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2Oophaga
Oophaga
Genus
Oophaga Bauer, 1994
e generic epithet Oophaga is derived from the Greek oon combined with phagos,
which together mean ‘egg eater’. e genus name is based on the egg-provisioning
behavior seen in this genus, where females provide infertile food eggs to their larvae.
Oophaga contains nine species; four (O. lehmanni, O. occultator, O. histrionica, and
O. sylvatica) are Andean. A fth Colombian species has been detected (A. Amézquita,
unpubl. results). In the Andean region, Oophaga occur in the Colombian Chocó
eco-region southward into northwestern Ecuador below about 1200 meters (Grant
et al., 2006).
e sister taxon of Oophaga is Dendrobates (Grant et al., 2006). Diagnosis of Oophaga is
based on the following external and other characteristics. 1) Typically brightly colored,
and 2) pattern is blotched, banded or spotted, lacking dened stripes. 3) e skin is
smooth. 4) e rst nger is shorter than the second nger with 5) only moderately
expanded nger disks. 6) Lipophilic alkaloids are sequestered in the skin (Grant et al.,
2006). 7) Cloacal amplexus is used in some species’ populations or reproduction occurs
with no amplexus. Note: similar diagnostic characteristics occur in other poison frog
taxa.
24.4 mm 0 mm
38.0 mm
Oophaga 3
Harlequin Poison Frog, Oophaga histrionica
Chapter 12
Harlequin Poison Frog,
Oophaga histrionica
Harlequin Poison Frog
Oophaga histrionica (Berthold, 1845)
Adolpho Amézquita¹, Daniel Mejía-Vargas¹ and Ted R. Kahn²
¹Universidad de los Andes, Bogotá, Colombia
²Neotropical Conservation Foundation, Washington, DC, USA
Names
Harlequin Poison Frog (Frank and Ramus, 1995); Rana Arlequín Venenosa; Rana
Común de Dardo Venenoso; Chico (Chocó Emberá) from the combination of Chico
(Spanish) and Bocorro (indigenous language) which means ‘Little Frog.’ Indigenous
peoples of the Quebrada Bochorama call this species Koe-Ko-Pa or Kokõe-pá. e
vernacular name Harlequin refers to the bright coloration of this species. e specic
epithet histrionica is Latin for ‘actor’, and makes reference to the spectacular clown-
like and costume-like colors and patterns found in this species. Interestingly, its
major alkaloids are the bicyclic histrionicotoxins, named after the species, for the
typically histrionic over-reaction when exposed to these noxious alkaloids (J.W. Daly
pers. comm.).
Taxonomic Comments
Oophaga histrionica was rst named as Hyaplesia de cocteau (Duméril and Bibron,
1841). Berthold (1845) placed this species in the genus Dendrobates, naming it
histrionicus. Steindachner (1864) kept the genus placement as a variation within
the tinctorius clade, Dendrobates tinctorius var. cocteani [sic] (misspelled cocteaui).
Boulenger (1913) used a trinomial to include this species as a subspecies of tinctorius,
Dendrobates tinctorius cocteaui. Later that same year, Boulenger revised his placement
as a subspecies to that of a variation of tinctorius, Dendrobates tinctorius var.
coctaei [sic], and again cocteaui was misspelled. Laurent (1942) used the trinomial
Dendrobates tinctorius wittei, but later that year (1942) revised this to Dendrobates
tinctorius histrionicus. Dunn (1944) allocated this species to Dendrobates histrionica;
subsequently corrected by Funkhouser (1956) to histrionicus, who also added the
subspecies confluens. Cochran and Goin (1970) continued the trend to place this
species in the tinctorius clade as Dendrobates tinctorius confluens. Silverstone (1975)
revised the taxonomy of this species to Dendrobates histrionicus. Bauer (1994) placed
this poison frog in the then-newly proposed genus Oophaga as histrionica. Grant et al.
(2006) recognized and kept Bauer’s placement in Oophaga.
Identification and Description
Within the range of Oophaga histrionica, individuals vary in size and are considered
medium-sized to large poison frogs with a snout-vent length (SVL) of about
4Oophaga
Chapter 12
26.8–34.1 mm in males, 26.3–34.8 mm in females (Myers and Daly, 1976a), and
overall 24.0–38.0 mm SVL (Lötters et al., 2007). e skin is smooth; the tarsal
tubercle is absent. e color and the pattern are highly variable, with >30 morphs
known that are often extremely localized (polytypic vs. polymorphic). e ground
color is various shades of tan, brown, dark brown or black. e color pattern can be
nely reticulated on the body, nely spotted, large-spotted, single-spotted, blotched
or banded in white, greenish-white, pale yellow, bright yellow, pale orange, bright
orange and pinkish-red to brilliant red or sky blue. ese colors and the corresponding
patterns appear on individuals throughout their range (polytypic) and vary among
localities (polymorphic).
