1059:
884:
567:
712:
597:
458:). 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,
582:
552:
1282:
1260:
893:
67:
1293:
1150:
42:
1271:
634:
173:
612:
466:, 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.
307:
929:(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
1228:
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
657:
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
357:
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
752:
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.
453:
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
1245:
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.
984:
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
725:
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
649:
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
2521:
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".
321:, 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
694:
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
681:
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
753:
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
822:
999:
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.
351:. 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.
865:
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
1967:
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".
3324:
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)".
2006:
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)".
3157:
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).
536:
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.
242:
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
2721:
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).
3735:
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)".
2348:
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).
358:
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
430:
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.
3048:
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".
827:
826:
823:
1032:, 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
828:
2908:
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".
2103:
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".
749:
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.
976:
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
2808:"New altitudinal record for Brachycephalus actaeus Monteiro, Condez, Garcia, Comitti, Amaral & Haddad, 2018 (Anura, Brachycephalidae), with comments on its habitats of occurrence"
2724:"Morphological and ecological convergence at the lower size limit for vertebrates highlighted by five new miniaturised microhylid frog species from three different Madagascan genera"
765:
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,
2200:
3627:
Pombal, J.P. Jr. (2010). "A posição taxonômica das "variedades" de
Brachycephalus ephippium (Spix, 1824) descritas por Miranda-Ribeiro, 1920 (Amphibia, Anura, Brachycephalidae)".
3837:
825:
989:(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
3919:
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).
726:
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.
968:
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
673:
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
374:
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
3360:
1058:
4182:
2318:
Campos, L.A..; H.R. da Silva; A. Sebben (2010). "Morphology and development of additional bony elements in the genus Brachycephalus (Anura: Brachycephalidae)".
611:
1073:, 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 (
450:) 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.
414:. 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
566:
4228:
4238:
4143:
824:
729:
Overall the saddleback toads are mostly active during the day, but some activity also occurs during the night and some species possibly are mostly
4169:
596:
745:
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
2283:
Pombal Jr., J.P.; I. Sazima; C.F.B. Haddad (1994). "Breeding Behavior of the Pumpkin Toadlet, Brachycephalus ephippium (Brachycephalidae)".
3646:
Moura-Campos, Diego; Greenspan, Sasha E.; DiRenzo, Graziella V.; Neely, Wesley J.; Toledo, LuĂs Felipe; Becker, C. Guilherme (2021-09-01).
2946:
Pombal Jr, J.P.; E.M. Wistuba; M.R. Bornschein (1998). "A New Species of Brachycephalid (Anura) from the Atlantic Rain Forest of Brazil".
4086:
1101:. The pumpkin toadlets, which largely are restricted to highlands and foothills, can be further divided into two subgroups: the southern
396:
approaching their tiny dimensions. In other continents, the only similar-sized or slightly smaller frogs are a few species in the genera
2208:
3892:
Condez, T.H.; J.P.d.C. Monteiro; C.F.B. Haddad (2017). "Comments on the current taxonomy of Brachycephalus (Anura: Brachycephalidae)".
3980:
Garey, Michel V.; Lima, AndrĂ© M. X.; Hartmann, MarĂlia T.; Haddad, CĂ©lio F. B. (2012). "A new species of miniaturized toadlet, genus
3757:"Geographical and altitudinal distribution of Brachycephalus (Anura: Brachycephalidae) endemic to the Brazilian Atlantic Rainforest"
3391:
2449:
Ribeiro, Luiz F.; Bornschein, Marcos R.; Belmonte-Lopes, Ricardo; Firkowski, Carina R.; Morato, Sergio A.A.; Pie, Marcio R. (2015).
1221:(one of the variants that had been described in 1920) was instated as a valid species, bringing the number of generally recognised
581:
4233:
489:, which has a much more subdued colouring but still with some yellow-orange, appears to be non-toxic. It is unknown how certain
551:
4081:
3299:
1241:. The many recently discovered species have mostly been the result of scientists visiting isolated highland peaks, so-called
995:
313:
is one of the smallest species in the genus; with its frog-like appearance and overall brown colour it is a typical flea frog
4174:
1801:"New World direct-developing frogs (Anura: Terrarana): Molecular phylogeny, classification, biogeography, and conservation"
3921:"Hidden by the name: A new fluorescent pumpkin toadlet from the Brachycephalus ephippium group (Anura: Brachycephalidae)"
66:
2572:"Two new species of the Brachycephalus pernix group (Anura: Brachycephalidae) from the state of Paraná, southern Brazil"
1575:
1485:
1190:
266:, ranging from near sea level to an altitude of 1,900 m (6,200 ft), with most species restricted to highland
4104:
1374:
392:
3704:
4187:
1737:
1677:
1525:
1040:
studies have shown that most of these genera belong in other families, but the—in morphology quite different—genus
3536:"Semicircular Canal Size Determines the Developmental Onset of Angular Vestibuloocular Reflexes in Larval Xenopus"
949:, have not been recorded since their original scientific descriptions a century ago and they are unknown in life.
2235:"Diet of Brachycephalus brunneus (Anura: Brachycephalidae) in the Atlantic Rainforest of Paraná, southern Brazil"
2163:"Conservation Status of Brachycephalus Toadlets (Anura: Brachycephalidae) from the Brazilian Atlantic Rainforest"
2051:"Conservation Status of Brachycephalus Toadlets (Anura: Brachycephalidae) from the Brazilian Atlantic Rainforest"
1727:
1667:
1565:
1545:
1475:
1454:
1344:
1717:
883:
306:
1687:
1606:
1595:
1585:
1555:
1464:
1404:
1384:
1364:
1065:
relationship of four pumpkin toadlet species showing that colours alone are not a good indicator of relatedness
386:
49:
3602:
2854:"A new species of Brachycephalus (Anura: Brachycephalidae) from the Quiriri mountain range of southern Brazil"
2640:
Pie, Marcio R.; Ribeiro, Luiz F.; Confetti, André E.; Nadaline, Mário J.; Bornschein, Marcos R. (2018-10-03).
1697:
1515:
1495:
1414:
833:
1535:
1444:
4223:
1505:
1434:
1424:
1394:
1334:
1172:
677:
that inflates when calling, mouth gaping and waving of their arms. It is speculated that their calling is a
3407:
985:
species do not. Their small distributions make them highly vulnerable to habitat loss and degradation from
4021:
3489:"Quantifying the Ontogeny of Optokinetic and Vestibuloocular Behaviors in Zebrafish, Medaka, and Goldfish"
1637:
1626:
1354:
1110:
380:
239:
4109:
2049:
Rebouças, R.; A.B. Carollo; M.d.O. Freitas; C. Lambertini; R.M. Nogueira dos Santos; L.F. Toledo (2019).
1707:
1657:
325:, and they mostly walk (they are unable to jump far). The second group are the flea frogs or flea toads (
3831:
1616:
938:
909:
683:
289:
species have only been discovered in the 21st century. Most species have tiny ranges and are seriously
3433:
1925:
Fitzinger (Amphibia: Anura: Brachycephalidae), from the Atlantic rainforest of southern Bahia, Brazil"
711:
528:. It was initially speculated that the fluorescent colour also is aposematic or that it is related to
4130:
4068:
3934:
3445:
3241:
3171:
2735:
2699:
2366:
1647:
1202:
1118:
322:
654:
from other males, but in most cases voice or visual displays are sufficient and fights are avoided.
