1048:
873:
556:
701:
586:
447:). However, in many species the yellow or orange is partial and incomplete. On the underparts, the yellow or orange may form a spotted or mottled pattern on a dull and dark background. On the upperparts, there can be spots, mottling or patches in black, brown, greenish, whitish or reddish. This results in some species where most but not all of the upperparts are yellow or orange and others where only small sections are yellow or orange. Two particularly dull-coloured species,
571:
541:
1271:
1249:
882:
56:
1282:
1139:
31:
1260:
623:
162:
601:
455:, are overall brown, while a few others are essentially all-brown or greenish above (sometimes with whitish on the mid-back), limiting yellow or orange to the underparts. In species where known, including ones that are bright orange-yellow as adults, the newly hatched juveniles are mostly brown overall and may have white markings below.
296:
918:(insufficient data for rating them). Since then, many new species have been recognised and more data has become available for evaluating their conservation status. In a review in 2019 that included all species recognised up until then and used IUCN's definitions, 10 species were data deficient, 5 least concern, 10
1217:
Starting in 1998, this changed dramatically. Since that year, there has been a couple of years at most between the description of new species and some years where several were described, peaking at eight species in 2015 alone. Additionally, the three remaining variants that had been described in 1920
646:
The advertising calls by males are performed either from a hidden or an exposed position. Whether a species is dull or brightly coloured does not accurately predict if it prefers an exposed or hidden calling position. In addition to advertising calls, there are aggressive calls directed towards other
346:
are very small frogs and adults range from 7.4 to 19.7 mm (0.29–0.78 in) in snout–to–vent length. Females tend to be larger than males of the same species. The flea frogs or flea toads are generally the smallest at 7.4–10.8 mm (0.29–0.43 in) in adults, but overlapping to some
741:
Based on the few species where the breeding behavior is known, the female saddleback toad lays about five or less yellow or yellow-white eggs, which are relatively large compared to the size of the adult. They are deposited in a hidden location (for example, under leaf litter or a tree log) on land.
442:
Among the remaining species in the genus, almost all have both underparts and upperparts that are at least partially yellow or orange. They are collectively known as pumpkin toadlets, which is a reference to the colour. In some species, this colour is almost uniform throughout, earning them the name
1234:
that are separated from each other by valleys. Although now generally recognised, the distinction of some of these recently described species has been questioned by some authorities since they often have been separated mainly by colour and skin texture, features that can be variable in frogs/toads.
973:
mountaintops and most species have completely separate distribution, isolated from each other by valleys and unsuitable habitat. Where found, they can however be common, sometimes even abundant. Although some of the range-restricted species have well-protected distributions in reserves, many of the
714:
Saddleback toads live in the leaf litter on forest floors, but on occasion may move to higher perches up to 1 m (3.3 ft) off the ground. During dry conditions, both sexes tend to stay hidden in the leaf litter and there is little noticeable activity, but during the wet season and high
638:
are most often located by their call. The primary calls are by adult males, but females and young may also call. A single square meter (slightly more than 10 sq ft) can host as many as five calling males, and at a single location one may hear tens of males at once. Males will defend their small
2510:
Dorigo, T.A.; C.C. Siqueira; D. Vrcibradic; T. Maia-Carneiro; M. Almeida-Santos; C.F.D. Rocha (2012). "Ecological aspects of the pumpkin toadlet, Brachycephalus garbeanus
Miranda-Ribeiro, 1920 (Anura: Neobatrachia: Brachycephalidae), in a highland forest of southeastern Brazil".
310:, but these are not present in all species in the genus and their function is unclear in the species that do have them. The genus can be divided into two main subgroups: The pumpkin toadlets comprises most species in the genus. They have an overall toad-like appearance and
683:
and some others, many species in the genus always call from a hidden position, making any signalling by movements inefficient, and it is likely they can hear their own calls. Direct studies of the medium-brightly coloured (mostly dull above, bright below) pumpkin toadlet
670:
from the ancestral form of the genus, whereas their reduced hearing ability (they do have some hearing ability in frequencies outside their call) is a novel change in these species. Sounds make them more vulnerable to predators, but there has likely been little direct
742:
The female rolls them until they are covered in particles, making them almost indistinguishable from the soil where they have been laid, but it may also help to preserve their water content. After about two months, they hatch into juvenile toads/frogs. There is no
811:
988:
despite the fungus largely affecting aquatic frogs; these outbreaks have been linked to dry years where low water availability poses a physiological stress on frogs and also forces them to retreat to refugia where they are at a high population density.
340:. The skin of the saddleback toads can be rugose (bumpy) or smooth. Their toes/fingers are reduced compared to most other frogs, with typically only two to four being clearly visible on each foot/hand. Some of the bones in their skull are also reduced.
854:
to sense changes in angular acceleration during a jump. For the same reason they are moving slowly when they walk. Because of the uncontrolled jumping and slow walking, they have evolved defense mechanisms such as toxicity and bony plates
1956:
Monteiro, J.P.D.C.; Condez, T.H.; Garcia, P.C.D.A.; Comitti, E.J.; Amaral, I.B.; Haddad, C.F.B. (2018). "A new species of
Brachycephalus (Anura, Brachycephalidae) from the coast of Santa Catarina State, southern Atlantic Forest, Brazil".
3313:
Verdade, V.K.; M.T. Rodrigues; J. Cassimiro; D. Pavan; N. Liou; M.C. Lange (2008). "Advertisement Call, Vocal
Activity, and Geographic Distribution of Brachycephalus hermogenesi (Giaretta and Sawaya, 1998) (Anura, Brachycephalidae)".
1995:
Pires Jr., O.R.; A. Sebben; E.F. Schwartz; S.W. Largura; C. Bloch Jr.; R.A. Morales; C.A. Schwartz (2002). "Occurrence of tetrodotoxin and its analogues in the
Brazilian frog Brachycephalus ephippium (Anura: Brachycephalidae)".
3146:
Sandra Goutte; Matthew J. Mason; Marta M. Antoniazzi; Carlos Jared; Didier Merle; Lilian Cazes; LuĂs Felipe Toledo; Hanane el-Hafci; StĂ©phane Pallu; Hugues
Portier; Stefan Schramm; Pierre Gueriau; Mathieu Thoury (2019).
525:
or determining fitness of a potential partner), but later studies indicate that the former explanation is unlikely, as predation attempts on the toadlets appear to be unaffected by the presence/absence of fluorescence.
231:
in southeastern Brazil. The genus includes two main groups, the often brightly coloured pumpkin toadlets, and the overall brown and more frog-like flea frogs (or flea toads), which once were placed in their own genus
2710:
Scherz, Mark D.; Hutter, Carl D.; Rakotoarison, Andolalao; Riemann, Jana C.; Rödel, Mark-Oliver; Ndriantsoa, Serge H.; Glos, Julian; Roberts, Sam Hyde; Crottini, Angelica; Vences, Miguel; Glaw, Frank (2019-03-27).
3724:
Kaplan, M. (2002). "Histology of the anteroventral part of the breast-shoulder apparatus of
Brachycephalus ephippium (Brachycephalidae) with comments on the validity of the genus Psyllophryne (Brachycephalidae)".
2337:
Lourenço-de-Moraes, Ricardo; Dias, Iuri R.; Mira-Mendes, Caio V.; Oliveira, Renan M. de; Barth, Adriane; Ruas, Danilo S.; Vences, Miguel; Solé, Mirco; Bastos, Rogério P. & Lötters, Stefan (19 September 2018).
347:
extent with the smallest species of pumpkin toadlets. Adults of the pumpkin toadlets are at least 8.3 mm (0.33 in), with most species about 9.5–13.5 mm (0.37–0.53 in), and the largest like
419:
vary greatly in colour and this is often useful for separating the species from each other. Within each species, there are often also some individual variations, especially in the details of their pattern.
3037:
Pires Jr., O.R.; A. Sebben; E.F. Schwartz; R.A.V. Morales; C. Bloch Jr.; C.A. Schwartz (2005). "Further report of the occurrence of tetrodotoxin and new analogues in the Anuran family
Brachycephalidae".
816:
815:
812:
1021:, with the latter being the generally accepted treatment in recent decades. Initially some authorities placed numerous genera in Brachycephalidae, while others limited it to only contain the genus
817:
2897:
Alves, A.C.R.; R.J. Sawaya; S.F. Dos Reis; C.F.B. Haddad (2009). "New
Species of Brachycephalus (Anura: Brachycephalidae) from the Atlantic Rain Forest in SĂŁo Paulo State, Southeastern Brazil".
2092:
Condez, T.H.; Monteiro, J.P.D.; Comitti, E.J.; Garcia, P.C.A.; Amaral, I.B.; Haddad, C.F.B. (2016). "A new species of flea-toad (Anura: Brachycephalidae) from southern
Atlantic Forest, Brazil".
