-
ultad de Biología (CH-FACBAC) (Córdoba) at Universidad Veracru-
zana, in Veracruz, Mexico.
Results
The bibliographic review provided scarce information on the
Variable Coral Snake diet because the recent taxonomic
change proposed by Reyes-Velasco et al. (2020) substantially
decreased the number of documented records of their diet.
We were able to collect only records of the diet of M. diaste-
ma from two references (Greene 1973, West et al. 2019) that
report four species of snake consumed from 1973 to 2020
(Table 1).
The sampling effort at the three collection sizes was 270
person-hours for 12 kilometers, with a low capture success,
Figure 1. Micrurus diastema diet records in Veracruz, Mexico (pub-
lished and this study). Observations: #1 – Geophis semidoliatus, #2 –
Chersodromus liebmanni, #3 – Holcosus amphigrammus, #4 – Dry-
marchon melanurus, #5 – Amerotyphlops tenuis, #6 –Coniophanes impe-
rialis).
and only three specimens located. The first individual we
found and collected was on 21 October 2011 at 15:00 h, 1.5
km northeast of Amatlán de Los Reyes, Veracruz, Mexico
(datum WGS84; 18.8584° N, 96.9007° W; 737 m a.s.l., Fig. 1).
The adult male M. diastema (snout-vent length, SVL = 900
mm; tail length = 105 mm) was found dead with many inju-
ries on the body (likely inflicted by local farmers) in a sugar
cane field near a group of trees (Fig. 2A). The examination of
the stomach contents revealed an adult female Rainbow
Ameiva, Holcosus amphigrammus (Smith & Laufe, 1945) (SVL
= 101 mm; Fig. 2B). Both specimens were conserved with
the same catalog number (CH-FACBAC-000667). We record-
ed the second snake next to the village of Caballo Blanco in
the municipality of Atoyac in Veracruz, Mexico (datum
WGS84; 18.8760° N, 96.7404° W; 423 m a.s.l.). An adult fe-
male, M. diastema (SVL = 686 mm; tail length = 80 mm), was
captured by local farmers during the surveys on 06 October
2013 at 01:02 h on a path near a tropical dry forest and "aca-
Table 1. Prey species recorded in the diet of the Variable Coral Snake (M. diastema) in Mexico.
Prey type
Cat. Number
Record site (coordinates)
References
Lizards
Holcosus amphigrammus
CH-FACBAC-000667
18.8584°N, 96.9007°W
This study
Snakes
Amerotyphlops tenuis
IIB-UV H00598
19.5019°N, 96.3897°W
This study
Chersodromus liebmann
UIMNH 19194
18.8840°N, 96.9260°W
West et al. 2019
Coniophanes imperialis
UCM 49376
Locality unspecified
Greene 1973, West et al. 2019
IIB-UV H00598B
19.5019°N, 96.3897°W
This study
Drymarchon melanurus
IIB-UVRe 0053f IIB-UVRe 0054f
18.8760°N, 96.7404°W
This study
Geophis semidoliatus
UIMNH 19193
18.8770°N, 97.0230°W
West et al. 2019
Ninia sebae
UIMNH 48822
Locality unspecified

E.A. Bello-Sánchez et al.
102
Figure 2. (A) An adult male Micrurus diastema collected in Amatlán de Los Reyes, Veracruz. (B) A Rainbow Ameiva (Holcosus amphigram-
mus) was found as part of the stomach contents of M. diastema.
hual" area (secondary vegetation). The individual was pho-
tographed by the locals (Fig. 3A) and placed in a cotton bag.
The snake's belly was distended with prey (Fig. 3A); while in
the bag for about 30 minutes, the snake regurgitated a likely
immature Western Indigo Snake, Drymarchon melanurus
(Duméril, Bibron, & Duméril, 1854) with an SVL= 605 mm,
tail length = 120 mm (Fig. 3B). The Variable Coral Snake
was released at the capture site; however, the prey (D.
melanurus) was prepared in a bottle with alcohol (70% con-
centration) and was preserved by local farmers as a souve-
nir; they could not donate it to herpetofauna collections. We
inspected the prey and identified lesions on its left side of
the Western Indigo; the marks on the parietal scale could be
coral snake teeth marks (Fig. 3C). The pictures were deposit-
ed at the Photographic Collection of Vertebrates "Alvar
González Christen" of the Universidad Veracruzana (cata-
logue numbers IIB-UVRe 0053f and 0054f).
