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Horned frog

The horned frog, genus Ceratophrys, comprises eight species of robust, ground-dwelling amphibians in the family Ceratophryidae, endemic to South America and distinguished by their large, triangular heads, expansive mouths, and prominent horn-like flaps of skin above each eye that aid in camouflage and threat displays.^[1]^[2] These frogs, also known as Pacman frogs due to their round bodies and voracious feeding habits, typically measure 8–15 cm in length, with females often larger than males, and exhibit cryptic coloration ranging from earthy browns and greens to aid in blending with leaf litter or soil.^[3]^[4] Horned frogs are sit-and-wait ambush predators, burying themselves partially in substrate within grasslands, rainforests, and semi-arid regions from Argentina and Brazil to the Amazon Basin, where they await prey using exceptional bite forces—up to 500 Newtons in species like Ceratophrys cranwelli—and adhesive, mucus-covered tongues to capture insects, small mammals, fish, and even conspecifics.^[1]^[4] Their diet is highly opportunistic and carnivorous, supported by sharp, curved teeth and vomerine odontophores, while unique behaviors such as pedal luring—wiggling toes to mimic prey—enhance their hunting efficiency.^[1] During dry periods, adults form a protective cocoon from shed skin to retain moisture, emerging after heavy rains for explosive breeding events in temporary pools.^[1] Reproduction in horned frogs is rapid and prolific, with females laying up to several thousand eggs in temporary pools, left unguarded after deposition; the resulting tadpoles are obligate carnivores, capable of producing underwater vocalizations to avoid cannibalism among siblings, and metamorphose within 90 days under favorable conditions.^[1]^[3] While several species thrive in the wild, two are declining due to habitat loss and overcollection for the pet trade—as of 2025, including Ceratophrys ornata now classified as Vulnerable—highlighting conservation concerns for this charismatic group.^[1]^[5]

Taxonomy and classification

Genus overview

The term "horned frog" primarily refers to members of the genus Ceratophrys, which belongs to the family Ceratophryidae in the order Anura.^[2] This genus encompasses South American species noted for their distinctive morphology, while the family Ceratophryidae comprises three extant genera and 12 species distributed across South America.^[2] Members of Ceratophryidae exhibit a robust build adapted for predatory lifestyles, including large heads and expansive mouths suited for ambushing prey.^[6] The genus Ceratophrys is classified within the subfamily Ceratophryinae, which also includes the genera Chacophrys and Lepidobatrachus.^[7] The etymology of Ceratophrys originates from the Greek words kéras (horn) and ophrys (brow), alluding to the prominent supratympanic projections that give the appearance of horns above the eyes.^[8] Historically, the genus Ceratophrys was first established by Maximilian zu Wied-Neuwied in 1824, with the family Ceratophryidae formally described by Johann Jakob von Tschudi in 1838.^[7] A key early species, Ceratophrys ornata, was described by Thomas Bell in 1843 as Uperodon ornatum, marking an important contribution to the taxonomic understanding of these frogs during the 19th century.^[9] Subsequent revisions, including phylogenetic analyses, have refined the classification while confirming the monophyly of the group.

Species diversity

The genus Ceratophrys includes eight recognized species of South American horned frogs, distributed across diverse habitats from the Amazon basin to the grasslands of Argentina and Brazil. These species are characterized by their robust bodies, wide mouths, and prominent superoocular spines, though individual differences in coloration, skin texture, and cranial features distinguish them.^[2]^[10]

