Diversity
Geographic Range
All 26 species of
Theloderma
can be found throughout southeastern Asia, northeastern India, Myanmar, southern
China, Indochina, Malaya, Sumatra, and Borneo. All of southeast Asia is within the
tropical and subtropical climatic zone. The region is subject to a monsoonal weather
system with wet and dry periods, but overall has a significant annual rainfall. The
terrain includes mountain ranges, planes, and plateaus, but species of
Theloderma
are most abundant in mountainous areas (Leinbach, 2020).
- Biogeographic Regions
- oriental
Habitat
- Habitat Regions
- tropical
- terrestrial
- Terrestrial Biomes
- forest
- rainforest
- mountains
- Aquatic Biomes
- lakes and ponds
- rivers and streams
- temporary pools
- Other Habitat Features
- agricultural
- riparian
Systematic and Taxonomic History
Theloderma
is within family
Rhacophoridae
and order
Anura
. Many species within this genus are still being identified, with the most recent
being in 2021. There has been some debate about the distinction between the three
subfamilies of
Rhacophoridae
,
Buergeriinae
, and
Rhacophorinae
, the latter to which
Theloderma
belongs. There have not been many name changes in
Theloderma
's history, as it is a recently-identified genus with the earliest species,
T. leporosum
, being identified in 1838. Some other closely-related groups to
Theloderma
are
Chiromantis
,
Ghatixalus
, and
Philautus
(Species, 2000; Species, 2020).
Physical Description
Limbs: In a study comparing the physical features of six Theloderma species, they all had many basic characteristics in common. Generally, Theloderma species have strong limbs with their hind legs being about twice the length of their forelimbs (Mian et al., 2017).
Mouths: Theloderma frogs have round snouts with distinct ridges and oval-shaped nostrils that are closer to their mouths than their eyes. They have large, heart-shaped tongues with vomerine teeth that are only located in the front, upper part of their mouth to assist in holding onto prey (rather than chewing) in order to swallow their prey whole.
Size: Size varies between species. T. asperum is one of the smallest species, with adult males as small as 1 inch and adult females around 1.75 inches. There are also larger species such as T. corticale , with males being around 3 inches and females as large as 3.5 inches (Amphibiaweb).
Vision: Theloderma frogs have large eyes with circular pupils. Eyelid coverage varies between species.
Methods of Movement: When hunting or escaping, Theloderma frogs use their muscular hind legs to leap across large distances, but they also crawl and use short hops to travel short distances.
Physical Differentiations Some characteristics that help distinguish between the physical appearances of Theloderma species are coloration, dorsal patterns, and the presence of vocal sacs. Cryptic coloration varies depending on the location and habitat of each species. For example, T. corticale individuals appear to resemble a clump of moss due to their green coloration that is decorated with black spots and tubercles. This camouflage helps them hide in water basins (Smithsonian's National Zoo and Conservation Biology Institute, 2022).
Sexual Dimorphism:
Males tend to be smaller and have nuptial pads on the inner side of their forelimbs
to assist in grip during copulation. Males also have smaller vocal sacs - or in some
species such as
T. kwangsiense
, have no vocal sacs at all.
- Other Physical Features
- ectothermic
- bilateral symmetry
- Sexual Dimorphism
- female larger
Development
Eggs hatch anywhere between 7-14 days, and the newly-hatched tadpoles drop into the
water directly below them. Metamorphosis from tadpole to frog takes about three months.
The exact lifespan of these frogs is unknown, but is estimated to be around 10 years
(Smithsonian's National Zoo and Conservation Biology Institute, 2022).
- Development - Life Cycle
- metamorphosis
Reproduction
Very little is known about
Theloderma
's mating systems. Research on mate competition is also sparse. Females are typically
larger than males, so it is thought that this may be a contributing factor in copulation
success (Mian et al., 2017).