A third M. diastema was found dead on the road near a
sugar cane field on 08 March 2016, at 09:00 h, at Paso del
Cedro locality in the municipality of Actopan, Veracruz,
Mexico (datum WGS84; 19.5019° N, 96.3897° W; 21 m a.s.l.;
Fig. 4A). The adult female snake (SVL = 802 mm; tail length
= 83 mm) was taken to the laboratory, where the stomach
content was examined, and we found the remains of a Cof-
fee Blind Snake, Amerotyphlops tenuis (Salvin, 1860), and a
Black-striped Snake, Coniophanes imperialis (Baird & Girard,
1859) (Fig. 4B). The specimens were deposited as IIB-UV
H00598 (M. diastema) and IIB-UV H00598B (prey).
Discussion
Literature reviews provide us with invaluable compendiums
of a particular topic, helping to condense important infor-
mation for decision making (Snyder 2019). In this sense, the
literature review of what is published about the diet of the
Variable Coral Snake M. diastema is no exception as it helps
substantially to understand a part of the trophic ecology of
this species. Our literature review shows that from 1973 to
2020, at least 28 species have been reported in the diet of
what was previously identified as the Variable Coral Snake
M. diastema, including fishes and specimens of M. diastema
(Blaney & Blaney 1978, Campbell 1998, Carbajal-Márquez et
al. 2019, Greene 1973, Köhler et al. 2016, Roze 1996, Seib
1985, West et al. 2019). However, the contemporary taxo-
nomic changes in the M. diastema group (Reyes-Velasco et al.
2020), significantly modify the results on prey data reported
previously for this species. We identified seven species in the
Variable Coral Snake diet (H. undulatus=H. amphigrammus,
Pseudoelaphe phaescens (Dowling, 1952), Ficimia publia Cope,
1866, Ninia sebae (Duméril, Bibron, & Duméril, 1854), Sibon
sanniolus (Cope, 1866), Stenorrhina freminvillei (Duméril,
Bibron, & Duméril, 1854), and Tantillita canula (Cope, 1875).
Thus, the number of prey species in the diet of the Variable
Coral Snake (as per Reyes-Velasco et al. 2020) has been re-
duced by 77% (from 28 prey species to 7) with the new taxo-
nomic arrangement.
Our results show that locating these snakes is difficult,
probably due to their abundance but also dependent on the
collectors’ expertise and other factors, leading to sampling
effort with high costs (e.g., economic resources and time)
and low success of capture (Rodríguez et al. 2018). Thus, we
suggest combining methods to increase detection and thus
obtain the much-needed data on the trophic ecology of this
snake with secretive habits.
Given the eight prey items recorded so far, the diet of M.
diastema includes seven vertebrate taxa: one lizard and six
snakes (Table 1; without considering prey consumed in cap-
tivity; Rodríguez-García et al. 1998). Our results suggest that
M. diastema prefers other snakes (ophiophagy); this is not
conclusive and further studies are necessary to support or
reject our statement. However, the genus Micrurus has been
documented to consume a wide variety of prey (i.e., Camp-
bell & Lamar 2004, West et al. 2019), such as Yellow-spotted
Night Lizard Lepidophyma flavimaculatum Duméril, 1851
(Roze 1996), West Forest Caecilian Gymnopis syntrema (Cope,
1866) (Campbell 1998), and the Blind Swamp Eel Ophisternon
infernale (Hubbs, 1938) (Köhler et al. 2016).
In this sense, the diet of M. diastema resembles those of
other North American elapids such as M. fulvius and M. ten-
er; 70 to 80% of the 25 reptile species documented in their di-
et were snakes (M. fulvius diet: Loveridge 1938, Obrecht
1946, Myers 1965, Jackson & Franz 1981, Greene 1984, Hein-
rich 1996, Roze 1996, Campbell & Lamar 2004, Wallin et al.
2013, Enge & Mays 2016; M. tener diet: Strecker 1908, Ruik
1948, Curtis 1952, Greene 1984, Roze 1996, Reams et al. 1999,

Dietary records of the Variable Coral Snake
103
Figure 3. (A) A female Micrurus diastema captured by a group of locals in Caballo Blanco, Atoyac, Veracruz. (B) A young Western Indigo
Snake (Drymarchon melanurus) regurgitated by M. diastema. (C) Probable marks infringed by coral snake teeth on the left side of the head
of D. melanurus.