469:
The bright yellow, orange or reddish colours of many species in this genus are generally considered
4059:
1168:
1089:). Until 2002, the latter group, which is relatively widespread, was regarded as a separate genus,
858:
533:
263:
345:
frogs, and they are better jumpers. The name "flea frog" is also sometimes used for the unrelated
4001:
3488:
3342:
3275:
3136:
3027:
2963:
2925:
2829:
2611:
2539:
2300:
934:
930:
651:
375:
290:
256:
189:
61:
2988:"A new species of Brachycephalus (Anura: Brachycephalidae) from Santa Catarina, southern Brazil"
278:
and breeding is by direct development, with eggs laid on land and hatching into young frogs (no
1281:
4195:
4117:
3962:
3788:
3669:
3573:
3555:
3516:
3508:
3469:
3461:
3387:
3267:
3197:
3128:
3066:
3019:
2885:
2763:
2681:
2663:
2603:
2494:
2476:
2394:
2120:
2024:
1985:
1877:
1869:
1259:
1186:
1106:
1094:
1017:
892:
850:
473:(warning colours), but toxicity has only been studied in a few species. The brightly coloured
145:
3095:
Bornschein, M.R.; L.F. Ribeiro; M.M. Rollo Jr.; A.E. Confetti; M.R. Pie; et al. (2018).
4200:
3993:
3952:
3942:
3901:
3819:
3778:
3768:
3659:
3563:
3547:
3500:
3453:
3334:
3257:
3249:
3187:
3179:
3118:
3108:
3058:
3009:
2999:
2955:
2917:
2875:
2865:
2819:
2753:
2743:
2671:
2653:
2593:
2583:
2531:
2484:
2466:
2384:
2374:
2327:
2292:
2246:
2174:
2112:
2062:
2016:
1977:
1936:
1861:
1845:
1815:
1149:
1029:
502:
224:
221:
128:
41:
1292:
3823:
3700:
1270:
969:
659:
228:
2355:(Anura: Eleutherodactylidae): A new species from the Atlantic Forest of northeast Brazil"
3938:
3867:
3449:
3245:
3175:
2739:
2370:
633:
4073:
3997:
3957:
3920:
3783:
3756:
3568:
3262:
3229:
3192:
3159:
3123:
3096:
3014:
2987:
2880:
2853:
2758:
2723:
2676:
2641:
2598:
2571:
2489:
2450:
2389:
2350:
1772:
986:
926:
798:
172:
2020:
1844:
GuimarĂŁes, Carla Silva; Luz, Sofia; Rocha, Pedro Carvalho; Feio, Renato Neves (2017).
641:
and at least one other pumpkin toadlet species are deaf to their own advertising calls
4217:
3807:
3590:
3279:
2833:
2331:
1314:
are four of the thirteen species that were scientifically described from 2015 to 2017
922:
506:
342:
4005:
3346:
3140:
3031:
2929:
2615:
2543:
2251:
2234:
3551:
1920:
1122:
1062:
977:
678:
498:
482:
275:
267:
4122:
3062:
1193:, but in 1955 a review did not recognise them, arguing that they were synonyms of
1121:
and north). Among the pumpkin toadlets, colours alone are not a good indicator of
3947:
3664:
3647:
2748:
2535:
2379:
908:(right), two recently described species that, according to a review in 2019, are
658:
males when close. The calls of the different species differ to various extent in
4156:
4053:
3648:"Fungal disease cluster in tropical terrestrial frogs predicted by low rainfall"
3534:
Lambert, François M.; Beck, James C.; Baker, Robert; Straka, Hans (2008-08-06).
2642:"A new species of Brachycephalus (Anura: Brachycephalidae) from southern Brazil"
1042:
778:
529:
521:
404:
398:
347:
247:. Some pumpkin toadlets are toxic and their often bright colours are considered
3905:
3535:
3253:
3183:
2986:
Bornschein, M.R.; L.F. Ribeiro; D,C. Blackburn; E.L. Stanley; M.R. Pie (2016).
2921:
2116:
1981:
1940:
1865:
1819:
1242:
981:
786:
494:
470:
317:
The common name "saddleback toad" refers to the bony shields that cover their
271:
248:
17:
4044:
3673:
3559:
3512:
3465:
3230:"Evidence of auditory insensitivity to vocalization frequencies in two frogs"
3097:"Advertisement call of Brachycephalus albolineatus (Anura: Brachycephalidae)"
2667:
2480:
1873:
861:— is not detected by sensory hairs and makes it difficult for individuals of
741:
In general, saddleback toads are reported to use axillary or nearly axillary
3591:
Semicircular canal size constrains vestibular function in miniaturized frogs
3487:
Beck, James C.; Gilland, Edwin; Tank, David W.; Baker, Robert (2004-12-01).
1021:
867:
854:
802:
730:
674:
410:
366:
at least 14.8 mm (0.58 in). At 8–8.4 mm (0.31–0.33 in),
108:
78:
3966:
3792:
3577:
3520:
3473:
3457:
3271:
3201:
3132:
3070:
3023:
2889:
2767:
2685:
2607:
2498:
2398:
2124:
2028:
1989:
1881:
645:
Although not very loud for a frog/toad (even relative to their tiny size),
4096:
3504:
4038:
3925:
3361:"Unidades de conservação reafirman sua importância para a biodiversidade"
1037:
770:
742:
719:
525:
318:
98:
1850:(Anura: Brachycephalidae) from Serra do Brigadeiro, southeastern Brazil"
1800:
4148:
3806:
Heyer, W.R.; A.S. Rand; C.A.G. Cruz; O.L. Peixoto; C.E. Nelson (1990).
3773:
3160:"Intense bone fluorescence reveals hidden patterns in pumpkin toadlets"
3113:
3004:
2967:
2870:
2824:
2807:
2784:
2658:
2588:
2471:
2304:
1702:
Mângia, Santana, Drummond, Sabagh, Ugioni, Costa & Wachlevski, 2023
993:
species have also been found to be afflicted by recurrent outbreaks of
754:
279:
2179:
2162:
2067:
2050:
917:
Only 11 species of saddleback toads have been officially rated by the
853:
in the genus are too small for dependable locomotion, and in turn the
4161:
3363:. Infraestrutura e Meio Ambiente, Government of SĂŁo Paulo state. 2018
3338:
845:
are hindered in their ability to land reliably after jumping. Due to
810:
794:
774:
454:
Brazilian gold frog (sometimes misapplied to the all-brown flea frog
252:
88:
4015:
2959:
2570:
Ribeiro LF; Blackburn DC; Stanley EL; Pie MR; Bornschein MR (2017).
2296:
1520:
Folly, Condez, Vrcibradic, Rocha, Machado, Lopes & Pombal, 2024
524:, which is visible through the skin of the toadlet when exposed to
418:, a species where adults are at least 12.5 mm (0.49 in).
1148:
1057:
1050:
and they belong together as the only members of Brachycephalidae.
806:
782:
710:
632:
305:
235:
217:
370:
has the smallest maximum adult size reported in the genus, while
3382:
Zweifel, Richard G. (1998). Cogger, H.G.; Zweifel, R.G. (eds.).
1177:
918:
790:
231:
118:
4019:
1919:
Napoli, M.F.; Caramaschi, U.; Cruz, C.A.G.; Dias, I.R. (2011).
4135:
2806:
Bornschein, M.R.; L.F. Ribeiro; L. Teixeira; M.R. Pie (2019).
