Andinobates bombetes

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AMPHIBIA: ANURA: DENDROBATIDAE Andinobates bombetes
Minyobates bombetes: Myers 1987:304.
Dendrobates bombetes: Jungfer, Lötters, and
Jörgens 2000:11.
Ranitomeya bombetes: Grant, Frost, Caldwell,
Gagliardo, Haddad, Kok, Means, Noonan,
Schargel, and Wheeler 2006:171.
Andinobates bombetes: Brown, Twomey,
Amézquita, de Souza, Caldwell, Lötters,
May, Melo-Sampaio, Mejía-Vargas, Pe-
rez-Peña, Pepper, Poelman, Sanchez-Ro-
driguez, and Summers 2011:36.
CONTENT. No subspecies are recognized.
DESCRIPTION. Individuals of Andinobates
bombetes have a body size (snout-vent length,
SVL) between 16.7– 21.5 mm, with no sexual
dimorphism in body size (males: mean SVL
= 17.8 ± 0.1 mm SD, range: 16.7–21.5 mm;
N = 28; females: mean SVL = 18.6 ± 0.1 mm
SD, range: 17.2–19.8 mm; N = 19) (Myers and
Daly 1980; Suárez-Mayorga 1999; Vargas-Sa-
linas and Amézquita 2013). In adults of Andi-
F 1. Male of Andinobates bombetes from Finca El Placer, in the municipality of Filandia, depart-
ment of Quindío, Colombia. Picture by Cristian González-Acosta.
Catalogue of American Amphibians and
Reptiles 926
F. Vargas-Salinas, M. A. Atehortua-Vallejo, L.
F. Arcila-Pérez, G. M. Jiménez-Vargas,
C. González-Acosta, S. Casas-Cardona,
and A. Grajales-Echeverry. 2020.
Andinobates bombetes.
Andinobates bombetes (Myers and Daly)
Rubí poison frog
Dendrobates bombetes Myers and Daly
1980:2. Type locality: “mountains above
south side of Lago de Calima, 1580–1600
meters elevation, about 2 km.,{sic} airline
southwest of Puente Tierra (village), De-
partment of Valle del Cauca, Colombia.”
Holotype, American Museum of Natural
History, AMNH 102601, adult female,
collected by C. W. Myers, J. W. Daly and
E. B. de Bernal on 21 November 1976 (not
examined by authors).

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M. Geographic distribution of Andinobates bombetes in Colombia, South America. Red
and yellow dots indicate populations where individuals exhibit red and yellow dorsolateral
stripes, respectively. e type locality is highlighted with a black dot in the center of the circle.

Account 926 3
nobates bombetes, the head is usually narrow-
er than the body, especially in gravid females;
greatest head width averaging about 32–33%
of SVL in adults. Nostrils are located near the
tip of the snout and are angled in a posterolat-
eral direction, the canthus rostralis is round-
ed, the loreal region is virtually at, the eyes
are positioned dorsally, and the eardrum is
hidden posterodorsally. Relative length of the
ngers is III>IV>II>I with an expanded ter-
minal disk on all digits except Finger I which
is short; in juveniles, terminal discs are rel-
atively less expanded. Relative length of the
toes is IV>III>V>II>I.
In life, the upper part of the body and
limbs are dark brown. Individuals possess
bright red or gold yellow (rarely light orange)
stripes that begin at the snout and extend over
the eyes, typically ending at midbody. Skin is
granular with granulation being particularly
coarse and strong on lower back and hind
limbs. e ventral surface is cream white with
brown blotches. e iris is dark brown in life
with little contrast from the pupil.
e morphology of seven tadpoles at
Gosner stage 25 (Gosner 1960) transported
on the back of parents was described by My-
ers and Daly (1980), and the morphology of
four tadpoles at Gosner stage 25, ve tadpoles
at Gosner stages 25–30, and two tadpoles at
Gosner stages 42–43 was described by Sán-
chez (2013). At Gosner stage 25, following
the description of Myers and Daly (1980), the
tadpole has a globular shape from a dorsal
view, with the body width averaging 79.4%
(74–92%) of the head-body length. e head
F 2. Dorsal view (A), lateral view (B), and mouth (C) of a tadpole of Andinobates
bombetes (Instituto de Ciencias Naturales [ICN] 42287, Museo de Historia Natural, Univer-
sidad Nacional de Colombia, Bogotá). Photographs courtesy of David Sánchez.

