González, Sibira, Barros, Fernández del Valle, Sormani & Rivas
16
ANARTIA
Publicación del Museo de Biología de la Universidad del Zulia
ISSN 1315-642X (impresa) / ISSN 2665-0347 (digital)
https://doi.org/10.5281/zenodo.18111972 / Anartia, 41 (diciembre 2025): 16-26
A case of evolutionary convergence? Striking resemblance
between a cockroach (Blattodea) and a frog (Anura) living in
bromeliads on the Paria Peninsula, northeastern Venezuela
¿Un caso de convergencia evolutiva? Notable semejanza entre una cucaracha
(Blattodea) y una rana (Anura) habitantes de bromelias en la península de Paria,
noreste de Venezuela
Jorge M. González1, Luis E. Sibira2, Tito R. Barros2, Ángel Fernández del Valle3,
Carlo G. Sormani4 & Gilson A. Rivas2
1Austin Achieve Public Schools, Austin, Texas, USA. Research Associate, McGuire Center for Lepidoptera and Biodiversity, USA.
gonzalez.jorge.m@gmail.com; https://orcid.org/0000-0001-7208-7166
2Museo de Biología, Facultad Experimental de Ciencias, Universidad del Zulia, Maracaibo, Venezuela.
anolis30@hotmail.com; https://orcid.org/0000-0001-8003-5075
3Herbario IVIC. Instituto Venezolano de Investigaciones Científicas, Caracas. Venezuela.
angelfern56@yahoo.com
4Musée dHistoire Naturelle de Genève, Geneva, Switzerland.
sormanihc@gmail.com; https://orcid.org/0000-0001-6661-6244
Correspondence: gonzalez.jorge.m@gmail.com
(Received: 15-04-2025 / Accepted: 15-11-2025 / On line: 31-12-2025)
ABSTRACT
Recent herpetological explorations carried out in the Serranía de Paria, northeastern Venezuela, allowed the identica-
tion of the bromeliad Glomeropitcairnia erectiora Mez as a vital refuge for amphibians of the species Phytotriades auratus
(Boulenger, 1917), and for cockroaches of the genus Dryadoblatta Rehn, 1930. It should be noted that the presence of the
frog Scinax ruber (Laurenti, 1768), a species typically of lowlands, has been previously reported in bromeliads of Cerro El
Copey (Margarita Island), but it is likely that the record is a misidentication of P. auratus. e present study also reveals an
association between G. erectiora and a cockroach of the genus Pelmatosilpha Dohrn, 1887. e similar color patterns of P.
auratus and a species of Dryadoblatta suggest possible evolutionary convergence or mimicry between an anuran amphibian
and a blattodean insect, indicating complex ecological relationships in the region.
Keywords: Dryadoblatta sp., evolutionary convergence, Glomeropitcairnia erectiora, mimicry, Phytotriades auratus, Sci-
nax ruber.
RESUMEN
Exploraciones herpetológicas recientes realizadas en la Serranía de Paria, noreste de Venezuela, permitieron identicar a
la bromelia Glomeropitcairnia erectiora Mez como refugio vital de anbios de la especie Phytotriades auratus (Boulenger,
1917) y de cucarachas del género Dryadoblatta Rehn, 1930. Cabe destacar que se ha informado previamente la presencia
de la rana Scinax ruber (Laurenti, 1768), una especie típicamente de tierras bajas, en bromelias del Cerro El Copey (Isla de
Margarita), pero es probable que el registro se trate de una identicación errónea de P. auratus. El presente estudio también
revela una asociación entre G. erectiora y una cucaracha del género Pelmatosilpha Dohrn, 1887. Los patrones de color
Color pattern convergence between a cockroach and a frog in Venezuela
17
similares de P. auratus y de una especie de Dryadoblatta sugieren una posible convergencia evolutiva o mimetismo entre un
anbio anuro y un insecto blattodeo, indicando relaciones ecológicas complejas en la región.