766:
1799:
Hedges, S. B.; Duellman, W. E. & Heinicke, M. P. (2008).
1156:, one of the species initially placed in the flea frog genus
703:
strongly suggest that they are able to hear their own calls.
1213:
was the only widely recognised species in the genus; it was
1141:, but these species pairs are more distant from each other.
665:
Peculiarly, at least the brightly coloured pumpkin toadlets
542:
Primary examples of colours and patterns in pumpkin toadlets
3711:. Version 6.0. American Museum of Natural History, New York
341:) that have an appearance and morphology that is more like
1662:
Almeida-Silva, Silva-Soares, Rodrigues & Verdade, 2021
2444:
2442:
2440:
2438:
2436:
2434:
2432:
2430:
2428:
1237:
in 2002. As of 2020, there were 36 recognised species of
1020:, but were historically included in the true toad family
2779:
2777:
2426:
2424:
2422:
2420:
2418:
2416:
2414:
2412:
2410:
2408:
1189:. In 1920, four additional "variants" were described by
2233:
Fontoura, P.L.; L.F. Ribeiro; M.R. Pie. Haddad (2011).
251:. At about 1 cm (0.4 in) or less in snout–to–
1229:
were instated as valid species in 2010 and the former
1024:, where one of the defining features is its presence.
921:, all between 2004 and 2010, with 3 species listed as
3408:"Here's why pumpkin toadlets are such clumsy jumpers"
1860:(4). Auckland, New Zealand: Magnolia Press: 327–344.
485:
and similar toxins in their skin and organs, whereas
3300:"These tiny frogs can't hear their own mating songs"
1846:"The dark side of pumpkin toadlet: a new species of
1325:, the following species are recognised in the genus
4028:
3709:
Amphibian Species of the World, an Online Reference
1767:
1765:
1763:
1761:
1759:
1757:
378:in the Americas, with only the northeast Brazilian
262:The saddleback toads live among leaf litter in the
3984:(Anura: Brachycephalidae), from southern Brazil".
3868:"Seven tiny frog species found on seven mountains"
1429:Condez, Clemente-Carvalho, Haddad & Reis, 2014
3836:: CS1 maint: DOI inactive as of September 2024 (
3090:
3088:
3086:
3084:
3082:
3080:
2981:
2979:
2977:
2201:"Seven New Mini-Frogs Found—Among Smallest Known"
2156:
2154:
2161:Bornschein, M.R.; M.R. Pie; L. Teixeira (2019).
2152:
2150:
2148:
2146:
2144:
2142:
2140:
2138:
2136:
2134:
497:, but they are possibly absorbed from the small
3434:"Size Limitations in Semicircular Duct Systems"
2455:(Anura: Brachycephalidae) from southern Brazil"
2228:
2226:
1839:
1837:
1835:
1833:
1831:
1829:
3293:
3291:
3289:
2565:
2563:
2561:
2559:
2557:
2555:
2553:
2351:"Diversity of miniaturized frogs of the genus
2044:
2042:
2040:
2038:
1962:
1960:
1958:
1956:
1954:
1952:
1950:
3730:
3728:
3726:
3622:
3620:
3223:
3221:
3219:
3217:
3215:
3213:
3211:
3152:
3150:
2941:
2939:
2516:
2514:
2512:
2510:
2508:
2343:
2341:
2278:
2276:
2274:
2272:
2270:
2268:
2266:
2264:
2262:
2098:
686:to lose it because of their toxicity. Unlike
509:, or produced by bacteria inside their body.
8:
3043:
3041:
2903:
2901:
2899:
2194:
2192:
2190:
2096:
2094:
2092:
2090:
2088:
2086:
2084:
2082:
2080:
2078:
1914:
1912:
1910:
1908:
1906:
1904:
1902:
1900:
1898:
1794:
1792:
1790:
2001:
1999:
1028:was subsequently placed in Atelopodidae or
4016:
3861:
3859:
3857:
3855:
3853:
3851:
3849:
3847:
3386:. San Diego: Academic Press. p. 103.
2847:
2845:
2843:
2700:"10 Smallest Frogs and Toads in the World"
2635:
2633:
2631:
2629:
2627:
2625:
171:
40:
31:
3956:
3946:
3782:
3772:
3663:
3567:
3261:
3191:
3122:
3112:
3013:
3003:
2879:
2869:
2823:
2787:. Berkeley, California: AmphibiaWeb. 2019
2757:
2747:
2675:
2657:
2597:
2587:
2488:
2470:
2388:
2378:
2320:Biological Journal of the Linnean Society
2250:
2178:
2066:
1775:. Berkeley, California: AmphibiaWeb. 2020
1652:Napoli, Caramaschi, Cruz & Dias, 2011
820:
512:At least two brightly coloured species,
3695:
3693:
3691:
3689:
3687:
3685:
3683:
3384:Encyclopedia of Reptiles and Amphibians
1753:
662:, speed, structure and other features.
547:
255:length, the flea frogs are some of the
3829:
3755:Bornschein, M.R.; et al. (2016).
1632:Alves, Sawaya, Reis & Haddad, 2009
1621:Pombal, Wistuba & Bornschein, 1998
1449:GuimarĂŁes, Luz, Rocha & Feio, 2017
1125:relationships. For example, the brown
849:' extremely small head and body size,
3824:10.11606/issn.2176-7793.v31i4p231-410
1175:in 1824, but originally in the genus
941:. Two of the data deficient species,
7:
2702:. gonefroggin.com. 18 December 2017
2451:"Seven new microendemic species of
1921:"A new species of flea-toad, genus
1599:(Serra Cantareira saddleback toad)
972:. Each of the remaining species of
925:(not threatened) and the remaining
870:) as protection against predators.
3998:10.1655/herpetologica-d-11-00074.1
3705:"Search for Taxon: Brachycephalus"
837:jumping, showing a lack of balance
25:
4229:Amphibians of the Atlantic Forest
3298:Supriya, L. (26 September 2017).
1137:is closely related to the golden
1129:is closely related to the golden
699:and the dull-coloured flea frog
605:(dull with bright markings below)
575:(bright with dull sections above)
3228:Goutte, S.; et al. (2017).
2852:Pie, M.R.; L.F. Ribeiro (2015).
2332:10.1111/j.1095-8312.2010.01375.x
1291:
1280:
1269:
1258:
891:
882:
610:
595:
580:
565:
550:
65:
4239:Taxa named by Leopold Fitzinger
3605:. IUCN Redlist. 2 February 2020
2252:10.1590/S1984-46702011000500019
220:of tiny toads and frogs in the
3866:Webb, Jonathan (4 June 2015).
3552:10.1523/JNEUROSCI.1288-08.2008
3438:Journal of Theoretical Biology
1510:Clemente-Carvalho et al., 2012
1323:Amphibian Species of the World
1205:, but placed in its own genus
996:Batrachochytrium dendrobatidis
434:The flea frogs or flea toads (
1:
3063:10.1016/j.toxicon.2004.09.016
2021:10.1016/s0041-0101(01)00279-3
1530:(Giaretta & Sawaya, 1998)
1093:, but it is now considered a
620:(fluorescence under UV-light)
501:they eat, as known from some
3948:10.1371/journal.pone.0244812
3665:10.1016/j.biocon.2021.109246
2749:10.1371/journal.pone.0213314
2536:10.1080/00222933.2012.713525
2380:10.1371/journal.pone.0201781
1576:Brachycephalus mariaeterezae
1570:Pombal & Izecksohn, 2011
1486:Brachycephalus fuscolineatus
1359:Pombal & Gasparini, 2006
1181:. Two years later the genus
1004:Classification and discovery
2199:Bates, Mary (4 June 2015).