738:
will use two, beginning with an inguinal amplexus (where the male holds the female around the lower waist, just in front of her hindlimbs) and only later changing to a nearly axillary position.
965:
are only known from seven or less locations and many at only one or two locations covering less than 1 km (0.4 sq mi) at a very narrow altitude range. These are typically in
2797:"New altitudinal record for Brachycephalus actaeus Monteiro, Condez, Garcia, Comitti, Amaral & Haddad, 2018 (Anura, Brachycephalidae), with comments on its habitats of occurrence"
2713:"Morphological and ecological convergence at the lower size limit for vertebrates highlighted by five new miniaturised microhylid frog species from three different Madagascan genera"
754:
Relatively little is known about the feeding behavior of saddleback toads, but they will take a wide range of minute invertebrates. They are known to feed on insects (among others,
2189:
3616:
Pombal, J.P. Jr. (2010). "A posição taxonômica das "variedades" de
Brachycephalus ephippium (Spix, 1824) descritas por Miranda-Ribeiro, 1920 (Amphibia, Anura, Brachycephalidae)".
3826:
814:
978:(changing temperature and rain patterns), fire, grazing by livestock, and deforestation for tree extraction and to make room for plantations, farming and cattle ranching. Some
3908:
Ivan Nunes; Carla S. GuimarĂŁes; Pedro Henrique A. G. Moura; Mariana Pedrozo; Matheus de Toledo Moroti; Leandro M. Castro; Daniel R. Stuginski; Edelcio Muscat (28 April 2021).
715:
humidity the males and in some species also females may move to the top of the leaf litter. In some species, they often take up quite conspicuous positions during this time.
957:
are relatively widespread in both distribution and altitude range (lowlands to about 1,100–1,250 m ), although it is possible that the last of these actually is a
662:
are unable to hear the frequency of their own advertising calls, as their ears are underdeveloped. Instead their communication appears to rely on certain movements like the
363:
at 7.4–10.8 mm (0.29–0.43 in) has the smallest minimum adult size in the genus, but its maximum size surpasses that reported for a few others. This makes them the
3349:
1047:
4171:
2307:
Campos, L.A..; H.R. da Silva; A. Sebben (2010). "Morphology and development of additional bony elements in the genus Brachycephalus (Anura: Brachycephalidae)".
600:
1062:, there are two rather distinctive groups: the typical members are the pumpkin toadlets, while the remaining four species are the flea frogs or flea toads (
439:) are well-camouflaged and overall brown, often with sections that are greyer or near-black, and sometimes with a few poorly-defined yellowish spots below.
403:. Newly hatched saddleback toads are much smaller than the already tiny adults. For example, they are typically 5.25–5.45 mm (0.207–0.215 in) in
555:
4217:
4227:
4132:
813:
718:
Overall the saddleback toads are mostly active during the day, but some activity also occurs during the night and some species possibly are mostly
4158:
585:
734:
where a male uses his front limbs to grab a female at her front limbs. However, whereas most frogs typically only use one technique, at least
2272:
Pombal Jr., J.P.; I. Sazima; C.F.B. Haddad (1994). "Breeding Behavior of the Pumpkin Toadlet, Brachycephalus ephippium (Brachycephalidae)".
3635:
Moura-Campos, Diego; Greenspan, Sasha E.; DiRenzo, Graziella V.; Neely, Wesley J.; Toledo, LuĂs Felipe; Becker, C. Guilherme (2021-09-01).
2935:
Pombal Jr, J.P.; E.M. Wistuba; M.R. Bornschein (1998). "A New Species of Brachycephalid (Anura) from the Atlantic Rain Forest of Brazil".
4075:
1090:. The pumpkin toadlets, which largely are restricted to highlands and foothills, can be further divided into two subgroups: the southern
385:
approaching their tiny dimensions. In other continents, the only similar-sized or slightly smaller frogs are a few species in the genera
2197:
3881:
Condez, T.H.; J.P.d.C. Monteiro; C.F.B. Haddad (2017). "Comments on the current taxonomy of Brachycephalus (Anura: Brachycephalidae)".
3969:
Garey, Michel V.; Lima, AndrĂ© M. X.; Hartmann, MarĂlia T.; Haddad, CĂ©lio F. B. (2012). "A new species of miniaturized toadlet, genus
3746:"Geographical and altitudinal distribution of Brachycephalus (Anura: Brachycephalidae) endemic to the Brazilian Atlantic Rainforest"
3380:
2438:
Ribeiro, Luiz F.; Bornschein, Marcos R.; Belmonte-Lopes, Ricardo; Firkowski, Carina R.; Morato, Sergio A.A.; Pie, Marcio R. (2015).
1210:(one of the variants that had been described in 1920) was instated as a valid species, bringing the number of generally recognised
570:
4222:
478:, which has a much more subdued colouring but still with some yellow-orange, appears to be non-toxic. It is unknown how certain
540:
4070:
3288:
1230:. The many recently discovered species have mostly been the result of scientists visiting isolated highland peaks, so-called
984:
302:
is one of the smallest species in the genus; with its frog-like appearance and overall brown colour it is a typical flea frog
4163:
1790:"New World direct-developing frogs (Anura: Terrarana): Molecular phylogeny, classification, biogeography, and conservation"
3910:"Hidden by the name: A new fluorescent pumpkin toadlet from the Brachycephalus ephippium group (Anura: Brachycephalidae)"
55:
2561:"Two new species of the Brachycephalus pernix group (Anura: Brachycephalidae) from the state of Paraná, southern Brazil"
1564:
1474:
1179:
255:, ranging from near sea level to an altitude of 1,900 m (6,200 ft), with most species restricted to highland
4093:
1363:
381:
3693:
4176:
1726:
1666:
1514:
1029:
studies have shown that most of these genera belong in other families, but the—in morphology quite different—genus
3525:"Semicircular Canal Size Determines the Developmental Onset of Angular Vestibuloocular Reflexes in Larval Xenopus"
938:, have not been recorded since their original scientific descriptions a century ago and they are unknown in life.
2224:"Diet of Brachycephalus brunneus (Anura: Brachycephalidae) in the Atlantic Rainforest of Paraná, southern Brazil"
2152:"Conservation Status of Brachycephalus Toadlets (Anura: Brachycephalidae) from the Brazilian Atlantic Rainforest"
2040:"Conservation Status of Brachycephalus Toadlets (Anura: Brachycephalidae) from the Brazilian Atlantic Rainforest"
1716:
1656:
1554:
1534:
1464:
1443:
1333:
1706:
872:
295:
1676:
1595:
1584:
1574:
1544:
1453:
1393:
1373:
1353:
1054:
relationship of four pumpkin toadlet species showing that colours alone are not a good indicator of relatedness
375:
38:
3591:
2843:"A new species of Brachycephalus (Anura: Brachycephalidae) from the Quiriri mountain range of southern Brazil"
2629:
Pie, Marcio R.; Ribeiro, Luiz F.; Confetti, André E.; Nadaline, Mário J.; Bornschein, Marcos R. (2018-10-03).
1686:
1504:
1484:
1403:
822:
1524:
1433:
4212:
1494:
1423:
1413:
1383:
1323:
1161:
666:
that inflates when calling, mouth gaping and waving of their arms. It is speculated that their calling is a
3396:
974:
species do not. Their small distributions make them highly vulnerable to habitat loss and degradation from
4010:
3478:"Quantifying the Ontogeny of Optokinetic and Vestibuloocular Behaviors in Zebrafish, Medaka, and Goldfish"
1626:
1615:
1343:
1099:
369:
228:
4098:
2038:
Rebouças, R.; A.B. Carollo; M.d.O. Freitas; C. Lambertini; R.M. Nogueira dos Santos; L.F. Toledo (2019).
1696:
1646:
314:, and they mostly walk (they are unable to jump far). The second group are the flea frogs or flea toads (
3820:
1605:
927:
898:
672:
278:
species have only been discovered in the 21st century. Most species have tiny ranges and are seriously
3422:
1914:
Fitzinger (Amphibia: Anura: Brachycephalidae), from the Atlantic rainforest of southern Bahia, Brazil"
700:
517:. It was initially speculated that the fluorescent colour also is aposematic or that it is related to
4119:
4057:
3923:
3434:
3230:
3160:
2724:
2688:
2355:
1636:
1191:
1107:
311:
643:
from other males, but in most cases voice or visual displays are sufficient and fights are avoided.
