Common toad: Difference between revisions

From Wikipedia, the free encyclopedia
Content deleted Content added
→‎Development and growth: Added information on lungworms.
→‎Development and growth: Fertiliser information.
Line 100: Line 100:


===Development and growth===
===Development and growth===
The common toad reaches [[Sexual maturity|maturity]] at three to seven years old but there is great variability between populations.<ref name=Collins/> Juveniles are often parasitised by the lung nematode ''[[Rhabdias bufonis]]''. This slows growth rates and reduces stamina and fitness. Larger juveniles at metamorphosis always outgrow smaller ones that have been reared in more crowded ponds. Even when they have heavy worm burdens, large juveniles grow faster than smaller individuals with light burdens.<ref>{{cite journal |author=Goater, Cameron P.; Semlitsch, Raymond D.; Bernasconi, Marco V. |year=1993 |title=Effects of body size and parasite infection on the locomotory performance of juvenile toads, ''Bufo bufo'' |journal=Oikos |volume=16 |issue=1 |pages=129&#8209;136 |jstor=3545205 }}</ref> After several months of heavy worm infection, some juveniles in a study were only half the mass of [[Scientific control|controls]]. Their parasite-induced anorexia caused a decrease in dietary intake and some died.<ref>{{cite journal |author=Goater, C. P.; Ward, P. I. |year=1992 |title=Negative effects of ''Rhabdias bufonis'' (Nematoda) on the growth and survival of toads (''Bufo bufo'') |journal=Oecologia |volume=89 |issue=2 |pages=161&#8209;165 |jstor=4219866 }}</ref>
The common toad reaches [[Sexual maturity|maturity]] at three to seven years old but there is great variability between populations.<ref name=Collins/> Juveniles are often parasitised by the lung nematode ''[[Rhabdias bufonis]]''. This slows growth rates and reduces stamina and fitness. Larger juveniles at metamorphosis always outgrow smaller ones that have been reared in more crowded ponds. Even when they have heavy worm burdens, large juveniles grow faster than smaller individuals with light burdens.<ref>{{cite journal |author=Goater, Cameron P.; Semlitsch, Raymond D.; Bernasconi, Marco V. |year=1993 |title=Effects of body size and parasite infection on the locomotory performance of juvenile toads, ''Bufo bufo'' |journal=Oikos |volume=16 |issue=1 |pages=129&#8209;136 |jstor=3545205 }}</ref> After several months of heavy worm infection, some juveniles in a study were only half the mass of [[Scientific control|controls]]. Their parasite-induced anorexia caused a decrease in dietary intake and some died.<ref>{{cite journal |author=Goater, C. P.; Ward, P. I. |year=1992 |title=Negative effects of ''Rhabdias bufonis'' (Nematoda) on the growth and survival of toads (''Bufo bufo'') |journal=Oecologia |volume=89 |issue=2 |pages=161&#8209;165 |jstor=4219866 }}</ref> Another study investigated whether the use of nitrogenous fertilisers affects the development of common toad tadpoles. Different strength solutions of [[ammonium nitrate]] were used and it was found that at certain concentrations, well above normal field limits, growth was increased and metamorphosis accelerated, but at others, there was no significant difference between the experimental larvae and controls. Nevertheless, certain unusual swimming patterns and a few deformities were found among the experimental animals.<ref>{{cite journal |author=Xu, Q.; Oldham, R. S. |year=1997 |title=Lethal and sublethal effects of nitrogen fertilizer ammonium nitrate on common toad (''Bufo bufo'') tadpoles |journal=Archives of Environmental Contamination and Toxicology |volume=32 |issue=3 |pages=298&#8209;303 |doi=10.1007/s002449900188 }}</ref>