Palabras clave: convergencia evolutiva, Dryadoblatta sp., Glomeropitcairnia erectiora, mimetismo, Phytotriades auratus,
Scinax ruber.
moths, beetles, wasps and bees, and ies) and vertebrates
(i.e., sh, snakes, birds) (Wickler 1968, Pasteur 1982,
Schmied et al. 2012, Kunte et al. 2021). Aposematism,
through bright or ashy color patterns, is a known warn-
ing sign in frogs, which exhibit it to “advertise” their toxic-
ity or lack of palatability to potential predators (Stuckert
et al. 2014a, Lorioux-Chevalier et al. 2023). It is worth
mentioning that although several types of insects mimick-
ing vertebrates have been documented, we know of only
one recent record of an insect (Cratosomus sp.; Curculion-
idae) mimicking a frog [Ameerega trivittata (Spix, 1824):
Dendrobatidae] (Ferreira et al. 2024).
Convergence is another ecological phenomenon (not
mutually exclusive with mimicry or camouage) that op-
erates at both the species and community levels. It may
also be observed in what we perceive as mimicry or cam-
ouage among several organisms. Convergence in ecology
is the independent evolution of similar traits in unrelated
organisms due to similar environmental pressures or eco-
logical roles (Matthews & Matthews 2010). is process
happens because natural selection favors adaptations that
are best suited for survival and reproduction in a particular
environment, leading distantly related species to develop
analogous structures, behaviors, or even behavioral and
physiological processes (Bittleston et al. 2016, Sackton &
Clark 2019).
Several frog groups are well known for including spe-
cies resembling other frogs, either to share a warning signal
with a toxic species (Müllerian mimicry), or to deter pred-
ators (Batesian mimicry), forming mimicry rings where
several species benet from an aposematic coloration
(Darst & Cummings 2006, Prates et al. 2012, Stuckert et
al. 2014b, Ferreira Souza et al. 2024).
Among cockroaches, however, their scavenging and
cryptic habits, as well as the high speed of some, allow
them to frequently escape predators (Evans 1968, Bastidas
Pérez & Zavala Gómez 1995, González 2005, Marshall
2017). Although many cockroach species are cryptic and
dark-colored, allowing them to “disappear” visually in their
environment, it appears to be rare for certain cockroach
species to become members of mimetic rings by exhibit-
ing characteristics that allow them to resemble unpleasant
or poisonous models providing them protection from po-
tential predators (Shelford 1912, Roth & Naskrecki 2001,
INTRODUCTION
Coloration patterns in animals serve several functions,
one of which is protecting them from predators (Badejo
et al. 2020). Two basic types of protective coloration pat-
terns are known. e rst type is camouage, which con-
sists of faint or discrete colors that imitate those of the
environment, substrate, and plants, allowing the animal
to blend in and remain unnoticed by potential predators.
is category includes contrasting and striking colors that
conceal internal features or disrupt the outline of the po-
tential prey (a phenomenon known as disruptive color-
ation; Adams et al. 2019, Hinkelmann 2023), interfering
with visual perception of the predator, thus reducing the
likelihood of an attack. e other type is aposematism, a
conspicuous and colorful coloration serving as a warning
that the potential prey is toxic or inedible, thus inhibiting
predation (Eisner & Grant 1981, Santos et al. 2003, Hin-
kelmann 2023).
In the case of aposematism, some organisms mimic
other aposematic ones, hence taking advantage of the pro-
tective coloration. e imitators (mimics) could be either
more or less toxic/repulsive than the species that they imi-
tate (Müllerian mimicry), or the mimic could be harm-
less (Batesian mimicry) (Bates 1862, 2020, Müller 1878,
Wickler 1968).
Mimicry, in general, is one of the most striking phe-
nomena in evolutionary ecology, occurring in a wide
range of organisms, including unrelated ones (Wickler
1968, Matthews & Matthews 2010, Schmied et al. 2012).
Mimicry systems are ecologically dened as assemblages in
which at least two organisms should be able to play up to
three possible roles: being a model, being a mimic, or be-
ing a deceiver (Wickler 1968, Schmied et al. 2012). Such
systems necessarily contain at least one defended prey that
exhibits a warning signal, which helps reduce predation
pressure, and at least one associated species that derives a
benet from mimicking the aposematic organism (Kunte
et al. 2021).