Identification and Description of Larvae and Froglets
e larvae are blackish-brown, tending towards black. Large teeth appear in labial
tooth rows. e labial tooth row formula (LTRF) is 1(1)/(1) or 1/1 depending on
developmental stage. Tooth rows vary during development, hence the LTRF variation
at dierent stages (R. Altig, pers. comm.). A large, heavily serrated beak sheath
distinguishes Oophaga larvae from other Andean dendrobatid larvae. Both teeth and
beak are adaptations to oophagy (egg feeding). e oral apparatus (OA) is oriented
anteroventrally; the oral disk (OD) is not emarginated and is surrounded by large,
rounded marginal papillae (MP) that are few in number. e spiracle is sinistral;
the vent is medial. Tadpoles in later stages of development, nearing metamorphosis,
have the pattern of the parent adults (Silverstone, 1976) and transform into froglets
similar in appearance to their founder stock.
Similar Species
Oophaga lehmanni and O. histrionica display clinal morphology toward one another.
Where their ranges overlap, genes are exchanged by interbreeding (hybridization)
(Vargas-Salinas and Amézquita, 2013). is results in similarly colored and
patterned O. histrionica that occur on the fringes of the small disjunct ranges of O.
lehmanni. In the southern limits of its range, O. histrionica can be confused with O.
sylvatica. Oophaga sylvatica generally has a less well-dened pattern, often reticulated
or mottled (vs. the very well-dened pattern often seen in O. histrionica), as well as a
distinctly dierent sounding call than O. histrionica.
Alkaloid Profile
e three major alkaloids found in this species of poison frog’s skins are
the bicyclic histrionicotoxin 283A, isodihydrohistrionictoxin 285A, and
allodihydrohistrionicotoxin 285C in most populations from western Colombia (J.W.
Daly, unpubl. results). Gephyrotoxins 287C and 289B are tricyclic alkaloids known
also to occur in millipedes, which are presumed to be the dietary source for the
two previously mentioned gephyrotoxins found in these poison frogs’ skins (J.W.
Daly et al., unpubl. results). e monocyclic alkaloid 2,5 disubstituted pyrrolidine
trans-2-butyl-5-pentylpyrrolidine (197B) was found to be the major alkaloid in
populations from El Valle, Chocó, Colombia, and as a trace alkaloid was found in a
Oophaga 5
Harlequin Poison Frog, Oophaga histrionica
population from a small mountain: Altos del Buey, about 25 km from the El Valle,
Chocó population (J.W. Daly, unpubl. results). Another minor monocyclic alkaloid
(225C) from the same sample is proposed to be a 2-butyl-5-heptyl-pyrrolidine (J.W.
Daly, unpubl. results). See Appendix 4.1 for a complete listing of alkaloids found in
the skins of O. histrionica.
Natural History and Ecology
Harlequin Poison Frogs are diurnal and terrestrial. ey climb up to 10 m up trees
and upon fallen logs (Lötters et al., 2007), and often sleep above ground, perching
on low-lying leaves. ey inhabit premontane primary and secondary lowland wet
and rain forests of the Chocó eco-regions (Fig. 12.1.). In Valle del Cauca they also
occur in dry forests with various cactus species (Fig. 12.2.). Harlequin Poison Frogs
show some ability to adapt to modied habitats if intact forest remains nearby
with adequate reproductive resources, such as bromeliads. Male O. histrionica are
territorial, and call from elevated perches. ey wrestle with intruding males over
mates and defend their territories. ese confrontations are accompanied be vigorous
repeated vocalizations.
Calls and Vocalizations
e advertisement call is a low-pitched ‘chirp’ (Myers and Daly, 1976a) with a
duration of 125 milliseconds, repeated at irregular intervals of 2–3.5 notes per second.
e dominant frequency is 2000–4000 Hz, with recordings taken at an ambient air
temperature of 27.0° Celsius (Myers and Daly, 1976a). e call is “reminiscent of a
buprestid beetle knocking in wood” (Funkhouser, 1956) or likened to the “quack of a
duck” (Silverstone, 1976). e call of the “Bullseye morph” (polytype) sounds distinctly
like a quiet ‘buzz’ (T.R. Kahn, pers. obs.).
Figure 12.1. Lowland wet forest in Baudó, Chocó eco-region, Colombia where Oophaga
histrionica, Phyllobates aurotenia, and Andinobates fulguritus co-occur.