4
edges; the lower jaw sheath is V-shaped. Tad-
poles reported by Sanchez (2013) were de-
scribed as having a U-shaped jaw sheath, with
short guts with visible organs, and the nostrils
without a projection on the inner margin of
the nasal rim.
Individuals of Andinobates bombetes have
toxins in the skin (Myers and Daly 1980).
ose toxins are rich in piperidine alkaloids,
with at least 22 kinds; three of those alka-
loids (247, 251F, and 265B) are present only
in Andinobates bombetes; the fourth alkaloid
(217) is shared with other species within the
family Dendrobatidae. With the exception of
two unclassied compounds, all the alkaloids
found in Andinobates bombetes were assigned
to several classes of pumiliotoxins. Despite
the abundance of alkaloids in their skin tox-
and body are slightly convex above and at-
tened below. e eyes and the nostrils are in
a dorsal position. e spiracle is sinistral and
situated low on the body, and the cloacal tube
is dextral. e low-nned tail is 64.3% of the
total length; the end of the tail is rounded. e
average head-body length is 4.3 mm (range:
4.0–4.6 mm); the average greatest body width
is 3.4 mm (range = 3.1–3.7 mm); the average
total length of the tadpole is 12.2 mm (range
= 11.1–13.7 mm); the average greatest tail
depth from upper edge dorsal n to lower
edge ventral n is 1.7 mm (range = 1.6–1.8
mm). e morphology of the mouth consists
of an oral disc that is oriented anteroventrally
and the labial tooth rows are 2/3; the second
upper row of teeth has a central gap. e up-
per jaw sheath is solid with serrated cutting
F 3. Types of habitats where individuals of Andinobates bombetes have been recorded. A) Humid
montane forest; in this habitat individuals lives in places away from streams. B) Tropical dry forest; in this
habitat individuals are found alongside streams. Photographs by Fernando Vargas-Salinas.
A B