A comparison was made between the growth rate of newly [[Metamorphosis|metamorphosed]] juveniles from different altitudes and latitudes, the specimens studied being from Norway, Germany, Switzerland, the Netherlands and France. All were subspecies ''B. b. bufo'' except for the French specimens which may have been ''B. b. spinosus''. At first the growth rates for males and females was identical. By the time they became mature their growth rate had slowed down to about 21% of the initial rate and they had reached 95% of their expected adult size. Some females that were on a biennial breeding cycle carried on growing rapidly for a longer time. Adjusting for differences in temperature and the length of the growing season, the toads grew and matured at much the same rate from the four colder localities. These juveniles reached maturity after 1.09 years for males and 1.55 years for females. However, the young toads from lowland France grew faster and longer to a much greater size taking an average 1.77 years for males and 2.49 years for females before reaching maturity.<ref>{{cite journal |author=Hemelaar, Agnes |year=1988 |title=Age, Growth and Other Population Characteristics of ''Bufo bufo'' from Different Latitudes and Altitudes |journal=Journal of Herpetology |volume=22 |issue=4 |pages=369&#8209;388 |jstor=1564332 }}</ref>
A comparison was made between the growth rate of newly [[Metamorphosis|metamorphosed]] juveniles from different altitudes and latitudes, the specimens studied being from Norway, Germany, Switzerland, the Netherlands and France. All were subspecies ''B. b. bufo'' except for the French specimens which may have been ''B. b. spinosus''. At first the growth rates for males and females was identical. By the time they became mature their growth rate had slowed down to about 21% of the initial rate and they had reached 95% of their expected adult size. Some females that were on a biennial breeding cycle carried on growing rapidly for a longer time. Adjusting for differences in temperature and the length of the growing season, the toads grew and matured at much the same rate from the four colder localities. These juveniles reached maturity after 1.09 years for males and 1.55 years for females. However, the young toads from lowland France grew faster and longer to a much greater size taking an average 1.77 years for males and 2.49 years for females before reaching maturity.<ref>{{cite journal |author=Hemelaar, Agnes |year=1988 |title=Age, Growth and Other Population Characteristics of ''Bufo bufo'' from Different Latitudes and Altitudes |journal=Journal of Herpetology |volume=22 |issue=4 |pages=369&#8209;388 |jstor=1564332 }}</ref>

Revision as of 13:31, 23 August 2012

Common toad
A common toad
Scientific classification
Kingdom:
Phylum:
Class:
Order:
Family:
Genus:
Species:
B. bufo
Binomial name
Bufo bufo
Range map of common toad
Synonyms[2]
  • Bufo alpinus
  • Bufo carbunculus
  • Bufo communis
  • Bufo minutus
  • Bufo palmarum
  • Bufo spinosus
  • Bufo ventricosus
  • Bufo vinearum
  • Bufo vulgaris
  • Rana bufo
  • Rana pluvialis
  • Rana ventricosa
  • Rana vulgaris

The common toad or European toad (Bufo bufo) is an amphibian that is found throughout most of Europe, with the exception of Ireland, Iceland and some Mediterranean islands. It is one of a group of closely related taxa that are descended from a common ancestral line of toads and which form a species complex. There are several subspecies, the ranges of which intergrade where the populations meet. The toad is an inconspicuous animal as it usually lies hidden during the day. It becomes active at dusk and spends the night hunting for the invertebrates on which it feeds. It moves with a slow ungainly walk or a short jump and has a greyish brown skin covered with wart-like lumps.

Although usually a solitary animal, in the breeding season large numbers of toads converge on certain breeding ponds. Here there is great competition among the males for females with which to mate. The eggs are laid in gelatinous strings in the water and hatch out into tadpoles. After several months of growth and development, these sprout limbs and undergo metamorphosis into tiny toads. These emerge from the water and remain largely terrestrial for the rest of their lives.

The common toad has long been associated in popular culture with witchcraft, a fact recorded in literature. Touching a toad was at one time thought by superstitious people to be the cause of warts on human skin.

Taxonomy

The common toad was first given the binomial name Rana bufo by the Swedish biologist Carl Linnaeus in the 10th edition of Systema Naturae in 1758.[3] In this work, he placed all the frogs and toads in the single genus Rana. It later became apparent that this genus should be subdivided and in 1768, the Austrian naturalist Josephus Nicolaus Laurenti placed the common toad in the genus Bufo, naming it Bufo bufo. The toads in this genus are known as true toads and are characterised by having no teeth, a dry warty skin and horizontal pupils to the eyes.[4]

Bufo bufo is now considered to be a species complex, a group of closely related species the demarcation between which is unclear. It is believed that the modern subspecies are descended from a common, preglacial ancestral form. These subspecies include B. b. spinosus, B. b. gredosicola, B. b. verrucosissima and B. b. japonicus. The subspecies B. b. bufo seems to have arisen more recently.[5] It is believed that the range of the ancestral form extended into Asia but that isolation between an eastern and western race occurred as a result of greater aridity and desertification in the Middle East during the Middle Miocene.[6] The exact systematic relationships between the subspecies remains unclear.[5] Some authorities consider B. verrucosissimus is a species in its own right while others support it being a subspecies of the common toad, B. bufo verrucosissimus. Similarly, there is confusion as to whether B. spinosus should be recognised as a species or as a subspecies.[7]

A study published in 2012 cast new light on this problem. It examined the phylogenetic relationships between the Eurasian and north African species in the B. bufo group and indicated a long evolutionary history for the group. The initial split saw Bufo eichwaldi, a recently described species from south Azerbaijan and Iran, split from the main lineage. This took place nine to thirteen million years ago. Further divisions occurred with B. spinosus splitting off about five million years ago at the time that the Pyrenees were being uplifted, an event which isolated the populations in the Iberian Peninsula from those in the rest of Europe. The remaining European lineage split into B. bufo and B. verrucosissimus less than three million years ago during the Pleistocene.[8]