Much of Theloderma 's reproduction method(s) is still unknown due to the cryptic appearances and arboreal lifestyles of this genus. T. asperum have been observed in small aggregations at breeding sites, which include water-filled tree cavities, small rainwater pools, or in artificial water containers (AmphibiaWeb, 2022). Some species, like T. stellatum , are phytotelm breeders; they attach their eggs to tree trunks above water where the larvae then develop (IUCN, 2022). Other species, such as T. corticale , breed in rock cavities where water has flooded the floor (Smithsonian's National Zoo and Conservation Biology Institute, 2022). During breeding, eggs usually measure around 8mm and are attached 2–3cm above the water surface in pairs by a clear, viscous gelatin.
Egg deposition of
T. asperum
appears similar to
T. corticale
. Clutches are small, averaging between 4-15 eggs, and survival rate is unknown. Egg-laying
has been observed during the months of April-June, but could vary between species
(IUCN, 2022).
- Key Reproductive Features
- seasonal breeding
- gonochoric/gonochoristic/dioecious (sexes separate)
- sexual
- fertilization
- oviparous
There is very little parental care after the eggs have been fertilized and deposited
(IUCN, 2022).
- Parental Investment
- no parental involvement
Lifespan/Longevity
The exact lifespan of
Theloderma
species in the wild is unknown due to the challenges faced when observing the frogs
in their natural habitat. The Smithsonian National Zoo and Conservation Biology Institute
(2022) estimates the species' lifespans to be about ten years.
Behavior
Amphibians are known to exhibit a wide range of defensive behaviors, and
Theloderma
species are no exception.
T. asperum
exhibit “shrinking or contracting behavior," during which they curl into an arched/rigid
posture, keep their eyes closed, pull extremities close to the body, and remain motionless.
No defensive calls/smells have been observed (Barrionuevo, 2015).
Theloderma
species are arboreal, making behavioral studies challenging. Most observational and
controlled experiments have been performed in captivity.
Theloderma
species are phytotelm breeders, and larvae go through metamorphosis in water (IUCN,
2022). There is still much to learn about
Theloderma
's intraspecific behaviors.
- Key Behaviors
- arboreal
- nocturnal
- motile
- territorial
- social
- dominance hierarchies
Communication and Perception
Theloderma use anuran vocalizations. When studying the calls of six species ( T. albopunctatum , T. corticale , T. licin , T. auratum , T. stellatum , and T. vietnamense ), the calls were all species-specific, differing in temporal features such as arrangement in call groups and call repetition rate. Some calls do overlap in dominant frequencies, but are otherwise distinguishable.
These advertisement calls can be used to attract conspecific females for mating purposes
or to deter other vocalizing males.
T. asperum
individuals' aggressive calls tend to have a staccato-like sound when two males are
fighting. It should also be noted that since frogs are ectotherms, calls will vary
with change in ambient temperature. The difference in temporal features of calls may
be due to varying snout length, body mass, and distance between frogs. Vocalizations
seem to be the main form of intraspecific communication within
Theloderma
species, although there is still much to be studied (Ginal et al., 2021).
Many amphibians have tetra-chromatic color vision, which includes ultraviolet cones. This enables them to perceive ultraviolet wavelengths. Many predators of frog species, such as pit vipers, have developed the ability to perceive infrared wavelengths. Several anuran species have shown increased reflectance in the infrared spectrum, possibly evolving in response to predator vision as a cryptic tactic (Toledo & Celio, 2009).
Theloderma
species exhibit cryptic coloration. This can make it hard to differentiate between
species and therefore is another challenge to overcome when determining how many species
there are within
Theloderma
without genetic information. Many species within
Theloderma
also exhibit mimicry, tricking predators into thinking they are plants/objects within
the ecosystem (Smithsonian's National Zoo and Conservation Biology Institute, 2022).
- Other Communication Modes
- mimicry
- Perception Channels
- visual
- infrared/heat
- ultraviolet
- tactile
- acoustic
Food Habits
After researchers studied tadpoles' morphology and gut contents,
Theloderma
larvae seem to be omnivorous. They seek a wide variety of food items such as fungal
spores, lepidoptera scales, and exoskeletons of macroscopic arthropods. It is also
possible that
Theloderma
females deposit “nutritive” or unfertilized eggs into arboreal habitats along with
the larvae as another source of food. Morphology of branchial food traps, branchial
baskets, dense gill filters, and floor depressors suggest that
Theloderma
feeds on microscopic plankton, but their IH/OH ratio is rather small for this to
be the case. Therefore, there is still much to learn about larvae consumption because
they seem to possess both dietary extremes of microphagy and macrophagy (Wasserug
et al., 1981).
