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The cane toad (''Rhinella marina''), also known as the giant neotropical toad or marine toad, is a large, terrestrial
true toad A true toad is any member of the family Bufonidae, in the order Anura (frogs and toads). This is the only family of anurans in which all members are known as toads, although some may be called frogs (such as harlequin frogs). The bufonids now ...
native to
South South is one of the cardinal directions or compass points. The direction is the opposite of north and is perpendicular to both west and east. Etymology The word ''south'' comes from Old English ''sūþ'', from earlier Proto-Germanic ''*sunþa ...
and mainland
Central America Central America is a subregion of North America. Its political boundaries are defined as bordering Mexico to the north, Colombia to the southeast, the Caribbean to the east, and the Pacific Ocean to the southwest. Central America is usually ...
, but which has been introduced to various islands throughout
Oceania Oceania ( , ) is a region, geographical region including Australasia, Melanesia, Micronesia, and Polynesia. Outside of the English-speaking world, Oceania is generally considered a continent, while Mainland Australia is regarded as its co ...
and the
Caribbean The Caribbean ( , ; ; ; ) is a region in the middle of the Americas centered around the Caribbean Sea in the Atlantic Ocean, North Atlantic Ocean, mostly overlapping with the West Indies. Bordered by North America to the north, Central America ...
, as well as
Northern Australia The unofficial geographic term Northern Australia includes those parts of Queensland and Western Australia north of latitude 26th parallel south, 26° and all of the Northern Territory. Those local government areas of Western Australia and Q ...
. It is a member of the genus '' Rhinella'', which includes many true toad
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
found throughout Central and South America, but it was formerly assigned to the genus '' Bufo''. A fossil toad (specimen UCMP 41159) from the La Venta fauna of the late
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
in
Colombia Colombia, officially the Republic of Colombia, is a country primarily located in South America with Insular region of Colombia, insular regions in North America. The Colombian mainland is bordered by the Caribbean Sea to the north, Venezuel ...
is morphologically indistinguishable from modern cane toads from northern South America. It was discovered in a floodplain deposit, which suggests the ''R. marina'' habitat preferences have long been for open areas. The cane toad is a prolific breeder; females lay single-clump spawns with thousands of eggs. Its reproductive success is partly because of opportunistic feeding: it has a diet, unusual among anurans, of both dead and living matter. Adults average in length; the largest recorded specimen had a snout-vent length of . The cane toad has poison
gland A gland is a Cell (biology), cell or an Organ (biology), organ in an animal's body that produces and secretes different substances that the organism needs, either into the bloodstream or into a body cavity or outer surface. A gland may also funct ...
s, and the
tadpole A tadpole or polliwog (also spelled pollywog) is the Larva, larval stage in the biological life cycle of an amphibian. Most tadpoles are fully Aquatic animal, aquatic, though some species of amphibians have tadpoles that are terrestrial animal, ...
s are highly toxic to most animals if ingested. Its toxic skin can kill many animals, both wild and domesticated, and cane toads are particularly dangerous to dogs. Because of its voracious appetite, the cane toad has been introduced to many regions of the Pacific and the Caribbean islands as a method of agricultural
pest control Pest control is the regulation or management of a species defined as a pest (organism), pest; such as any animal, plant or fungus that impacts adversely on human activities or environment. The human response depends on the importance of the da ...
. The common name of the species is derived from its use against the
cane beetle ''Dermolepida albohirtum'', the cane beetle, is a native Australian beetle and a pest of sugarcane. Adult beetles eat the leaves of sugarcane, but greater damage is done by their larvae hatching underground and eating the roots, which either ki ...
(''Dermolepida albohirtum''), which damages
sugar cane Sugarcane or sugar cane is a species of tall, Perennial plant, perennial grass (in the genus ''Saccharum'', tribe Andropogoneae) that is used for sugar Sugar industry, production. The plants are 2–6 m (6–20 ft) tall with stout, jointed, fib ...
. The cane toad is now considered a pest and an
invasive species An invasive species is an introduced species that harms its new environment. Invasive species adversely affect habitats and bioregions, causing ecological, environmental, and/or economic damage. The term can also be used for native spec ...
in many of its introduced regions. The 1988 film '' Cane Toads: An Unnatural History'' documented the trials and tribulations of the introduction of
cane toads in Australia The cane toad in Australia is regarded as an exemplary case of an invasive species. Australia's relative isolation prior to History of Australia (1788–1850), European colonisation and the Industrial Revolution, both of which dramatically in ...
.