Account 926 5
ins, Andinobates bombetes is not highly toxic.
is species shows geographic variation in
the skin toxins through qualitative and quan-
titative dierences of alkaloids found in the
skin across dierent populations (Myers and
Daly 1980).
e advertisement call of Andinobates
bombetes is a ‘buzz’ with an average duration
of 1.28 s (range = 0.9–1.7 s), consists of ap-
proximately 160 pulses (pulse emission rate =
113–134 per second), and its dominant fre-
quency varies between 4000–5500 Hz (Brown
et al. 2011; Myers and Daly 1980; Vargas-Sali-
nas et al. 2014a). See Etymology.
Andinobates bombetes is easily distin-
guished from congeneric species based on
small size, and especially because of the dis-
tinctive color pattern and the rst nger be-
ing shorter than the second nger (Myers and
Daly 1980). Despite the descriptions of new
species and the subsequent rearrangement of
the phylogenetic relationships among species
(Amézquita et al. 2013; Brown et al. 2011;
Márquez et al. 2017), these characters are still
useful. For instance, Andinobates bombetes
exhibits a black or dark brown color with red
and yellow dorsolateral stripes, while Andi-
nobates opisthomelas is largely red (Brown et
al. 2011; Stuart et al. 2008), the posterior two-
thirds of the dorsal surface of Andinobates
virolinensis is dark brown (Ruiz-Carranza
and Ramírez-Pinilla 1992), Andinobates do-
risswansonae is dark brown or black with ir-
regular red blotches (Rueda-Almonacid et al.
2006), Andinobates tolimensis is a metallic yel-
low-bronze color anteriorly that fades to dark
brown by midbody (Bernal et al. 2007, Bernal
& Luna-Mora 2014), and Andinobates cassi-
dyhornae has a bright red dorsum with a ven-
ter that is black with well-dened red blotch-
es or spots (Amézquita et al. 2013). Moreover,
Andinobates bombetes is not sympatric with
any other species of the genus Andinobates.
Some populations are syntopic with species of
the genus Leucostethus, but those species are
easily distinguished because of their mostly
brown cryptic coloration (Grant and Castro
1998; Marin et al. 2018).
e tadpole of Andinobates bombetes can
be dierentiated from other congeners be-
cause of the presence of a U-shaped upper
jaw sheath, a dierentiated short gut with
visible organs, the absence of a projection
on the inner margin of the nasal rim, and
the presence of a median gap that interrupts
the papillate fringe on the lower (posterior)
edge of the oral disc (Myers and Daly 1980;
Sánchez 2013). Tadpoles of Andinobates
bombetes develop in small pools of bromeli-
ads, distinguishing them from other syntopic
dendrobatid tadpoles (Leucostethus) that de-
velop in terrestrial bodies of water (Lötters et
al. 2007).
DIAGNOSIS. Andinobates bombetes has the
following diagnostic characters: vomerine
teeth are absent; width of head is usually less
than width of the body; dorsal and ventral
skin are granular, particularly coarse on lower
back and the hind limbs; snout rounded from
a lateral view and bluntly rounded to truncate
from a dorsal or ventral view; external nares
situated near the tip of the snout and directed
posterolaterally; canthus rostralis is rounded;
the loreal region is at; tympanum is hidden
posterodorsally; the relative length of the n-
gers have the following order III>IV≥II>I;
distal discs of the ngers are expanded on
all of them except the rst one; a large circu-
lar-to-elliptical outer metacarpal tubercle on
median base of palm; the relative length of
the toes have the following order IV>III>V
>II>I; rst toe short and without expanded
distal disc; the coloration of the back and
limbs is black or dark brown with two red
dorsolateral stripes (in some populations the
stripes are yellow), which begins at the snout
and extends to midbody although sometimes
extending the full length of the body; ventral
coloration of the head, body, and hind limbs
is motted black on a pale yellow or pale green.