1668:Brachycephalus quiririensis
1566:Brachycephalus margaritatus
1375:Brachycephalus auroguttatus
1345:Brachycephalus albolineatus
1201:, was described in 1971 by
4255:
3906:10.11646/zootaxa.4290.2.12
3493:Journal of Neurophysiology
3254:10.1038/s41598-017-12145-5
3184:10.1038/s41598-019-41959-8
2922:10.1670/0022-1511-43.2.212
2785:"Brachycephalus ephippium"
2524:Journal of Natural History
1738:Brachycephalus vertebralis
1718:Brachycephalus tridactylus
1678:Brachycephalus rotenbergae
1526:Brachycephalus hermogenesi
1217:pumpkin toadlet. In 1990,
637:Unique among all animals,
493:species attain the strong
3432:Muller, M. (1999-06-07).
2117:10.11646/zootaxa.4083.1.2
1982:10.11646/zootaxa.4407.4.2
1941:10.11646/zootaxa.2739.1.3
1866:10.11646/zootaxa.4258.4.2
1820:10.11646/zootaxa.1737.1.1
1728:Brachycephalus verrucosus
1688:Brachycephalus sulfuratus
1586:Brachycephalus mirissimus
1546:Brachycephalus izecksohni
1476:Brachycephalus ferruginus
1468:(Spix’s saddleback toad)
1455:Brachycephalus didactylus
1405:Brachycephalus bufonoides
1365:Brachycephalus atelopoide
1191:AlĂpio de Miranda-Ribeiro
195:
188:
179:
170:
162:
158:
153:
62:Scientific classification
60:
48:
39:
34:
2205:National Geographic News
1698:Brachycephalus tabuleiro
1607:Brachycephalus olivaceus
1596:Brachycephalus nodoterga
1556:Brachycephalus leopardus
1516:Brachycephalus herculeus
1496:Brachycephalus garbeanus
1465:Brachycephalus ephippium
1415:Brachycephalus coloratus
1385:Brachycephalus boticario
1233:species were moved into
1165:Brachycephalus ephippium
1054:Genus and species groups
964:and the pumpkin toadlet
841:Unlike most amphibians,
757:or other aquatic stage.
387:Eleutherodactylus iberia
54:Brachycephalus ephippium
4234:Endemic frogs of Brazil
3652:Biological Conservation
3540:Journal of Neuroscience
1672:Pie & Ribeiro, 2015
1536:Brachycephalus ibitinga
1445:Brachycephalus darkside
1435:Brachycephalus curupira
1395:Brachycephalus brunneus
1349:Bornschein et al., 2016
1173:Johann Baptist von Spix
1046:is a close relative of
857:— which is regarded by
3826:(inactive 2024-09-12).
3458:10.1006/jtbi.1999.0922
3327:Journal of Herpetology
2948:Journal of Herpetology
2910:Journal of Herpetology
2285:Journal of Herpetology
1638:Brachycephalus pombali
1630:(Red pumpkin toadlet)
1627:Brachycephalus pitanga
1506:Brachycephalus guarani
1425:Brachycephalus crispus
1355:Brachycephalus alipioi
1335:Brachycephalus actaeus
1160:
1066:
838:
722:
642:
520:, have bones that are
314:
50:Spix's saddleback toad
3505:10.1152/jn.00311.2004
1617:Brachycephalus pernix
1601:Miranda-Ribeiro, 1920
1500:Miranda-Ribeiro, 1920
1409:Miranda-Ribeiro, 1920
1369:Miranda-Ribeiro, 1920
1339:Monteiro et al., 2018
1152:
1061:
952:The three flea frogs
939:critically endangered
910:critically endangered
831:
714:
684:evolutionary pressure
636:
309:
234:, ranging from south
3812:Arquivos de Zoologia
2530:(39–40): 2497–2507.
1732:Ribeiro et al., 2015
1648:Brachycephalus pulex
1611:Ribeiro et al., 2015
1580:Ribeiro et al., 2015
1560:Ribeiro et al., 2015
1550:Ribeiro et al., 2005
1490:Ribeiro et al., 2015
1439:Ribeiro et al., 2017
1419:Ribeiro et al., 2017
1399:Ribeiro et al., 2005
1389:Ribeiro et al., 2015
1379:Ribeiro et al., 2015
1209:. As a consequence,
785:) and their larvae,
381:Adelophryne michelin
285:The majority of the
3939:2021PLoSO..1644812N
3808:"Frogs of Boracéia"
3629:Zoologia (Curitiba)
3450:1999JThBi.198..405M
3246:2017NatSR...712121G
3176:2019NatSR...9.5388G
2740:2019PLoSO..1413314S
2371:2018PLoSO..1301781L
2239:Zoologia (Curitiba)
1712:Haddad et al., 2010
1708:Brachycephalus toby
1692:Condez et al., 2016
1658:Brachycephalus puri
1540:Condez et al., 2021
1197:. Another species,
1163:The first species,
1113:) and the northern
874:Conservation status
534:species recognition
422:Colour and toxicity
302:Morphology and size
264:Atlantic rainforest
27:Genus of amphibians
3774:10.7717/peerj.2490
3234:Scientific Reports
3164:Scientific Reports
3114:10.7717/peerj.5273
3005:10.7717/peerj.2629
2871:10.7717/peerj.1179
2825:10.15560/15.6.1031
2659:10.7717/peerj.5683
2589:10.7717/peerj.3603
2472:10.7717/peerj.1011
1773:"Brachycephalidae"
1722:Garey et al., 2012
1682:Nunes et al., 2021
1642:Alves et al., 2006
1480:Alves et al., 2006
1161:
1067:
851:vestibular systems
839:
723:
643:
315:
270:. They are mostly
4211:
4210:
4196:Open Tree of Life
4022:Taxon identifiers
3738:Amphibia-Reptilia
3546:(32): 8086–8095.
2180:10.3390/d11090150
2068:10.3390/d11090150
1743:
1733:
1723:
1713:
1703:
1693:
1683:
1673:
1663:
1653:
1643:
1633:
1622:
1612:
1602:
1591:
1581:
1571:
1561:
1551:
1541:
1531:
1521:
1511:
1501:
1491:
1481:
1471:
1460:
1459:(Izecksohn, 1971)
1450:
1440:
1430:
1420:
1410:
1400:
1390:
1380:
1370:
1360:
1350:
1340:
1203:EugĂŞnio Izecksohn
1187:Leopold Fitzinger
829:
629:Voice and hearing
503:poison dart frogs
206:
205:
149:
35:Saddleback toads
16:(Redirected from
4246:
4204:
4203:
4191:
4190:
4178:
4177:
4165:
4164:
4152:
4151:
4139:
4138:
4126:
4125:
4113:
4112:
4100:
4099:
4090:
4089:
4077:
4076:
4064:
4063:
4062:
4049:
4048:
4047:
4017:
4010:
4009:
3977:
3971:
3970:
3960:
3950:
3916:
3910:
3909:
3889:
3883:
3882:
3880:
3878:
3863:
3842:
3841:
3835:
3827:
3803:
3797:
3796:
3786:
3776:
3752:
3746:
3745:
3732:
3721:
3720:
3718:
3716:
3701:Frost, Darrel R.