Taxonomy

Historically, the cane toad was used to eradicate pests from
sugarcane Sugarcane or sugar cane is a species of tall, Perennial plant, perennial grass (in the genus ''Saccharum'', tribe Andropogoneae) that is used for sugar Sugar industry, production. The plants are 2–6 m (6–20 ft) tall with stout, jointed, fib ...
, giving rise to its common name. The cane toad has many other common names, including "giant toad" and "marine toad"; the former refers to its size, and the latter to the
binomial name In taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin grammatical forms, altho ...
, ''R. marina''. It was one of many species described by
Carl Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
in his 18th-century work ''
Systema Naturae ' (originally in Latin written ' with the Orthographic ligature, ligature æ) is one of the major works of the Sweden, Swedish botanist, zoologist and physician Carl Linnaeus (1707–1778) and introduced the Linnaean taxonomy. Although the syste ...
'' (1758). Linnaeus based the
specific epithet In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...
''marina'' on an illustration by Dutch zoologist Albertus Seba, who mistakenly believed the cane toad to inhabit both terrestrial and marine environments. Other common names include "giant neotropical toad", "Dominican toad", "giant marine toad", and "South American cane toad". In
Trinidad Trinidad is the larger, more populous island of the Republic of Trinidad and Tobago, the country. The island lies off the northeastern coast of Venezuela and sits on the continental shelf of South America. It is the southernmost island in ...
ian English, they are commonly called ''crapaud'', the French word for toad. The genus ''Rhinella'' is considered to constitute a distinct
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of its own, thus changing the
scientific name In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...
of the cane toad. In this case, the specific name ''marinus'' (
masculine Masculinity (also called manhood or manliness) is a set of attributes, behaviors, and roles generally associated with men and boys. Masculinity can be theoretically understood as socially constructed, and there is also evidence that some beh ...
) changes to ''marina'' (feminine) to conform with the rules of gender agreement as set out by the
International Code of Zoological Nomenclature The International Code of Zoological Nomenclature (ICZN) is a widely accepted Convention (norm), convention in zoology that rules the formal scientific name, scientific naming of organisms treated as animals. It is also informally known as the I ...
, changing the binomial name from ''Bufo marinus'' to ''Rhinella marina''; the binomial ''Rhinella marinus'' was subsequently introduced as a synonym through misspelling by Pramuk, Robertson, Sites, and Noonan (2008). Though controversial (with many traditional herpetologists still using ''Bufo marinus'') the binomial ''Rhinella marina'' is gaining in acceptance with such bodies as the
IUCN The International Union for Conservation of Nature (IUCN) is an international organization working in the field of nature conservation and sustainable use of natural resources. Founded in 1948, IUCN has become the global authority on the status ...
, Encyclopaedia of Life, Amphibian Species of the World and increasing numbers of scientific publications adopting its usage. Since 2016, cane toad populations native to
Mesoamerica Mesoamerica is a historical region and cultural area that begins in the southern part of North America and extends to the Pacific coast of Central America, thus comprising the lands of central and southern Mexico, all of Belize, Guatemala, El S ...
and northwestern South America are sometimes considered to be a separate species, '' Rhinella horribilis''. In Australia, the adults may be confused with large native frogs from the genera '' Limnodynastes'', '' Cyclorana'', and '' Mixophyes''. These species can be distinguished from the cane toad by the absence of large
parotoid gland The parotoid gland (alternatively, paratoid gland) is an external skin gland on the back, neck, and shoulder of some frogs (especially toads), and salamanders. It can secrete a number of milky alkaloid substances (depending on the species) known ...
s behind their eyes and the lack of a ridge between the nostril and the eye. Cane toads have been confused with the giant burrowing frog (''Heleioporus australiacus''), because both are large and warty in appearance; however, the latter can be readily distinguished from the former by its vertical pupils and its silver-grey (as opposed to gold) irises. Juvenile cane toads may be confused with species of the genus '' Uperoleia'', but their adult colleagues can be distinguished by the lack of bright colouring on the groin and thighs. In the United States, the cane toad closely resembles many bufonid species. In particular, it could be confused with the southern toad (''Bufo terrestris''), which can be distinguished by the presence of two bulbs in front of the parotoid glands.


Taxonomy and evolution

The cane toad genome has been sequenced and certain Australian academics believe this will help in understanding how the toad can quickly evolve to adapt to new environments, the workings of its infamous toxin, and hopefully provide new options for halting this species' march across Australia and other places it has spread as an invasive pest. Studies of the genome confirm its evolutionary origins in northern part of South America and its close genetic relation to '' Rhinella diptycha'' and other similar species of the genus. Recent studies suggest that ''R. marina'' diverged between 2.75 and 9.40 million years ago. A recent split in the species into further subspecies may have occurred approximately 2.7 million years ago following the isolation of population groups by the rising Venezuelan Andes.


Description

Considered the largest species in the Bufonidae, the cane toad is very large; the females are significantly longer than males, reaching a typical length of , with a maximum of . Larger toads tend to be found in areas of lower population density. They have a life expectancy of 10 to 15 years in the wild, and can live considerably longer in captivity, with one specimen reportedly surviving for 35 years. The skin of the cane toad is dry and warty. Distinct ridges above the eyes run down the snout. Individual cane toads can be grey, yellowish, red-brown, or olive-brown, with varying patterns. A large parotoid gland lies behind each eye. The ventral surface is cream-coloured and may have blotches in shades of black or brown. The pupils are horizontal and the irises golden. The toes have a fleshy webbing at their base, and the fingers are free of webbing. Typically, juvenile cane toads have smooth, dark skin, although some specimens have a red wash. Juveniles lack the adults' large parotoid glands, so they are usually less poisonous. The
tadpole A tadpole or polliwog (also spelled pollywog) is the Larva, larval stage in the biological life cycle of an amphibian. Most tadpoles are fully Aquatic animal, aquatic, though some species of amphibians have tadpoles that are terrestrial animal, ...
s are small and uniformly black, and are bottom-dwellers, tending to form
schools A school is the educational institution (and, in the case of in-person learning, the building) designed to provide learning environments for the teaching of students, usually under the direction of teachers. Most countries have systems of ...
. Tadpoles range from in length.