© 2012 Dennis Nilsson
6Oophaga
Chapter 12
Reproduction
In nature, Oophaga histrionica reproduce year round (Lötters et al., 2007). Small
egg clutches (about 1–4) are laid terrestrially every few weeks throughout the
year. Amplexus is conducted via cloacal apposition, though, with equal frequency,
no amplexus occurs. In these cases the male will clean and fertilize the site of egg
deposition during courtship (or occasionally, after egg deposition, will fertilize the
clutch). e female then lays her eggs on this site. In the case of cloacal apposition,
spermatoposition takes place as the eggs are laid. Courtship is elaborate and involves
tactile, vocal, visual, and positioning keys (E. Zimmermann and H. Zimmerman,
1986). Typically, a male leads a female to an egg deposition site he has chosen, where
courtship and egg deposition take place. e female cares for the eggs, transports
the larvae to phytotelmata and deposits them individually. She returns every few
days to feed them infertile nutritive eggs, which she produces specically to feed her
ospring. Oophaga histrionica is considered an obligate egg feeder; infertile food eggs
are the primary food source for the developing larvae, though they are also known to
be cannibalistic.
Distribution
Endemic to the Pacic lowlands of Colombia, O. histrionica occurs only in the
departments of Antioquia, Chocó, Risaralda, and Valle del Cauca, sea level to
1000 meters (IUCN Red List of reatened Species™, version 2011.2).
Figure 12.2. Dry forest in Valle de Cauca, Colombia, where Oophaga histrionica occurs.
© 2011 Dennis Nilsson
Oophaga 7
Harlequin Poison Frog, Oophaga histrionica
Threats
e major threats to this poison frog are deforestation for agricultural development,
logging, illegal crops, human settlements, and pollution of water sources as a result of
aerial spraying of herbicides on illicit crops. e species is also collected illegally for
the international pet trade (IUCN Red List of reatened Species™, version 2011.2).
Conservation Status
e Harlequin Poison Frog occurs in several protected areas. It is classied as Least
Concern (LC) on the IUCN Red List of reatened Species™ (version 2011.2).
However, IUCN does not take into consideration the varied polytypic forms that
are disjunct and highly localized and consequently vulnerable to extirpation. It is
recommended that this species be reevaluated considering this information. Oophaga
histrionica is highly sought after in the international pet trade, and recommendations
have been made that some sustainable harvest be conducted (IUCN Red List of
reatened Species™, version 2011.2). However, the basis for the harvest of poison
frogs from the wild is poorly documented. is species is protected in Colombia by
decree INDERENA No. 39 of 9 July, 1985, that forbids the collection of Oophaga
spp. from the wild for breeding (or other) purposes. It is listed in Appendix II of
CITES.
Figure 12.3. Lowland tropical forest habitat of O. histrionica, Cerro Mecano, Bahia Solano,
Department of Chocó, Colombia.
© 2009 omas Ostrowski
8Oophaga
Chapter 12
Oophaga histrionica Morphs
= 20 mm
Oophaga 9
Harlequin Poison Frog, Oophaga histrionica
Morphs Oophaga histrionica
= 20 mm
Naturally occuring hybrids between Oophaga lehmanni and O. histrionica
10 Oophaga
Chapter 12
Oophaga histrionica Morphs
= 20 mm
Oophaga 11
Harlequin Poison Frog, Oophaga histrionica
Oophaga histrionica Map
12 Oophaga
Chapter 12
Lehmann’s Poison Frog,
Oophaga lehmanni
Lehmann’s Poison Frog
Oophaga lehmanni (Myers and Daly, 1976)
Adolfo Amézquita
Universidad de los Andes, Bogotá, Colombia
Names
Lehmann’s Poison Frog (Frank and Ramus, 1995); Rana Coral Candelilla. e
vernacular name Lehmann’s Poison Frog and specic epithet lehmanni are eponyms
honoring the late Federico Carlos Lehmann (1914–1974), a renowned Colombian
ornithologist who contributed substantially to the knowledge of the Colombian
Pacic fauna and to the creation of Parque Nacional Natural Farallones de Cali, to
which the distribution of O. lehmanni is restricted.
Taxonomic Comments
Oophaga lehmanni was considered a “morph” of the highly polytypic O. histrionica
(see Silverstone 1975). Myers and Daly (1976a) described it as a valid species based
on the distinctive coloration and a pattern of dorsal cross bands, as well as the
lack of histrionicotoxins (considered diagnostic at the time, it is now known that
alkaloid proles are not species specic), and placed it in the genus Dendrobates.
Bauer (1994) proposed this species be placed in his genus Oophaga. Grant et al.