Subspecies

Subspecies Bufo bufo spinosus
Subspecies Bufo bufo spinosus

Bufo bufo bufo (Poche, 1912) is the nominate species and is found in northern Europe.[9]

Bufo bufo spinosus (Mertens, 1925) is a subspecies found in the Mediterranean area. It grows to a larger size and has spinier skin than its more northern counterpart. There is no hard and fast line demarcating the edge of its range, instead there is a gradual grading of one subspecies into the other.[9]

Bufo bufo gredosicola (Müller and Hellmich, 1935) is restricted to the Sierra de Gredos, a mountain range in central Spain. It has exceptionally large paratoid glands and tends to be less uniform in colour than the other subspecies.[9]

Bufo bufo verrucosissima (Pallas, 1814; Caucasian toad), also called Bufo colchicus and Rana caucasica .[10] It is found in the mountainous regions of the Caucasus. It has a larger genome and differs from the other subspecies morphologically.[5]

Very occasionally the common toad hybridizes with the natterjack toad, (Bufo calamita) or the European green toad (Bufo viridis).[9]

Description

Common toad walking
Common toad walking

The common toad can grow to about 15 cm (6 in) in length. Females grow larger than males and southern specimens tend to be larger than northern ones. The head is broad with a wide mouth below the terminal snout which has two small nostrils. There are no teeth. The large, protruding eyes have yellow or copper coloured irises and horizontal slit-shaped pupils. Just behind the eyes are two bulging regions, the paratoid glands, which are positioned obliquely. They contain a noxious substance, bufotoxin, which is used to deter potential predators. The head joins the body without a noticeable neck and there is no external vocal sac. The body is broad and squat and positioned close to the ground. The fore limbs are short with the toes of the fore feet turning inwards. At breeding time, the male develops nuptial pads on the first three fingers with which to grasp the female. The hind legs are short relative to a frog's legs and the hind feet have long, unwebbed toes. There is no tail. The skin is dry and covered with small warts. The colour is a fairly uniform shade of brown, olive-brown or greyish-brown, sometimes partly blotched or banded with a darker shade. The common toad tends to be sexually dimorphic with the females being browner and the males greyer. [7] The underside is a dirty white speckled with grey and black patches.[9][11]

Other species with which the common toad could be confused include the natterjack toad (Bufo calamita) and the European green toad (Bufo viridis). The former is usually smaller and has a yellow band running down its back while the latter has a distinctive mottled pattern. The paratoid glands of both are parallel rather than slanting as in the common toad.[9] It also resembles the common frog (Rana temporaria) but that species has a less rounded snout, a more angular shape, a damp wart-free skin and usually moves in leaps.[12]

Common toads can live for many years and have survived for fifty years in captivity.[13] In the wild, common toads are thought to live for about ten to twelve years. Their age can be determined by counting the number of annual growth rings in the bones of their phalanges.[14]

Distribution and habitat

After the common frog (Rana temporaria), the edible frog (Pelophylax kl. esculentus) and the smooth newt (Lissotriton vulgaris), the common toad is the fourth commonest amphibian in Europe.[9] It is found throughout the continent with the exception of Iceland, the cold northern parts of Scandinavia, Ireland and a number of Mediterranean islands. These include Malta, Crete, Corsica, Sardinia and the Balearic Islands. Its easterly range extends to Irkutsk in Siberia and its southerly range includes parts of northwestern Africa in the northern mountain ranges of Morocco, Algeria and Tunisia. A closely related variant lives in eastern Asia including Japan.[9] The common toad is found at altitudes of up to 2,500 metres (8,200 ft) in the southern part of its range. It inhabits open countryside, fields, the edges of woods, parks and gardens, often in dry areas well away from standing water.[9]

Behaviour

A common toad adopting a defensive stance
A common toad adopting a defensive stance
Common toad infected with larvae of Lucilia bufonivora
A common toad infected with larvae of Lucilia bufonivora. The nostrils of the toad have already been destroyed by the larvae.

The common toad usually moves by walking rather slowly or in short shuffling jumps involving all four legs. It spends the day concealed in a lair that it has hollowed out under foliage or beneath a root or a stone where its colouring makes it inconspicuous. It emerges at dusk and may travel some distance in the dark while hunting. It is most active in wet weather. By morning it has returned to its base and may occupy the same place for several months. It has a large appetite and eats woodlice, slugs, beetles, caterpillars, flies, worms and even small mice.[15][16] Small, fast moving prey may be caught by a flick of the tongue while larger items are grabbed with the jaws. Having no teeth, it swallows food whole in a series of gulps.[15] It does not recognise its prey as such but will try to consume any small, dark coloured, moving object it encounters at night. A research study showed that it would snap at a moving 1 cm (0.4 in) piece of black paper as if it were prey but would disregard a larger moving piece.[17] Toads seem to use visual cues for feeding and can see their prey at very low light intensities where humans are unable to discern anything.[18] Periodically, the common toad sheds its skin which comes away in tattered pieces. The sloughed skin is eaten.[15]