3697:
3678:
3677:
3667:
3643:
3637:
3636:
3624:
3615:
3614:
3612:
3610:
3603:"Brachycephalus"
3599:
3593:
3588:
3582:
3581:
3571:
3531:
3525:
3524:
3499:(6): 3546–3561.
3484:
3478:
3477:
3429:
3423:
3422:
3420:
3419:
3404:
3398:
3397:
3379:
3373:
3372:
3370:
3368:
3357:
3351:
3350:
3339:10.1670/07-287.1
3321:
3315:
3314:
3312:
3310:
3295:
3284:
3283:
3265:
3225:
3206:
3205:
3195:
3154:
3145:
3144:
3126:
3116:
3092:
3075:
3074:
3045:
3036:
3035:
3017:
3007:
2983:
2972:
2971:
2943:
2934:
2933:
2905:
2894:
2893:
2883:
2873:
2849:
2838:
2837:
2827:
2818:(6): 1031–1036.
2803:
2797:
2796:
2794:
2792:
2781:
2772:
2771:
2761:
2751:
2718:
2712:
2711:
2709:
2707:
2696:
2690:
2689:
2679:
2661:
2637:
2620:
2619:
2601:
2591:
2567:
2548:
2547:
2518:
2503:
2502:
2492:
2474:
2446:
2403:
2402:
2392:
2382:
2345:
2336:
2335:
2315:
2309:
2308:
2280:
2257:
2256:
2254:
2230:
2221:
2220:
2218:
2216:
2207:. Archived from
2196:
2185:
2184:
2182:
2158:
2129:
2128:
2100:
2073:
2072:
2070:
2046:
2033:
2032:
2003:
1994:
1993:
1964:
1945:
1944:
1916:
1893:
1892:
1890:
1888:
1841:
1824:
1823:
1805:
1796:
1785:
1784:
1782:
1780:
1769:
1741:
1731:
1721:
1711:
1701:
1691:
1681:
1671:
1661:
1651:
1641:
1631:
1620:
1610:
1600:
1590:Pie et al., 2018
1589:
1579:
1569:
1559:
1549:
1539:
1529:
1519:
1509:
1499:
1489:
1479:
1469:
1458:
1448:
1438:
1428:
1418:
1408:
1398:
1388:
1378:
1368:
1358:
1348:
1338:
1308:B. mariaeterezae
1295:
1284:
1273:
1262:
1225:species to two.
1030:Brachycephalidae
895:
886:
859:Poiseuille's law
830:
614:
599:
590:(overall bright)
584:
573:B. fuscolineatus
569:
554:
225:Brachycephalidae
210:saddleback toads
180:Distribution of
175:
144:
129:Brachycephalidae
70:
69:
44:
32:
21:
4254:
4253:
4249:
4248:
4247:
4245:
4244:
4243:
4214:
4213:
4212:
4207:
4199:
4194:
4186:
4181:
4173:
4168:
4160:
4155:
4147:
4142:
4134:
4129:
4121:
4116:
4108:
4103:
4095:
4093:
4085:
4080:
4072:
4067:
4058:
4057:
4052:
4043:
4042:
4037:
4024:
4014:
4013:
3979:
3978:
3974:
3933:(4): e0244812.
3918:
3917:
3913:
3891:
3890:
3886:
3876:
3874:
3865:
3864:
3845:
3828:
3805:
3804:
3800:
3754:
3753:
3749:
3734:
3733:
3724:
3714:
3712:
3699:
3698:
3681:
3645:
3644:
3640:
3626:
3625:
3618:
3608:
3606:
3601:
3600:
3596:
3589:
3585:
3533:
3532:
3528:
3486:
3485:
3481:
3431:
3430:
3426:
3417:
3415:
3406:
3405:
3401:
3394:
3381:
3380:
3376:
3366:
3364:
3359:
3358:
3354:
3323:
3322:
3318:
3308:
3306:
3297:
3296:
3287:
3227:
3226:
3209:
3156:
3155:
3148:
3094:
3093:
3078:
3047:
3046:
3039:
2985:
2984:
2975:
2960:10.2307/1565481
2945:
2944:
2937:
2907:
2906:
2897:
2864:(1792): e1179.
2851:
2850:
2841:
2805:
2804:
2800:
2790:
2788:
2783:
2782:
2775:
2734:(3): e0213314.
2720:
2719:
2715:
2705:
2703:
2698:
2697:
2693:
2639:
2638:
2623:
2569:
2568:
2551:
2520:
2519:
2506:
2448:
2447:
2406:
2365:(9): e0201781.
2347:
2346:
2339:
2317:
2316:
2312:
2297:10.2307/1564972
2282:
2281:
2260:
2232:
2231:
2224:
2214:
2212:
2211:on June 6, 2015
2198:
2197:
2188:
2160:
2159:
2132:
2102:
2101:
2076:
2048:
2047:
2036:
2005:
2004:
1997:
1966:
1965:
1948:
1918:
1917:
1896:
1886:
1884:
1843:
1842:
1827:
1803:
1798:
1797:
1788:
1778:
1776:
1771:
1770:
1755:
1750:
1319:
1318:
1317:
1316:
1315:
1300:B. auroguttatus
1298:Top to bottom:
1296:
1287:
1286:
1285:
1276:
1275:
1274:
1265:
1264:
1263:
1252:
1147:
1056:
1011:
1006:
970:species complex
915:
914:
913:
912:
906:B. quiririensis
898:
897:
896:
888:
887:
876:
821:
819:
799:pseudoscorpions
763:
739:
709:
631:
626:
625:
624:
621:
615:
606:
600:
591:
585:
576:
570:
561:
555:
544:
543:
424:
364:B. margaritatus
304:
299:
274:, feed on tiny
166:
143:
64:
28:
23:
22:
15:
12:
11:
5:
4252:
4250:
4242:
4241:
4236:
4231:
4226:
4224:Brachycephalus
4216:
4215:
4209:
4208:
4206:
4205:
4192:
4179:
4166:
4153:
4140:
4127:
4114:
4101:
4091:
4087:Brachycephalus
4078:
4074:Brachycephalus
4065:
4060:Brachycephalus
4050:
4034:
4032:
4030:Brachycephalus
4026:
4025:
4020:
4012:
4011:
3992:(2): 266–271.
3982:Brachycephalus
3972:
3911:
3900:(2): 395–400.
3884:
3843:
3818:(4): 231–410.
3798:
3747:
3722:
3679:
3638:
3616:
3594:
3583:
3526:
3479:
3444:(3): 405–437.
3424:
3399:
3392:
3374:
3352:
3333:(3): 542–549.
3316:
3285:
3207:
3146:
3076:
3037:
2973:
2935:
2916:(2): 212–219.
2895:
2839:
2798:
2773:
2713:
2691:
2621:
2549:
2504:
2453:Brachycephalus
2404:
2337:
2326:(4): 752–767.
2310:
2291:(4): 516–519.
2258:
2245:(5): 687–689.
2222:
2186:
2130:
2074:
2034:
1995:
1976:(4): 483–505.