(2006) placed lehmanni in Bauer’s Oophaga. e species’ status is strongly supported
by microsatellite, mitochondrial, and nuclear DNA analyses (Grant et al., 2006;
I. Medina et al., unpubl. results; A. Amézquita et al., unpubl. results). Bioacoustic
data analyses (Vargas-Salinas and Amézquita, 2013) indicates that O. lehmanni is
the sister species of O. sylvatica, and that their ancestry is the sister lineage of O.
histrionica (see Grant et al., 2006). Oophaga histrionica and O. lehmanni hybridize in
the wild (Vargas-Salinas and Amézquita, 2013).
Identification and Description
Oophaga lehmanni is a medium-sized to large poison frog with a snout-vent length
(SVL) of 31.3–35.3 mm in males, and 31.3–34.3 mm in females (Myers and Daly,
1976a). e skin is smooth. e coloration and pattern are variable. All the known
morphs of O. lehmanni typically exhibit two dorsal, transverse, yellow, red, orange
or intermediately-colored bands. A similarly colored blotch occurs on the snout. In
some individuals, these blotches or bands may be incomplete, reduced or enlarged.
Narrower, similarly colored bands (‘limb bracelets’) are usually present on the
forearms and hind limbs. Ventral coloration and pattern are similar to that of the
dorsum, but the bands can be distorted, and the brilliant color somewhat faded or
obscured by the black ground color. Oophaga lehmanni has white or bluish-white
ngertip disks (Myers and Daly, 1976a).
Oophaga 13
Lehmann’s Poison Frog, Oophaga lehmanni
Lehmann’s Poison Frog,
Oophaga lehmanni
Identification and Description of Larvae and Froglets
e larvae of O. lehmanni have not been formally described. e froglets appear to be
colored and patterned as miniature adults.
Similar Species
Oophaga lehmanni can easily be confused with two forms of O. histrionica that
occur less than 20 km away from the localities of O. lehmanni. eir coloration
consists of yellow blotches that gradually become orange or reddish towards the head.
Also, an O. lehmanni × O. histrionica hybrid form (Vargas-Salinas and Amézquita,
2013) occurs near O. lehmanni populations, but individuals from this population
usually bear a single, narrower dorsal band accompanied by several blotches and/
or highly variable irregular bands (Lötters, 1992). e only other species exhibiting
dorsal bands is O. histrionica from the Department of Chocó, Colombia. It lacks,
however, the white color of the ngertip discs that characterize O. lehmanni (D.
Mejía et al., unpubl. results). Four other populations of O. histrionica exhibit dorsal
bands, in orange, pale yellow or blue (D. Mejía et al., unpubl. results); however,
those occur in the foothills of the western versant of the Pacic Andes, Department
of Chocó, Colombia, about 250 km north of the known distribution of O. lehmanni,
and molecular data unambiguously support their very close phylogenetic anity to
other northern forms of O. histrionica (A. Amézquita et al. unpubl. results). Oophaga
lehmanni can be readily distinguished from banded populations of O. histrionica and
hybrids by the white or bluish-white dorsal coloration of their ngertip discs, which is
present only in O. lehmanni (D. Mejía et al., unpubl. results).
Alkaloid Profile
Myers and Daly (1976a) noted that O. lehmanni skins contain pumiliotoxin A and
B. Garrao et al. (2001) identied the most distinctive alkaloid of O. lehmanni, an
izidine (at the time a new class of alkaloids), containing lehmizidine 275A. Two
other alkaloids found in O. lehmanni are the homopumiliotoxins 251R and 265N
(Daly et al., 1999). See Appendix 4.1 for additional alkaloids found in the skins of
this species.
Natural History and Ecology
Oophaga lehmanni is diurnal and terrestrial, and can be found in primary cloud
forests (see Fig. 12.6.) and associated riparian habitats, often on steep hillsides with
profuse vegetation and a high density of bromeliads (Velásquez et al., 2009). It hides
in crevices formed by tree roots or fallen trees when inactive or when disturbed.
Oophaga lehmanni is an ant specialist. e stomach contents of ten individuals
(Arenas, 2010) revealed ants (about 80%) and mites (about 10%). Oophaga lehmanni
showed the narrowest dietary niche when compared with three populations of O.
histrionica and one of O. sylvatica within 20–150 km of O. lehmanni localities.
14 Oophaga
Chapter 12
Figure 12.4. An illegal coca (Erythroxylum coca) plantation in the Anchicaya Valley,
Departamento Valle del Cauca, Colombia, 800–850 meters (foreground) with intact mid-
elevation cloud forest (background) where O. lehmanni occurs. e spraying of herbicides,
for example, defoliants, by international and national law enforcement agencies is an ongoing
threat to this Critically Endangered poison frog.