Ecology, behaviour and life history

The common name "marine toad" and the scientific name ''Rhinella marina'' suggest a link to
marine life Marine life, sea life or ocean life is the collective ecological communities that encompass all aquatic animals, aquatic plant, plants, algae, marine fungi, fungi, marine protists, protists, single-celled marine microorganisms, microorganisms ...
, but cane toads do not live in the sea. However, laboratory experiments suggest that tadpoles can tolerate salt concentrations equivalent to 15% of
seawater Seawater, or sea water, is water from a sea or ocean. On average, seawater in the world's oceans has a salinity of about 3.5% (35 g/L, 35 ppt, 600 mM). This means that every kilogram (roughly one liter by volume) of seawater has approximat ...
(~5.4‰), and recent field observations found living tadpoles and toadlets at salinities of 27.5‰ on Coiba Island,
Panama Panama, officially the Republic of Panama, is a country in Latin America at the southern end of Central America, bordering South America. It is bordered by Costa Rica to the west, Colombia to the southeast, the Caribbean Sea to the north, and ...
. The cane toad inhabits open grassland and woodland, and has displayed a "distinct preference" for areas modified by humans, such as gardens and drainage ditches. In their native habitats, the toads can be found in subtropical forests, although dense foliage tends to limit their dispersal. The cane toad begins life as an egg, which is laid as part of long strings of jelly in water. A female lays 8,000–25,000 eggs at once and the strings can stretch up to in length. The black eggs are covered by a membrane and their diameter is about . The rate at which an egg grows into a tadpole increases with temperature. Tadpoles typically hatch within 48 hours, but the period can vary from 14 hours to almost a week. This process usually involves thousands of tadpoles—which are small, black, and have short tails—forming into groups. Between 12 and 60 days are needed for the tadpoles to develop into juveniles, with four weeks being typical. Similarly to their adult counterparts, eggs and tadpoles are toxic to many animals. When they emerge, toadlets typically are about in length, and grow rapidly. While the rate of growth varies by region, time of year, and sex, an average initial growth rate of per day is seen, followed by an average rate of per day. Growth typically slows once the toads reach sexual maturity. This rapid growth is important for their survival; in the period between metamorphosis and subadulthood, the young toads lose the toxicity that protected them as eggs and tadpoles, but have yet to fully develop the parotoid glands that produce
bufotoxin Bufotoxins are a family of toxic steroid lactones or substituted Substituted tryptamine, tryptamines of which some are toxic. They occur in the parotoid glands, skin, and poison of many toads (True toad, Bufonidae family) and other amphibians, an ...
. Only an estimated 0.5% of cane toads reach adulthood, in part because they lack this key defense—but also due to tadpole cannibalism. Although cannibalism does occur in the native population in South America, the rapid evolution occurring in the unnaturally large population in Australia has produced tadpoles 30x more likely to be ''interested'' in cannibalising their siblings, and 2.6x more likely to actually ''do so''. They have also evolved to shorten their tadpole phase in response to the presence of older tadpoles. These changes are likely genetic, although no genetic basis has been determined. As with rates of growth, the point at which the toads become sexually mature varies across different regions. In New Guinea, sexual maturity is reached by female toads with a snout–vent length between , while toads in
Panama Panama, officially the Republic of Panama, is a country in Latin America at the southern end of Central America, bordering South America. It is bordered by Costa Rica to the west, Colombia to the southeast, the Caribbean Sea to the north, and ...
achieve maturity when they are between in length. In tropical regions, such as their native habitats, breeding occurs throughout the year, but in subtropical areas, breeding occurs only during warmer periods that coincide with the onset of the
wet season The wet season (sometimes called the rainy season or monsoon season) is the time of year when most of a region's average annual rainfall occurs. Generally, the season lasts at least one month. The term ''green season'' is also sometimes used a ...
. The cane toad is estimated to have a critical thermal maximum of and a minimum of around . The ranges can change due to adaptation to the local environment. Cane toads from some populations can adjust their thermal tolerance within a few hours of encountering low temperatures. The toad is able to rapidly acclimate to the cold using physiological plasticity, though there is also evidence that more northerly populations of cane toads in the United States are better cold-adapted than more southerly populations. These adaptations have allowed the cane toad to establish invasive populations across the world. The toad's ability to rapidly acclimate to thermal changes suggests that current models may underestimate the potential range of habitats that the toad can populate. The cane toad has a high tolerance to water loss; some can withstand a 52.6% loss of body water, allowing them to survive outside tropical environments.


Diet

Most frogs identify prey by movement, and vision appears to be the primary method by which the cane toad detects prey; however, it can also locate food using its sense of smell. They eat a wide range of material; in addition to the normal prey of small
rodent Rodents (from Latin , 'to gnaw') are mammals of the Order (biology), order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and Mandible, lower jaws. About 40% of all mammal specie ...
s, other small
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s,
reptile Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...
s, other
amphibian Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
s,
bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s, and even
bat Bats are flying mammals of the order Chiroptera (). With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most birds, flying with their very long spread-out ...
s and a range of
invertebrate Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...
s (such as
ant Ants are Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Cre ...
s,
beetle Beetles are insects that form the Taxonomic rank, order Coleoptera (), in the superorder Holometabola. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 40 ...
s,
earwig Earwigs make up the insect order (biology), order Dermaptera. With about 2,000 species in 12 families, they are one of the smaller insect orders. Earwigs have characteristic cercus, cerci, a pair of forceps-like pincer (biology), pincers on ...
s,
dragonflies A dragonfly is a flying insect belonging to the infraorder Anisoptera below the order Odonata. About 3,000 extant species of dragonflies are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat threate ...
,
grasshopper Grasshoppers are a group of insects belonging to the suborder Caelifera. They are amongst what are possibly the most ancient living groups of chewing herbivorous insects, dating back to the early Triassic around 250 million years ago. Grassh ...
s, true bugs,
crustacean Crustaceans (from Latin meaning: "those with shells" or "crusted ones") are invertebrate animals that constitute one group of arthropods that are traditionally a part of the subphylum Crustacea (), a large, diverse group of mainly aquatic arthrop ...
s, and
gastropod Gastropods (), commonly known as slugs and snails, belong to a large Taxonomy (biology), taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, freshwater, and fro ...
s), they also eat plants, dog food, cat food, feces, and household refuse.