Common toads in amplexus
Common toads in amplexus

When attacked, the common toad adopts a characteristic stance, inflating its body and standing with its hindquarters raised and its head lowered. Its chief means of defence lies in the foul tasting secretion that is produced by its paratoid glands and the warts on its skin. This contains a bufotoxin called bufagin and is enough to deter many predators although grass snakes seem to be unaffected by it.[9] Other predators of adult toads include hedgehogs, rats and mink, and even domestic cats.[16] Birds that feed on toads include herons, crows and birds of prey. Crows have been observed to puncture the skin with their beak and then peck out the animal's liver, thus avoiding the toxin.[16] The tadpoles also exude noxious substances which deter fishes from eating them but not the great crested newt. Aquatic invertebrates that feed on toad tadpoles include dragonfly larvae, diving beetles and water boatmen. These usually avoid the noxious secretion by puncturing the tadpole's skin and sucking out its juices.[16]

A parasitic fly, Lucilia bufonivora, attacks adult common toads. It lays its eggs on the toad's skin and when these hatch, the larvae crawl into the toad's nostrils and eat its flesh internally with lethal consequences.[19]

The European fingernail clam (Sphaerium corneum) is unusual in that it can climb up water plants and move around on its muscular foot. It sometimes clings to the toe of a common toad and this is believed to be one of the means by which it disperses to new locations.[20]

In 2007, researchers using a remotely operated underwater vehicle to search for the monster in Loch Ness, Scotland, were surprised to find a common toad hopping along the bottom of the lake at a depth of 324 feet (99 m). They questioned how an air-breathing animal could survive at such depths.[21]

Reproduction

Tadpole development
Toad tadpole development over a two week period
Egg strings of common toad
Egg strings of common toad
Tadpoles of common toad
Tadpoles of common toad, some with hind legs

The common toad emerges from hibernation in spring and there is a mass migration towards the breeding ponds. The toads converge on certain ponds that they favour while avoiding other stretches of water that seem eminently suitable.[15] Adults use the same pond year after year and over 80% of males marked as juveniles have been found to return to the pond at which they were spawned.[22] They find their way to these ponds mainly by using olfactory and magnetic cues. Toads experimentally moved elsewhere and fitted with tracking devices have been found to be able to locate their chosen breeding pond when the displacement exceeded three kilometres (two miles).[23]

The males arrive first and remain in the location for several weeks while the females only stay long enough to mate and spawn. Rather than fighting for the right to mate with a female, male toads may settle disputes by means of the pitch of their voice. Croaking provides a reliable sign of body size and hence of prowess.[24] Nevertheless, fights occur in some instances. In a study at one pond where males outnumbered females by four or five to one, it was found that 38% of the males won the right to mate by defeating rivals in combat or by displacing other males already mounted on females.[25] Male toads generally outnumber female toads at breeding ponds. A Swedish study found that female mortality was higher than that of males and that 41% of females did not come to the breeding pond in the spring and missed a year before reproducing again.[26]

The males mount on the females' backs, grasping them with their fore limbs in a grip that is known as amplexus. The males are very enthusiastic, will try to grasp fish or inanimate objects and often mount on the backs of other males. Sometimes several toads form a heap, each male trying to grasp the female at the base. It is a stressful period and mortality is high among breeding toads.[9] A successful male stays in amplexus for several days and, as the female lays a long, double string of small black eggs, he fertilises them with his sperm. As the pair wander piggyback around the shallow edges of the pond, the gelatinous egg strings, which may contain 3000 to 6000 eggs and be 3 to 4.5 metres (10 to 15 ft) in length, get tangled in plant stalks.

The strings of eggs absorb water and swell in size, and small tadpoles hatch out after a fortnight to three weeks. At first they cling to the remains of the strings and feed on the jelly. They later attach themselves to the underside of the leaves of water weed before becoming free swimming. The tadpoles at first look similar to those of the common frog (Rana temporaria) but they are a darker colour, being blackish above and dark grey below. They can be distinguished from the tadpoles of other species by the fact that the mouth is the same width as the space between the eyes, and this is twice as large as the distance between the nostrils. Over the course of a few weeks their legs develop and their tail gradually gets reabsorbed. By twelve weeks of age they are miniature toads measuring about 1.5 cm (0.6 in) long and ready to leave the pond.[15]