© 2012 Andreas Zarling
Figure 12.5. Partially cleared mid-elevation cloud forest in the Anchicaya Valley, Departamento
Valle del Cauca, Colombia, 800–850 meters. Oophaga lehmanni was found in the intact,
marginal, still-forested habitat.
© 2012 Andreas Zarling
Oophaga 15
Lehmann’s Poison Frog, Oophaga lehmanni
Calls and Vocalizations
Males of O. lehmanni utter ‘chirp’ advertisement calls (Myers and Daly, 1976a). e
notes of O. lehmanni are signicantly shorter (122 vs. 181 milliseconds) and consist
of slightly fewer pulses (29 vs. 33) than the vocalizations of the closely related O.
histrionica (Vargas-Salinas and Amézquita, 2013). e notes of O. lehmanni are
produced at slower repetition rates (Lötters et al., 1999; Myers and Daly, 1976a).
Call frequencies are 1500–3500 Hz recorded at 28.0° Celsius (Myers and Daly,
1976a). Interestingly, individuals inhabiting riparian zones (both O. lehmanni and O.
histrionica and hybrids) call at higher frequencies than individuals inhabiting forested
habitats away from noisy streams (Vargas-Salinas and Amézquita, 2013).
Figure 12.6. Intact mid-elevation cloud forest where O. lehmanni occurs in the
Anchicaya Valley, Departamento Valle del Cauca, Colombia, 800–850 meters.
© 2012 Andreas Zarling
16 Oophaga
Chapter 12
Reproduction
Oophaga lehmanni reproduces year-round. e primary calling and breeding activity
seem to depend upon a combination of coinciding high humidity and warm
temperatures. Males announce territory with advertisement calls. ey chase and
wrestle other calling males in their territories, and combat durations are longer when
opponent males are of similar size (Rojas, 2002). e female initiates courtship. Male
and female will then engage in elaborate courtship, exchanging visual and tactile
signals (H. Zimmermann and E. Zimmermann, 1981). Typically, four to six eggs
are laid under cover, probably in the leaf litter or on leaves in the axils of bromeliads
on or close to the ground. As in other Oophaga species, the larvae of O. lehmanni are
obligate egg feeders. Metamorphosis occurs within about 60 days after egg deposition
(see chapter 5, for additional information on this species’ reproduction).
Distribution
Endemic to Colombia, the species’ distribution is restricted to the Pacic lowlands in
an area of <200 km² in the upper Anchicayá valley, Department of Valle del Cauca,
600–1200 meters. An intensive survey detected only nine reproductive colonies, several
consisting of very few individuals (Velásquez et al., 2009). Some specimens from the
Department of Chocó that actually represent O. histrionica were erroneously assigned
to O. lehmanni (see Lynch and Suárez-Mayorga, 2004).
Threats
Oophaga lehmanni is currently considered the most threatened dendrobatid species in
Colombia, likely because of the very intensive illegal tracking over the past 30 years
(D. Mejía et al., unpubl. results). In the 1970s, dozens of farmers were hired to collect
thousands of individuals per year for the pet trade and for scientic research (Myers
and Daly, 1976a). Illegal commercial pressure on this species for the international
pet trade continues. Illegal drug crops and subsequent forest clearing pose serious
ongoing threats (see Figs. 13.3. and 13.4.). e conscation of about one hundred
Oophaga lehmanni and O. histrionica (hybrids also) occurred in 2009 (see Fig. 2.16.
chapter 2). ere are only a handful of small and disjunct localities where this species
is relatively abundant.
Conservation Status
Oophaga lehmanni is classied as Critically Endangered (CR) in the Libro Rojo de
Anfibios de Colombia (Rueda et al., 2004) and on the IUCN Red List of reatened
Species™ (version 2011.2). A few reproductive colonies occur in the Parque Nacional
Natural Farallones de Cali, but even there they are vulnerable to poaching. Rapid
and multiple conservation actions are needed to conserve this species. It is protected
in Colombia by decree INDERENA No. 39 of 9 July, 1985, which forbids the
collection of Oophaga spp. from the wild for breeding (or other) purposes. It is listed
in Appendix II of CITES.
Lehmann’s Poison Frog, Oophaga lehmanni
Oophaga 17
Morphs Oophaga lehmanni
= 20 mm
18 Oophaga
Chapter 12
Oophaga lehmanni MorphsOophaga lehmanni Map
Oophaga 19
La Brea Poison Frog, Oophaga occultator
La Brea Poison Frog,
Oophaga occultator
La Brea Poison Frog
Oophaga occultator (Myers and Daly, 1976)
Francisco José López-López¹, Nathalia Suescún R.² and Ted R. Kahn³
¹Grupo de Estudios en Biodiversidad de Anbios y Reptiles Neotropicales, Popayán, Colombia
²Universidad del Rosario, Bogotá, Colombia
³Neotropical Conservation Foundation, Washington, DC, USA
Names
La Brea Poison Frog (Frank and Ramus, 1995); Rana Venenosa del Saija (Amézquita
in Rueda-Almonacid et al., 2004). e specic epithet occultator is derived from
Latin, meaning ‘a hider’, referring to the secretive nature of this poison frog (Myers
and Daly, 1976a). e vernacular name La Brea refers to the small town of that name
near the type locality.