458:
The bright yellow, orange or reddish colours of many species in this genus are generally considered
4048:
1157:
1078:). Until 2002, the latter group, which is relatively widespread, was regarded as a separate genus,
847:
522:
252:
334:
frogs, and they are better jumpers. The name "flea frog" is also sometimes used for the unrelated
3990:
3477:
3331:
3264:
3125:
3016:
2952:
2914:
2818:
2600:
2528:
2289:
923:
919:
640:
364:
279:
245:
178:
50:
2977:"A new species of Brachycephalus (Anura: Brachycephalidae) from Santa Catarina, southern Brazil"
267:
and breeding is by direct development, with eggs laid on land and hatching into young frogs (no
1270:
4184:
4106:
3951:
3777:
3658:
3562:
3544:
3505:
3497:
3458:
3450:
3376:
3256:
3186:
3117:
3055:
3008:
2874:
2752:
2670:
2652:
2592:
2483:
2465:
2383:
2109:
2013:
1974:
1866:
1858:
1248:
1175:
1095:
1083:
1006:
881:
839:
462:(warning colours), but toxicity has only been studied in a few species. The brightly coloured
134:
3084:
Bornschein, M.R.; L.F. Ribeiro; M.M. Rollo Jr.; A.E. Confetti; M.R. Pie; et al. (2018).
4189:
3982:
3941:
3931:
3890:
3808:
3767:
3757:
3648:
3552:
3536:
3489:
3442:
3323:
3246:
3238:
3176:
3168:
3107:
3097:
3047:
2998:
2988:
2944:
2906:
2864:
2854:
2808:
2742:
2732:
2660:
2642:
2582:
2572:
2520:
2473:
2455:
2373:
2363:
2316:
2281:
2235:
2163:
2101:
2051:
2005:
1966:
1925:
1850:
1834:
1804:
1138:
1018:
491:
213:
210:
117:
30:
1281:
3812:
3689:
1259:
958:
648:
217:
2344:(Anura: Eleutherodactylidae): A new species from the Atlantic Forest of northeast Brazil"
3927:
3856:
3438:
3234:
3164:
2728:
2359:
622:
4062:
3986:
3946:
3909:
3772:
3745:
3557:
3251:
3218:
3181:
3148:
3112:
3085:
3003:
2976:
2869:
2842:
2747:
2712:
2665:
2630:
2587:
2560:
2478:
2439:
2378:
2339:
1761:
975:
915:
787:
161:
2009:
1833:
GuimarĂŁes, Carla Silva; Luz, Sofia; Rocha, Pedro Carvalho; Feio, Renato Neves (2017).
630:
and at least one other pumpkin toadlet species are deaf to their own advertising calls
4206:
3796:
3579:
3268:
2822:
2320:
1303:
are four of the thirteen species that were scientifically described from 2015 to 2017
911:
495:
331:
3994:
3335:
3129:
3020:
2918:
2604:
2532:
2240:
2223:
3540:
1909:
1111:
1051:
966:
667:
487:
471:
264:
256:
4111:
3051:
1182:, but in 1955 a review did not recognise them, arguing that they were synonyms of
1110:
and north). Among the pumpkin toadlets, colours alone are not a good indicator of
3936:
3653:
3636:
2737:
2524:
2368:
897:(right), two recently described species that, according to a review in 2019, are
647:
males when close. The calls of the different species differ to various extent in
4145:
4042:
3637:"Fungal disease cluster in tropical terrestrial frogs predicted by low rainfall"
3523:
Lambert, François M.; Beck, James C.; Baker, Robert; Straka, Hans (2008-08-06).
2631:"A new species of Brachycephalus (Anura: Brachycephalidae) from southern Brazil"
1031:
767:
518:
510:
393:
387:
336:
236:. Some pumpkin toadlets are toxic and their often bright colours are considered
3894:
3524:
3242:
3172:
2975:
Bornschein, M.R.; L.F. Ribeiro; D,C. Blackburn; E.L. Stanley; M.R. Pie (2016).
2910:
2105:
1970:
1929:
1854:
1808:
1231:
970:
775:
483:
459:
306:
The common name "saddleback toad" refers to the bony shields that cover their
260:
237:
4033:
3662:
3548:
3501:
3454:
3219:"Evidence of auditory insensitivity to vocalization frequencies in two frogs"
3086:"Advertisement call of Brachycephalus albolineatus (Anura: Brachycephalidae)"
2656:
2469:
1862:
850:— is not detected by sensory hairs and makes it difficult for individuals of
730:
In general, saddleback toads are reported to use axillary or nearly axillary
3580:
Semicircular canal size constrains vestibular function in miniaturized frogs
3476:
Beck, James C.; Gilland, Edwin; Tank, David W.; Baker, Robert (2004-12-01).
1010:
856:
843:
791:
719:
663:
399:
355:
at least 14.8 mm (0.58 in). At 8–8.4 mm (0.31–0.33 in),
97:
67:
3955:
3781:
3566:
3509:
3462:
3446:
3260:
3190:
3121:
3059:
3012:
2878:
2756:
2674:
2596:
2487:
2387:
2113:
2017:
1978:
1870:
634:
Although not very loud for a frog/toad (even relative to their tiny size),
4085:
3493:
4027:
3914:
3350:"Unidades de conservação reafirman sua importância para a biodiversidade"
1026:
759:
731:
708:
514:
307:
87:
1839:(Anura: Brachycephalidae) from Serra do Brigadeiro, southeastern Brazil"
1789:
4137:
3795:
Heyer, W.R.; A.S. Rand; C.A.G. Cruz; O.L. Peixoto; C.E. Nelson (1990).
3762:
3149:"Intense bone fluorescence reveals hidden patterns in pumpkin toadlets"
3102:
2993:
2956:
2859:
2813:
2796:
2773:
2647:
2577:
2460:
2293:
1691:
Mângia, Santana, Drummond, Sabagh, Ugioni, Costa & Wachlevski, 2023
982:
species have also been found to be afflicted by recurrent outbreaks of
743:
268:
2168:
2151:
2056:
2039:
906:
Only 11 species of saddleback toads have been officially rated by the
842:
in the genus are too small for dependable locomotion, and in turn the
4150:
3352:. Infraestrutura e Meio Ambiente, Government of SĂŁo Paulo state. 2018
3327:
834:
are hindered in their ability to land reliably after jumping. Due to
799:
783:
763:
443:
Brazilian gold frog (sometimes misapplied to the all-brown flea frog
241:
77:
4004:
2948:
2559:
Ribeiro LF; Blackburn DC; Stanley EL; Pie MR; Bornschein MR (2017).
2285:
1509:
Folly, Condez, Vrcibradic, Rocha, Machado, Lopes & Pombal, 2024
513:, which is visible through the skin of the toadlet when exposed to
407:, a species where adults are at least 12.5 mm (0.49 in).
1137:
1046:
1039:
and they belong together as the only members of Brachycephalidae.
795:
771:
699:
621:
294:
224:
206:
359:
has the smallest maximum adult size reported in the genus, while
3371:
Zweifel, Richard G. (1998). Cogger, H.G.; Zweifel, R.G. (eds.).
1166:
907:
779:
220:
107:
4008:
1908:
Napoli, M.F.; Caramaschi, U.; Cruz, C.A.G.; Dias, I.R. (2011).
4124:
2795:
Bornschein, M.R.; L.F. Ribeiro; L. Teixeira; M.R. Pie (2019).
755:
1788:
Hedges, S. B.; Duellman, W. E. & Heinicke, M. P. (2008).
1145:, one of the species initially placed in the flea frog genus
692:
strongly suggest that they are able to hear their own calls.
1202:
was the only widely recognised species in the genus; it was
1130:, but these species pairs are more distant from each other.
654:
Peculiarly, at least the brightly coloured pumpkin toadlets
531:
Primary examples of colours and patterns in pumpkin toadlets
3700:. Version 6.0. American Museum of Natural History, New York
330:) that have an appearance and morphology that is more like
1651:
Almeida-Silva, Silva-Soares, Rodrigues & Verdade, 2021
2433:
2431:
2429:
2427:
2425:
2423:
2421:
2419:
2417:
1226:
in 2002. As of 2020, there were 36 recognised species of
1009:, but were historically included in the true toad family
2768:
2766:
2415:
2413:
2411:
2409:
2407:
2405:
2403:
2401:
2399:
2397:
1178:. In 1920, four additional "variants" were described by
2222:
Fontoura, P.L.; L.F. Ribeiro; M.R. Pie. Haddad (2011).
240:. At about 1 cm (0.4 in) or less in snout–to–
1218:
were instated as valid species in 2010 and the former
1013:, where one of the defining features is its presence.
910:, all between 2004 and 2010, with 3 species listed as
3397:"Here's why pumpkin toadlets are such clumsy jumpers"
1849:(4). Auckland, New Zealand: Magnolia Press: 327–344.
474:
and similar toxins in their skin and organs, whereas
3289:"These tiny frogs can't hear their own mating songs"
1835:"The dark side of pumpkin toadlet: a new species of
1314:, the following species are recognised in the genus
4017:
3698:
Amphibian Species of the World, an Online Reference
1756:
1754:
1752:
1750:
1748:
1746:
367:in the Americas, with only the northeast Brazilian
251:The saddleback toads live among leaf litter in the
3973:(Anura: Brachycephalidae), from southern Brazil".