Defences

The skin of the adult cane toad is toxic, as well as the enlarged parotoid glands behind the eyes, and other glands across its back. When the toad is threatened, its glands secrete a milky-white fluid known as
bufotoxin Bufotoxins are a family of toxic steroid lactones or substituted Substituted tryptamine, tryptamines of which some are toxic. They occur in the parotoid glands, skin, and poison of many toads (True toad, Bufonidae family) and other amphibians, an ...
. Components of bufotoxin are toxic to many animals; even human deaths have been recorded due to the consumption of cane toads. Dogs are especially prone to be poisoned by licking or biting toads. Pets showing excessive drooling, extremely red gums, head-shaking, crying, loss of coordination, and/or convulsions require immediate veterinary attention. Bufotenin, one of the chemicals excreted by the cane toad, is classified as a schedule 9 drug under Australian law, alongside
heroin Heroin, also known as diacetylmorphine and diamorphine among other names, is a morphinan opioid substance synthesized from the Opium, dried latex of the Papaver somniferum, opium poppy; it is mainly used as a recreational drug for its eupho ...
and LSD. The effects of bufotenin are thought to be similar to those of mild poisoning; the stimulation, which includes mild
hallucination A hallucination is a perception in the absence of an external stimulus that has the compelling sense of reality. They are distinguishable from several related phenomena, such as dreaming ( REM sleep), which does not involve wakefulness; pse ...
s, lasts less than an hour. As the cane toad excretes bufotenin in small amounts, and other toxins in relatively large quantities, toad licking could result in serious illness or death. In addition to releasing toxin, the cane toad is capable of inflating its lungs, puffing up, and lifting its body off the ground to appear taller and larger to a potential predator. Since 2011, experimenters in the Kimberley region of Western Australia have used poisonous sausages containing toad meat in an attempt to protect native animals from cane toads' deadly impact. The Western Australian Department of Environment and Conservation, along with the University of Sydney, developed these sausage-shaped baits as a tool in order to train native animals not to eat the toads. By blending bits of toad with a nausea-inducing chemical, the baits train the animals to stay away from the amphibians. Young cane toads that aren't lethal upon ingestion have also been used to teach native predators avoidance, namely yellow-spotted monitors. 200,000 metamorphs, tadpoles, and eggs in total were released in areas ahead of inevitable invasion fronts. Following invasion by wild cane toads, yellow-spotted monitors in control areas bereft of the "teacher toads" were virtually wiped out, but experimental areas still contained substantial populations of yellow-spotted monitors.


Predators

Many species prey on the cane toad and its tadpoles in its native habitat, including the broad-snouted caiman (''Caiman latirostris''), the banded cat-eyed snake ('' Leptodeira annulata''), eels (family Anguillidae), various species of killifish, and '' Paraponera clavata'' (bullet ants). Predators outside the cane toad's native range include the rock flagtail ('' Kuhlia rupestris''), some species of
catfish Catfish (or catfishes; order (biology), order Siluriformes or Nematognathi) are a diverse group of ray-finned fish. Catfish are common name, named for their prominent barbel (anatomy), barbels, which resemble a cat's whiskers, though not ...
(order Siluriformes), some species of
ibis The ibis () (collective plural ibises; classical plurals ibides and ibes) are a group of long-legged wading birds in the family Threskiornithidae that inhabit wetlands, forests and plains. "Ibis" derives from the Latin and Ancient Greek word f ...
(subfamily Threskiornithinae), the whistling kite (''Haliastur sphenurus''), the rakali (''Hydromys chrysogaster''), the
black rat The black rat (''Rattus rattus''), also known as the roof rat, ship rat, or house rat, is a common long-tailed rodent of the stereotypical rat genus ''Rattus'', in the subfamily Murinae. It likely originated in the Indian subcontinent, but is n ...
(''Rattus rattus'') and the water monitor (''Varanus salvator''). The tawny frogmouth (''Podargus strigoides'') and the Papuan frogmouth (''Podargus papuensis'') have been reported as feeding on cane toads; some Australian crows ('' Corvus'' spp.) have also learned strategies allowing them to feed on cane toads, such as using their beak to flip toads onto their backs.
Kookaburra Kookaburras (pronounced ) are terrestrial animal, terrestrial tree kingfishers of the genus ''Dacelo'' native to Australia and New Guinea, which grow to between in length and weigh around . The name is a loanword from Wiradjuri language, Wira ...
s also prey on the amphibians. Opossums of the genus '' Didelphis'' likely can eat cane toads with impunity.
Meat ant The meat ant (''Iridomyrmex purpureus''), also known as the gravel ant or southern meat ant, is a species of ant Endemism, endemic to Australia (island), Australia. A member of the genus ''Iridomyrmex'' in the subfamily Dolichoderinae, it was ...
s are unaffected by the cane toads' toxins, so are able to kill them without reaction. The cane toad's normal response to attack is to stand still and let its toxin kill or repel the attacker, which allows the ants to attack and eat the toad. Saw-shelled turtles have also been seen successfully and safely eating cane toads. In Australia rakali (Australian water rats) in two years learnt how to eat cane toads safely. They select the largest toads, turn them over, remove the poisonous gallbladder, and eat the heart and other organs with "surgical precision". They remove the toxic skin and eat the thigh muscle. Other animals such as crows and kites turn cane toads inside out and eat non-poisonous organs, also thus avoiding the skin.


Distribution

The cane toad is native to the Americas, and its range stretches from the Rio Grande Valley in South Texas to the central
Amazon Amazon most often refers to: * Amazon River, in South America * Amazon rainforest, a rainforest covering most of the Amazon basin * Amazon (company), an American multinational technology company * Amazons, a tribe of female warriors in Greek myth ...
and southeastern
Peru Peru, officially the Republic of Peru, is a country in western South America. It is bordered in the north by Ecuador and Colombia, in the east by Brazil, in the southeast by Bolivia, in the south by Chile, and in the south and west by the Pac ...
, and some of the continental islands near
Venezuela Venezuela, officially the Bolivarian Republic of Venezuela, is a country on the northern coast of South America, consisting of a continental landmass and many Federal Dependencies of Venezuela, islands and islets in the Caribbean Sea. It com ...
(such as
Trinidad Trinidad is the larger, more populous island of the Republic of Trinidad and Tobago, the country. The island lies off the northeastern coast of Venezuela and sits on the continental shelf of South America. It is the southernmost island in ...
and
Tobago Tobago, officially the Ward of Tobago, is an List of islands of Trinidad and Tobago, island and Regions and municipalities of Trinidad and Tobago, ward within the Republic of Trinidad and Tobago. It is located northeast of the larger islan ...
). This area encompasses both
tropical The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...
and semiarid environments. The density of the cane toad is significantly lower within its native distribution than in places where it has been introduced. In South America, the density was recorded to be 20 adults per of shoreline, 1 to 2% of the density in Australia.