6
PUBLISHED DESCRIPTIONS. Descrip-
tions of Andinobates bombetes and taxonom-
ic reviews of the family Dendrobatidae were
published by Brown et al. (2011), Clough and
Summers (2000), Grant et al. (2006, 2017),
Myers (1987), Myers and Daly (1980), and
Santos et al. (2009). Observations of breeding
behavior and parental care of this species were
recorded by Herkrath (2006), Myers and Daly
(1980), and Súarez-Mayorga (1999, 2004);
the diet was described by Gómez-Hoyos et al.
(2014); the eectiveness of red and yellow col-
or in dorsolateral stripes as aposematic signal
was discussed by Casas-Cardona et al. (2018).
e acoustic communication behavior of in-
dividuals in anthropogenic noisy habitats and
the distribution of individuals according to
habitat features (e.g., number of bromeliads)
were studied by Vargas-Salinas and Amézqui-
ta et al. (2013). Geographic variation in the
advertisement call as a possible adaptation to
reduce acoustic masking of the call by stream
noise was studied by Vargas-Salinas et al.
(2014a). e advertisement call of Andino-
bates bombetes was described by Myers and
Daly (1980) and Súarez-Mayorga (1999).
ILLUSTRATIONS. Color photographs of
dorsal and lateral views of individuals with
red stripes were published by Amézqui-
ta and Vélez [2010], Arcila-Pérez et al. (in
press), Brown et al. (2011), Castro-Herrera
and Bolívar-García (2010), Castro-Herrera
et al. (2007), Daly et al. (1997), Diaz-Pulido
et al. (2016), Herrmann (2005), Lötters et al.
(2007), Renjifo [1997], Rodríguez-Suárez and
Corredor-Londoño (2012), Rueda-Almonac-
id et al. (2004), Ruiz-Carranza et al. (1996),
Suárez-Mayorga (2004), Vanegas-Guerrero et
al. (2016), Vargas-Salinas et al. (2014b), and
Walls (1994); dorsal views of individuals with
yellow stripes were published by Brown et
al (2011), Suárez-Mayorga et al. (2016), and
Vargas-Salinas et al. (2014b); a ventral view of
an individual with red stripes was published
PHYLOGENETIC RELATIONSHIPS. An-
dinobates bombetes was originally assigned
to the genus Dendrobates (Myers and Daly
1980) inside of the “minutus group” sensu
Silverstone (1975). Within this group, Den-
drobates bombetes, Dendrobates abditus, and
Dendrobates opisthomelas were hypothesized
to form a monophyletic subgroup based on
the presence of a median gap that interrupts
the papillate fringe on the posterior (lower)
edge of the oral disc in tadpoles. Later, Den-
drobates bombetes was included in the genus
Minyobates by Myers (1987) based on the
presence of cephalic amplexus, small body
size, presence of an oblique lateral stripe, lar-
vae with the cloaca in dextral position, and
lateral indentations of the oral disc. e spe-
cies was included in the genus Ranitomeya by
Grant et al. (2006) aer taking into account a
reduction in the length of Finger I, and oth-
er characteristics such as conspicuous dorsal
coloration, the absence of toe webbing, and
the presence of lipophilic alkaloids. Final-
ly, the genus Andinobates was proposed by
Brown et al. (2011) using a compendium of
morphological, behavioral, ecological, and
molecular characters. Currently, there are 15
species recognized within the genus Andino-
bates (Frost 2020); Andinobates bombetes is
the type species of the genus.
CONSERVATION STATUS. Andinobates
bombetes is considered Vulnerable by the
IUCN (IUCN SSC Amphibian Specialist
Group 2017). e threats to this species are
associated with loss of forest habitats that are
converted for agriculture and cattle grazing.
is species is listed in Appendix II of CITES
because individuals are collected illegally for
pet trade (CITES 2020). Populations of And-
inobates bombetes are present in several nat-
ural areas protected by Colombia, including
Bosque de Yotoco in the western Andes (Val-
le del Cauca), and Parque Regional Natural
Barbas-Bremem in Central Andes (Quindío).

Account 926 7
F 4. Males of Andinobates bombetes carrying tadpoles on their backs. A) Individual with red dor-
solateral stripes recorded in the Parque Nacional Natural Los Catíos, municipality of Toro, Valle del Cau-
ca, Colombia. Photograph by Fernando Vargas-Salinas. B) Individual with yellow dorsolateral stripes
recorded in Reserva Natural Altamira, municipality of El Cairo, Valle del Cauca, Colombia. Photograph
by Alejandro Grajales-Echeverry.
A
B

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Brown et al. (2011). A color photograph of
an individual with red stripes was mistakenly
labeled as Ameerega picta by Marent (2008,
2010). Color photographs demonstrating the
ontogenetic changes in dorsal coloration (red
morph) were published by Rodríguez-Suárez
and Corredor-Londoño (2012); color photo-
graphs of eggs, embryos, and tadpoles in dif-
ferent developmental stages were published
by Rodríguez-Suárez and Corredor-Lon-
by Rodríguez-Suárez and Corredor-Londoño
(2012), and similar ventral photographs of
individuals with yellow stripes were pub-
lished by Brown et al. (2011). Photographs
of a red morph male carrying tadpoles on
its back were presented by Castaño and Car-
ranza-Quiceno (2015), Stevens and Ruxton
(2012), and Vargas-Salinas et al. (2014b); a
photograph of a yellow morph male carry-
ing tadpoles on its back was published by
F 5. Oscillogram (A) and spectrogram (B) of the advertisement call of Andinobates bombe-
tes. Male body size = 20.09 mm; body temperature of male = 21.6ºC; locality: Parque Nacional
Natural Los Catíos, municipality of Toro, Valle del Cauca in the western Andes of Colombia.