1946:
1923:Brachycephalus
1894:
1848:Brachycephalus
1825:
1786:
1752:
1751:
1749:
1746:
1745:
1744:
1734:
1724:
1714:
1704:
1694:
1684:
1674:
1664:
1654:
1644:
1634:
1623:
1613:
1603:
1592:
1582:
1572:
1562:
1552:
1542:
1532:
1522:
1512:
1502:
1492:
1482:
1472:
1461:
1451:
1441:
1431:
1421:
1411:
1401:
1391:
1381:
1371:
1361:
1351:
1341:
1327:Brachycephalus
1297:
1290:
1289:
1288:
1279:
1278:
1277:
1268:
1267:
1266:
1257:
1256:
1255:
1254:
1253:
1251:
1248:
1239:Brachycephalus
1235:Brachycephalus
1223:Brachycephalus
1185:was coined by
1183:Brachycephalus
1167:, was already
1154:B. hermogenesi
1146:
1143:
1133:and the brown
1111:Santa Catarina
1099:Brachycephalus
1079:B. hermogenesi
1071:Brachycephalus
1055:
1052:
1048:Brachycephalus
1034:Brachycephalus
1026:Brachycephalus
1018:Bidder's organ
1014:Brachycephalus
1010:
1007:
1005:
1002:
991:Brachycephalus
987:climate change
974:Brachycephalus
966:B. ephippiumsi
958:B. hermogenesi
927:data deficient
900:
899:
890:
889:
881:
880:
879:
878:
877:
875:
872:
863:Brachycephalus
847:Brachycephalus
843:Brachycephalus
818:
815:
762:
759:
738:
735:
708:
705:
701:B. hermogenesi
647:Brachycephalus
630:
627:
623:
622:
616:
609:
607:
601:
594:
592:
586:
579:
577:
571:
564:
562:
560:(overall dull)
556:
549:
546:
545:
541:
540:
539:
507:mantella frogs
491:Brachycephalus
440:B. hermogenesi
428:Brachycephalus
426:The different
423:
420:
376:smallest frogs
355:Brachycephalus
331:B. hermogenesi
303:
300:
298:
295:
287:Brachycephalus
259:in the world.
257:smallest frogs
240:Santa Catarina
214:Brachycephalus
204:
203:
202:
201:
193:
192:
186:
185:
182:Brachycephalus
177:
176:
168:
167:
161:More than 35,
159:
156:
155:
151:
150:
140:Brachycephalus
136:
132:
131:
126:
122:
121:
116:
112:
111:
106:
102:
101:
96:
92:
91:
86:
82:
81:
76:
72:
71:
58:
57:
46:
45:
37:
36:
26:
24:
18:Brachycephalus
14:
13:
10:
9:
6:
4:
3:
2:
4251:
4240:
4237:
4235:
4232:
4230:
4227:
4225:
4222:
4221:
4219:
4202:
4197:
4193:
4189:
4184:
4180:
4176:
4171:
4167:
4163:
4158:
4154:
4150:
4145:
4141:
4137:
4132:
4128:
4124:
4119:
4115:
4111:
4106:
4102:
4098:
4092:
4088:
4083:
4079:
4075:
4070:
4066:
4061:
4055:
4051:
4046:
4040:
4036:
4035:
4033:
4031:
4027:
4023:
4018:
4007:
4003:
3999:
3995:
3991:
3987:
3986:Herpetologica
3983:
3976:
3973:
3968:
3964:
3959:
3954:
3949:
3944:
3940:
3936:
3932:
3928:
3927:
3922:
3915:
3912:
3907:
3903:
3899:
3895:
3888:
3885:
3873:
3869:
3862:
3860:
3858:
3856:
3854:
3852:
3850:
3848:
3844:
3839:
3833:
3825:
3821:
3817:
3813:
3809:
3802:
3799:
3794:
3790:
3785:
3780:
3775:
3770:
3766:
3762:
3758:
3751:
3748:
3744:(2): 225–227.
3743:
3739:
3731:
3729:
3727:
3723:
3710:
3706:
3702:
3696:
3694:
3692:
3690:
3688:
3686:
3684:
3680:
3675:
3671:
3666:
3661:
3657:
3653:
3649:
3642:
3639:
3634:
3630:
3623:
3621:
3617:
3604:
3598:
3595:
3592:
3587:
3584:
3579:
3575:
3570:
3565:
3561:
3557:
3553:
3549:
3545:
3541:
3537:
3530:
3527:
3522:
3518:
3514:
3510:
3506:
3502:
3498:
3494:
3490:
3483:
3480:
3475:
3471:
3467:
3463:
3459:
3455:
3451:
3447:
3443:
3439:
3435:
3428:
3425:
3413:
3409:
3403:
3400:
3395:
3393:0-12-178560-2
3389:
3385:
3378:
3375:
3362:
3356:
3353:
3348:
3344:
3340:
3336:
3332:
3328:
3320:
3317:
3305:
3301:
3294:
3292:
3290:
3286:
3281:
3277:
3273:
3269:
3264:
3259:
3255:
3251:
3247:
3243:
3239:
3235:
3231:
3224:
3222:
3220:
3218:
3216:
3214:
3212:
3208:
3203:
3199:
3194:
3189:
3185:
3181:
3177:
3173:
3169:
3165:
3161:
3153:
3151:
3147:
3142:
3138:
3134:
3130:
3125:
3120:
3115:
3110:
3106:
3102:
3098:
3091:
3089:
3087:
3085:
3083:
3081:
3077:
3072:
3068:
3064:
3060:
3056:
3052:
3044:
3042:
3038:
3033:
3029:
3025:
3021:
3016:
3011:
3006:
3001:
2998:(10): e2629.