3857:"Seven tiny frog species found on seven mountains"
1418:Condez, Clemente-Carvalho, Haddad & Reis, 2014
3825:: CS1 maint: DOI inactive as of September 2024 (
3079:
3077:
3075:
3073:
3071:
3069:
2970:
2968:
2966:
2190:"Seven New Mini-Frogs Found—Among Smallest Known"
2145:
2143:
2150:Bornschein, M.R.; M.R. Pie; L. Teixeira (2019).
2141:
2139:
2137:
2135:
2133:
2131:
2129:
2127:
2125:
2123:
486:, but they are possibly absorbed from the small
3423:"Size Limitations in Semicircular Duct Systems"
2444:(Anura: Brachycephalidae) from southern Brazil"
2217:
2215:
1828:
1826:
1824:
1822:
1820:
1818:
3282:
3280:
3278:
2554:
2552:
2550:
2548:
2546:
2544:
2542:
2340:"Diversity of miniaturized frogs of the genus
2033:
2031:
2029:
2027:
1951:
1949:
1947:
1945:
1943:
1941:
1939:
3719:
3717:
3715:
3611:
3609:
3212:
3210:
3208:
3206:
3204:
3202:
3200:
3141:
3139:
2930:
2928:
2505:
2503:
2501:
2499:
2497:
2332:
2330:
2267:
2265:
2263:
2261:
2259:
2257:
2255:
2253:
2251:
2087:
675:to lose it because of their toxicity. Unlike
498:, or produced by bacteria inside their body.
8:
3032:
3030:
2892:
2890:
2888:
2183:
2181:
2179:
2085:
2083:
2081:
2079:
2077:
2075:
2073:
2071:
2069:
2067:
1903:
1901:
1899:
1897:
1895:
1893:
1891:
1889:
1887:
1783:
1781:
1779:
1990:
1988:
1017:was subsequently placed in Atelopodidae or
4005:
3850:
3848:
3846:
3844:
3842:
3840:
3838:
3836:
3375:. San Diego: Academic Press. p. 103.
2836:
2834:
2832:
2689:"10 Smallest Frogs and Toads in the World"
2624:
2622:
2620:
2618:
2616:
2614:
160:
29:
20:
3945:
3935:
3771:
3761:
3652:
3556:
3250:
3180:
3111:
3101:
3002:
2992:
2868:
2858:
2812:
2776:. Berkeley, California: AmphibiaWeb. 2019
2746:
2736:
2664:
2646:
2586:
2576:
2477:
2459:
2377:
2367:
2309:Biological Journal of the Linnean Society
2239:
2167:
2055:
1764:. Berkeley, California: AmphibiaWeb. 2020
1641:Napoli, Caramaschi, Cruz & Dias, 2011
809:
501:At least two brightly coloured species,
3684:
3682:
3680:
3678:
3676:
3674:
3672:
3373:Encyclopedia of Reptiles and Amphibians
1742:
651:, speed, structure and other features.
536:
244:length, the flea frogs are some of the
3818:
3744:Bornschein, M.R.; et al. (2016).
1621:Alves, Sawaya, Reis & Haddad, 2009
1610:Pombal, Wistuba & Bornschein, 1998
1438:GuimarĂŁes, Luz, Rocha & Feio, 2017
1114:relationships. For example, the brown
838:' extremely small head and body size,
3813:10.11606/issn.2176-7793.v31i4p231-410
1164:in 1824, but originally in the genus
930:. Two of the data deficient species,
7:
2691:. gonefroggin.com. 18 December 2017
2440:"Seven new microendemic species of
1910:"A new species of flea-toad, genus
1588:(Serra Cantareira saddleback toad)
961:. Each of the remaining species of
914:(not threatened) and the remaining
859:) as protection against predators.
3987:10.1655/herpetologica-d-11-00074.1
3694:"Search for Taxon: Brachycephalus"
826:jumping, showing a lack of balance
14:
4218:Amphibians of the Atlantic Forest
3287:Supriya, L. (26 September 2017).
1126:is closely related to the golden
1118:is closely related to the golden
688:and the dull-coloured flea frog
594:(dull with bright markings below)
564:(bright with dull sections above)
3217:Goutte, S.; et al. (2017).
2841:Pie, M.R.; L.F. Ribeiro (2015).
2321:10.1111/j.1095-8312.2010.01375.x
1280:
1269:
1258:
1247:
880:
871:
599:
584:
569:
554:
539:
54:
4228:Taxa named by Leopold Fitzinger
3594:. IUCN Redlist. 2 February 2020
2241:10.1590/S1984-46702011000500019
209:of tiny toads and frogs in the
3855:Webb, Jonathan (4 June 2015).
3541:10.1523/JNEUROSCI.1288-08.2008
3427:Journal of Theoretical Biology
1499:Clemente-Carvalho et al., 2012
1312:Amphibian Species of the World
1194:, but placed in its own genus
985:Batrachochytrium dendrobatidis
423:The flea frogs or flea toads (
1:
3052:10.1016/j.toxicon.2004.09.016
2010:10.1016/s0041-0101(01)00279-3
1519:(Giaretta & Sawaya, 1998)
1082:, but it is now considered a
609:(fluorescence under UV-light)
490:they eat, as known from some
3937:10.1371/journal.pone.0244812
3654:10.1016/j.biocon.2021.109246
2738:10.1371/journal.pone.0213314
2525:10.1080/00222933.2012.713525
2369:10.1371/journal.pone.0201781
1565:Brachycephalus mariaeterezae
1559:Pombal & Izecksohn, 2011
1475:Brachycephalus fuscolineatus
1348:Pombal & Gasparini, 2006
1170:. Two years later the genus
993:Classification and discovery
2188:Bates, Mary (4 June 2015).
1657:Brachycephalus quiririensis
1555:Brachycephalus margaritatus
1364:Brachycephalus auroguttatus
1334:Brachycephalus albolineatus
1190:, was described in 1971 by
4244:
3895:10.11646/zootaxa.4290.2.12
3482:Journal of Neurophysiology
3243:10.1038/s41598-017-12145-5
3173:10.1038/s41598-019-41959-8
2911:10.1670/0022-1511-43.2.212
2774:"Brachycephalus ephippium"
2513:Journal of Natural History
1727:Brachycephalus vertebralis
1707:Brachycephalus tridactylus
1667:Brachycephalus rotenbergae
1515:Brachycephalus hermogenesi
1206:pumpkin toadlet. In 1990,
626:Unique among all animals,
482:species attain the strong
3421:Muller, M. (1999-06-07).
2106:10.11646/zootaxa.4083.1.2
1971:10.11646/zootaxa.4407.4.2
1930:10.11646/zootaxa.2739.1.3
1855:10.11646/zootaxa.4258.4.2
1809:10.11646/zootaxa.1737.1.1
1717:Brachycephalus verrucosus
1677:Brachycephalus sulfuratus
1575:Brachycephalus mirissimus
1535:Brachycephalus izecksohni
1465:Brachycephalus ferruginus
1457:(Spix’s saddleback toad)
1444:Brachycephalus didactylus
1394:Brachycephalus bufonoides
1354:Brachycephalus atelopoide
1180:AlĂpio de Miranda-Ribeiro
184:
177:
168:
159:
151:
147:
142:
51:Scientific classification
49:
37:
28:
23:
2194:National Geographic News
1687:Brachycephalus tabuleiro
1596:Brachycephalus olivaceus
1585:Brachycephalus nodoterga
1545:Brachycephalus leopardus
1505:Brachycephalus herculeus
1485:Brachycephalus garbeanus
1454:Brachycephalus ephippium
1404:Brachycephalus coloratus
1374:Brachycephalus boticario
1222:species were moved into
1154:Brachycephalus ephippium
1043:Genus and species groups
953:and the pumpkin toadlet
830:Unlike most amphibians,
746:or other aquatic stage.
376:Eleutherodactylus iberia
43:Brachycephalus ephippium
4223:Endemic frogs of Brazil
3641:Biological Conservation
3529:Journal of Neuroscience
1661:Pie & Ribeiro, 2015
1525:Brachycephalus ibitinga
1434:Brachycephalus darkside
1424:Brachycephalus curupira
1384:Brachycephalus brunneus
1338:Bornschein et al., 2016
1162:Johann Baptist von Spix
1035:is a close relative of
846:— which is regarded by
3815:(inactive 2024-09-12).