Species	Common Name	Key Distinguishing Features
C. ornata	Argentine horned frog	Ornate dorsal patterns with bright green and black mottling; octoploid chromosome number.^[10]
C. cornuta	Amazon horned frog	Pronounced sexual dimorphism in size, with males smaller than females; elongate tubercles on upper eyelids.^[10]^[11]
C. cranwelli	Cranwell's horned frog	Diploid chromosome number; dorsal bony shield; found in arid Chaco regions.^[10]
C. stolzmanni	Pacific horned frog	Adapted to xeric environments with variable skin texture; lacks dorsal shield in some populations.^[10]
C. aurita	Brazilian horned frog	Prominent cranial morphology with reduced horns; dorsal shield present.^[10]
C. calcarata	Brazilian fat toad	Aggressive behavior and pedal luring in juveniles; hyperossified skeleton.^[10]
C. joazeirensis	Joazeiro horned frog	Restricted to northeast Brazil; dorsal shield similar to C. cranwelli.^[10]
C. testudo	Ecuadorian horned frog	Known only from type specimen; validity debated, possibly synonymous with C. cornuta.^[2]^[10]

Taxonomic debates within Ceratophrys have centered on species boundaries and polyploidy, with C. ornata historically encompassing populations now recognized as C. cranwelli based on molecular data. Post-2000 phylogenetic studies using mitochondrial and nuclear DNA have confirmed the monophyly of the genus and resolved two major clades: one featuring a dorsal bony shield (C. aurita, C. cranwelli, C. joazeirensis, C. ornata) and another including C. calcarata, C. cornuta, and C. stolzmanni. These analyses also suggest complex ploidy evolution, with a single origin of octoploidy in the C. ornata lineage and potential diploid reversals.^[10] It is important to distinguish Ceratophrys from unrelated "horned frogs" in other genera, such as the Asian Pelobatrachus nasutus (formerly Megophrys nasuta), which belongs to the family Megophryidae and exhibits convergent horn-like projections for camouflage rather than predatory adaptations.

Physical description

Morphology and adaptations

Horned frogs of the genus Ceratophrys exhibit a robust, dorsoventrally flattened body form that facilitates burrowing and stationary ambush postures in their environment. This squat physique is characterized by a disproportionately large head relative to the body, often comprising a significant portion of the overall length, with a wide mouth that can extend nearly the full width of the head.^[12] The expansive oral gape, supported by a heavily ossified skull and fused mandibular symphysis, allows these frogs to ingest prey items substantially larger than their own head, including vertebrates up to half their body size.^[12] Key adaptations in the cranial region include powerful jaws bolstered by large adductor muscles and a rigid skeletal structure, enabling exceptional bite forces that scale allometrically with head width—for example, up to 32.9 N in adults with ~46 mm head width, and extrapolated to ~500 N in larger individuals.^[12]^[1] The dentition features non-pedicellate, monocuspid teeth on the upper jaw with sharp, recurved tips, complemented by a pair of robust, fang-like odontoids on the lower jaw's symphysis for piercing and securing prey integument.^[12] Additionally, the tongue is a muscular, viscoelastic structure coated in mucus that generates adhesive forces up to 17.7 kPa, primarily through hydrostatic coupling and rapid projection via jaw depression, rather than solely mucus viscosity.^[13] Supratympanic "horns," consisting of fleshy eyelid projections or ridges above the eyes, contribute to defensive posturing and visual camouflage by mimicking environmental protrusions.^[14] The skin of Ceratophrys species is notably thick and glandular, with a well-differentiated dermis and compact stratum featuring early maturation of multiple epidermal layers, providing structural reinforcement and protection against desiccation. This integument often incorporates co-ossification with underlying bones, such as a dermal shield over presacral vertebrae in some taxa, enhancing overall rigidity for survival in variable substrates.^[14] Limbs are short and sturdy, with the prehallux modified into a keratinous spade—comprising spherical proximal and axehead-shaped distal elements—for efficient burrowing into soil or leaf litter. Sensory adaptations include large, dorsally positioned eyes with lenticular pupils, optimizing detection of overhead or approaching threats during immobile phases. Males possess a subgular vocal sac, a flexible membrane of pigmented skin that amplifies acoustic signals, though its morphology varies slightly across species with a darkly pigmented gular region.