As an introduced species

The cane toad has been introduced to many regions of the world—particularly the Pacific—for the biological control of agricultural pests. These introductions have generally been well documented, and the cane toad may be one of the most studied of any
introduced species An introduced species, alien species, exotic species, adventive species, immigrant species, foreign species, non-indigenous species, or non-native species is a species living outside its native distributional range, but which has arrived ther ...
. Before the early 1840s, the cane toad had been introduced into
Martinique Martinique ( ; or ; Kalinago language, Kalinago: or ) is an island in the Lesser Antilles of the West Indies, in the eastern Caribbean Sea. It was previously known as Iguanacaera which translates to iguana island in Carib language, Kariʼn ...
and
Barbados Barbados, officially the Republic of Barbados, is an island country in the Atlantic Ocean. It is part of the Lesser Antilles of the West Indies and the easternmost island of the Caribbean region. It lies on the boundary of the South American ...
, from
French Guiana French Guiana, or Guyane in French, is an Overseas departments and regions of France, overseas department and region of France located on the northern coast of South America in the Guianas and the West Indies. Bordered by Suriname to the west ...
and
Guyana Guyana, officially the Co-operative Republic of Guyana, is a country on the northern coast of South America, part of the historic British West Indies. entry "Guyana" Georgetown, Guyana, Georgetown is the capital of Guyana and is also the co ...
. An introduction to
Jamaica Jamaica is an island country in the Caribbean Sea and the West Indies. At , it is the third-largest island—after Cuba and Hispaniola—of the Greater Antilles and the Caribbean. Jamaica lies about south of Cuba, west of Hispaniola (the is ...
was made in 1844 in an attempt to reduce the rat population. Despite its failure to control the rodents, the cane toad was introduced to
Puerto Rico ; abbreviated PR), officially the Commonwealth of Puerto Rico, is a Government of Puerto Rico, self-governing Caribbean Geography of Puerto Rico, archipelago and island organized as an Territories of the United States, unincorporated territo ...
in the early 20th century in the hope that it would counter a beetle infestation ravaging the sugarcane plantations. The Puerto Rican scheme was successful and halted the economic damage caused by the beetles, prompting scientists in the 1930s to promote it as an ideal solution to agricultural pests. As a result, many countries in the Pacific region emulated the lead of Puerto Rico and introduced the toad in the 1930s. Introduced populations are in
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
,
Florida Florida ( ; ) is a U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders the Gulf of Mexico to the west, Alabama to the northwest, Georgia (U.S. state), Georgia to the north, the Atlantic ...
,
Papua New Guinea Papua New Guinea, officially the Independent State of Papua New Guinea, is an island country in Oceania that comprises the eastern half of the island of New Guinea and offshore islands in Melanesia, a region of the southwestern Pacific Ocean n ...
, the
Philippines The Philippines, officially the Republic of the Philippines, is an Archipelagic state, archipelagic country in Southeast Asia. Located in the western Pacific Ocean, it consists of List of islands of the Philippines, 7,641 islands, with a tot ...
, the Ogasawara, Ishigaki Island and the
Daitō Islands The are an archipelago consisting of three isolated coral islands, administered by Japan, in the Philippine Sea southeast of Okinawa Prefecture, Okinawa. The islands have a total area of and a population of 2,107. Administratively, the whole gr ...
of Japan,
Taiwan Taiwan, officially the Republic of China (ROC), is a country in East Asia. The main geography of Taiwan, island of Taiwan, also known as ''Formosa'', lies between the East China Sea, East and South China Seas in the northwestern Pacific Ocea ...
Nantou Caotun, most Caribbean islands,
Fiji Fiji, officially the Republic of Fiji, is an island country in Melanesia, part of Oceania in the South Pacific Ocean. It lies about north-northeast of New Zealand. Fiji consists of an archipelago of more than 330 islands—of which about ...
and many other Pacific islands, including
Hawaii Hawaii ( ; ) is an island U.S. state, state of the United States, in the Pacific Ocean about southwest of the U.S. mainland. One of the two Non-contiguous United States, non-contiguous U.S. states (along with Alaska), it is the only sta ...
. Since then, the cane toad has become a pest in many host countries, and poses a serious threat to native animals.


Australia

Following the apparent success of the cane toad in eating the beetles threatening the sugarcane plantations of Puerto Rico, and the fruitful introductions into Hawaiʻi and the Philippines, a strong push was made for the cane toad to be released in Australia to negate the pests ravaging the Queensland cane fields. As a result, 102 toads were collected from Hawaiʻi and brought to Australia. Queensland's sugar scientists released the toad into cane fields in August 1935. After this initial release, the Commonwealth Department of Health decided to ban future introductions until a study was conducted into the feeding habits of the toad. The study was completed in 1936 and the ban lifted, when large-scale releases were undertaken; by March 1937, 62,000 toadlets had been released into the wild. The toads became firmly established in Queensland, increasing exponentially in number and extending their range into the
Northern Territory The Northern Territory (abbreviated as NT; known formally as the Northern Territory of Australia and informally as the Territory) is an states and territories of Australia, Australian internal territory in the central and central-northern regi ...
and
New South Wales New South Wales (commonly abbreviated as NSW) is a States and territories of Australia, state on the Eastern states of Australia, east coast of :Australia. It borders Queensland to the north, Victoria (state), Victoria to the south, and South ...
. In 2010, one was found on the far western coast in
Broome, Western Australia Broome, also known as Rubibi by the Yawuru, Yawuru people, is a coastal Pearl hunting, pearling and tourist town in the Kimberley (Western Australia), Kimberley region of Western Australia, north of Perth. The town recorded a population of 14,6 ...
. However, the toad was generally unsuccessful in reducing the targeted grey-backed cane beetles ('' Dermolepida albohirtum''), in part because the cane fields provided insufficient shelter for the predators during the day, and in part because the beetles live at the tops of sugar cane—and cane toads are not good climbers. Since its original introduction, the cane toad has had a particularly marked effect on Australian
biodiversity Biodiversity is the variability of life, life on Earth. It can be measured on various levels. There is for example genetic variability, species diversity, ecosystem diversity and Phylogenetics, phylogenetic diversity. Diversity is not distribut ...
. The population of a number of native predatory reptiles has declined, such as the varanid lizards '' Varanus mertensi'', '' V. mitchelli'', and '' V. panoptes'', the land snakes '' Pseudechis australis'' and '' Acanthophis antarcticus'', and the freshwater crocodile species '' Crocodylus johnstoni''; in contrast, the population of the agamid lizard '' Amphibolurus gilberti''—known to be a prey item of ''V. panoptes''—has increased.. On snake populations see .
Meat ant The meat ant (''Iridomyrmex purpureus''), also known as the gravel ant or southern meat ant, is a species of ant Endemism, endemic to Australia (island), Australia. A member of the genus ''Iridomyrmex'' in the subfamily Dolichoderinae, it was ...
s, however, are able to kill cane toads.''Meat Ant.''
Australian Environmental Pest Managers Association (AEPMA) (accessed July 2022)
The cane toad has also been linked to decreases in northern quolls in the southern region of Kakadu National Park and even their
local extinction Local extinction, also extirpation, is the termination of a species (or other taxon) in a chosen geographic area of study, though it still exists elsewhere. Local extinctions are contrasted with extinction, global extinctions. Local extinctions ...
.