Account 926 9
doño (2012). Black-and-white photographs
of the dorsal view of red-morph adults were
published by Myers and Daly (1980), and the
ornamentation of the margin of the naris at
stage 27 and the ventral view of the U-shaped
larval jaw sheaths at stage 28 were published
by Sánchez (2013). Color illustrations of an
adult individual with red stripes was present-
ed by Amézquita et al. (2013), Arcila-Pérez et
al. (in press), and Brown et al. (2011); color
illustrations of red and yellow morphs were
published by Casas-Cardona et al. (2018).
A black-and-white illustration of tadpole
mouth parts was presented by Walls (1994)
and an illustration of the mouth of a tadpole
at stage 25 was published by Myers and Daly
(1980).
DISTRIBUTION. Andinobates bombetes is
found in leaf litter or under fallen logs lo-
cated along the forest edge and in the forest
interior, oen adjacent to streams.. Although
populations in humid montane forests can be
found away from streams, some populations
established in tropical dry forests are found
alongside small streams. is species is en-
demic to Colombia and has been found be-
tween 867–2100 m elevation on both slopes
of the Western Andes and on the western
slope of the Central Andes in the departments
of Quindío, Risaralda, and Valle del Cauca
(Ruiz-Carranza et al. 1996; Súarez-Mayorga
2004; Vargas-Salinas et al. 2014b).
FOSSIL RECORD. None.
PERTINENT LITERATURE. Published ref-
erences to the species are listed by topic:
advertisement call (Myers and Daly 1980;
Myers 1982), antipredatory behavior in
adults (Casas-Cardona et al. 2018), breeding
behavior and parental care (Herkrath 2006;
Hesselhaus 1994; Suárez-Mayorga 1999),
checklists (Anonymous 1993a; Castro-Her-
rera and Vargas-Salinas 2008; Duellman 1993;
Frank and Ramus 1995; Frost 1985; Glaw et al.
1998, 2000a, 2000b; Harding 1983; Hutchins
et al. 2003; Renjifo [1997]; Ruiz-Carranza et
al. 1996), conservation status (Castro-Herre-
ra and Bolívar-García 2010; Furrer and Corre-
dor 2008; Marín Gómez and Gómez Hoyos
2011; Rueda-Almonacid et al. 2004; Stuart et
al. 2008; Suárez-Mayorga 2004), development
of embryos and tadpoles (Rodríguez-Suárez
and Corredor-Londoño 2012; Sánchez 2013),
diet (Gómez-Hoyos 2010; Gómez-Hoyos et
al. 2014; Herrmann 2005), geographic dis-
tribution (Acosta-Galvis 2000; Castro-Her-
rera and Bolívar-García 2010; Ruiz-Carranza
et al. 1996; Suárez-Mayorga 2004; Vargas-Sa-
linas et al. 2014b), habitat use and acoustic
communication (Vanegas-Guerrero et al.
2016; Vargas-Salinas and Amézquita 2013;
Vargas-Salinas et al. 2014a), homing behav-
ior (Arcila-Pérez et al. in press), husbandry
(Herrmann 2005), natural history (Hessel-
haus 1994; Mattison 1987, 2002; Walls 1994),
pet trade (Anonymous 1993a, 1993b), pop-
ulation ecology (Gómez-Hoyos 2010), skin
toxins (Daly 1988; Daly et al. 1987, 1993;
Daly et al. 1997; Erspamer 1994; Myers and
Daly 1976, 1980; Myers et al. 1995), and tax-
onomy (Brown et al. 2011; Grant et al. 2006;
Ruiz-Carranza and Ramírez-Pinilla 1992; Sil-
verstone 1975).
ETYMOLOGY. e specic epithet bombe-
tes refers to the loud buzzing sounds made
by calling males, similar to those made by
insects. e specic epithet is Latinized from
the Greek βομβητηϛ meaning ‘buzzer’ (Myers
and Daly 1980).
COMMENTS. Two studies of Andinobates
bombetes are being performed by students
at the University of Quindío, Colombia. One
study is focused on the intraspecic geo-
graphic variation of temporal and spectral
features of the advertisement call, including
comparisons between morphs (i.e., popula-
tions whose individuals exhibit red vs. yel-
low dorsolateral stripes) and populations