Over the past few decades, there has been a wealth of
research showing “mimicry rings” (assemblages of species
sharing a similar appearance which serves as an eective
signal to potential predators), Müllerian and Batesian,
involving various groups of insects (i.e., butteries and
González, Sibira, Barros, Fernández del Valle, Sormani & Rivas
18
Deans & Roth 2003, Schmied et al. 2012). At the same
time, some cockroaches are known to produce defensive
secretions and produce pungent-smelling compounds
that potentially make them become models in mimicry
rings (Roth & Willis 1960, Evans 1968, Farine et al. 1997,
O’Connel & Reagle 2002).
To our knowledge, the case presented here may be the
rst reported case of a frog and a cockroach involved in a
possible mimicry ring, in what appears to be an interesting
case of evolutionary convergence.
MATERIALS AND METHODS
On July 2016, a week of eldwork was conducted in
the Cordillera de Paria (sensu Rivas et al. 2021), the east-
ernmost portion of the Cordillera de la Costa in north-
ern Venezuela, specically at Cerro Humo, the highest
summit of the Paria Peninsula (~1,250 m elevation). is
eldwork was carried out as part of a project on the con-
servation status of endemic frog species inhabiting the
region. Invertebrate and small vertebrate specimens were
found in the phytotelmata of the bromeliad Glomeropit-
cairnia erectiora Mez (Bromeliaceae) (Fig. 1).
Several individuals of the tank bromeliad G. erectiora
from Cerro Humo were examined for small vertebrates
such as lizards and frogs. e plants were epiphytes on
branches of an unidentied tree, probably of the genus
Miconia (Melastomataceae).
Although there are no meteorological data from the
summits of Paria, it has been mentioned that in Cerro
Humo there are two thermal oors: the subtropical one
from 400 to 1,000 m above sea level, and the temperate
one restricted to the mountain tops. In these thermal
oors, mean annual temperatures could respectively be
24 and 19 °C, while precipitation is around 2,000 mm per
year (Motta 2001, Fernández & Michelangeli 2003).
RESULTS AND DISCUSSION
The host plant
e genus Glomeropitcairnia Harms (Bromeliaceae)
contains two species native to the Lesser Antilles, Trini-
dad, and Venezuela (Smith 1971, Smith & Downs 1977,
Howard 1979, Hoyos 1985, Ulloa et al. 2018). One of
these species, Glomeropitcairnia penduliora (Griseb.)
Mez, is an endemic epiphyte to the islands of Montserrat,
Guadeloupe, Martinique, and Dominica. e other one,
G. erectiora Mez, which is also epiphytic and occasionally
terrestrial, and is endemic to the montane cloud forests of
northeastern Venezuela (including Margarita Island) and
northern Trinidad (Smith & Downs 1977, Oliva Esteva &
Steyermark 1987, Holst 1994, Hokche et al. 2008). Stey-
ermark (1974, 1976) highlighted that the cloud forests
of the Cordillera de la Costa in Venezuela host a unique
and endemic Amazonian-Guyanese ora element, con-
ned to elevations between 800 and 1,500 meters, which
is crucial for conserving these ecosystems and their biodi-
versity. Such environment and ora (and associated fauna)
are currently threatened, despite being located within two
National Parks (Cerro El Copey, Margarita Island, Nueva
Esparta State; and Paria Peninsula, Sucre State).
It is well known that many bromeliad species generate
microhabitats for arthropods and amphibians that partial-
ly or totally depend on the ecological niche so created to
complete their life cycle (Richardson 1999, Kitching 2000,
Srivastava & Kortright 2006). Such symbiotic relationship
allows plants to obtain nutrients from the activity of the
associated fauna while the animals obtain food, shelter, and
substrate (Frank & Lounibos 2009, Sabagh et al. 2017).
is bromeliad, originally described from Margarita Is-
land, Venezuela, from an altitude of 700 m (Mez 1904),
has been found in other areas of northeastern Venezuela,
in the state of Sucre. It inhabits cloud forests above 700
m above sea level and up to 1,250 m. e plant can also
be found along the Northern Coast Range of the island
of Trinidad (Jowers et al. 2008). is bromeliad was con-
sidered Vulnerable due to habitat destruction caused by
anthropogenic interference in the forests where it is found
(Llamozas et al. 2003), but is currently considered Near
reatened (Huérfano et al. 2020). It might be categorized
as Vulnerable again due to increased forest destruction.