Size and sexual dimorphism

Horned frogs in the genus Ceratophrys exhibit considerable variation in adult body size, with snout-vent length (SVL) typically ranging from 10 to 13 cm across species, though some individuals reach up to 20 cm in larger forms.^[15]^[16] Females are consistently larger than males, displaying female-biased sexual size dimorphism that averages 6-10% greater SVL in several species.^[17]^[18] This dimorphism supports differences in resource allocation, with females developing a bulkier, more rounded body shape compared to the slimmer build of males.^[3] Species-specific sizes highlight this variability; for instance, in C. cornuta, adult females attain up to 120 mm SVL and 130 g body mass, while males reach a maximum of 72 mm SVL and 60 g.^[15] In C. ornata, the average SVL is 112.4 mm (±13.4 mm) for adults of both sexes, though females tend toward the upper end of this range.^[16] Similarly, C. cranwelli adults vary widely from 42 to 130 mm SVL, with females occupying the larger size classes.^[19] Males across the genus often exhibit additional dimorphic traits, such as pronounced subgular vocal sacs used in calling, which are absent in females.^[3]^[20] Growth patterns in Ceratophrys are characterized by rapid juvenile development, with accelerated rates in the first year post-metamorphosis leading to sexual maturity within 1-2 years.^[18]^[21] This fast growth enables quick attainment of predatory size, though no major ontogenetic shifts in coloration occur, maintaining consistent patterns from juvenile to adult stages.^[22]

Distribution and habitat

Geographic range

Horned frogs of the genus Ceratophrys are endemic to South America, with distributions spanning diverse regions from the Amazon Basin to the grasslands of the southern cone. The genus occupies a broad latitudinal range across the continent, primarily in lowland and foothill areas below 1,000 meters elevation, though some populations extend into Andean foothills up to approximately 600 meters in southwestern Ecuador and northwestern Peru. Species distributions vary regionally, reflecting ecological specialization within the genus. Ceratophrys cornuta, the Amazon horned frog, is widespread in the Amazon Basin, occurring in Colombia, Ecuador, the Guianas, Venezuela, Peru, Bolivia, and Brazil.^[4] Ceratophrys cranwelli inhabits the Gran Chaco region, found in Argentina, Bolivia, Paraguay, and western Brazil.^[23] Ceratophrys ornata ranges through the Pampean grasslands of central and eastern Argentina (including Buenos Aires, Córdoba, Entre Ríos, La Pampa, Mendoza, and Santa Fe provinces), Uruguay, and southern Brazil.^[16] Ceratophrys aurita is restricted to eastern Brazil, from Bahia and Minas Gerais southward to Rio Grande do Sul, primarily in the Atlantic Forest region.^[24] Ceratophrys stolzmanni, the Pacific horned frog, occurs along the Pacific coast in seasonally dry environments of southwestern Ecuador (Manabí, Santa Elena, Guayas, Loja, and El Oro provinces) and northwestern Peru (Tumbes department).^[25] Ceratophrys calcarata occurs in the Caribbean lowlands of Colombia and Venezuela.^[26] Ceratophrys joazeirensis is endemic to the Caatinga region of northeastern Brazil.^[27] Ceratophrys testudo is endemic to Ecuador.^[28] Historically, the ranges of Ceratophrys species have shown relative stability since their description in the 19th and early 20th centuries, with no evidence of major latitudinal or elevational shifts. However, habitat fragmentation has increased since the 1980s due to agricultural expansion and urbanization, particularly affecting peripheral populations; for instance, C. ornata has not been recorded in Uruguay since 1982 or in Brazil since 1992, resulting in discontinuous distributions within its former continuous grassland range.^[29] Introduced populations of horned frogs outside their native South American range are rare and unestablished. While species like C. cranwelli and C. ornata are commonly collected for the international pet trade, escapes or releases have not led to viable wild populations elsewhere, with individuals occasionally reported as vagrants in North America and Europe but failing to persist.^[16]