As for mature individuals of
Theloderma
, they eat similarly to other anuran species. Frogs are generalist predators so they
can adapt their diet to whatever is most abundant at any given time.
Theloderma
are known to preferentially hunt large insects such as crickets and cockroaches.
When given the opportunity, they will also eat aquatic invertebrates (Smithsonian's
National Zoo and Conservation Biology Institute, 2022).
- Primary Diet
-
carnivore
- eats eggs
- insectivore
- herbivore
- planktivore
Predation
Predators of
Theloderma
include tree-dwelling mammals and snakes. Vietnamese mossy frogs (
T. corticale
) are known to fold into a ball when frightened and play dead. Curling up may help
them appear inedible to a snake.
Theloderma
species are known for their anti-predator, cryptic coloration (Smithsonian National
Zoo and Conservation Biology Institute, 2022).
Theloderma
frogs are also nocturnal, so they avoid predation during the day by hiding in water,
under rocks, or by attaching to rock outcroppings where their coloration camouflages
them ("Aquarium of the Pacific", 2022).
- Anti-predator Adaptations
- cryptic
Ecosystem Roles
Frogs play an important role in the food chain as both a predator and prey. As tadpoles,
they eat microorganisms and help regulate algal blooms. Tadpoles are also a food source
for turtles, alligators, crocodiles, and birds. When mature,
Theloderma
frogs are a food source for arboreal mammals and snakes. Adults also help control
insect populations (Gonzalez, 2017). Without insect population control, transmissible
diseases such as malaria could become rampant in human populations.
Theloderma
species also have novel defensin-like antimicrobial peptides found in their skin
secretions that make them 1) susceptible to environmental changes and 2) good indicators
of ecological health (Shen et al., 2016). This becomes especially important as climate
change, pollution, anthropogenic change, and other ecological threats continue to
create lasting impacts on environments.
Economic Importance for Humans: Positive
Theloderma
frogs may not have a direct impact on the economy, but they do have many positive,
indirect impacts. They play an important role as mesopredators, regulating prey species
and serving as food for higher trophic predators (Gonzalez, 2017). Without them, there
would be an accumulation of primary producers and primary consumers. Additionally,
there would be more algal blooms, which are harmful to the environment because they
can block aquatic species from receiving light (Kudela & Gobler, 2012). All of these
issues could lead to the complete failure of ecosystems and, ultimately, species extinctions.
Additionally, multiple species of
Theloderma
, including
T. corticale
,
T. vietnamese
, and
T. auratum
, are used and traded as pets, displays, and for use in horticulture. Their unique
coloration makes them highly attractive to people. Many zoos have also begun to house
species of
Theloderma
(IUCN, 2022).
- Positive Impacts
- pet trade
- ecotourism
- research and education
- controls pest population
Economic Importance for Humans: Negative
Amphibian populations can be costly to maintain, especially when they become overly
abundant. For example, poisonous cane toads (
Rhinella marina
) overpopulated Australia and not only became a nuisance, but a serious concern for
humans (Beckmann & Shine, 2009). Overpopulation can be managed via extermination,
relocation, and other costly tactics. Another way amphibians can become economically
costly is when they are underpopulated. Conservationists recognize
Theloderma
's long-term importance and have attempted to conserve their populations. Sometimes,
these efforts are in vain - or if they are successful, they are very costly.