Caribbean

The cane toad was introduced to various Caribbean islands to counter a number of pests infesting local crops. While it was able to establish itself on some islands, such as
Barbados Barbados, officially the Republic of Barbados, is an island country in the Atlantic Ocean. It is part of the Lesser Antilles of the West Indies and the easternmost island of the Caribbean region. It lies on the boundary of the South American ...
,
Jamaica Jamaica is an island country in the Caribbean Sea and the West Indies. At , it is the third-largest island—after Cuba and Hispaniola—of the Greater Antilles and the Caribbean. Jamaica lies about south of Cuba, west of Hispaniola (the is ...
,
Hispaniola Hispaniola (, also ) is an island between Geography of Cuba, Cuba and Geography of Puerto Rico, Puerto Rico in the Greater Antilles of the Caribbean. Hispaniola is the most populous island in the West Indies, and the second-largest by List of C ...
and
Puerto Rico ; abbreviated PR), officially the Commonwealth of Puerto Rico, is a Government of Puerto Rico, self-governing Caribbean Geography of Puerto Rico, archipelago and island organized as an Territories of the United States, unincorporated territo ...
, other introductions, such as in
Cuba Cuba, officially the Republic of Cuba, is an island country, comprising the island of Cuba (largest island), Isla de la Juventud, and List of islands of Cuba, 4,195 islands, islets and cays surrounding the main island. It is located where the ...
before 1900 and in 1946, and on the islands of
Dominica Dominica, officially the Commonwealth of Dominica, is an island country in the Caribbean. It is part of the Windward Islands chain in the Lesser Antilles archipelago in the Caribbean Sea. The capital, Roseau, is located on the western side of t ...
and
Grand Cayman Grand Cayman is the largest of the three Cayman Islands and the location of the territory's capital, George Town, Cayman Islands, George Town. In relation to the other two Cayman Islands, it is approximately 75 miles (121 km) southwest of L ...
, were unsuccessful. The earliest recorded introductions were to Barbados and
Martinique Martinique ( ; or ; Kalinago language, Kalinago: or ) is an island in the Lesser Antilles of the West Indies, in the eastern Caribbean Sea. It was previously known as Iguanacaera which translates to iguana island in Carib language, Kariʼn ...
. The Barbados introductions were focused on the biological control of pests damaging the sugarcane crops, and while the toads became abundant, they have done even less to control the pests than in Australia.Kennedy, Anthony quoted in The toad was introduced to Martinique from
French Guiana French Guiana, or Guyane in French, is an Overseas departments and regions of France, overseas department and region of France located on the northern coast of South America in the Guianas and the West Indies. Bordered by Suriname to the west ...
before 1944 and became established. Today, they reduce the
mosquito Mosquitoes, the Culicidae, are a Family (biology), family of small Diptera, flies consisting of 3,600 species. The word ''mosquito'' (formed by ''Musca (fly), mosca'' and diminutive ''-ito'') is Spanish and Portuguese for ''little fly''. Mos ...
and
mole cricket Mole (or Molé) may refer to: Animals * Mole (animal) or "true mole" * Golden mole, southern African mammals * Marsupial mole, Australian mammals Other common meanings * Nevus, a growth on human skin ** Melanocytic nevus, a specific type of ...
populations. A third introduction to the region occurred in 1884, when toads appeared in Jamaica, reportedly imported from Barbados to help control the rodent population. While they had no significant effect on the rats, they nevertheless became well established. Other introductions include the release on
Antigua Antigua ( ; ), also known as Waladli or Wadadli by the local population, is an island in the Lesser Antilles. It is one of the Leeward Islands in the Caribbean region and the most populous island of the country of Antigua and Barbuda. Antigua ...
—possibly before 1916, although this initial population may have died out by 1934 and been reintroduced at a later date—and
Montserrat Montserrat ( , ) is a British Overseas Territories, British Overseas Territory in the Caribbean. It is part of the Leeward Islands, the northern portion of the Lesser Antilles chain of the West Indies. Montserrat is about long and wide, wit ...
, which had an introduction before 1879 that led to the establishment of a solid population, which was apparently sufficient to survive the Soufrière Hills volcano eruption in 1995. In 1920, the cane toad was introduced into Puerto Rico to control the populations of white grub (''Phyllophaga'' spp.), a sugarcane pest. Before this, the pests were manually collected by humans, so the introduction of the toad eliminated labor costs. A second group of toads was imported in 1923, and by 1932, the cane toad was well established.. "After a completely successful method of killing white grubs by chemical means was found, the only opportunities for its use in Puerto Rico have been limited to small areas in pineapple plantations at elevations where the toad is even yet not present in sufficient abundance." The population of white grubs dramatically decreased, and this was attributed to the cane toad at the annual meeting of the International Sugar Cane Technologists in Puerto Rico. However, there may have been other factors. The six-year period after 1931—when the cane toad was most prolific, and the white grub had a dramatic decline—had the highest-ever rainfall for Puerto Rico. Nevertheless, the cane toad was assumed to have controlled the white grub; this view was reinforced by a ''
Nature Nature is an inherent character or constitution, particularly of the Ecosphere (planetary), ecosphere or the universe as a whole. In this general sense nature refers to the Scientific law, laws, elements and phenomenon, phenomena of the physic ...
'' article titled "Toads save sugar crop", and this led to large-scale introductions throughout many parts of the Pacific. The cane toad has been spotted in Carriacou and
Dominica Dominica, officially the Commonwealth of Dominica, is an island country in the Caribbean. It is part of the Windward Islands chain in the Lesser Antilles archipelago in the Caribbean Sea. The capital, Roseau, is located on the western side of t ...
, the latter appearance occurring in spite of the failure of the earlier introductions. On September 8, 2013, the cane toad was also discovered on the island of
New Providence New Providence is the most populous island in The Bahamas, containing more than 70% of the total population. On the eastern side of the island is the national capital, national capital city of Nassau, Bahamas, Nassau; it had a population of 246 ...
in the Bahamas.