Ecological preferences

Horned frogs of the genus Ceratophrys primarily inhabit humid grasslands, tropical rainforests, savannas, and areas adjacent to temporary pools, where they exhibit semi-aquatic behaviors during the breeding season.^[15]^[19]^[16] These environments provide the moist conditions necessary for their fossorial lifestyle, with species like C. cornuta favoring the leaf-litter floors of Amazonian rainforests and C. ornata occurring in temperate grasslands near ephemeral water bodies in the Pampas region.^[15]^[16] In savanna-like habitats, such as those occupied by C. calcarata, individuals are often found in shrublands and inland wetlands that support seasonal flooding.^[26] Microhabitat preferences center on burrowing into leaf litter, loose soil, or sandy substrates to create ambush sites, allowing these frogs to remain concealed while awaiting prey.^[15]^[30] Most species avoid strictly arid zones, opting instead for areas with periodic moisture; however, C. stolzmanni tolerates drier Pacific coastal environments, including seasonally dry forests and disturbed agricultural lands with sparse vegetation.^[30] During dry periods, individuals aestivate underground, forming protective cocoons from shed skin to minimize water loss.^[16] Climate tolerances for Ceratophrys species align with tropical and subtropical regimes, typically ranging from 20–30°C with high humidity levels of 60–80%, particularly elevated at night or during wet seasons.^[31]^[32] Activity is strongly seasonal, peaking during rainy periods when temperatures and moisture facilitate emergence, breeding, and foraging, as seen in the Gran Chaco where C. cranwelli activates post-rainfall in semi-arid settings.^[19] Sympatric interactions involve co-occurrence with other amphibians such as Rhinella toads and Leptodactylus frogs in shared breeding sites, where competitive exclusion may occur due to aggressive tadpole behaviors and resource competition in temporary pools.^[30]^[31] These dynamics are evident in overlapping distributions in subtropical wetlands, though horned frog tadpoles often dominate through cannibalism, reducing conspecific and heterospecific densities.^[19]

Behavior and life cycle

Predatory strategies

Horned frogs of the genus Ceratophrys are specialized sit-and-wait predators that employ an ambush strategy, burying themselves in leaf litter or soil with only their faces exposed to blend seamlessly with the environment through cryptic coloration and dorsal patterns resembling foliage.^[33] This stationary camouflage allows them to remain motionless for extended periods, relying on patience to detect passing prey via visual cues, particularly at night when their activity peaks. Upon detection, they lunge explosively, using their protrusible tongue to adhere to the prey before clamping down with a powerful bite that can generate forces up to approximately 500 N in larger adults, sufficient to subdue vertebrates comparable in size to the frog itself.^[33] Some species employ pedal luring, wiggling their toes to attract prey by mimicking insects or worms.^[34] Their diet is broadly carnivorous and opportunistic, consisting primarily of invertebrates such as crickets, beetles, and other insects, alongside small vertebrates including other frogs, lizards, and rodents. Juveniles frequently exhibit cannibalism, preying on smaller conspecifics or tadpoles in temporary pools, which supplements their intake of insects and reinforces their generalist feeding habits. To consume large prey, they utilize their expansive gape and robust jaws to swallow items whole, often employing forelimbs to manipulate and position struggling victims during ingestion.^[33] When threatened, horned frogs display aggressive defensive behaviors, including rapid body inflation via lung expansion to appear larger and deter attackers, erection of their prominent cranial horns to enhance intimidation, and direct biting with their strong jaws to inflict injury. These tactics, combined with their nocturnal lifestyle and physical adaptations like a rigid skull and recurved teeth, enable effective prey capture while minimizing risks from predators or rivals.^[33]