2997:
2993:
2989:
2982:
2980:
2978:
2974:
2969:
2965:
2961:
2957:
2953:
2949:
2942:
2940:
2936:
2931:
2927:
2923:
2919:
2915:
2911:
2904:
2902:
2900:
2896:
2891:
2887:
2882:
2877:
2872:
2867:
2863:
2859:
2855:
2848:
2846:
2844:
2840:
2835:
2831:
2826:
2821:
2817:
2813:
2809:
2802:
2799:
2786:
2780:
2778:
2774:
2769:
2765:
2760:
2755:
2750:
2745:
2741:
2737:
2733:
2729:
2725:
2717:
2714:
2701:
2695:
2692:
2687:
2683:
2678:
2673:
2669:
2665:
2660:
2655:
2651:
2647:
2643:
2636:
2634:
2632:
2630:
2628:
2626:
2622:
2617:
2613:
2609:
2605:
2600:
2595:
2590:
2585:
2581:
2577:
2573:
2566:
2564:
2562:
2560:
2558:
2556:
2554:
2550:
2545:
2541:
2537:
2533:
2529:
2525:
2517:
2515:
2513:
2511:
2509:
2505:
2500:
2496:
2491:
2486:
2482:
2478:
2473:
2468:
2464:
2460:
2456:
2454:
2445:
2443:
2441:
2439:
2437:
2435:
2433:
2431:
2429:
2427:
2425:
2423:
2421:
2419:
2417:
2415:
2413:
2411:
2409:
2405:
2400:
2396:
2391:
2386:
2381:
2376:
2372:
2368:
2364:
2360:
2356:
2354:
2344:
2342:
2338:
2333:
2329:
2325:
2321:
2314:
2311:
2306:
2302:
2298:
2294:
2290:
2286:
2279:
2277:
2275:
2273:
2271:
2269:
2267:
2265:
2263:
2259:
2253:
2248:
2244:
2240:
2236:
2229:
2227:
2223:
2210:
2206:
2202:
2195:
2193:
2191:
2187:
2181:
2176:
2172:
2168:
2164:
2157:
2155:
2153:
2151:
2149:
2147:
2145:
2143:
2141:
2139:
2137:
2135:
2131:
2126:
2122:
2118:
2114:
2110:
2106:
2099:
2097:
2095:
2093:
2091:
2089:
2087:
2085:
2083:
2081:
2079:
2075:
2069:
2064:
2060:
2056:
2052:
2045:
2043:
2041:
2039:
2035:
2030:
2026:
2022:
2018:
2014:
2010:
2002:
2000:
1996:
1991:
1987:
1983:
1979:
1975:
1971:
1963:
1961:
1959:
1957:
1955:
1953:
1951:
1947:
1942:
1938:
1934:
1930:
1926:
1924:
1915:
1913:
1911:
1909:
1907:
1905:
1903:
1901:
1899:
1895:
1883:
1879:
1875:
1871:
1867:
1863:
1859:
1855:
1851:
1849:
1840:
1838:
1836:
1834:
1832:
1830:
1826:
1821:
1817:
1813:
1809:
1802:
1795:
1793:
1791:
1787:
1774:
1768:
1766:
1764:
1762:
1760:
1758:
1754:
1747:
1740:
1739:
1735:
1730:
1729:
1725:
1720:
1719:
1715:
1710:
1709:
1705:
1700:
1699:
1695:
1690:
1689:
1685:
1680:
1679:
1675:
1670:
1669:
1665:
1660:
1659:
1655:
1650:
1649:
1645:
1640:
1639:
1635:
1629:
1628:
1624:
1619:
1618:
1614:
1609:
1608:
1604:
1598:
1597:
1593:
1588:
1587:
1583:
1578:
1577:
1573:
1568:
1567:
1563:
1558:
1557:
1553:
1548:
1547:
1543:
1538:
1537:
1533:
1528:
1527:
1523:
1518:
1517:
1513:
1508:
1507:
1503:
1498:
1497:
1493:
1488:
1487:
1483:
1478:
1477:
1473:
1467:
1466:
1462:
1457:
1456:
1452:
1447:
1446:
1442:
1437:
1436:
1432:
1427:
1426:
1422:
1417:
1416:
1412:
1407:
1406:
1402:
1397:
1396:
1392:
1387:
1386:
1382:
1377:
1376:
1372:
1367:
1366:
1362:
1357:
1356:
1352:
1347:
1346:
1342:
1337:
1336:
1332:
1331:
1330:
1328:
1324:
1321:According to
1313:
1312:B. verrucosus
1309:
1305:
1301:
1294:
1283:
1272:
1261:
1249:
1247:
1244:
1240:
1236:
1232:
1226:
1224:
1220:
1216:
1212:
1208:
1204:
1200:
1199:B. didactylus
1196:
1192:
1188:
1184:
1180:
1179:
1174:
1170:
1166:
1159:
1155:
1151:
1144:
1142:
1140:
1136:
1132:
1131:B. izecksohni
1128:
1124:
1120:
1116:
1112:
1108:
1104:
1100:
1096:
1092:
1088:
1087:B. sulfuratus
1084:
1080:
1076:
1075:B. didactylus
1072:
1064:
1060:
1053:
1051:
1049:
1045:
1044:
1039:
1035:
1031:
1027:
1023:
1019:
1015:
1008:
1003:
1001:
998:
997:
992:
988:
983:
979:
975:
971:
967:
963:
962:B. sulfuratus
959:
955:
954:B. didactylus
950:
948:
947:B. bufonoides
944:
943:B. atelopoide
940:
936:
932:
928:
924:
923:least concern
920:
911:
907:
903:
894:
885:
873:
871:
869:
864:
860:
856:
852:
848:
844:
836:
835:
816:
814:
812:
808:
804:
800:
796:
792:
788:
784:
780:
776:
772:
768:
760:
758:
756:
750:
748:
744:
736:
734:
732:
727:
721:
717:
713:
706:
704:
702:
698:
693:
689:
685:
680:
676:
672:
668:
663:
661:
655:
653:
648:
640:
635:
628:
619:
613:
608:
604:
598:
593:
589:
588:B. izecksohni
583:
578:
574:
568:
563:
559:
553:
548:
538:
535:
531:
527:
523:
519:
515:
510:
508:
504:
500:
499:invertebrates
496:
492:
488:
484:
480:
476:
472:
467:
465:
461:
457:
456:B. didactylus
451:
449:
448:B. sulfuratus
445:
441:
437:
436:B. didactylus
432:
429:
421:
419:
417:
413:
412:
407:
406:
401:
400:
395:
394:
389:
388:
383:
382:
377:
373:
372:B. sulfuratus
369:
365:
361:
356:
352:
350:
349:
344:
343:leptodactylid
340:
339:B. sulfuratus
336:
332:
328:
327:B. didactylus
324:
320:
312:
308:
301:
296:
294:
292:
288:
283:
281:
277:
276:invertebrates
273:
269:
265:
260:
258:
254:
250:
246:
241:
237:
233:
230:
226:
223:
219:
215:
211:
200:
197:
196:
194:
191:
187:
183:
178:
174:
169:
165:
164:
157:
152:
147:
142:
141:
137:
134:
133:
130:
127:
124:
123:
120:
117:
114:
113:
110:
107:
104:
103:
100:
97:
94:
93:
90:
87:
84:
83:
80:
77:
74:
73:
68:
63:
59:
55:
51:
47:
43:
38:
33:
30:
19:
4029:
3989:
3985:
3981:
3975:
3930:
3924:
3914:
3897:
3893:
3887:
3875:. Retrieved
3871:
3832:cite journal
3815:
3811:
3801:
3764:
3760:
3750:
3741:
3737:
3713:. Retrieved
3708:
3655:
3651:
3641:
3632:
3628:
3607:. Retrieved
3597:
3586:
3543:
3539:
3529:
3496:
3492:
3482:
3441:
3437:
3427:
3416:. Retrieved
3414:. 2022-06-15
3412:Science News
3411:
3402:
3383:
3377:
3365:. Retrieved
3355:
3330:
3326:
3319:
3307:. Retrieved
3303:
3240:(1): 12121.
3237:
3233:
3167:
3163:
3104:
3100:
3057:(1): 73–79.
3054:
3050:
2995:
2991:
2954:(1): 70–74.
2951:
2947:
2913:
2909:
2861:
2857:
2815:
2811:
2801:
2789:. Retrieved
2731:
2727:
2716:
2704:. Retrieved
2694:
2649:
2645:
2579:
2575:
2527:
2523:
2462:
2458:
2452:
2362:
2358:
2352:
2323:
2319:
2313:
2288:
2284:
2242:
2238:
2213:. Retrieved
2209:the original
2204:
2170:
2166:
2111:(1): 40–56.
2108:
2104:
2061:(1): 39–47.
2058:
2054:
2015:(6): 761–6.