The Philippines

The cane toad was first introduced deliberately into the
Philippines The Philippines, officially the Republic of the Philippines, is an Archipelagic state, archipelagic country in Southeast Asia. Located in the western Pacific Ocean, it consists of List of islands of the Philippines, 7,641 islands, with a tot ...
in 1930 as a biological control agent of pests in sugarcane plantations, after the success of the experimental introductions into Puerto Rico. It subsequently became the most ubiquitous amphibian in the islands. It still retains the common name of ''bakî'' or ''kamprag'' in the Visayan languages, a
corruption Corruption is a form of dishonesty or a criminal offense that is undertaken by a person or an organization that is entrusted in a position of authority to acquire illicit benefits or abuse power for one's gain. Corruption may involve activities ...
of 'American frog', referring to its origins. It is also commonly known as "bullfrog" in Philippine English.


Fiji

The cane toad was introduced into
Fiji Fiji, officially the Republic of Fiji, is an island country in Melanesia, part of Oceania in the South Pacific Ocean. It lies about north-northeast of New Zealand. Fiji consists of an archipelago of more than 330 islands—of which about ...
to combat insects that infested sugarcane plantations. The introduction of the cane toad to the region was first suggested in 1933, following the successes in Puerto Rico and Hawaiʻi. After considering the possible side effects, the national government of Fiji decided to release the toad in 1953, and 67 specimens were subsequently imported from Hawaiʻi. Once the toads were established, a 1963 study concluded, as the toad's diet included both harmful and beneficial invertebrates, it was considered "economically neutral". Today, the cane toad can be found on all major islands in Fiji, although they tend to be smaller than their counterparts in other regions.


New Guinea

The cane toad was introduced into
New Guinea New Guinea (; Hiri Motu: ''Niu Gini''; , fossilized , also known as Papua or historically ) is the List of islands by area, world's second-largest island, with an area of . Located in Melanesia in the southwestern Pacific Ocean, the island is ...
to control the hawk moth larvae eating
sweet potato The sweet potato or sweetpotato (''Ipomoea batatas'') is a dicotyledonous plant in the morning glory family, Convolvulaceae. Its sizeable, starchy, sweet-tasting tuberous roots are used as a root vegetable, which is a staple food in parts of ...
crops. The first release occurred in 1937 using toads imported from Hawaiʻi, with a second release the same year using specimens from the Australian mainland. Evidence suggests a third release in 1938, consisting of toads being used for human pregnancy tests—many species of toad were found to be effective for this task, and were employed for about 20 years after the discovery was announced in 1948. Initial reports argued the toads were effective in reducing the levels of
cutworm Cutworms are moth larvae that hide under litter or soil during the day, coming out in the dark to feed on plants. A larva typically attacks the first part of the plant it encounters, namely the stem, often of a seedling, and consequently cuts it ...
s and sweet potato yields were thought to be improving. As a result, these first releases were followed by further distributions across much of the region, although their effectiveness on other crops, such as cabbages, has been questioned; when the toads were released at Wau, the cabbages provided insufficient shelter and the toads rapidly left the immediate area for the superior shelter offered by the forest. A similar situation had previously arisen in the Australian cane fields, but this experience was either unknown or ignored in New Guinea. The cane toad has since become abundant in rural and urban areas.


United States

The cane toad naturally exists in South Texas, but attempts (both deliberate and accidental) have been made to introduce the species to other parts of the country. These include introductions to
Florida Florida ( ; ) is a U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders the Gulf of Mexico to the west, Alabama to the northwest, Georgia (U.S. state), Georgia to the north, the Atlantic ...
and to Hawaiʻi, as well as largely unsuccessful introductions to
Louisiana Louisiana ( ; ; ) is a state in the Deep South and South Central regions of the United States. It borders Texas to the west, Arkansas to the north, and Mississippi to the east. Of the 50 U.S. states, it ranks 31st in area and 25 ...
. Initial releases into Florida failed. Attempted introductions before 1936 and 1944, intended to control sugarcane pests, were unsuccessful as the toads failed to proliferate. Later attempts failed in the same way. However, the toad gained a foothold in the state after an accidental release by an importer at
Miami International Airport Miami International Airport , also known as MIA and historically as Wilcox Field, is the primary international airport serving Miami and its Miami metropolitan area, surrounding metropolitan area, in the U.S. state of Florida. It hosts over 1, ...
in 1957, and deliberate releases by animal dealers in 1963 and 1964 established the toad in other parts of Florida. Today, the cane toad is well established in the state, from the Keys to north of
Tampa Tampa ( ) is a city on the Gulf Coast of the United States, Gulf Coast of the U.S. state of Florida. Tampa's borders include the north shore of Tampa Bay and the east shore of Old Tampa Bay. Tampa is the largest city in the Tampa Bay area and t ...
, and they are gradually extending further northward. In Florida, the toad is a regarded as a threat to native species and pets; so much so, the Florida Fish and Wildlife Conservation Commission recommends residents to kill them. Around 150 cane toads were introduced to Oʻahu in Hawaiʻi in 1932, and the population swelled to 105,517 after 17 months. The toads were sent to the other islands, and more than 100,000 toads were distributed by July 1934; eventually over 600,000 were transported.


