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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 True toads

anurans in which all members are known as toads, although Temporal range: Late Paleocene –
some may be called frogs (such as harlequin frogs). The Recent[1]
bufonids now comprise more than 35 genera, Bufo being the
best known.

Contents
History
Taxonomy
Characteristics Common toad or European toad, Bufo

Reproduction bufo
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Taxonomy
References Territorial call of an Atelopus

External links franciscus male

Scientific classification

History Kingdom: Animalia

Phylum: Chordata
Bufonidae is thought to have originated in South America.
Class: Amphibia
Some studies date the origin of the group to after the
breakup of Gondwana, about 78 - 98 million years ago in the Order: Anura
Late Cretaceous.[2] In contrast, other studies have dated the
Superfamily: Hyloidea
origin of the group to the early Paleocene.[3] The bufonids
likely radiated out of South America during the Eocene, with Family: Bufonidae

the entire radiation occurring during the Eocene to Gray, 1825

Oligocene, marking an extremely rapid divergence likely
facilitated by the Paleogene's changing climatic Genera
conditions.[3] Over 35 see text

Taxonomy
The following phylogeny of most genera in the family is
based on Portik and Papenfuss, 2015:,[4] Chan et al., 2016,[5]
Chandramouli et. al., 2016,[6] and Kok et. al., 2017[3]
Native distribution of Bufonidae (in
black)
 
  Melanophryniscus
     
        Atelopus
 
 
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  Oreophrynella
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Song of Common toad or European

 
Osornophryne toad, Bufo bufo.
 
 
 
 
  Frostius
 
 
  Amazophrynella
 
 
 
  Dendrophryniscus
 
 
    Nannophryne
 
  Common toad, female and male on
 
  Peltophryne her back.
 
 
    Rhaebo
 
 
   
Rhinella
 

   
Anaxyrus
     
   
  Incilius
 
 
 
  Didynamipus
 
   
  Poyntonophrynus

 
  Sclerophrys

 
Nimbaphrynoides
 
   
     
Vandijkophrynus
 
 
   
 
    undescribed lineage

   
    Capensibufo
 
 
 
  Mertensophryne
     
        Wolterstorffina
 

 
Werneria
 
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Nectophryne
 
 
    Barbarophryne
 
 
 
 
  Schismaderma

   
    Churamiti
 
 
 
  Nectophrynoides

 
Pedostibes
 

 
  Adenomus
 
 
 
  Blythophryne
 
     
  Xanthophryne
   
 
 
    Bufoides
 

  Duttaphrynus
  (paraphyletic)

 
Bufotes
 
 
   
  Epidalea

   
Strauchbufo
     
 
 
Bufo
 
 
 
  Leptophryne
 
  
 
    Ghatophryne
 
 
   
  Sabahphrynus

    Ansonia
   
 
 
 
  Pelophryne
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Ingerophrynus
 

   
Phrynoidis
   
 
   
  Rentapia

Ingerophrynus alongside Leptophryne was grouped as basal to the clade containing all other
Southeast Asian toad genera and Ghatophryne by Portik and Papenfuss, but was found to group
with Phrynoidis and Rentapia by Chan et al. Ghatophryne was grouped with Phrynoidis and
Rentapia by Portik and Papenfuss but was found to group with Pelophryne and Ansonia by Chan
et al. In addition, Sabahphrynus was grouped with Strauchbufo and Bufo by Portik and Papenfuss
but was found to group with Pelophryne, Ansonia, and Ghatophryne by Chan et al.

Characteristics
True toads are widespread and are native to every continent except Australia and Antarctica,
inhabiting a variety of environments, from arid areas to rainforest. Most lay eggs in paired strings
that hatch into tadpoles, although, in the genus Nectophrynoides, the eggs hatch directly into
miniature toads.[1]

All true toads are toothless and generally warty in appearance. They have a pair of parotoid glands
on the back of their heads. These glands contain an alkaloid poison which the toads excrete when
stressed. The poison in the glands contains a number of toxins causing different effects. Bufotoxin
is a general term. Different animals contain significantly different substances and proportions of
substances. Some, like the cane toad Rhinella marina, are more toxic than others. Some
"psychoactive toads", such as the Colorado River toad Incilius alvarius,[7] have been used
recreationally for the effects of their bufotoxin.