3447:10.1006/jtbi.1999.0922
3316:Journal of Herpetology
2937:Journal of Herpetology
2899:Journal of Herpetology
2274:Journal of Herpetology
1627:Brachycephalus pombali
1619:(Red pumpkin toadlet)
1616:Brachycephalus pitanga
1495:Brachycephalus guarani
1414:Brachycephalus crispus
1344:Brachycephalus alipioi
1324:Brachycephalus actaeus
1149:
1055:
827:
711:
631:
509:, have bones that are
303:
39:Spix's saddleback toad
3494:10.1152/jn.00311.2004
1606:Brachycephalus pernix
1590:Miranda-Ribeiro, 1920
1489:Miranda-Ribeiro, 1920
1398:Miranda-Ribeiro, 1920
1358:Miranda-Ribeiro, 1920
1328:Monteiro et al., 2018
1141:
1050:
941:The three flea frogs
928:critically endangered
899:critically endangered
820:
703:
673:evolutionary pressure
625:
298:
223:, ranging from south
3801:Arquivos de Zoologia
2519:(39–40): 2497–2507.
1721:Ribeiro et al., 2015
1637:Brachycephalus pulex
1600:Ribeiro et al., 2015
1569:Ribeiro et al., 2015
1549:Ribeiro et al., 2015
1539:Ribeiro et al., 2005
1479:Ribeiro et al., 2015
1428:Ribeiro et al., 2017
1408:Ribeiro et al., 2017
1388:Ribeiro et al., 2005
1378:Ribeiro et al., 2015
1368:Ribeiro et al., 2015
1198:. As a consequence,
774:) and their larvae,
370:Adelophryne michelin
274:The majority of the
3928:2021PLoSO..1644812N
3797:"Frogs of Boracéia"
3618:Zoologia (Curitiba)
3439:1999JThBi.198..405M
3235:2017NatSR...712121G
3165:2019NatSR...9.5388G
2729:2019PLoSO..1413314S
2360:2018PLoSO..1301781L
2228:Zoologia (Curitiba)
1701:Haddad et al., 2010
1697:Brachycephalus toby
1681:Condez et al., 2016
1647:Brachycephalus puri
1529:Condez et al., 2021
1186:. Another species,
1152:The first species,
1102:) and the northern
863:Conservation status
523:species recognition
411:Colour and toxicity
291:Morphology and size
253:Atlantic rainforest
16:Genus of amphibians
3763:10.7717/peerj.2490
3223:Scientific Reports
3153:Scientific Reports
3103:10.7717/peerj.5273
2994:10.7717/peerj.2629
2860:10.7717/peerj.1179
2814:10.15560/15.6.1031
2648:10.7717/peerj.5683
2578:10.7717/peerj.3603
2461:10.7717/peerj.1011
1762:"Brachycephalidae"
1711:Garey et al., 2012
1671:Nunes et al., 2021
1631:Alves et al., 2006
1469:Alves et al., 2006
1150:
1056:
840:vestibular systems
828:
712:
632:
304:
259:. They are mostly
4200:
4199:
4185:Open Tree of Life
4011:Taxon identifiers
3727:Amphibia-Reptilia
3535:(32): 8086–8095.
2169:10.3390/d11090150
2057:10.3390/d11090150
1732:
1722:
1712:
1702:
1692:
1682:
1672:
1662:
1652:
1642:
1632:
1622:
1611:
1601:
1591:
1580:
1570:
1560:
1550:
1540:
1530:
1520:
1510:
1500:
1490:
1480:
1470:
1460:
1449:
1448:(Izecksohn, 1971)
1439:
1429:
1419:
1409:
1399:
1389:
1379:
1369:
1359:
1349:
1339:
1329:
1192:EugĂŞnio Izecksohn
1176:Leopold Fitzinger
818:
618:Voice and hearing
492:poison dart frogs
195:
194:
138:
24:Saddleback toads
4235:
4193:
4192:
4180:
4179:
4167:
4166:
4154:
4153:
4141:
4140:
4128:
4127:
4115:
4114:
4102:
4101:
4089:
4088:
4079:
4078:
4066:
4065:
4053:
4052:
4051:
4038:
4037:
4036:
4006:
3999:
3998:
3966:
3960:
3959:
3949:
3939:
3905:
3899:
3898:
3878:
3872:
3871:
3869:
3867:
3852:
3831:
3830:
3824:
3816:
3792:
3786:
3785:
3775:
3765:
3741:
3735:
3734:
3721:
3710:
3709:
3707:
3705:
3690:Frost, Darrel R.
3686:
3667:
3666:
3656:
3632:
3626:
3625:
3613:
3604:
3603:
3601:
3599:
3592:"Brachycephalus"
3588:
3582:
3577:
3571:
3570:
3560:
3520:
3514:
3513:
3488:(6): 3546–3561.
3473:
3467:
3466:
3418:
3412:
3411:
3409:
3408:
3393:
3387:
3386:
3368:
3362:
3361:
3359:
3357:
3346:
3340:
3339:
3328:10.1670/07-287.1
3310:
3304:
3303:
3301:
3299:
3284:
3273:
3272:
3254:
3214:
3195:
3194:
3184:
3143:
3134:
3133:
3115:
3105:
3081:
3064:
3063:
3034:
3025:
3024:
3006:
2996:
2972:
2961:
2960:
2932:
2923:
2922:
2894:
2883:
2882:
2872:
2862:
2838:
2827:
2826:
2816:
2807:(6): 1031–1036.
2792:
2786:
2785:
2783:
2781:
2770:
2761:
2760:
2750:
2740:
2707:
2701:
2700:
2698:
2696:
2685:
2679:
2678:
2668:
2650:
2626:
2609:
2608:
2590:
2580:
2556:
2537:
2536:
2507:
2492:
2491:
2481:
2463:
2435:
2392:
2391:
2381:
2371:
2334:
2325:
2324:
2304:
2298:
2297:
2269:
2246:
2245:
2243:
2219:
2210:
2209:
2207:
2205:
2196:. Archived from
2185:
2174:
2173:
2171:
2147:
2118:
2117:
2089:
2062:
2061:
2059:
2035:
2022:
2021:
1992:
1983:
1982:
1953:
1934:
1933:
1905:
1882:
1881:
1879:
1877:
1830:
1813:
1812:
1794:
1785:
1774:
1773:
1771:
1769:
1758:
1730:
1720:
1710:
1700:
1690:
1680:
1670:
1660:
1650:
1640:
1630:
1620:
1609:
1599:
1589:
1579:Pie et al., 2018
1578:
1568:
1558:
1548:
1538:
1528:
1518:
1508:
1498:
1488:
1478:
1468:
1458:
1447:
1437:
1427:
1417:
1407:
1397:
1387:
1377:
1367:
1357:
1347:
1337:
1327:
1297:B. mariaeterezae
1284:
1273:
1262:
1251:
1214:species to two.
1019:Brachycephalidae
884:
875:
848:Poiseuille's law
819:
603:
588:
579:(overall bright)
573:
562:B. fuscolineatus
558:
543:
214:Brachycephalidae
199:saddleback toads
169:Distribution of
164:
133:
118:Brachycephalidae
59:
58:
33:
21:
4243:
4242:
4238:
4237:
4236:
4234:
4233:
4232:
4203:
4202:
4201:
4196:
4188:
4183:
4175:
4170:
4162:
4157:
4149:
4144:
4136:
4131:
4123:
4118:
4110:
4105:
4097:
4092:
4084:
4082:
4074:
4069:
4061:
4056:
4047:
4046:
4041:
4032:
4031:
4026:
4013:
4003:
4002:
3968:
3967:
3963:
3922:(4): e0244812.
3907:
3906:
3902:
3880:
3879:
3875:
3865:
3863:
3854:
3853:
3834:
3817:
3794:
3793:
3789:
3743:
3742:
3738:
3723:
3722:
3713:
3703:
3701:
3688:
3687:
3670:
3634:
3633:
3629:
3615:
3614:
3607:
3597:
3595:
3590:
3589:
3585:
3578:
3574:
3522:
3521:
3517:
3475:
3474:
3470:
3420:
3419:
3415:
3406:
3404:
3395:
3394:
3390:
3383:
3370:
3369:
3365:
3355:
3353:
3348:
3347:
3343:
3312:
3311:
3307:
3297:
3295:
3286:
3285:
3276:
3216:
3215:
3198:
3145:
3144:
3137:
3083:
3082:
3067:
3036:
3035:
3028:
2974:
2973:
2964:
2949:10.2307/1565481
2934:
2933:
2926:
2896:
2895:
2886:
2853:(1792): e1179.
2840:
2839:
2830:
2794:
2793:
2789:
2779:
2777:
2772:
2771:
2764:
2723:(3): e0213314.
2709:
2708:
2704:
2694:
2692:
2687:
2686:
2682:
2628:
2627:
2612:
2558:
2557:
2540:
2509:
2508:
2495:
2437:
2436:
2395:
2354:(9): e0201781.