Figure 1. Live individual of the golden tree frog, Phytotriades
auratus (Hylidae, Anura), from the Paria Peninsula, Venezuela
(A), and preserved type of the cockroach Dryadoblatta scotti
(Blaberidae, Blattodea) (B) from Trinidad showing their simi-
larity in dorsal coloration (“Horseshoe pattern”). Photos: M.De
Freitas (le); Amoret Spooner, Hope Entomological Collec-
tion, Oxford University Museum of Natural History (right).
Color pattern convergence between a cockroach and a frog in Venezuela
19
Anuran found inside the bromeliad tank
Phytotriades auratus (Boulenger, 1917) is a frog known
to occur on three small summits in Trinidad and on a peak
in the Paria Peninsula, northeastern Venezuela, where it is
considered endangered (IUCN SSC Amphibian Special-
ist Group 2020). e species thrives at an altitudinal range
from 700 to 1,250 m above sea level, and inhabits the tank
bromeliad G. erectifolia (Rivas & De Freitas 2015, Jowers
et al. 2024).
is rare and endangered anuran ranges in size from
small to medium; males can be up to 29–30 mm long,
while females can reach 35 mm in length from snout to
cloaca (Gray 2003, Jowers et al. 2008). eir base color is
chocolate brown with two distinctive iridescent golden
yellow dorsal stripes running from head to back (Jowers et
al. 2008). Curiously, other frogs exhibit a somewhat simi-
lar pattern (Lehtinen 2020, Ferreira Souza 2024; G. Bec-
caloni, pers. comm.)
Cockroaches inside a bromeliad
On July 2016, a specimen of a dark brown (almost
black) cockroach with translucent mustard-yellow prono-
tum and tegmina edges was collected inside the bromeliad
tank of a G. erectiora individual plant (Fig. 2). e plant
was found on the summit of Cerro Humo, at its highest
elevation (1,250 m above sea level).
e specimen was identied as belonging to Dryadob-
latta Rehn 1930. is genus has two recognized species.
One of them, D. mira Rehn 1937 was described from
the Venezuelan Amazon region, the Cerro Duida at an
elevation of 1,370 (Rehn 1937, Cazorla-Perfetti 2019).
e other one, D. scotti (Shelford, 1912), was initially de-
scribed within the genus Homalopteryx Brunner von Wat-
tenwyl, 1865, to be later placed as the type for the new
genus Dryadoblatta (Shelford 1912, Rehn 1930). e spe-
cies has been collected in northern Trinidad at an eleva-
tion of 950 m (Shelford 1912).
Dryadoblatta scotti is one of more than 60 cockroach
species associated with Bromeliaceae (Roth & Willis
1960; Rocha e Silva et al. 1976). is beautiful species is
considered amphibious or semi-aquatic, and lives at the
water surface in or near the bromeliad tank, diving into
it to collect food or to escape predators (Princis & Kevan
1955). e species has been found associated with Tilland-
sia sp. and G. erectiora, in the mountains of the northern
Cordillera of Trinidad (Shelford 1912, Rehn 1930). Dry-
adoblatta scotti has a chocolate brown coloration contrast-
ing with the edges of the pronotum and tegmina which are
of a very striking golden yellow color, and can measure up
to 30 mm.
Figure 2. Dorsal (A) and ventral (B) views of a specimen of Dry-
adoblatta sp. collected inside a Glomeropitcairnia erectiora from
Cerro Humo, Paria Peninsula, Venezuela. Photo: L. E. Sibira.
González, Sibira, Barros, Fernández del Valle, Sormani & Rivas
20
While visiting Cerro Copey, on Margarita Island
(Venezuela), a cockroach of the genus Pelmatosilpha was
photographed walking out of the tank of a G. erectiora
plant at 750 m elevation (Fig. 3). Pelmatosilpha has 24 rec-
ognized species, of which 21 have been found in Central
America (Costa Rica and Panama), the Caribbean islands
(Trinidad, Grenada, Barbados, St. Lucia, Martinique, An-
tigua, Dominica, and Puerto Rico), and in South America
(Peru, Ecuador, Colombia, Venezuela, Guyana, and Bra-
zil). ree species are known from Venezuela, and at least
one of them is also found in Trinidad (https://cockroach.
speciesle.org/).