Depending on the species, male or female toads may possess a Bidder's organ, a trait unique to all
bufonids except genera Melanophryniscus and Truebella.[8] Under the right conditions, the organ
becomes an active ovary.[9]

The loss of teeth has arisen in frogs independently over 20 times. Notably, all members of
Bufonidae are toothless. Another Anuran family with a comparable degree of edentulism is the
family Microhylidae.[10]

Reproduction
Internal fertilization occurs in four bufonid genera.[11]

Mertensophryne (some species)

Nectophrynoides (presumably all species)
Altiphrynoides malcolmi (one out of two species in the genus Altiphrynoides)
Nimbaphrynoides occidentalis (the sole species in the monotypic genus Nimbaphrynoides)

Ascaphus (all species) and Eleutherodactylus (two species, E. coqui and E. jasperi) are the only
other frog genera that have internal fertilization.[11] Limnonectes larvaepartus also has internal
fertilization.[12]

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Taxonomy
The family Bufonidae contains over 570 species among 52 genera.

Genus Latin name and author Common name Species

Adenomus Cope, 1861 Dwarf toads 2
Altiphrynoides Dubois, 1987 Ethiopian toads 2
Amazophrynella Fouquet et al., 2012 12
Anaxyrus Tschudi, 1845 23
Ansonia Stoliczka, 1870 Stream toads 34
Atelopus Duméril & Bibron, 1841 Stubfoot toads 96
Barbarophryne Beukema, de Pous, Donaire-Barroso, Bogaerts, Tiznit toad; Brongersma's
1
Garcia-Porta, Escoriza, Arribas, El Mouden, and Carranza, 2013 (1 sp.) toad
Blythophryne Chandramouli et al., 2016[13] Andaman bush toads 1
Bufo Garsault, 1764 Toads 18
Mawblang toads; Rock
Bufoides Pillai & Yazdani, 1973 2
toads
Bufotes Rafinesque, 1815 Palearctic green toads 15
Capensibufo Grandison, 1980 Cape toads 5
Churamiti Channing & Stanley, 2002 1
Dendrophryniscus Jiménez de la Espada, 1871 Tree toads 16
Didynamipus Andersson, 1903 Four-digit toad 1
Duttaphrynus Frost et al., 2006 Dutta's toads 27
Epidalea Cope, 1864 Natterjack toad 1
Frostius Cannatella, 1986 Frost's toads 2
Ghatophryne Biju, Van Bocxlaer, Giri, Loader, and Bossuyt, 2009 2
Central American toads;
Incilius Cope, 1863 Middle American toads; 39
Cerro Utyum toads
Ingerophrynus Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá,
Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Hainan toads 12
Drewes, Nussbaum, Lynch, Green, and Wheeler, 2006
Laurentophryne Tihen, 1960 Parker's tree toad 1
Leptophryne Fitzinger, 1843 Indonesia tree toads 3
South American redbelly
Melanophryniscus Gallardo, 1961 29
toads
Mertensophryne Tihen, 1960 Snouted frogs 14
Metaphryniscus Señaris, Ayarzagüena & Gorzula, 1994 1
Nannophryne Günther, 1870 4
Nectophryne Buchholz & Peters, 1875 African tree toads 2
Nectophrynoides Buchholz & Peters, 1875 African live-bearing toads 13
Nimbaphrynoides Dubois, 1987 Nimba toads 1
Oreophrynella Boulenger, 1895 Bush toads 8
Osornophryne Ruiz-Carranza & Hernández-Camacho, 1976 Plump toads 11
Parapelophryne Fei, Ye & Jiang, 2003 1
Pedostibes Günther, 1876 Asian tree toads 1
Pelophryne Barbour, 1938 Flathead toads 13
Peltophryne Fitzinger, 1843 Caribbean toads 14

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Phrynoidis Fitzinger in Treitschke, 1842 Rough toads 2

Poyntonophrynus Frost, Grant, Faivovich, Bain, Haas, Haddad, de
Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Pygmy toads 11
Moler, Drewes, Nussbaum, Lynch, Green, and Wheeler, 2006
Pseudobufo Tschudi, 1838 False toad 1
Rentapia Chan, Grismer, Zachariah, Brown, and Abraham, 2016 2
Rhaebo Cope, 1862 Cope toads 13
Rhinella Fitzinger, 1826 Beaked toads 92
Sabahphrynus Matsui, Yambun, and Sudin, 2007 Sabah earless toad 1
Schismaderma Smith, 1849 African split-skin toad 1
Sclerophrys Tschudi, 1838 44
Sigalegalephrynus Tschudi, 1838 Puppet toads 5
Siberian toad; Mongolian
Strauchbufo Fei, Ye, and Jiang, 2012 1
toad
Truebella Graybeal & Cannatella, 1995 2
Vandijkophrynus Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá,
Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Van Dijk's toads 6
Drewes, Nussbaum, Lynch, Green, and Wheeler, 2006
Werneria Poche, 1903 Smalltongue toads 6
Wolterstorffina Mertens, 1939 Wolterstorff toads 3
Xanthophryne Biju, Van Bocxlaer, Giri, Loader & Bossuyt, 2009 2