Development and growth

The common toad reaches maturity at three to seven years old but there is great variability between populations.[9] Juveniles are often parasitised by the lung nematode Rhabdias bufonis. This slows growth rates and reduces stamina and fitness. Larger juveniles at metamorphosis always outgrow smaller ones that have been reared in more crowded ponds. Even when they have heavy worm burdens, large juveniles grow faster than smaller individuals with light burdens.[27] After several months of heavy worm infection, some juveniles in a study were only half the mass of controls. Their parasite-induced anorexia caused a decrease in dietary intake and some died.[28] Another study investigated whether the use of nitrogenous fertilisers affects the development of common toad tadpoles. Different strength solutions of ammonium nitrate were used and it was found that at certain concentrations, well above normal field limits, growth was increased and metamorphosis accelerated, but at others, there was no significant difference between the experimental larvae and controls. Nevertheless, certain unusual swimming patterns and a few deformities were found among the experimental animals.[29]

A comparison was made between the growth rate of newly metamorphosed juveniles from different altitudes and latitudes, the specimens studied being from Norway, Germany, Switzerland, the Netherlands and France. All were subspecies B. b. bufo except for the French specimens which may have been B. b. spinosus. At first the growth rates for males and females was identical. By the time they became mature their growth rate had slowed down to about 21% of the initial rate and they had reached 95% of their expected adult size. Some females that were on a biennial breeding cycle carried on growing rapidly for a longer time. Adjusting for differences in temperature and the length of the growing season, the toads grew and matured at much the same rate from the four colder localities. These juveniles reached maturity after 1.09 years for males and 1.55 years for females. However, the young toads from lowland France grew faster and longer to a much greater size taking an average 1.77 years for males and 2.49 years for females before reaching maturity.[30]

Conservation

The IUCN Red List of Threatened Species considers the common toad as being of Least Concern. This is because it has a wide distribution and, over most of its range, is a common species. It is not particularly threatened by habitat loss because it is adaptable and is found in deciduous and coniferous forests, scrubland, meadows, parks and gardens. It prefers damp areas with dense foliage. The major threats it faces include loss of habitat locally, the drainage of wetlands where it breeds, agricultural activities, pollution and mortality on roads. Chytridiomycosis, an infectious disease of amphibians, has been reported in common toads in Spain and the United Kingdom and may affect some populations.[1]

Locally, some populations of common toad are in decline. In Spain, increased aridity and habitat loss have led to a diminution in numbers and it is regarded as "Near Threatened". The subspecies Bufo bufo gredosicola is restricted to the Sierra de Gredos mountain range where it is facing predation by otters and increased competition from the frog Rana perezi which both seem to be extending their ranges to higher altitudes.[1] The common toad cannot be legally sold or traded in the United Kingdom [31] but there is still a slow decline in toad numbers,[1] and it has therefore been declared a Biodiversity Action Plan priority species.[11] In Russia, it is considered to be a "Rare Species" in the provinces of Bashkiria, Tataria, Yamal-Nenets Autonomous County and Irkutsk,[11] but during the 1990s, it became more abundant in Moscow Province.[11]

It has been found that urban populations of common toad occupying small areas and isolated by development show a lower level of genetic diversity and reduced fitness as compared to nearby rural populations. The researchers demonstrated this by genetic analysis and by noting the greater number of physical abnormalities among urban as against rural tadpoles when raised in a controlled environment. It was considered that long term depletion in numbers and habitat fragmentation can reduce population persistence in such urban environments.[32]

Roadkill

Tunnel for toads
Tunnel under road for toads, Germany

Many toads are killed by traffic while migrating to their breeding grounds. In Europe they have the highest rate of mortality from roadkill among amphibians. Many of the deaths take place on stretches of road where streams flow underneath showing that migration routes often follow water courses.[33] In some places in Germany, Great Britain, Northern Italy and Poland, special tunnels have been constructed so that toads can cross under roads in safety. In other places, local wildlife groups run "toad patrols", carrying the amphibians across roads at busy crossing points in buckets. The toads start moving at dusk and for them to travel far, the temperature needs to remain above 5°C. On a warm wet night they may continue moving all night but if it cools down, they may stop earlier.[34] An estimate was made of the significance of roadkill in toad populations in Holland. The number of females killed in the spring migration on a quiet country road (10 vehicles per hour) was compared with the number of strings of eggs laid in nearby fens. A 30% mortality rate was found, with the rate for deaths among males likely to be of a similar order.[35]

Bufotoxin

The main toxic substance found in the parotoid gland and skin of the common toad is called bufotoxin. It was first isolated by Heinrich Wieland and his colleagues in 1922 and they succeeded in identifying its structure about 20 years later.[36] Meanwhile other workers succeeded in isolating the same compound and its parent steroid bufotalin from the Japanese toad (Bufo japonicus). By 1986, researchers at the Arizona State University had succeeded in synthesizing the toad venom constituents bufotalin, cinobufagin, bufalitoxin and bufotoxin.[37] The chemical formula of bufotoxin is C40H60N4O10. Its physical effects resemble those of digitalis [38] which in small doses increases the strength with which the heart muscle contracts and which is used in the treatment of congestive heart failure. The skin of one toad contains enough toxin to cause serious symptoms or even death in animals and man.[39] Clinical effects include severe irritation and pain to eyes, mouth, nose and throat, cardiovascular and respiratory symptoms, paralysis and seizures, increased salivation, vomiting, hyperkalemia, cyanosis and hallucinations.[39] There is no known anti-venom.[39] Treatment consists of supporting respiratory and cardiovascular functions, prevention of absorption and electrocardiography to monitor the condition. Atropine, phenytoin, cholestyramine and lidocaine may prove useful in its management.[39]