Most known cockroach species are cryptic and appear
to be palatable to predators (Evans 1968, Marshall 2017).
However, some are known to be unpalatable, while others,
aer being “crushed” or “torn,” release a foul odor (Evans
1968, Eisner & Grant 1981, Santos et al. 2003, Hinkel-
mann 2023).
The “horseshoe” pattern of roaches and frogs
e coloration pattern of Dryadoblatta scotti, Dryadob-
latta sp. (from the Paria Peninsula), and Pelmatosilpha sp.,
consists of brown shades contrasting with light borders.
is pattern resembles that of frogs such as Phytotriades
auratus, that have a similar size and share the same envi-
ronment. Perhaps one should refer to the frog resembling
the cockroach in terms of the evolutionary scenario result-
ing in these similarities. is type of coloration on these
and other cockroaches is known as the “horseshoe pat-
tern,” which has also been observed in at least one beetle
(see Ferreira Souza et al. 2024), and may be disruptive
and/or aposematic due to its contrasting nature (Fig. 4).
e association of such pattern with some degree of tox-
icity is observed in the cockroach, Pelmatosilpha coriacea
Rehn, 1903, from Puerto Rico. is species is known to
release a repellent secretion which is eective against ants
(Blum 1964). A closely related species, Eurycotis oridana
(Walker, 1868), has been studied for its repellent ecacy
against mice and some insects (Turnbull & Fashing 2002).
Although the “horseshoe” pattern observed involves
cockroach species associated with phytotelmata, this pat-
tern might not be a recent adaptation. It appears to be an
ancestral coloration shared by several other cockroach
species, including terrestrial ones such as Methana mar-
ginalis (Saussure, 1864) from Australia and Dorylaea spp.
from Southeast Asia (Mackerras 1968; G. Beccaloni, pers.
Figure 3. Pelmatosilpha sp. (male) on a leaf of Glomeropitcairnia erectifolia, from Cerro Copey, Margarita Island, Venezuela, photo-
graphed on 2017. Photo: G. A. Rivas.
Color pattern convergence between a cockroach and a frog in Venezuela
21
Figure 4. Dryadoblatta and Pelmatosilpha species showing the “horseshoe pattern” coloration: A. Pelmatosilpha purpurascens, Puerto
Rico [Beccaloni, 2025. Cockroach Species File. Cockroach Species File - Pelmatosilpha purpurascens Kirby, 1903]; B. P. larifuga [Bec-
caloni, 2025. Cockroach Species File. Cockroach Species File - Pelmatosilpha larifuga Gurney, 1965]; C. P. marginalis [Beccaloni,
2025. Cockroach Species File. Cockroach Species File - Pelmatosilpha marginalis Brunner von Wattenwyl, 1893]; D. P. coriacea, Puerto
Rico [Beccaloni, 2025. Cockroach Species File. Cucaracha Arbórea (Pelmatosilpha coriacea) Arboreal Cockroach]; E. Pelmatosilpha
sp. Venezuela (Cerro Copey, Margarita Island) [this work]; F. P. erytrocephala Colombia (Cerro Pintado) [Pelmatosilpha erythrocepha-
la. Photo: ©nmoorhatch. Licenced under Creative Common License Attribution 4.0 International (CC BY 4.0). URL: https://www.
inaturalist.org/observations/172533741 and https://www.gbif.org/occurrence/5167126518]; G.Dryadoblatta mira (male), Venezu-
ela (Cerro Duida) [Taken from Rehn 1937: Pl. XV: g. 21]; H. Dryadoblatta sp. (female), Venezuela (Cerro Humo, Paria Peninsula)
[this work]; I. Dryadoblatta scotti (female), northern Trinidad [Beccaloni, 2025. Cockroach Species File. Cockroach Species File - Dry-
adoblatta scotti (Shelford, 1912)].