References
1. Zweifel, Richard G. (1998). Cogger, H.G.; Zweifel, R.G. (eds.). Encyclopedia of Reptiles and
Amphibians. San Diego: Academic Press. pp. 91–92. ISBN 978-0-12-178560-4.
2. Pramuk, Jennifer B.; Robertson, Tasia; Sites, Jack W.; Noonan, Brice P. (2008). "Around the
world in 10 million years: biogeography of the nearly cosmopolitan true toads (Anura:
Bufonidae)" (https://onlinelibrary.wiley.com/doi/abs/10.1111/j.1466-8238.2007.00348.x). Global
Ecology and Biogeography. 17 (1): 72–83. doi:10.1111/j.1466-8238.2007.00348.x (https://doi.o
rg/10.1111%2Fj.1466-8238.2007.00348.x). ISSN 1466-8238 (https://www.worldcat.org/issn/14
66-8238).
3. Kok, Philippe J. R.; Ratz, Sebastian; MacCulloch, Ross D.; Lathrop, Amy; Dezfoulian,
Raheleh; Aubret, Fabien; Means, D. Bruce (2018). "Historical biogeography of the
palaeoendemic toad genus Oreophrynella (Amphibia: Bufonidae) sheds a new light on the
origin of the Pantepui endemic terrestrial biota" (https://onlinelibrary.wiley.com/doi/abs/10.1111/j
bi.13093). Journal of Biogeography. 45 (1): 26–36. doi:10.1111/jbi.13093 (https://doi.org/10.111
1%2Fjbi.13093). ISSN 1365-2699 (https://www.worldcat.org/issn/1365-2699).
4. Portik, Daniel M.; Papenfuss, Theodore J. (2015-08-06). "Historical biogeography resolves the
origins of endemic Arabian toad lineages (Anura: Bufonidae): Evidence for ancient vicariance
and dispersal events with the Horn of Africa and South Asia" (https://www.ncbi.nlm.nih.gov/pm
c/articles/PMC4527211). BMC Evolutionary Biology. 15 (1): 152. doi:10.1186/s12862-015-
0417-y (https://doi.org/10.1186%2Fs12862-015-0417-y). ISSN 1471-2148 (https://www.worldc
at.org/issn/1471-2148). PMC 4527211 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC452721
1). PMID 26245197 (https://pubmed.ncbi.nlm.nih.gov/26245197).