2336:
2335:
2328:
2306:
2305:
2301:
2286:10.2307/1564972
2271:
2270:
2249:
2221:
2220:
2213:
2203:
2201:
2200:on June 6, 2015
2187:
2186:
2177:
2149:
2148:
2121:
2091:
2090:
2065:
2037:
2036:
2025:
1994:
1993:
1986:
1955:
1954:
1937:
1907:
1906:
1885:
1875:
1873:
1832:
1831:
1816:
1792:
1787:
1786:
1777:
1767:
1765:
1760:
1759:
1744:
1739:
1308:
1307:
1306:
1305:
1304:
1289:B. auroguttatus
1287:Top to bottom:
1285:
1276:
1275:
1274:
1265:
1264:
1263:
1254:
1253:
1252:
1241:
1136:
1045:
1000:
995:
959:species complex
904:
903:
902:
901:
895:B. quiririensis
887:
886:
885:
877:
876:
865:
810:
808:
788:pseudoscorpions
752:
728:
698:
620:
615:
614:
613:
610:
604:
595:
589:
580:
574:
565:
559:
550:
544:
533:
532:
413:
353:B. margaritatus
293:
288:
263:, feed on tiny
155:
132:
53:
17:
12:
11:
5:
4241:
4239:
4231:
4230:
4225:
4220:
4215:
4213:Brachycephalus
4205:
4204:
4198:
4197:
4195:
4194:
4181:
4168:
4155:
4142:
4129:
4116:
4103:
4090:
4080:
4076:Brachycephalus
4067:
4063:Brachycephalus
4054:
4049:Brachycephalus
4039:
4023:
4021:
4019:Brachycephalus
4015:
4014:
4009:
4001:
4000:
3981:(2): 266–271.
3971:Brachycephalus
3961:
3900:
3889:(2): 395–400.
3873:
3832:
3807:(4): 231–410.
3787:
3736:
3711:
3668:
3627:
3605:
3583:
3572:
3515:
3468:
3433:(3): 405–437.
3413:
3388:
3381:
3363:
3341:
3322:(3): 542–549.
3305:
3274:
3196:
3135:
3065:
3026:
2962:
2924:
2905:(2): 212–219.
2884:
2828:
2787:
2762:
2702:
2680:
2610:
2538:
2493:
2442:Brachycephalus
2393:
2326:
2315:(4): 752–767.
2299:
2280:(4): 516–519.
2247:
2234:(5): 687–689.
2211:
2175:
2119:
2063:
2023:
1984:
1965:(4): 483–505.
1935:
1912:Brachycephalus
1883:
1837:Brachycephalus
1814:
1775:
1741:
1740:
1738:
1735:
1734:
1733:
1723:
1713:
1703:
1693:
1683:
1673:
1663:
1653:
1643:
1633:
1623:
1612:
1602:
1592:
1581:
1571:
1561:
1551:
1541:
1531:
1521:
1511:
1501:
1491:
1481:
1471:
1461:
1450:
1440:
1430:
1420:
1410:
1400:
1390:
1380:
1370:
1360:
1350:
1340:
1330:
1316:Brachycephalus
1286:
1279:
1278:
1277:
1268:
1267:
1266:
1257:
1256:
1255:
1246:
1245:
1244:
1243:
1242:
1240:
1237:
1228:Brachycephalus
1224:Brachycephalus
1212:Brachycephalus
1174:was coined by
1172:Brachycephalus
1156:, was already
1143:B. hermogenesi
1135:
1132:
1122:and the brown
1100:Santa Catarina
1088:Brachycephalus
1068:B. hermogenesi
1060:Brachycephalus
1044:
1041:
1037:Brachycephalus
1023:Brachycephalus
1015:Brachycephalus
1007:Bidder's organ
1003:Brachycephalus
999:
996:
994:
991:
980:Brachycephalus
976:climate change
963:Brachycephalus
955:B. ephippiumsi
947:B. hermogenesi
916:data deficient
889:
888:
879:
878:
870:
869:
868:
867:
866:
864:
861:
852:Brachycephalus
836:Brachycephalus
832:Brachycephalus
807:
804:
751:
748:
727:
724:
697:
694:
690:B. hermogenesi
636:Brachycephalus
619:
616:
612:
611:
605:
598:
596:
590:
583:
581:
575:
568:
566:
560:
553:
551:
549:(overall dull)
545:
538:
535:
534:
530:
529:
528:
496:mantella frogs
480:Brachycephalus
429:B. hermogenesi
417:Brachycephalus
415:The different
412:
409:
365:smallest frogs
344:Brachycephalus
320:B. hermogenesi
292:
289:
287:
284:
276:Brachycephalus
248:in the world.
246:smallest frogs
229:Santa Catarina
203:Brachycephalus
193:
192:
191:
190:
182:
181:
175:
174:
171:Brachycephalus
166:
165:
157:
156:
150:More than 35,
148:
145:
144:
140:
139:
129:Brachycephalus
125:
121:
120:
115:
111:
110:
105:
101:
100:
95:
91:
90:
85:
81:
80:
75:
71:
70:
65:
61:
60:
47:
46:
35:
34:
26:
25:
15:
13:
10:
9:
6:
4:
3:
2:
4240:
4229:
4226:
4224:
4221:
4219:
4216:
4214:
4211:
4210:
4208:
4191:
4186:
4182:
4178:
4173:
4169:
4165:
4160:
4156:
4152:
4147:
4143:
4139:
4134:
4130:
4126:
4121:
4117:
4113:
4108:
4104:
4100:
4095:
4091:
4087:
4081:
4077:
4072:
4068:
4064:
4059:
4055:
4050:
4044:
4040:
4035:
4029:
4025:
4024:
4022:
4020:
4016:
4012:
4007:
3996:
3992:
3988:
3984:
3980:
3976:
3975:Herpetologica
3972:
3965:
3962:
3957:
3953:
3948:
3943:
3938:
3933:
3929:
3925:
3921:
3917:
3916:
3911:
3904:
3901:
3896:
3892:
3888:
3884:
3877:
3874:
3862:
3858:
3851:
3849:
3847:
3845:
3843:
3841:
3839:
3837:
3833:
3828:
3822:
3814:
3810:
3806:
3802:
3798:
3791:
3788:
3783:
3779:
3774:
3769:
3764:
3759:
3755:
3751:
3747:
3740:
3737:
3733:(2): 225–227.
3732:
3728:
3720:
3718:
3716:
3712:
3699:
3695:
3691:
3685:
3683:
3681:
3679:
3677:
3675:
3673:
3669:
3664:
3660:
3655:
3650:
3646:
3642:
3638:
3631:
3628:
3623:
3619:
3612:
3610:
3606:
3593:
3587:
3584:
3581:
3576:
3573:
3568:
3564:
3559:
3554:
3550:
3546:
3542:
3538:
3534:
3530:
3526:
3519:
3516:
3511:
3507:
3503:
3499:
3495:
3491:
3487:
3483:
3479:
3472:
3469:
3464:
3460:
3456:
3452:
3448:
3444:
3440:
3436:
3432:
3428:
3424:
3417:
3414:
3402:
3398:
3392:
3389:
3384:
3382:0-12-178560-2
3378:
3374:
3367:
3364:
3351:
3345:
3342:
3337:
3333:
3329:
3325:
3321:
3317:
3309:
3306:
3294:
3290:
3283:
3281:
3279:
3275:
3270:
3266:
3262:
3258:
3253:
3248:
3244:
3240:
3236:
3232:
3228:
3224:
3220:
3213:
3211:
3209:
3207:
3205:
3203:
3201:
3197:
3192:
3188:
3183:
3178:
3174:
3170:
3166:
3162:
3158:
3154:
3150:
3142:
3140:
3136:
3131:
3127:
3123:
3119:
3114:
3109:
3104:
3099:
3095:
3091:
3087:
3080:
3078:
3076:
3074:
3072:
3070:
3066:
3061:
3057:
3053:
3049:
3045:
3041:
3033:
3031:
3027:
3022:
3018:
3014:
3010:
3005:
3000:
2995:
2990:
2987:(10): e2629.