Taxonomic Comments
Myers and Daly (1976a) initially described Oophaga occultator in the genus
Dendrobates, based on specimens collected by Malkin in 1971, and additional
specimens collected by Myers and Daly in 1973. Bauer (1994) proposed the genus
Oophaga for these and eight other species that have subsequently been accepted and
adopted by Grant et al. (2006).
Identification and Description
A medium-sized poison frog, O. occultator has a snout-vent length (SVL) of 24.4
27.1 millimeters. Females are more robust than males. e skin is smooth. e
ground color is black with a brick red dorsum that is ‘marred’ with irregular black
‘scratches’ (Myers and Daly, 1976a). e red color extends onto the forearms and
terminates at the elbows. e remainder of the lower forearms and lower hind limbs
are black. e anks are spotted golden yellow on the black ground color. e venter
is spotted and/or mottled in golden yellow on black. e yellow mottling extends
onto the gular. e rst nger is slightly shorter than the second. e feet have tarsal
tubercles; vocal slits are present in males (Myers and Daly, 1976a).
Identification and Description of Larvae and Froglets
Unknown.
20 Oophaga
Chapter 12
Similar Species
Due to its unique vocalizations (Lötters et al., 1997a), distinctive color pattern,
and smaller size compared to the sympatric O. histrionica (Myers and Daly, 1976a),
Oophaga occultator cannot be confused with any other poison frog within its range.
Alkaloid Profile
e bicyclic pumiliotoxin B is one of the major alkaloids found in the skin of this
frog, together with several histrionicotoxins and isodihydrohistrionicotoxins, as well
as the 1,4-disubstituted quinolizidine 231 (Myers and Daly, 1976a). See Appendix
4.1 for additional alkaloids found in the skin of this poison frog.
Natural History and Ecology
A diurnal, terrestrial and semiarboreal poison frog—it is active on low vegetation
and sleeps above ground—this species inhabits steep terrain in lowlands of primary
monsoon forest hillsides, where both terrestrial and arboreal bromeliads thrive under
daily-moistened leeward winds. Oophaga occultator is commonly found amongst
bromeliads and leaf litter on the forest oor. Individuals are occasionally seen
climbing about on low vegetation at >3 m above ground level (Lötters et al., 2007),
where they deposit and care for their larvae, not unlike O. histrionica, although the
reproductive mode of O. occultator has yet to be documented. Similar parental care
to that of other Oophaga species is likely (Lötters et al., 2007).
Reproduction
Unknown, but likely to be similar to other species of Oophaga, for example, O.
histrionica (see Lötters et al., 1999, 2007; Amézquita in Rueda-Almonacid et al.,
2004).
Calls and Vocalizations
e calls of Oophaga occultator include a ‘Chirp’ (Myers and Daly, 1976a) with a note
length of 100 milliseconds and a repetition of ve notes per second (Lötters et al.,
1999), and, interestingly, a unique warning call. Apparently, males have territorial
vocalizations and their warnings are a long series of calls, each of which sounds
similar to the ‘quack’ of ducks (Amézquita in Rueda-Almonacid et al., 2004). Lötters
et al. (1996) indicated that the call of O. occultator is distinguished by being shorter
and repeated at a higher rate than the populations of O. histrionica in the northern
Chocó eco-region of Colombia.
Distribution
Oophaga occultator is endemic to a small area on the western versant of the Cordillera
Occidental of the Colombian Andes. is poison frog is known from the upper Río
Gorge basin and Río Saija and Quebrada Guanguí in the Department of Cauca,
50–200 m; in a range of about 10 km² (Amézquita in Rueda-Almonacid et al., 2004;
Oophaga 21
La Brea Poison Frog, Oophaga occultator
Lötters et al., 2007). e IUCN Red List of reatened Species™ (version 2011.2)
map depicts its range as considerably less than a few km².
Threats
Deforestation and destruction of habitat by expanding agriculture, pasture and
human settlements are ongoing threats. Illegal commercial timber extraction in
Colombia continues unabated and deleteriously aects this species’ habitat. Illicit
crops (Erythroxylon coca) and the resulting contamination from fumigation adversely
aect local water resources with toxic chemical residues, negatively impacting this
species of poison frog’s long- and short-term survival in the riparian zones of streams
where it is known to occur (IUCN Red List of reatened Species™ version 2011.2).