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5. Chan, Kin Onn; Grismer, L. Lee; Zachariah, Anil; Brown, Rafe M.; Abraham, Robin Kurian
(2016-01-20). "Polyphyly of Asian Tree Toads, Genus Pedostibes Günther, 1876 (Anura:
Bufonidae), and the Description of a New Genus from Southeast Asia" (https://www.ncbi.nlm.ni
h.gov/pmc/articles/PMC4720419). PLOS ONE. 11 (1): e0145903.
Bibcode:2016PLoSO..1145903C (https://ui.adsabs.harvard.edu/abs/2016PLoSO..1145903C).
doi:10.1371/journal.pone.0145903 (https://doi.org/10.1371%2Fjournal.pone.0145903).
ISSN 1932-6203 (https://www.worldcat.org/issn/1932-6203). PMC 4720419 (https://www.ncbi.n
lm.nih.gov/pmc/articles/PMC4720419). PMID 26788854 (https://pubmed.ncbi.nlm.nih.gov/2678
8854).
6. Chandramouli, S. R.; Vasudevan, Karthikeyan; Harikrishnan, S.; Dutta, Sushil Kumar; Janani,
S. Jegath; Sharma, Richa; Das, Indraneil; Aggarwal, Ramesh (2016-01-20). "A new genus and
species of arboreal toad with phytotelmonous larvae, from the Andaman Islands, India
(Lissamphibia, Anura, Bufonidae)" (https://zookeys.pensoft.net/article/6522/). ZooKeys (555):
57–90. doi:10.3897/zookeys.555.6522 (https://doi.org/10.3897%2Fzookeys.555.6522).
ISSN 1313-2970 (https://www.worldcat.org/issn/1313-2970). PMC 4740822 (https://www.ncbi.n
lm.nih.gov/pmc/articles/PMC4740822). PMID 26877687 (https://pubmed.ncbi.nlm.nih.gov/2687
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Threatened Species. Retrieved 2019-01-19.
8. Piprek, Rafal P., et al. “Bidder’s Organ – Structure, Development and Function.” The
International Journal of Developmental Biology, vol. 58, no. 10-11–12, 2014, pp. 819–27.
Crossref, doi:10.1387/ijdb.140147rp.
9. Brown, Federico D.; Del Pino, Eugenia M.; Krohne, Georg (December 2002). "Bidder's organ
in the toad Bufo marinus: Effects of orchidectomy on the morphology and expression of
lamina-associated polypeptide 2" (https://doi.org/10.1046%2Fj.1440-169X.2002.00665.x).
Development, Growth & Differentiation. 44 (6): 527–535. doi:10.1046/j.1440-
169X.2002.00665.x (https://doi.org/10.1046%2Fj.1440-169X.2002.00665.x). ISSN 1440-169X
(https://www.worldcat.org/issn/1440-169X). PMID 12492511 (https://pubmed.ncbi.nlm.nih.gov/
12492511). S2CID 44753338 (https://api.semanticscholar.org/CorpusID:44753338).
10. Paluh, Daniel J., et al. “Rampant Tooth Loss Across 200 Million Years of Frog Evolution.”
BioRxiv, 2021. Crossref, doi:10.1101/2021.02.04.429809.
11. Vitt, Laurie J.; Caldwell, Janalee P. (2014). Herpetology: An Introductory Biology of Amphibians
and Reptiles (4th ed.). Academic Press. p. 122.
12. Iskandar, D. T.; Evans, B. J.; McGuire, J. A. (2014). "A novel reproductive mode in frogs: a new
species of fanged frog with internal fertilization and birth of tadpoles" (https://www.ncbi.nlm.nih.
gov/pmc/articles/PMC4281041). PLOS ONE. 9 (12): e115884. Bibcode:2014PLoSO...9k5884I
(https://ui.adsabs.harvard.edu/abs/2014PLoSO...9k5884I). doi:10.1371/journal.pone.0115884
(https://doi.org/10.1371%2Fjournal.pone.0115884). PMC 4281041 (https://www.ncbi.nlm.nih.go
v/pmc/articles/PMC4281041). PMID 25551466 (https://pubmed.ncbi.nlm.nih.gov/25551466).
13. S. R. Chandramouli, Karthikeyan, Vasudevan, S Harikrishnan, Sushil Kumar Dutta, S Jegath
Janani, Richa Sharma, Indraneil Das, Ramesh Aggarwal. “A new genus and species of
arboreal toad with phytotelmonous larvae, from the Andaman Islands, India (Lissamphibia,
Anura, Bufonidae)” ZooKeys (2016) 555: 57-90, https://doi.org/10.3897/zookeys.555.6522
"Amphibian Species of the World 5.1 - Bufonidae" (https://archive.today/20120715231112/htt
p://research.amnh.org/herpetology/amphibia/names.php?taxon=&family=Bufonidae&subfamily
=&genus=&commname=&authority=&year=&geo=0&dist=&comment=). Archived from the
original (http://research.amnh.org/herpetology/amphibia/names.php?taxon=&family=Bufonidae
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15. Retrieved 2008-04-10.
Stebbins, Robert. Western Reptiles & Amphibians (3rd ed.). Houghton Mifflin Co., 2003.
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Facts on File, New York, 2002.

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External links
Tolweb.org: Bufonidae (http://tolweb.org/Bufonidae)
Bufonidae.com (http://www.bufonidae.com)
Amphibian and Reptiles of Peninsular Malaysia - Family Bufonidae (http://www.amphibia.my/p
age.php?pageid=Browse%20Species%20Guide&family=Bufonidae%20(Toads))
FED.us (http://www.fs.fed.us/r4/amphibians/family_bufonidae.htm)
Bufonidae recordings (http://sounds.bl.uk/Browse.aspx?category=Environment&collection=Am
phibians&browseby=Browse+by+family&choice=Bufonidae) from the British Library Sound
Archive

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