- Negative Impacts
- injures humans
Conservation Status
Currently, four
Theloderma
species are listed as Endangered, three are listed as Vulnerable, two are listed
as Near-Threatened, fifteen are listed as Least Concern, and five are listed as Data
Deficient. Their populations are threatened by habitat fragmentation, habitat loss,
land conversion, anthropogenic development, and more (IUCN, 2022). These amphibians
can only be found in tropical and subtropical environments, which are some of the
most damaged and threatened ecosystems in the world due to climate change, pollution,
and other anthropogenic impacts. Due to our inability to determine the exact number
of individuals in each
Theloderma
species, it is important to protect as many of these species as possible.
Other Comments
Multiple species of
Theloderma
, including
T. corticale
,
T. vietnamese
, and
T. auratum
, are used and traded as pets, displays, and for use in horticulture. Their unique
coloration makes them attractive to people. Many zoos have also begun to house species
of
Theloderma
(IUCN, 2022).
Additional Links
Contributors
Hannah Grant (author), Colorado State University, Audrey Bowman (editor), Colorado State University, Sydney Collins (editor), Colorado State University.
- oriental
-
found in the oriental region of the world. In other words, India and southeast Asia.
- introduced
-
referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- tropical
-
the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.
- terrestrial
-
Living on the ground.
- forest
-
forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.
- rainforest
-
rainforests, both temperate and tropical, are dominated by trees often forming a closed canopy with little light reaching the ground. Epiphytes and climbing plants are also abundant. Precipitation is typically not limiting, but may be somewhat seasonal.
- mountains
-
This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.
- marsh
-
marshes are wetland areas often dominated by grasses and reeds.
- swamp
-
a wetland area that may be permanently or intermittently covered in water, often dominated by woody vegetation.
- bog
-
a wetland area rich in accumulated plant material and with acidic soils surrounding a body of open water. Bogs have a flora dominated by sedges, heaths, and sphagnum.
- agricultural
-
living in landscapes dominated by human agriculture.
- riparian
-
Referring to something living or located adjacent to a waterbody (usually, but not always, a river or stream).
- seasonal breeding
-
breeding is confined to a particular season
- sexual
-
reproduction that includes combining the genetic contribution of two individuals, a male and a female
- fertilization
-
union of egg and spermatozoan
- internal fertilization
-
fertilization takes place within the female's body
- oviparous
-
reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.
- arboreal
-
Referring to an animal that lives in trees; tree-climbing.
- nocturnal
-
active during the night
- motile
-
having the capacity to move from one place to another.
- territorial
-
defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement
- social
-
associates with others of its species; forms social groups.
- dominance hierarchies
-
ranking system or pecking order among members of a long-term social group, where dominance status affects access to resources or mates
- visual
-
uses sight to communicate
- acoustic
-
uses sound to communicate
- mimicry
-
imitates a communication signal or appearance of another kind of organism
- visual
-
uses sight to communicate
- infrared/heat
-
(as keyword in perception channel section) This animal has a special ability to detect heat from other organisms in its environment.
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- cryptic
-
having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.
- pet trade
-
the business of buying and selling animals for people to keep in their homes as pets.
- ecotourism
-
humans benefit economically by promoting tourism that focuses on the appreciation of natural areas or animals. Ecotourism implies that there are existing programs that profit from the appreciation of natural areas or animals.
- poisonous
-
an animal which has a substance capable of killing, injuring, or impairing other animals through its chemical action (for example, the skin of poison dart frogs).
- carnivore
-
an animal that mainly eats meat
- insectivore
-
An animal that eats mainly insects or spiders.
- herbivore
-
An animal that eats mainly plants or parts of plants.
- folivore
-
an animal that mainly eats leaves.
- planktivore
-
an animal that mainly eats plankton
- ectothermic
-
animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature
- bilateral symmetry
-
having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.
- metamorphosis
-
A large change in the shape or structure of an animal that happens as the animal grows. In insects, "incomplete metamorphosis" is when young animals are similar to adults and change gradually into the adult form, and "complete metamorphosis" is when there is a profound change between larval and adult forms. Butterflies have complete metamorphosis, grasshoppers have incomplete metamorphosis.
References
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Beckmann, C., R. Shine. 2009. Impact of Invasive Cane Toads on Australian Birds. Conservation Biology , 23 (6): 1544-1549.
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