Conservation Status
Oophaga occultator is classied as Data Decient (DD) on the IUCN Red List of
reatened Species™ (version 2011.2). In the national category, Oophaga occultator is
classied as Vulnerable (VU) because its range is small and poorly dened and includes
areas that have an observed increase in illegal crop (Erythroxylon coca) cultivation/
fumigation that, together with deforestation, constitute major immediate threats to
this unique poison frog species (see above; A. Amézquita in Rueda-Almonacid et al.,
2004). Decree INDERENA No. 39 of 9 July, 1985, forbids collection of Oophaga spp.
for commercial (or other) purposes. is species is listed in Appendix II of CITES.
22 Oophaga
Chapter 12
Oophaga occultator Morphs
= 10 mm
Oophaga 23
La Brea Poison Frog, Oophaga occultator
Oophaga occultator Map
24 Oophaga
Chapter 12
Forest Harlequin Poison Frog
Oophaga sylvatica (Funkhouser, 1956)
Stefan Lötters1 and Ted R. Kahn2
1Universität Trier, Trier, Germany
2Neotropical Conservation Foundation, Washington, DC, USA
Names
Forest Harlequin Poison Frog (rst use here); Harlequin Poison Frog; Diablito (‘little
devil’ in Spanish). Indigenous peoples of the Quebrada Bochorama Colombia call
this species Koe-Ko-Pa or Kokõe-pá. e specic epithet sylvatica is from the Latin
word silva; sylvatica means “of the forest”.
Taxonomic Comments
Oophaga sylvatica was for a long time treated as color variations of Oophaga histrionica.
Lötters et al. (1999), studying advertisement calls of O. histrionica sensu Myers and
Daly (1976a), established considerable evidence of dierences between northern
and southern populations suggesting the existence of distinct species. e former
population, henceforth, was attributed to O. histrionica sensu stricto, while Lötters et
al. (1999) resurrected the species name sylvatica (originally described as Dendrobates
histrionicus sylvaticus in 1956 by Funkhouser from Ecuador, and treated as a junior
synonym of O. histrionica). eir taxonomic treatment is still provisional and the
authors do not rule out the possibility that O. sylvatica is a dierent, and as yet
unidentied species. Additionally, the advertisement calls of O. occultator and O.
lehmanni from the same general area show high overlap with that of O. sylvatica.
Signicantly, Funkhouser (1956) also described Dendrobates histrionicus confluens
from Ecuador, currently a junior synonym of O. sylvatica (see Lötters et al., 1999).
Grant et al. (2006) considered the status of O. histrionica sensu stricto, O. lehmanni
and O. sylvatica and concluded that the three Andean Oophaga are distinct species.
Concerning the taxonomy of O. sylvatica, Lötters et al. (1999) noted that all the
southern species (Colombia and Ecuador) studied in these early expeditions were
in fact Oophaga sylvatica, mistaken as O. histrionica and inadvertently misidentied
by not only Berthold (1845), but also Daly and Myers, (1976a) who pioneered
investigations into what is now called Oophaga sylvatica in southwestern Colombia
in the 1970s.
Identification and Description
A medium-sized poison frog with an adult snout-vent length (SVL) of 26.0 in
males and 38.0 mm in females. e skin is smooth (Silverstone, 1975). Like the
similar Oophaga histrionica sensu stricto, O. sylvatica is highly polytypic in its colors
and patterns. On a gray, brown, orange or black ground color, there are whitish,
Oophaga 25
Forest Harlequin Poison Frog, Oophaga sylvatica
Forest Harlequin Poison Frog,
Oophaga sylvatica
yellow, orange, red and or greenish, usually asymmetrically arranged spots, dots, or
a clearly reticulated or faded marbled pattern. e venter often corresponds to the
dorsal coloration or is reddish (Lötters et al., 1999, 2007). A sky blue form, splashed
with scarlet red dorsally is also known from the Department of Nariño in southern
Colombia. e characteristics of the advertisement call are important identication
features (Lötters et al., 1999).
Identification and Description of Larvae and Froglets
e larvae are blackish-brown, tending towards black. Large teeth appear in labial
tooth rows. e labial tooth row formula (LTRF) is 1(1)/(1) and or 1/1. Tooth rows
vary during development, hence the LTRF variation (R. Altig, pers. comm.). A large
heavily serrated beak distinguishes Oophaga larvae from other Andean aposematic
dendrobatid larvae. Both teeth and beak are adaptations to oophagy (egg feeding).
e oral apparatus (OA) is oriented anteroventrally; the oral disk (OD) is not
emarginated and is edged with large rounded marginal papillae (MP) that are few
in number. e spiracle is sinistral; the vent is medial or dextral. Tadpoles have the
pattern of adults in later stages of metamorphosis (Silverstone, 1976), and transform
into froglets similar in appearance to the parent stock.