González, Sibira, Barros, Fernández del Valle, Sormani & Rivas
22
comm.). As noted above, a similar pattern has also been re-
ported in at least one member of the Curculionidae (Co-
leoptera) (Ferreira Souza et al. 2024) (Fig. 5).
General remarks
In recent years, several localities outside the Paria Pen-
insula have been surveyed for anurans, encompassing the
distribution of the tank bromeliad G. erectiora, a host
plant of P. auratus (Rivas in litt.). e recent discovery
of P. auratus in Venezuela allows us to solve a puzzling
mystery of herpetology in the country. In 1950, two frogs
matching the description of P. auratus were collected in
Cerro Copey, Margarita Island (Roze 1964). Roze (1964)
stated that aer checking 70 bromeliads from the sum-
mit of that mountain, he and his team were able to nd
two frogs that they identied as Scinax ruber (Laurenti,
1768). Unfortunately, the two specimens seem to be lost
(we could not nd them in the Museo de Historia Natu-
ral La Salle, Caracas, where they were originally housed).
ose frogs might represent Phytotriades auratus and
not Scinax ruber. Both species have a vague resemblance,
but the former is normally found in lowlands, although
at least one specimen is known from 800 m above sea
level in a highly anthropogenic environment on the Paria
peninsula (Fig. 6). In addition, G. erectifolia is the most
abundant bromeliad on the summit of Cerro Copey, and
P. auratus is closely associated with this plant species in
Trinidad and Paria Peninsula, and both (the plant and the
frog) appear to be relict species.
Several organisms linked to the bromeliad G. erectifolia
have also been found in the Northern Cordillera of Trini-
dad (Trinidad and Tobago), and Cerro Copey in Margar-
ita Island, and the Paria Peninsula in Sucre State (Venezu-
ela) (Mez 1904, Smith & Downs 1977, Jowers et al. 2008).
Among them, we collected in Paria Peninsula the endemic
lizard Euspondylus monsfumus Mijares-Urrutia, Señaris, &
Arends, 2001, as well as specimens of P. auratus, and some
isopods. Likewise, we found the two cockroach species
observed and mentioned in this study as being associated
with G. erectifolia.
Besides, sucient evidence exists for a biotic relation-
ship between amphibians and cockroaches as predator-
prey, as several families of amphibians are known to fre-
quently feed on cockroaches (Picado 1913, Princis &
Kevan 1955, Roth & Willis 1960). An example of this are
cockroaches of the genus Epilampra found in the stomach
contents of the frog, Eleutherodactylus maestrensis Díaz,
Cádiz & Navarro, 2005, in Cuba, in tropical mountain
and pine forests between 900 and 1,640 m above sea level
(Díaz et al. 2005).
Figure 5. A weevil (Cratosoma sp., Curculionidae, Coleoptera)
(A) and a poison dart frog (Ameerega trivitatata, Dendrobatidae,
Anura) (B) from Brazil, showing similar “horseshoe patterns” to
those of Dryadoblatta spp. and Phytotriades auratus (compare
with Figure 1). Photo: U. Ferreira Souza.
Color pattern convergence between a cockroach and a frog in Venezuela
23
Which arrived first, the cockroach or the frog?
e ancestors of cockroaches originated during the
Carboniferous period, approximately 350–320 million
years ago (McKittrich 1964, Djernæs et al. 2020). How-
ever, modern cockroach lineages emerged around 235 mil-
lion years ago, predating the earliest conrmed cockroach
fossils by about 95 million years (McKittrick 1964, We-
gener 1966, Djernaes et al. 2020, Jin-Lin et al. 2023).
e earliest protofrogs appeared around 250 million
years ago, with the lineage leading to modern frogs emerg-
ing over 150 million years ago (Blackburn & Wake 2011,
Feng et al. 2017, Portik et al. 2023). Most contemporary
lineages, including those of poison dart frogs, diversied
approximately 66 million years ago aer the dinosaur ex-
tinction (Grant et al. 2006, Blackburn & Wake 2011, Feng
et al. 2017, Portik et al. 2023). Interestingly, poison dart
frogs acquire their toxicity as a result of consuming poi-
sonous insects (Summers & Clough 2001, Vargas-Salinas
& Rojas 2024).