Uses

Other than the use as a biological control for pests, the cane toad has been employed in a number of commercial and noncommercial applications. Traditionally, within the toad's natural range in South America, the Embera-Wounaan would "milk" the toads for their toxin, which was then employed as an arrow poison. The toxins may have been used as an
entheogen Entheogens are psychoactive substances used in spiritual and religious contexts to induce altered states of consciousness. Hallucinogens such as the psilocybin found in so-called "magic" mushrooms have been used in sacred contexts since ancie ...
by the
Olmec The Olmecs () or Olmec were an early known major Mesoamerican civilization, flourishing in the modern-day Mexican states of Veracruz and Tabasco from roughly 1200 to 400 Before the Common Era, BCE during Mesoamerica's Mesoamerican chronolog ...
people. The toad has been hunted as a food source in parts of
Peru Peru, officially the Republic of Peru, is a country in western South America. It is bordered in the north by Ecuador and Colombia, in the east by Brazil, in the southeast by Bolivia, in the south by Chile, and in the south and west by the Pac ...
, and eaten after the careful removal of the skin and
parotoid gland The parotoid gland (alternatively, paratoid gland) is an external skin gland on the back, neck, and shoulder of some frogs (especially toads), and salamanders. It can secrete a number of milky alkaloid substances (depending on the species) known ...
s. When properly prepared, the meat of the toad is considered healthy and as a source of
omega-3 fatty acid Omega−3 fatty acids, also called omega−3 oils, ω−3 fatty acids or ''n''−3 fatty acids, are polyunsaturated fatty acids (PUFAs) characterized by the presence of a double bond three atoms away from the terminal methyl group in their ...
s. More recently, the toad's toxins have been used in a number of new ways: bufotenin has been used in Japan as an
aphrodisiac An aphrodisiac is a substance that increases libido, sexual desire, sexual attraction, sexual pleasure, or sexual behavior. These substances range from a variety of plants, spices, and foods to synthetic chemicals. Natural aphrodisiacs, such as ...
and a hair restorer, and in cardiac surgery in China to lower the heart rates of patients. New research has suggested that the cane toad's poison may have some applications in treating
prostate cancer Prostate cancer is the neoplasm, uncontrolled growth of cells in the prostate, a gland in the male reproductive system below the bladder. Abnormal growth of the prostate tissue is usually detected through Screening (medicine), screening tests, ...
. Other modern applications of the cane toad include pregnancy testing, as pets, laboratory research, and the production of
leather Leather is a strong, flexible and durable material obtained from the tanning (leather), tanning, or chemical treatment, of animal skins and hides to prevent decay. The most common leathers come from cattle, sheep, goats, equine animals, buffal ...
goods. Pregnancy testing was conducted in the mid-20th century by injecting urine from a woman into a male toad's lymph sacs, and if
spermatozoa A spermatozoon (; also spelled spermatozoön; : spermatozoa; ) is a motile sperm cell (biology), cell produced by male animals relying on internal fertilization. A spermatozoon is a moving form of the ploidy, haploid cell (biology), cell that is ...
appeared in the toad's urine, the patient was deemed to be pregnant. The tests using toads were faster than those employing mammals; the toads were easier to raise, and, although the initial 1948 discovery employed ''
Bufo arenarum ''Rhinella arenarum'' is a species of toad in the family True toad, Bufonidae that is found in southern Brazil, Uruguay, Paraguay, Argentina (Chubut Province northward) and Bolivia (east of the Andes). ''Rhinella arenarum'' inhabits small ponds o ...
'' for the tests, it soon became clear that a variety of anuran species were suitable, including the cane toad. As a result, toads were employed in this task for around 20 years. As a laboratory animal, the cane toad has numerous advantages: they are plentiful, and easy and inexpensive to maintain and handle. The use of the cane toad in experiments started in the 1950s, and by the end of the 1960s, large numbers were being collected and exported to high schools and universities. Since then, a number of Australian states have introduced or tightened importation regulations. There are several commercial uses for dead cane toads. Cane toad skin is made into leather and novelty items. Stuffed cane toads, posed and accessorised, are merchandised at souvenir shops for tourists. Attempts have been made to produce
fertiliser A fertilizer or fertiliser is any material of natural or synthetic origin that is applied to soil or to plant tissues to supply plant nutrition, plant nutrients. Fertilizers may be distinct from Liming (soil), liming materials or other non- ...
from toad carcasses.


References


Citations


Bibliography

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External links


Species Profile – Cane Toad (''Rhinella marina'')
National Invasive Species Information Center,
United States National Agricultural Library The United States National Agricultural Library (NAL) is one of the world's largest agricultural research libraries, and serves as a national library of the United States and as the library of the United States Department of Agriculture. Locate ...
. Lists general information and resources for cane toad. {{Authority control Rhinella Taxa named by Carl Linnaeus Agricultural pests Amphibians described in 1758 Amphibians of Central America Amphibians of the Dominican Republic Amphibians of Guyana Amphibians of Japan Amphibians of Mauritius Amphibians of New South Wales Amphibians of the Northern Territory Amphibians of Queensland Amphibians of Trinidad and Tobago Fauna of Barbados Fauna of the Rio Grande valleys Frogs of Australia Frogs of Brazil