Cultural significance

1579 woodcut showing a witch feeding her toad familiars

The toad has long been considered to be an animal of ill omen or a connection to the other world. This may have its origins in the fact that it is at home both on land and in the water. It may cause repugnance because of its blackish, warty skin, its slow movements and the way it emerges from some dark hole. One of the earliest references to toads as being associated with the forces of darkness was when Zoroaster issued an edict in 600 BC that all toads should be killed. In Europe in the Middle Ages, the toad was associated with the Devil, whose coat-of-arms has three toads emblazoned on it. It was thought that the toad could poison people and that it was the witch's familiar and thus possessed magical powers.[40] In France it was thought that witches could be identified by the fact that they were marked by a toad's foot somewhere on their body and in the Pyrenees, the mark was said to be at the back of their eye. In Dorset it was believed that if one found a toad in the house it should be removed without harming it or the householder would face the wrath of the witch whose familiar it was. In some areas, the finding of a toad in a house was considered as evidence that a witch was present.[40] The saliva of the toad was considered poisonous and was known as "sweltered venom" and it was believed that it could spit or vomit poisonous fire. It was thought unlucky for a toad to jump over one's foot and in some regions, this was a portent of death. Toads were associated with devils and demons and in "Paradise Lost", John Milton depicted Satan as a toad when he poured poison into Eve's ear.[40] The First Witch in Shakespeare's Macbeth gave instructions on using a toad in the concoction of spells:[41]

Round about the cauldron go;
In the poison'd entrails throw.
Toad, that under cold stone
Days and nights has thirty-one
Swelter'd venom sleeping got,
Boil thou first i' the charmed pot.

The toad is also associated with female fecundity. Models of toads were left at shrines by women who wanted to conceive. In Romania it was believed that someone who killed a toad was capable of killing their own mother and in Scottish folklore it was held that the tongue of a toad, carried by a man in his breast pocket, would enable him to have his way with any woman.[40] It was also believed that there was a jewel inside a toad's head, a "toadstone", that when worn as a necklace or ring would warn the wearer of attempts to poison them.[42] Shakespeare mentioned this in As you like it:[43]

Sweet are the uses of adversity
Which, like the toad, ugly and venomous,
Wears yet a precious jewel in his head.
Mr. Toad
Mr. Toad as illustrated by Paul Bransom, 1913

Touching a toad has long been suspected of causing warts. This has been shown to be a myth and warts are actually caused by the human papillomavirus and are transferred by direct contact from person to person.[44]

Mr. Toad Esq. is one of the main characters in the children's novel, The Wind in the Willows, by Kenneth Grahame.[45] This has been dramatized by several authors including A. A. Milne who called his play Toad of Toad Hall. Mr Toad is a very conceited, anthropomorphic toad and in the book he composes a ditty in his own praise which starts like this:[46]

The world has held great heroes,
As history books have showed;
But never a name went down to fame
Compared with that of Toad!
The clever men at Oxford
Know all there is to be knowed.
But none of them know half as much
As intelligent Mr. Toad!