2986:
2982:
2978:
2971:
2969:
2967:
2963:
2958:
2954:
2950:
2946:
2942:
2938:
2931:
2929:
2925:
2920:
2916:
2912:
2908:
2904:
2900:
2893:
2891:
2889:
2885:
2880:
2876:
2871:
2866:
2861:
2856:
2852:
2848:
2844:
2837:
2835:
2833:
2829:
2824:
2820:
2815:
2810:
2806:
2802:
2798:
2791:
2788:
2775:
2769:
2767:
2763:
2758:
2754:
2749:
2744:
2739:
2734:
2730:
2726:
2722:
2718:
2714:
2706:
2703:
2690:
2684:
2681:
2676:
2672:
2667:
2662:
2658:
2654:
2649:
2644:
2640:
2636:
2632:
2625:
2623:
2621:
2619:
2617:
2615:
2611:
2606:
2602:
2598:
2594:
2589:
2584:
2579:
2574:
2570:
2566:
2562:
2555:
2553:
2551:
2549:
2547:
2545:
2543:
2539:
2534:
2530:
2526:
2522:
2518:
2514:
2506:
2504:
2502:
2500:
2498:
2494:
2489:
2485:
2480:
2475:
2471:
2467:
2462:
2457:
2453:
2449:
2445:
2443:
2434:
2432:
2430:
2428:
2426:
2424:
2422:
2420:
2418:
2416:
2414:
2412:
2410:
2408:
2406:
2404:
2402:
2400:
2398:
2394:
2389:
2385:
2380:
2375:
2370:
2365:
2361:
2357:
2353:
2349:
2345:
2343:
2333:
2331:
2327:
2322:
2318:
2314:
2310:
2303:
2300:
2295:
2291:
2287:
2283:
2279:
2275:
2268:
2266:
2264:
2262:
2260:
2258:
2256:
2254:
2252:
2248:
2242:
2237:
2233:
2229:
2225:
2218:
2216:
2212:
2199:
2195:
2191:
2184:
2182:
2180:
2176:
2170:
2165:
2161:
2157:
2153:
2146:
2144:
2142:
2140:
2138:
2136:
2134:
2132:
2130:
2128:
2126:
2124:
2120:
2115:
2111:
2107:
2103:
2099:
2095:
2088:
2086:
2084:
2082:
2080:
2078:
2076:
2074:
2072:
2070:
2068:
2064:
2058:
2053:
2049:
2045:
2041:
2034:
2032:
2030:
2028:
2024:
2019:
2015:
2011:
2007:
2003:
1999:
1991:
1989:
1985:
1980:
1976:
1972:
1968:
1964:
1960:
1952:
1950:
1948:
1946:
1944:
1942:
1940:
1936:
1931:
1927:
1923:
1919:
1915:
1913:
1904:
1902:
1900:
1898:
1896:
1894:
1892:
1890:
1888:
1884:
1872:
1868:
1864:
1860:
1856:
1852:
1848:
1844:
1840:
1838:
1829:
1827:
1825:
1823:
1821:
1819:
1815:
1810:
1806:
1802:
1798:
1791:
1784:
1782:
1780:
1776:
1763:
1757:
1755:
1753:
1751:
1749:
1747:
1743:
1736:
1729:
1728:
1724:
1719:
1718:
1714:
1709:
1708:
1704:
1699:
1698:
1694:
1689:
1688:
1684:
1679:
1678:
1674:
1669:
1668:
1664:
1659:
1658:
1654:
1649:
1648:
1644:
1639:
1638:
1634:
1629:
1628:
1624:
1618:
1617:
1613:
1608:
1607:
1603:
1598:
1597:
1593:
1587:
1586:
1582:
1577:
1576:
1572:
1567:
1566:
1562:
1557:
1556:
1552:
1547:
1546:
1542:
1537:
1536:
1532:
1527:
1526:
1522:
1517:
1516:
1512:
1507:
1506:
1502:
1497:
1496:
1492:
1487:
1486:
1482:
1477:
1476:
1472:
1467:
1466:
1462:
1456:
1455:
1451:
1446:
1445:
1441:
1436:
1435:
1431:
1426:
1425:
1421:
1416:
1415:
1411:
1406:
1405:
1401:
1396:
1395:
1391:
1386:
1385:
1381:
1376:
1375:
1371:
1366:
1365:
1361:
1356:
1355:
1351:
1346:
1345:
1341:
1336:
1335:
1331:
1326:
1325:
1321:
1320:
1319:
1317:
1313:
1310:According to
1302:
1301:B. verrucosus
1298:
1294:
1290:
1283:
1272:
1261:
1250:
1238:
1236:
1233:
1229:
1225:
1221:
1215:
1213:
1209:
1205:
1201:
1197:
1193:
1189:
1188:B. didactylus
1185:
1181:
1177:
1173:
1169:
1168:
1163:
1159:
1155:
1148:
1144:
1140:
1133:
1131:
1129:
1125:
1121:
1120:B. izecksohni
1117:
1113:
1109:
1105:
1101:
1097:
1093:
1089:
1085:
1081:
1077:
1076:B. sulfuratus
1073:
1069:
1065:
1064:B. didactylus
1061:
1053:
1049:
1042:
1040:
1038:
1034:
1033:
1028:
1024:
1020:
1016:
1012:
1008:
1004:
997:
992:
990:
987:
986:
981:
977:
972:
968:
964:
960:
956:
952:
951:B. sulfuratus
948:
944:
943:B. didactylus
939:
937:
936:B. bufonoides
933:
932:B. atelopoide
929:
925:
921:
917:
913:
912:least concern
909:
900:
896:
892:
883:
874:
862:
860:
858:
853:
849:
845:
841:
837:
833:
825:
824:
805:
803:
801:
797:
793:
789:
785:
781:
777:
773:
769:
765:
761:
757:
749:
747:
745:
739:
737:
733:
725:
723:
721:
716:
710:
706:
702:
695:
693:
691:
687:
682:
678:
674:
669:
665:
661:
657:
652:
650:
644:
642:
637:
629:
624:
617:
608:
602:
597:
593:
587:
582:
578:
577:B. izecksohni
572:
567:
563:
557:
552:
548:
542:
537:
527:
524:
520:
516:
512:
508:
504:
499:
497:
493:
489:
488:invertebrates
485:
481:
477:
473:
469:
465:
461:
456:
454:
450:
446:
445:B. didactylus
440:
438:
437:B. sulfuratus
434:
430:
426:
425:B. didactylus
421:
418:
410:
408:
406:
402:
401:
396:
395:
390:
389:
384:
383:
378:
377:
372:
371:
366:
362:
361:B. sulfuratus
358:
354:
350:
345:
341:
339:
338:
333:
332:leptodactylid
329:
328:B. sulfuratus
325:
321:
317:
316:B. didactylus
313:
309:
301:
297:
290:
285:
283:
281:
277:
272:
270:
266:
265:invertebrates
262:
258:
254:
249:
247:
243:
239:
235:
230:
226:
222:
219:
215:
212:
208:
204:
200:
189:
186:
185:
183:
180:
176:
172:
167:
163:
158:
154:
153:
146:
141:
136:
131:
130:
126:
123:
122:
119:
116:
113:
112:
109:
106:
103:
102:
99:
96:
93:
92:
89:
86:
83:
82:
79:
76:
73:
72:
69:
66:
63:
62:
57:
52:
48:
44:
40:
36:
32:
27:
22:
19:
4018:
3978:
3974:
3970:
3964:
3919:
3913:
3903:
3886:
3882:
3876:
3864:. Retrieved
3860:
3821:cite journal
3804:
3800:
3790:
3753:
3749:
3739:
3730:
3726:
3702:. Retrieved
3697:
3644:
3640:
3630:
3621:
3617:
3596:. Retrieved
3586:
3575:
3532:
3528:
3518:
3485:
3481:
3471:
3430:
3426:
3416:
3405:. Retrieved
3403:. 2022-06-15
3401:Science News
3400:
3391:
3372:
3366:
3354:. Retrieved
3344:
3319:
3315:
3308:
3296:. Retrieved
3292:
3229:(1): 12121.
3226:
3222:
3156:
3152:
3093:
3089:
3046:(1): 73–79.
3043:
3039:
2984:
2980:
2943:(1): 70–74.
2940:
2936:
2902:
2898:
2850:
2846:
2804:
2800:
2790:
2778:. Retrieved
2720:
2716:
2705:
2693:. Retrieved
2683:
2638:
2634:
2568:
2564:
2516:
2512:
2451:
2447:
2441:
2351:
2347:
2341:
2312:
2308:
2302:
2277:
2273:
2231:
2227:
2202:. Retrieved
2198:the original
2193:
2159:
2155:
2100:(1): 40–56.
2097:
2093:
2050:(1): 39–47.
2047:
2043:
2004:(6): 761–6.