Similar Species
On the basis of morphology, Oophaga sylvatica is dicult to distinguish from
O. histrionica sensu stricto. In most O. sylvatica morphs the dorsal pattern fades
gradually into the ground color and often appears dull, (vs. in O. histrionica sensu
stricto it is usually sharply dened). e two species can best be distinguished by
locality and advertisement calls (Lötters et al., 1999, 2007).
Alkaloid Profile
Bicyclic and tricyclic alkaloids occur in these poison frogs, primarily of the
histrionicotoxin classes (Daly et al., 1999). e bicylic octahydrohistrionicotoxin
291A is the major alkaloid in Ecuadorian populations. e tricyclic gephyrotoxins
287C and 289B are the major alkaloids in Colombian populations (Daly et al.,
2005). See Appendix 4.1 for additional alkaloids found in the skin of these poison
frogs.
Natural History and Ecology
is species inhabits mesic lowland rain forests of the Chocó-Tumbesian eco-region.
It occurs in primary and secondary wet forest and along forest margins, though it is
usually found in primary forests, often on steep slopes, it also may be encountered
in forest remnants that sometimes border agricultural land (banana and other
plantations), but only as long as suitable forest with reproductive resources (forests
with ample bromeliads) is nearby. Oophaga sylvatica is diurnal and primarily terrestrial
with an inclination towards climbing. ese poison frogs are often found on shrubs
and trees or on large fallen tree trunks (especially advertising males). Populations
exhibit dierences in their escape behavior, and individuals are either bold or shy, and
26 Oophaga
Chapter 12
have a relatively large escape jump distance. Males establish territories and call from
exposed sites almost all day long, with breaks often being limited only to mid-day and
during heavy rain, when this poison frog tends to retreat to shelter. ey aggressively
defend their territories against conspecics that include the non-territorial females.
Fights over territory are a daily occurrence amongst males.
Calls and Vocalizations
e advertisement call of this species is typically a loud ‘chirp’ (Myers and Daly,
1976a) having a note length of ≤100 milliseconds at a repetition rate of ≥5 notes per
second. e similar call of Oophaga histrionica has a note length of ≥125 milliseconds
and repetition rates of 2–3.5 notes per second (Lötters et al., 1999).
Reproduction
is species reproduces from December to June in Ecuador (Lötters et al., 2007).
Either cloacal apposition or no amplexus is used in reproduction. Small clutches of
2–8 eggs are laid and the female cares for them. Females transport larvae to water-
holding phytotelmata of various bromeliad species, and the larvae are commonly
found singly in those phytotelma and feed on infertile food eggs deposited by their
mother every few days; this behavior is considered obligate oophagy (Lötters et al.,
2007).
Distribution
According to Lötters et al. (1999, 2007) this species is known from the Pacic
lowlands of southwestern Colombia (departments of Cauca and Nariño) south to
northwestern Ecuador (provinces of Esmeraldas, Pichincha, Imbabura, and Los
Rios), at about 100–800 m (A. Amézquita, pers. comm.), although elevations of
0–1000 m have been reported (Lötters et al., 2007).
Threats
Widespread habitat degradation due to deforestation, gold mining, illegal crops,
road building, human settlements and agricultural development have extirpated this
species from most of its former range in Ecuador. It is intensively collected (illegally)
for the international pet trade. Colombian populations face similar threats (IUCN
Red List of reatened Species™, version 2011.2).
Conservation Status
Oophaga sylvatica is classied as Near reatened (NT) on e IUCN Red List of
reatened Species™ (version 2011.2). Although it is still relatively widely distributed,
it has undergone a major decline in Ecuador, where its overall status is considered to be
of serious conservation concern. is species occurs in the Reserva Ecológica Cotacachi-
Cayapas, Parque Nacional Mache-Chindul and Reserva Bilsa, Fundación Jatun Sacha.
It is protected under Decree INDERENA No. 39, 9 July, 1985, of Colombia that
forbids collection of Oophaga spp. for commercial (or other) purposes and is listed in
Appendix II of CITES.
Forest Harlequin Poison Frog, Oophaga sylvatica
Oophaga 27
Morphs Oophaga sylvatica
= 20 mm
28 Oophaga
Chapter 12
Oophaga sylvatica Morphs
= 20 mm
Oophaga 29
Forest Harlequin Poison Frog, Oophaga sylvatica
Oophaga sylvatica Map
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