References

  1. ^ a b c d Agasyan, A.; Avisi, A.; Tuniyev, B.; Isailovic, J. C.; Lymberakis, P.; Andrén, C.; Cogalniceanu, D.; Wilkinson, J.; Ananjeva, N.; Üzüm, N.; Orlov, N.; Podloucky, R.; Tuniyev, S.; Kaya, U. (2009). "Bufo bufo (Common toad)". IUCN Red List of Threatened Species Version 2012.1. Retrieved 2012-08-23.{{cite web}}: CS1 maint: multiple names: authors list (link)
  2. ^ "Bufo bufo (Linnaeus, 1758)". African Amphibians Lifedesk. Retrieved 2012-06-21.
  3. ^ von Linné, Carl, translated by William Turton (1806). Volume 1. A general system of nature: through the three grand kingdoms of animals, vegetables, and minerals, systematically divided into their several classes, orders, genera, species, and varieties. Lackington, Allen, and Co.{{cite book}}: CS1 maint: multiple names: authors list (link)
  4. ^ "Amphibian Species of the World 5.1 - Bufonidae". Retrieved 2012-06-12.
  5. ^ a b c Birstein, V. J.; Mazin, A. L. (1982). "Chromosomal polymorphism of Bufo bufo: Karyotype and C-banding pattern of B. b. verrucosissima". Genetica. 59 (2): 93&#8209, 98. doi:10.1007/BF00133292.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  6. ^ Garcia-Porta, J.; Litvinchuk, S. N.; Crochet, P. A.; Romano, A.; Lo-Valvo, M.; Lymberakis, P.; Carranza, S. (2012). "Molecular phylogenetics and historical biogeography of the west-palearctic common toads (Bufo bufo species complex)". Molecular Phylogenetics and Evolution. 63 (1): 113&#8209, 130. PMID 22214922.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  7. ^ a b Naish, Darren. "Toads of the world: first, (some) toads of the north". Tetrapod zoology. Retrieved 2012-06-23.
  8. ^ Recuero E.; Canestrelli D.; Voeroes J.; Szabó, K.; Poyarkov, N. A.; Arntzen, J. W.; Crnobrnja-Isailovic, J.; Kidov A. A.; Cogălniceanu, D.; Caputo, F. P.; Nascetti, G.; Martínez-Solano, I. (2012). "Multilocus species tree analyses resolve the radiation of the widespread Bufo bufo species group (Anura, Bufonidae)". Molecular Phylogenetics and Evolution. 62 (1): 71&#8209, 86. doi:10.1016/j.ympev.2011.09.008.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  9. ^ a b c d e f g h i j k l Arnold, Nicholas (2002). Reptiles and Amphibians of Britain and Europe. Harper Collins Publishers. pp. 73&#8209, 74. ISBN 0002199645. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  10. ^ "Bufo verrucosissimus". IUCN Red List of Threatened Species. Retrieved 2009. {{cite web}}: Check date values in: |accessdate= (help)
  11. ^ a b c d "Bufo bufo: Common toad". AmphibiaWeb. Retrieved 2012-05-04.
  12. ^ Fairchild, G. J. (2003). "Common Toad ‑ Bufo bufo". Reptiles and Amphibians of the UK. Retrieved 2012-03-16.
  13. ^ "The common toad (Bufo bufo)". The Amphibian and Reptile Conservation Trust. Retrieved 2012-05-04.
  14. ^ Hemelaar, A. S. M.; van Gelder, J. J. (1979). "Annual growth rings in phalanges of Bufo bufo (Anura, Amphibia) from the Netherlands and their use for age determination". Netherlands Journal of Zoology. 30 (1): 129&#8209, 135. doi:10.1163/002829680X00069.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  15. ^ a b c d e Stokeo, W. J. (1980). The Observer's Book of British Wild Animals. Frederick Warne & Co. Ltd. pp. 213&#8209, 217. ISBN 0723215030.
  16. ^ a b c d Daniel Winchester. "Common Toad". Surrey Amphibian and Reptile Group. Retrieved 2012-05-03.
  17. ^ Ewart, J. P. (1987). "Neuroethology of releasing mechanisms: Prey-catching in toads". Behavioral and Brain Sciences. 10: 337&#8209, 405. doi:10.1017/S0140525X00023128.
  18. ^ Larsen, Lis Olesen; Pedersen, Jan Nyholm (1981). "The snapping response of the toad, Bufo bufo, towards prey dummies at very low light intensities". Amphibia-Reptilia. 2 (4): 321&#8209, 327. doi:10.1163/156853882X00248.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  19. ^ Strijbosch, H. (1980). "Mortality in a population of Bufo bufo resulting from the fly Lucilia bufonivora". Oecologia. 45 (2): 285&#8209, 286. doi:10.1007/BF00346472.
  20. ^ Petkeviciute, R.; Stunzenas, V.; Staneviciute, G. (2004). "Cytogenetic and sequence comparison of adult Phyllodistomum (Digenea: Gorgoderidae) from the three-spined stickleback with larvae from two bivalves". Parasitology. 129: 771&#8209, 778. doi:10.1017/S0031182004006109.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  21. ^ Simpson, Alan (2007-05-03). "The Loch Ness toad; Scientists stunned as 'miraculous' amphibian is found 324ft down". Daily Mail. via HighBeam Research. Retrieved 2012-04-18.
  22. ^ Reading, C. J. (1991). "Breeding pond fidelity in the common toad, Bufo bufo". Journal of Zoology. 225 (2): 201&#8209, 211. doi:10.1111/j.1469-7998.1991.tb03811.x.