Reproduction and development

Horned frogs of the genus Ceratophrys exhibit an explosive reproductive strategy synchronized with environmental cues, primarily heavy rainfall that fills temporary pools in their arid or semi-arid habitats. Breeding is triggered by these rains, prompting males to emerge from burrows and initiate chorusing with advertisement calls to attract females, often leading to intense, short-duration mating aggregations. During amplexus, males employ an axillary grasp to stimulate oviposition, with courtship and egg-laying typically occurring nocturnally on the same night.^[19]^[35]^[36] Females produce large clutches of 1,000–2,000 pigmented, gelatinous eggs, which are typically laid directly in shallow temporary pools, without foam nests; clutch size correlates positively with female body size and condition. Paternal care is absent in most species. Maternal guarding is not observed. Eggs hatch within 1–3 days at temperatures around 25–27°C, releasing tadpoles into the water.^[19]^[37]^[38] Tadpoles of Ceratophrys are specialized carnivores, featuring robust keratinized jaw sheaths adapted for tearing prey such as conspecifics, insects, and microcrustaceans, though incidental plant material may be ingested, and can produce underwater vocalizations during interactions with conspecifics, potentially aiding in predator avoidance or cannibalistic encounters. Development proceeds rapidly through free-swimming and primarily benthic phases to exploit ephemeral pools, with metamorphosis completing in 20–40 days depending on temperature and resource availability; Gosner stages 39–42 tadpoles measure 25–31 mm in snout-vent length before transforming into juveniles of 25–43 mm.^[39]^[40]^[19]^[41] Individuals reach sexual maturity at 1–2 years of age, with males often maturing slightly earlier than females, enabling annual breeding cycles tied to seasonal rains in most species. This fast life history supports high reproductive output but limits longevity, with adults typically breeding for only a few seasons.^[21]^[42]^[43]

Conservation and human interaction

Threats and status

Horned frogs of the genus Ceratophrys face several primary threats in the wild, with habitat destruction being the most significant. Agricultural expansion and urbanization have led to substantial losses of native grasslands, particularly in the Argentine Pampas, where approximately 30% of the original vegetation has been cleared since the 1980s for cropland and urban development.^[44] This fragmentation directly impacts species like C. ornata, whose burrowing and breeding habits rely on intact temperate grasslands.^[29] Additionally, collection for the international pet trade contributes to population declines, especially for C. ornata, where surveys indicate that a majority of traded specimens are wild-caught, exacerbating pressure on already vulnerable populations.^[45] Regarding conservation status, most Ceratophrys species are classified as Least Concern by the IUCN Red List, including C. cornuta, C. cranwelli, and C. aurita, due to their relatively wide distributions and adaptability in some Amazonian and Chacoan habitats.^[15] However, C. ornata was recently uplisted to Vulnerable in 2025, reflecting ongoing declines from habitat loss and trade, while C. stolzmanni remains Vulnerable owing to arid Pacific coastal habitat degradation in Ecuador and Peru.^[5] No species in the genus is currently listed as Endangered.^[46] Conservation efforts for C. ornata include the Giant of the Pampas Project, initiated in 2016, which focuses on ecological research and community engagement to support in-situ and ex-situ conservation. As of 2025, funding from organizations such as the Rufford Foundation and Amphibian Ark supports actions like habitat protection and captive breeding programs to mitigate declines.^[47]^[48] Emerging risks include climate change, which is projected to alter rainfall patterns and increase drought frequency across South American grasslands and forests, potentially disrupting breeding cycles that depend on seasonal flooding for Ceratophrys species.^[49] Diseases such as chytridiomycosis, caused by the fungus Batrachochytrium dendrobatidis, have been detected in some captive and wild horned frogs but are minimally reported as a major driver of declines compared to habitat threats.^[16] Population trends vary by region and protection status; recent 2020s surveys show stability in protected areas like national parks in Argentina and Brazil, where habitat remains intact, but declines in fragmented landscapes due to ongoing agricultural conversion.^[29] For instance, C. ornata populations have decreased in unprotected Pampas sites, with no records in parts of Uruguay and southern Brazil for decades.^[50]