2001:
1997:
1962:
1958:
1921:
1917:
1911:
1874:. Retrieved
1846:
1842:
1836:
1800:
1796:
1766:. Retrieved
1731:Pombal, 2001
1725:
1715:
1705:
1695:
1685:
1675:
1665:
1655:
1645:
1635:
1625:
1614:
1604:
1594:
1583:
1573:
1563:
1553:
1543:
1533:
1523:
1513:
1503:
1493:
1483:
1473:
1463:
1459:(Spix, 1824)
1452:
1442:
1432:
1422:
1412:
1402:
1392:
1382:
1372:
1362:
1352:
1342:
1332:
1322:
1315:
1311:
1309:
1300:
1296:
1293:B. coloratus
1292:
1288:
1227:
1223:
1220:Psyllophryne
1219:
1216:
1211:
1208:B. nodoterga
1207:
1203:
1200:B. ephippium
1199:
1196:Psyllophryne
1195:
1187:
1184:B. ephippium
1183:
1171:
1165:
1153:
1151:
1147:Psyllophryne
1146:
1142:
1128:B. leopardus
1127:
1123:
1119:
1115:
1112:phylogenetic
1104:B. ephippium
1103:
1091:
1087:
1080:Psyllophryne
1079:
1075:
1071:
1067:
1063:
1059:
1057:
1052:Phylogenetic
1036:
1030:
1022:
1014:
1002:
1001:
983:
979:
967:cloud forest
962:
954:
950:
946:
942:
940:
935:
931:
905:
894:
890:
851:
835:
831:
829:
823:B. coloratus
821:
768:grasshoppers
753:
740:
736:B. ephippium
735:
729:
717:
713:
705:B. leopardus
704:
689:
685:
680:
677:B. ephippium
676:
668:vestigiality
659:
656:B. ephippium
655:
653:
645:
635:
633:
627:
606:
592:B. olivaceus
591:
576:
561:
546:
506:
503:B. ephippium
502:
500:
479:
476:B. nodoterga
475:
472:tetrodotoxin
467:
464:B. ephippium
463:
457:
452:
448:
444:
441:
436:
432:
428:
424:
422:
416:
414:
405:B. ephippium
404:
398:
392:
386:
380:
374:
373:, and Cuban
368:
360:
356:
352:
348:
343:
342:
335:
327:
323:
319:
315:
305:
299:
275:
273:
257:cloud forest
250:
234:Psyllophryne
233:
202:
198:
196:
188:Psyllophryne
187:
170:
149:
128:
127:
42:
18:
4146:iNaturalist
4043:Wikispecies
3159:(1): 5388.
2342:Adelophryne
2204:11 February
1232:sky islands
1124:B. brunneus
1116:B. curupira
1032:Ischnocnema
893:(left) and
891:B. darkside
776:springtails
547:B. curupira
519:mate choice
511:fluorescent
484:neurotoxins
453:B. curupira
449:B. brunneus
394:Paedophryne
382:E. limbatus
349:B. darkside
337:Adelophryne
173:(in black)
4207:Categories
3704:2 February
3647:: 109246.
3598:2 February
3407:2022-09-11
3356:9 February
3298:9 February
3293:ScienceMag
2801:Check List
2780:5 February
2695:30 January
2162:(9): 150.
1768:2 February
1737:References
1106:subgroup (
1094:subgroup (
971:sky island
924:endangered
920:vulnerable
857:osteoderms
792:millipedes
686:B. actaeus
681:B. pitanga
660:B. pitanga
628:B. pitanga
607:B. pitanga
507:B. pitanga
460:aposematic
312:morphology
286:Appearance
280:threatened
238:aposematic
3756:: e2490.
3663:0006-3207
3549:0270-6474
3502:0022-3077
3455:0022-5193
3269:205604026
3096:: e5273.
2823:212771392
2657:2167-8359
2641:: e5683.
2571:: e3603.
2470:2167-8359
2454:: e1011.
2156:Diversity
2044:Diversity
1924:: 33–40.
1863:1175-5334
1803:: 1–182.
1158:described
1108:SĂŁo Paulo
1092:B. pernix
1011:Bufonidae
844:endolymph
760:true bugs
720:nocturnal
664:vocal sac
649:frequency
641:territory
468:B. pernix
400:Stumpffia
308:vertebrae
135:Fitzinger
74:Kingdom:
68:Eukaryota
4083:BioLib:
4028:Wikidata
3995:86572812
3956:33909613
3915:PLOS ONE
3861:BBC News
3782:27761312
3692:(2020).
3567:18685033
3510:15269231
3463:10366494
3336:86638006
3261:28935936
3191:30926879
3130:50776284
3122:30042895
3060:15581685
3021:23294169
3013:27812425
2919:86086374
2879:26339556
2757:30917162
2717:PLOS ONE
2675:30310742
2605:22414191
2597:28761788
2533:86664601
2488:26056613
2388:30231059
2348:PLOS ONE
2114:27394218
2018:12175613
1979:29690168
1871:28609910
1072:B. pulex
1005:lacks a
732:amplexus
726:Breeding
709:amplexus
707:pair in
696:Behavior
515:UV light
433:B. pulex
357:B. pulex
324:B. pulex
300:B. pulex
271:stage).
205:) are a
179:Synonyms
152:see text
143:Species
114:Family:
98:Amphibia
88:Chordata
84:Phylum:
78:Animalia
64:Domain:
4138:2424890
4034:Q634679
3947:8081201
3924:Bibcode
3883:Zootaxa
3773:5068354
3624:: 1–12.
3558:2647017
3435:Bibcode
3252:5608807
3231:Bibcode
3182:6441030
3161:Bibcode
3113:6055690
3040:Toxicon
3004:5088579
2957:1565481
2870:4558078
2748:6436692
2725:Bibcode
2666:6174073
2588:5534166
2479:4458131
2379:6145526
2356:Bibcode
2294:1564972
2094:Zootaxa
1998:Toxicon
1959:Zootaxa
1918:Zootaxa
1843:Zootaxa
1797:Zootaxa
1134:Species
1084:synonym
1058:Within
1027:Genetic
806:Jumping
800:isopods
784:spiders
764:beetles
750:Feeding
744:tadpole
269:tadpole
261:diurnal
216:in the
124:Genus:
104:Order:
94:Class:
4190:719425
4177:164301
4164:208123
4099:257746
4086:178258
3993:
3954:
3944:
3866:4 June
3780:
3770:
3661:
3565:
3555:
3547:
3508:
3500:
3461:
3453:
3379:
3334:
3267:
3259:
3249:
3189:
3179:
3128:
3120:
3110:
3058:
3019:
3011:
3001:
2955:
2917:
2877:
2867:
2821:
2755:
2745:
2673:
2663:
2655:
2603:
2595:
2585:
2531:
2486:
2476:
2468:
2386:
2376:
2292:
2112:
2016:
1977:
1869:
1861:
1096:Paraná
998:Family
926:and 6
796:snails
211:family
137:, 1826
4151:24486
4125:40616
3991:S2CID
3750:PeerJ
3332:S2CID
3265:S2CID
3126:S2CID
3090:PeerJ
3017:S2CID
2981:PeerJ
2953:JSTOR
2915:S2CID
2847:PeerJ
2819:S2CID
2635:PeerJ
2601:S2CID
2565:PeerJ
2529:S2CID
2448:PeerJ
2290:JSTOR
1876:1 May
1793:(PDF)
780:mites
772:flies
470:have
225:Bahia
221:Anura
218:order
207:genus
108:Anura
4172:NCBI
4159:ITIS
4133:GBIF
4112:3C9X
4094:BOLD
3952:PMID
3887:4290
3868:2015
3827:link
3778:PMID
3706:2020
3659:ISSN
3600:2020
3563:PMID
3545:ISSN
3506:PMID
3498:ISSN
3459:PMID
3451:ISSN
3377:ISBN
3358:2020
3300:2020
3257:PMID
3187:PMID
3118:PMID
3056:PMID
3009:PMID
2875:PMID
2782:2020
2753:PMID
2697:2020
2671:PMID
2653:ISSN
2593:PMID
2484:PMID
2466:ISSN
2384:PMID
2206:2020
2110:PMID
2098:4083
2014:PMID
1975:PMID
1963:4407
1922:2739
1878:2017
1867:PMID
1859:ISSN
1847:4258
1801:1737
1770:2020
1299:and
1239:List
1167:Bufo
1098:and
1074:and
949:and
934:and
922:, 5
908:IUCN
798:and
770:and
756:ants
658:and
505:and
494:and
466:and
451:and
435:and
397:and
388:Mini
379:and
351:and
326:and
242:vent
197:The
4120:EoL
4107:CoL
4071:ASW
4058:ADW
3983:doi
3942:PMC
3932:doi
3891:doi
3809:doi
3768:PMC
3758:doi
3649:doi
3645:261
3622:526
3553:PMC
3537:doi
3490:doi
3443:doi
3431:198
3324:doi
3247:PMC
3239:doi
3177:PMC
3169:doi
3108:PMC
3098:doi
3048:doi
2999:PMC
2989:doi
2945:doi
2907:doi
2865:PMC
2855:doi
2809:doi
2743:PMC
2733:doi
2661:PMC
2643:doi
2583:PMC
2573:doi
2521:doi
2474:PMC
2456:doi
2374:PMC
2364:doi
2317:doi
2282:doi
2236:doi
2164:doi
2102:doi
2052:doi
2006:doi
1967:doi
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