10
separated by geographic barriers (i.e., low-
lands between Western and Central Andes)
(C. Conzález-Acosta, unpublished data). A
second study examines how individuals deal
with acoustic anthropogenic disturbance in
their habitat; specically, whether males have
the capacity to increase the frequency of the
advertisement call when they are exposed to
noise by trac on roads (G. M. Jiménez-Var-
gas, unpublished data).
ADDITIONAL VERNACULAR NAMES.
Cauca-Baumsteiger (Herrmann 2005), Cauca
Poison Frog (Suárez-Mayorga et al. 2016; Wro-
bel 2004), Cauca Poison-arrow Frog (Wrobel
2004), Elevated Fondle-shun (Mitchell 2017),
rana rubí (Castro-Herrera and Bolívar-García
2010; Herkrath 2006), Rana venenosa del
Calima (Amézquita and Vélez [2010]), rana
venenosa del Cauca (Suárez-Mayorga 2004),
and Red-banded Poison Arrow Frog (as
Краснополосый Древолаз) (Sokolov 1988).
ACKNOWLEDGMENTS. We thank the Bi-
ology program of the University of Quindío,
Colombia, for logistic support during the
preparation of this manuscript. We thank
Dina Lucia Rivera for her help with building
the distribution map. anks to J. D. Lynch
for allowing access to tadpole vouchers in the
Collection of Amphibians of the ICN, Bogotá,
Colombia. Finally, we also give thanks to the
Asociación Colombiana de Herpetología
ACH (Grant Botas al Campo BC-2018-03)
for economic support for our studies with
Andinobates bombetes.
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See page 16 for author aliations and
contact information.

16
FERNANDO VARGAS-SALINAS, Pro-
grama de Biología, Universidad del Quindío,
Carrera 15 Calle 12 N, Armenia, Colombia
(fvargas@uniquindio.edu.co); MICHELLE
A. ATEHORTUA-VALLEJO, Grupo de In-
vestigación en Evolución, Ecología y Con-
servación EECO. Universidad del Quindío,
Carrera 15 Calle 12 N, Armenia, Colombia
(michelleatehortuav@gmail.com); LUISA F.
ARCILA-PÉREZ, Grupo de Investigación en
Evolución, Ecología y Conservación EECO.
Universidad del Quindío, Carrera 15 Cal-
le 12 N, Armenia, Colombia (arcilaluisap@
gmail.com); GINA M. JIMÉNEZ-VAR-
GAS, Grupo de Investigación en Evolución,
Ecología y Conservación EECO. Universidad
del Quindío, Carrera 15 Calle 12 N, Arme-
nia, Colombia (ginamarcelajimenez@gmail.
com); CRISTIAN GONZÁLEZ-ACOS-
TA, Grupo de Investigación en Evolución,
Ecología y Conservación EECO. Universidad
del Quindío, Carrera 15 Calle 12 N, Arme-
nia, Colombia (cristiandread11@gmail.com);
SANTIAGO CASAS-CARDONA, Colec-
ción de Anbios y Reptiles de la Universidad
del Quindío, Programa de Biología, Univer-
sidad del Quindío, Carrera 15 Calle 12 N,
Armenia, Colombia (santiagocasasc@gmail.
com); and ALEJANDRO GRAJALES-ECH-
EVERRY, Birding and Herping, Cra 6 #19-25
Apto 303 bloque 6, Torres del Río, Armenia,
Colombia (alejandrobirdingandherping@
gmail.com).
Primary editors for this account, Christopher
J. Bell and Travis J. LaDuc.
Published 29 October 2020 and Copyright
© 2020 by the Society for the Study of
Amphibians and Reptiles.