Captivity and pet trade

Horned frogs, particularly species in the genus Ceratophrys such as C. cranwelli, C. ornata, and C. cornuta, are popular in the international pet trade due to their distinctive appearance, large size, and aggressive feeding behavior, often earning them the nickname "Pacman frogs."^[4]^[3] These amphibians are valued for their relative ease of care compared to more active frog species, making them suitable for beginner hobbyists, though their sedentary nature limits interactive handling.^[51] In captivity, horned frogs require a minimum 10-20 gallon terrarium with high humidity (50-80%) maintained via damp substrate like sphagnum moss or coconut fiber, allowing for burrowing behavior, and a shallow water dish for soaking.^[3]^[51] Temperatures should range from 75-82°F (24-28°C) during the day, dropping to 65-78°F (18-26°C) at night, achieved with under-tank heaters rather than overhead lighting to avoid desiccation.^[3]^[51] Diet consists primarily of gut-loaded insects such as crickets and earthworms, supplemented with small vertebrates like fish or pinky mice for adults, dusted with calcium and vitamin supplements 1-3 times weekly to prevent metabolic bone disease.^[3]^[51] They must be housed solitarily to prevent cannibalism, as these ambush predators will consume tank mates, including conspecifics.^[51] Lifespan in captivity averages 10 years, with some reaching 15 years under optimal conditions.^[4] Breeding occurs readily in captivity, with sexual maturity reached at 18-24 months; males develop nuptial pads and vocalize during the breeding season, while females are larger and produce 1,000-2,000 eggs per clutch in shallow water setups.^[3] Captive-bred specimens dominate the trade in many regions, reducing pressure on wild populations, though wild-caught individuals remain common in areas like Argentina, where surveys indicate up to 77% of pet horned frogs (C. ornata) are sourced from the wild, contributing to local declines.^[3]^[45] Efforts to promote captive breeding, such as those by specialized ranches, have increased availability of color morphs and hybrids, enhancing trade sustainability.^[52] Common health issues in captive horned frogs include obesity from overfeeding, corneal lipidosis from high-fat diets, and infectious diseases like chytridiomycosis and red-leg syndrome, exacerbated by poor water quality or stress during transport.^[3]^[53] Quarantine periods of 30-90 days, regular veterinary checks, and biosecurity measures are essential to mitigate parasite loads and bacterial infections prevalent in the pet trade.^[53] While not currently listed under CITES Appendix I, the trade's impact underscores the need for sourcing from reputable breeders to support conservation.^[45]

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[41]
[PDF] Table 7: Species changing IUCN Red List Status (2024–2025)
Oct 10, 2025 · Ceratophrys ornata. Argentinean Horned Frog. NT. VU. N. 2025-2. AMPHIBIANS. Churamiti maridadi. Beautiful Metallic Toad. CR. CR(PE). N. 2025-2.
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The IUCN Red List of Threatened Species
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Effect of climate change on distribution of species of common ...
Our main objectives were to verify the effect of climate change on distribution of frogs of the family Ceratophryidae and if the legal protection areas in ...
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[PDF] In Search of the Horned Frog (Ceratophrys ornata) in Argentina
The Horned Frog (Ceratophrys ornata; Appendix 1) is an amphibian species that occurs in the South American temperate grasslands.
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A Guide to Caring for Pacman Frogs as Pets
Jun 5, 2025 · Pacman frogs are primarily terrestrial and require a humid environment with a substrate suitable for burrowing. They can grow to be about 6 ...
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Ornate Horned Frog for Sale
We have stunning captive bred Ornate Horned frogs for sale at really low prices. We get our frogs directly from The Frog Ranch, so you know their genetics are ...
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Infectious Diseases of Amphibians - Exotic and Laboratory Animals - Merck Veterinary Manual
### Summary of Infectious Diseases of Amphibians Relevant to Horned Frogs or Pet Trade