Figure 4. (A) An adult female Micrurus diastema found dead on the road at Paso del Cedro, Actopan, Veracruz. (B) Remains of a Coffee
Blind Snake (Amerotyphlops tenuis) and a Black-striped Snake (Coniophanes imperialis) extracted from the digestive cavity of the Micrurus.
Campbell & Lamar 2004).
Regarding the items reported in this study, the Black-
striped Snake (C. imperialis) was previously recorded
(Greene 1973) in M. diastema diet. However, we found novel
prey species for the Variable Coral Snake, such that Coffee
Blind Snake (A. tenuis) might be an ordinary component of
the coral snake diet for its fossorial habits (Pérez-Higareda et
al. 2007). The discovery of a Western Indigo Snake (D.
melanurus) in the stomach of M. diastema also suggests that
this coral snake can hunt and consume prey approximately
its length. Although this is the first record of interaction be-
tween these two species, another coral snake species (M. dis-
tans Kennicott, 1860) has been reported to prey on D.
melanurus (Warfel et al. 2015). In both cases, the coral snakes
were adult females that consumed juvenile D. melanurus.
The Variable Coral Snake M. diastema seems to capture
and consume prey of different body sizes, from small blind
snakes like A. tenuis to snakes nearly as large as the predator

E.A. Bello-Sánchez et al.
104
(e.g., juvenile D. melanurus). This behavior has also been rec-
orded in the Eastern Coral Snake (M. fulvius; Greene 1984);
this seems to suggest that coral snakes have a preference for
small prey and can occasionally capture and consume prey
that may exceed its body mass (Cundall & Greene 2000), a
situation that probably characterizes the feeding behavior of
the Micrurus genus.
Similarly, the Rainbow Ameiva (H. amphigrammus) we
recorded for the first time in the diet of M. diastema, and the
first record of lacertids in its diet, was reported for M. apiatus
(Greene 1973), a member of the M. diastema group, and sup-
porting their close relationship (Reyes-Velasco et al. 2020).
The bibliographic review showed that the diet of the M.
diastema group (M. apiatus and M. diastema) is poorly known,
being most evident with the recent taxonomic changes
(Reyes-Velasco et al. 2020). The available records on the diet
of M. diastema (before taxonomy changes) were located at the
extreme of its geographic distribution, in the Petén region,
Huehuetenango and Alta Verapaz in Guatemala, in the Yu-
catán Peninsula area in the state of Quintana Roo, and in the
Gulf Coastal Plain and the High Mountains in Central Vera-
cruz, Mexico (see Fig. 1). However, with the current taxo-
nomic changes (Reyes-Velasco et al. 2020), the diet records
for M. diastema were reduced to only three reports, including
the present contribution; therefore, a geographic gap in the
knowledge of the diet of the species is detected, which in-
cludes the states of Oaxaca, Tabasco, and Veracruz. This is
not yet conclusive since there is no updated model of the ge-
ographic distribution of M. diastema that includes the new
taxonomic changes. In this context of proposed species
change, it is necessary to delimit the geographic distribution
of M. diastema using novel tools such as ecological niche
models (e.g., Kass et al. 2018, Osorio-Olvera et al. 2020).
Our results update and extend the knowledge on the
trophic ecology of the Variable Coral Snake in the northern
extreme of its geographical distribution in Mexico and refine
its place in the food web mainly as a snake-eating predator
throughout it geographical distribution, but they also em-
phasize the lack of information on this taxonomic group.
Thus, we urge that such research be continued to provide
additional information about the genus Micrurus natural his-
tory.
Acknowledgments. We are grateful to the reviewers (Dr. Uri García
and Dr. Larry David Wilson) and the editor Yurii Kornilev for
improving the manuscript; additionally, we thank the inhabitants of
Caballo Blanco for the support during the surveys in the field and
for the photographs and the staff at Facultad de Biología Campus
Córdoba and the Laboratorio de Zoología at the Instituto de
Investigaciones Biológicas of the Universidad Veracruzana for
vouchering the specimens in their collection. Eduardo Morteo,
Yadira Peralta, Dominic Olver, and Alvaro Villafañe assisted with
the preparation of the manuscript. Field equipment and logistical
support (e.g., GPS, vehicle, and others) were provided by the Project
"PRODEP: 40857 DSA/103.5/14/10595/PTC-749".
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