  23. ^ Sinsch, Ulrich (1987). "Orientation behaviour of toads (Bufo bufo) displaced from the breeding site". Journal of Comparative Physiology A. 161 (5): 715&#8209, 727. doi:10.1007/BF00605013.
  24. ^ Davies, N. B.; Halliday, T. R. (1978). "Deep croaks and fighting assessment in toads Bufo bufo". Nature. 274: 683&#8209, 685. doi:10.1038/274683a0.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  25. ^ Davies, N. B.; Halliday, T. R. (1979). "Competitive mate searching in male common toads, Bufo bufo". Animal Behaviour. 27 (4): 1253&#8209, 1267. doi:10.1016/0003-3472(79)90070-8.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  26. ^ "Studies conducted at Lund University on amphibian research recently published". Science Letter. via HighBeam Research. 2010-12-21. Retrieved 2012-04-18.
  27. ^ Goater, Cameron P.; Semlitsch, Raymond D.; Bernasconi, Marco V. (1993). "Effects of body size and parasite infection on the locomotory performance of juvenile toads, Bufo bufo". Oikos. 16 (1): 129&#8209, 136. JSTOR 3545205.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  28. ^ Goater, C. P.; Ward, P. I. (1992). "Negative effects of Rhabdias bufonis (Nematoda) on the growth and survival of toads (Bufo bufo)". Oecologia. 89 (2): 161&#8209, 165. JSTOR 4219866.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  29. ^ Xu, Q.; Oldham, R. S. (1997). "Lethal and sublethal effects of nitrogen fertilizer ammonium nitrate on common toad (Bufo bufo) tadpoles". Archives of Environmental Contamination and Toxicology. 32 (3): 298&#8209, 303. doi:10.1007/s002449900188.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  30. ^ Hemelaar, Agnes (1988). "Age, Growth and Other Population Characteristics of Bufo bufo from Different Latitudes and Altitudes". Journal of Herpetology. 22 (4): 369&#8209, 388. JSTOR 1564332.
  31. ^ "Common Toad: Bufo bufo". Amphibian and Reptile Conservation Trust. Retrieved 2012-05-04.
  32. ^ Hitchings, S. P.; Beebee, T. J. C. (1998). "Loss of genetic diversity and fitness in Common Toad (Bufo bufo) populations isolated by inimical habitat". Journal of Evolutionary Biology. 11 (3): 269&#8209, 283. doi:10.1046/j.1420-9101.1998.11030269.x.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  33. ^ Santos, X.; Llorente, G. A.; Montori, A.; Carretero, M. A.; Franch, M.; Garriga, N.; Richter-Boix, A. (2007). "Evaluating factors affecting amphibian mortality on roads: the case of the Common Toad Bufo bufo, near a breeding place [[File:Open Access logo PLoS transparent.svg|9px|link=Open access|open access publication – free to read|alt=Open access icon]]". Animal Biodiversity and Conservation. 30 (1). The Natural Science Museum of Barcelona. {{cite journal}}: Text "pages97‑104" ignored (help); URL–wikilink conflict (help)CS1 maint: multiple names: authors list (link)
  34. ^ "Toad watch: Helping toads to survive". Retrieved 2012-04-30.
  35. ^ Gelder, J. J. (1973). "A quantitative approach to the mortality resulting from traffic in a population of Bufo bufo L." Oecologia. 13 (1): 93&#8209, 95. doi:10.1007/BF00379622.
  36. ^ Chen, K. K. (1932). "Action of Bufotoxins". Proceedings of the Society for Experimental Biology and Medicine. 7. 29 (907). New York. doi:10.3181/00379727-29-6141. ISSN 1535-3699. Wieland and Alles isolated bufotoxin from the skin of B. vulgaris or B. bufo bufo. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help); Unknown parameter |month= ignored (help)
  37. ^ Pettit, G. R.; Kamano, Y.; Drasar, P.; Inoue, M.; Knight, J. C. (1987). "Steroids and related natural products. 104. Bufadienolides. 36. Synthesis of bufalitoxin and bufotoxin". Journal of Organic Chemistry. 52 (16): 3573–3578. doi:10.1021/jo00392a014.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  38. ^ "Bufotoxin". Merriam-Webster Dictionary. Retrieved 2012-05-26.
  39. ^ a b c d "Toad toxins". Textfiles.com. Retrieved 2012-05-26.
  40. ^ a b c d Burns, William E. (2003). Witch Hunts in Europe and America: An Encyclopedia. Greenwood Publishing Group. p. 7. ISBN 0-313-32142-6.
  41. ^ Shakespeare, William ((1605-1606)). Macbeth. p. Act 4, Scene 1. {{cite book}}: Check date values in: |year= (help)
  42. ^ Wanner, Noel (2011). "Frogs: Frog myths across cultures". The Exploratorium. The museum of science, art and human perception. Retrieved 2012-08-23.
  43. ^ Shakespeare, William ((1599-1600)). As you like it. p. Act 2, Scene 1. {{cite book}}: Check date values in: |year= (help)
  44. ^ "Does touching toads gives you warts?". Health.ninemsn.com.au. Retrieved 2012-03-15.
  45. ^ Grahame, Kenneth (1908). The Wind in the Willows. Methuen. ISBN 978-0-674-03447-1.
  46. ^ Grahame, Kenneth (1908). "Chapter 10". The Wind in the Willows.

External links

Template:Link FA