Is this mimicry or just convergence?
ree species of the genus Dryadoblatta are now known
from northern South America: D. mira from the Ven-
ezuelan Amazon region, D. scotti from northern Trinidad
(Roth & Willis 1960, Rehn 1937, Shelford 1913), and
Dryadoblatta sp. from the Paria peninsula in Venezuela
(reported herein). is is particularly interesting because
the anuran genus Phytotriades is monotypic, and its sis-
ter genus Itapotihyla Faivovich, Haddad, Garcia, Frost,
Campbell & Wheeler, 2005 is also a monotypic, though
it is distributed in the Atlantic Forests of Brazil, with iso-
lated populations in eastern Paraguay and northeastern
Argentina (Blotto 2021, Frost 2024). Phytotriades and
Dryadoblatta could be relicts of the former Amazonian
refugia of northwestern Venezuela, as established for some
other plant and animal species (Steyermark 1974, 1976,
1982, Schargel et al. 2005). In turn, the tank bromeliad
G. erectifolia could also be considered a relict, and it is cur-
rently isolated on some peaks of northeastern Venezuela,
Margarita Island, and northern Trinidad (this work).
e similarity of color patterns and adult sizes between
the frogs and cockroaches that we studied (despite being
somewhat variable and alike to those observed in other
frog and cockroach groups associated with humid envi-
ronments; G. Beccaloni, pers. comm.), along with their
shared Glomeropitcairnia phytotelmata habitat, suggest
that we are likely observing a convergence system that
could be interpreted or associated with mimicry. It is
worth mentioning again that the only other insect, namely
Cratosomus sp. (Curculionidae) associated to a mimicry
system involving a frog also exhibits the horseshoe pat-
tern (see Ferreira Souza et al. 2024). is reinforces the
idea that such a pattern extends to a broader taxonomic
level in insect-mimicking frogs (or perhaps the other way
around) or even a simple ecological convergence (“a plain
coincidence,” G.Beccaloni, pers. comm.).
In the case of cockroaches of the subfamily Eury-
cotiinae, in which Pelmatosilpha sp. is included, they are
known to emit repellent secretions eective against other
insects and vertebrates (Blum 1964, Turnbull & Fashin
2002). Some frogs of the tribe Lophyohylini, to which P.
auratus belongs, are also known to exude toxic secretions
through serous glands (Blotto et al. 2021). Although simi-
lar secretions have not been recorded in P. auratus, this
frog might produce unpleasant or poisonous exudates.
ese and the coincidental presence of an aposematic col-
or pattern in both species lead us to suspect that we are in
the presence of a possible mimetic association. However,
we cannot rule out the idea that this is just a case of inde-
pendent evolutionary convergence.
e frogs and cockroaches in this study, as well as their
host plant, appear to be relicts of a wider past distribution
in northern South America (e.g., Jowers et al. 2024), and
are now isolated on some peaks in Venezuela and Trini-
dad, where they can still nd an appropriate microhabitat
to live and breed.
ACKNOWLEDGEMENTS
We are deeply indebted to Amoret Spooner and the
Hope Entomological Collection of the Oxford University
Museum of Natural History for allowing us access to the
images of the type specimens of Dryadoblatta scotti. Our
gratitude also goes to Ubiratã Ferreira Souza (State Uni-
Figure 6. Scinax ruber from Cachipal, Península de Paria, Ve-
nezuela. Note the general similarity to Phytotriades auratus.
Photo: L. A. Rodríguez J.
González, Sibira, Barros, Fernández del Valle, Sormani & Rivas
24
versity of Campinas) for providing and allowing us to use
his photographs in Figure 5. We are also grateful for the
thorough comments we received from George Beccaloni
(e Charles Darwin Trust – e Alfred Russel Wallace
Website), who enlightened us greatly, allowing us to im-
prove the original manuscript. We are indebted to the re-
viewers, Jesús Molinari and Marcelo R. Sánchez, for their
thoughtful comments, which allowed us to greatly im-
prove the original manuscript.
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