2012:
2008:
1973:
1969:
1932:
1928:
1922:
1885:. Retrieved
1857:
1853:
1847:
1811:
1807:
1777:. Retrieved
1742:Pombal, 2001
1736:
1726:
1716:
1706:
1696:
1686:
1676:
1666:
1656:
1646:
1636:
1625:
1615:
1605:
1594:
1584:
1574:
1564:
1554:
1544:
1534:
1524:
1514:
1504:
1494:
1484:
1474:
1470:(Spix, 1824)
1463:
1453:
1443:
1433:
1423:
1413:
1403:
1393:
1383:
1373:
1363:
1353:
1343:
1333:
1326:
1322:
1320:
1311:
1307:
1304:B. coloratus
1303:
1299:
1238:
1234:
1231:Psyllophryne
1230:
1227:
1222:
1219:B. nodoterga
1218:
1214:
1211:B. ephippium
1210:
1207:Psyllophryne
1206:
1198:
1195:B. ephippium
1194:
1182:
1176:
1164:
1162:
1158:Psyllophryne
1157:
1153:
1139:B. leopardus
1138:
1134:
1130:
1126:
1123:phylogenetic
1115:B. ephippium
1114:
1102:
1098:
1091:Psyllophryne
1090:
1086:
1082:
1078:
1074:
1070:
1068:
1063:Phylogenetic
1047:
1041:
1033:
1025:
1013:
1012:
994:
990:
978:cloud forest
973:
965:
961:
957:
953:
951:
946:
942:
916:
905:
901:
862:
846:
842:
840:
834:B. coloratus
832:
779:grasshoppers
764:
751:
747:B. ephippium
746:
740:
728:
724:
716:B. leopardus
715:
700:
696:
691:
688:B. ephippium
687:
679:vestigiality
670:
667:B. ephippium
666:
664:
656:
646:
644:
638:
617:
603:B. olivaceus
602:
587:
572:
557:
517:
514:B. ephippium
513:
511:
490:
487:B. nodoterga
486:
483:tetrodotoxin
478:
475:B. ephippium
474:
468:
463:
459:
455:
452:
447:
443:
439:
435:
433:
427:
425:
416:B. ephippium
415:
409:
403:
397:
391:
385:
384:, and Cuban
379:
371:
367:
363:
359:
354:
353:
346:
338:
334:
330:
326:
316:
310:
286:
284:
268:cloud forest
261:
245:Psyllophryne
244:
213:
209:
207:
199:Psyllophryne
198:
181:
160:
139:
138:
53:
29:
4157:iNaturalist
4054:Wikispecies
3170:(1): 5388.
2353:Adelophryne
2215:11 February
1243:sky islands
1135:B. brunneus
1127:B. curupira
1043:Ischnocnema
904:(left) and
902:B. darkside
787:springtails
558:B. curupira
530:mate choice
522:fluorescent
495:neurotoxins
464:B. curupira
460:B. brunneus
405:Paedophryne
393:E. limbatus
360:B. darkside
348:Adelophryne
184:(in black)
4218:Categories
3715:2 February
3658:: 109246.
3609:2 February
3418:2022-09-11
3367:9 February
3309:9 February
3304:ScienceMag
2812:Check List
2791:5 February
2706:30 January
2173:(9): 150.
1779:2 February
1748:References
1117:subgroup (
1105:subgroup (
982:sky island
935:endangered
931:vulnerable
868:osteoderms
803:millipedes
697:B. actaeus
692:B. pitanga
671:B. pitanga
639:B. pitanga
618:B. pitanga
518:B. pitanga
471:aposematic
323:morphology
297:Appearance
291:threatened
249:aposematic
3767:: e2490.
3674:0006-3207
3560:0270-6474
3513:0022-3077
3466:0022-5193
3280:205604026
3107:: e5273.
2834:212771392
2668:2167-8359
2652:: e5683.
2582:: e3603.
2481:2167-8359
2465:: e1011.
2167:Diversity
2055:Diversity
1935:: 33–40.
1874:1175-5334
1814:: 1–182.
1169:described
1119:SĂŁo Paulo
1103:B. pernix
1022:Bufonidae
855:endolymph
771:true bugs
731:nocturnal
675:vocal sac
660:frequency
652:territory
479:B. pernix
411:Stumpffia
319:vertebrae
146:Fitzinger
85:Kingdom:
79:Eukaryota
4094:BioLib:
4039:Wikidata
4006:86572812
3967:33909613
3926:PLOS ONE
3872:BBC News
3793:27761312
3703:(2020).
3578:18685033
3521:15269231
3474:10366494
3347:86638006
3272:28935936
3202:30926879
3141:50776284
3133:30042895
3071:15581685
3032:23294169
3024:27812425
2930:86086374
2890:26339556
2768:30917162
2728:PLOS ONE
2686:30310742
2616:22414191
2608:28761788
2544:86664601
2499:26056613
2399:30231059
2359:PLOS ONE
2125:27394218
2029:12175613
1990:29690168
1882:28609910
1083:B. pulex
1016:lacks a
743:amplexus
737:Breeding
720:amplexus
718:pair in
707:Behavior
526:UV light
444:B. pulex
368:B. pulex
335:B. pulex
311:B. pulex
282:stage).
216:) are a
190:Synonyms
163:see text
154:Species
125:Family:
109:Amphibia
99:Chordata
95:Phylum:
89:Animalia
75:Domain:
4149:2424890
4045:Q634679
3958:8081201
3935:Bibcode
3894:Zootaxa
3784:5068354
3635:: 1–12.
3569:2647017
3446:Bibcode
3263:5608807
3242:Bibcode
3193:6441030
3172:Bibcode
3124:6055690
3051:Toxicon
3015:5088579
2968:1565481
2881:4558078
2759:6436692
2736:Bibcode
2677:6174073
2599:5534166
2490:4458131
2390:6145526
2367:Bibcode
2305:1564972
2105:Zootaxa
2009:Toxicon
1970:Zootaxa
1929:Zootaxa
1854:Zootaxa
1808:Zootaxa
1145:Species
1095:synonym
1069:Within
1038:Genetic
817:Jumping
811:isopods
795:spiders
775:beetles
761:Feeding
755:tadpole
280:tadpole
272:diurnal
227:in the
135:Genus:
115:Order:
105:Class:
4201:719425
4188:164301
4175:208123
4110:257746
4097:178258
4004:
3965:
3955:
3877:4 June
3791:
3781:
3672:
3576:
3566:
3558:
3519:
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3464:
3390:
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3278:
3270:
3260:
3200:
3190:
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3030:
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2928:
2888:
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2596:
2542:
2497:
2487:
2479:
2397:
2387:
2303:
2123:
2027:
1988:
1880:
1872:
1107:Paraná
1009:Family
937:and 6
807:snails
222:family
148:, 1826
4162:24486
4136:40616
4002:S2CID
3761:PeerJ
3343:S2CID
3276:S2CID
3137:S2CID
3101:PeerJ
3028:S2CID
2992:PeerJ
2964:JSTOR
2926:S2CID
2858:PeerJ
2830:S2CID
2646:PeerJ
2612:S2CID
2576:PeerJ
2540:S2CID
2459:PeerJ
2301:JSTOR
1887:1 May
1804:(PDF)
791:mites
783:flies
481:have
236:Bahia
232:Anura
229:order
218:genus
119:Anura
4183:NCBI
4170:ITIS
4144:GBIF
4123:3C9X
4105:BOLD
3963:PMID
3898:4290
3879:2015
3838:link
3789:PMID
3717:2020
3670:ISSN
3611:2020
3574:PMID
3556:ISSN
3517:PMID
3509:ISSN
3470:PMID
3462:ISSN
3388:ISBN
3369:2020
3311:2020
3268:PMID
3198:PMID
3129:PMID
3067:PMID
3020:PMID
2886:PMID
2793:2020
2764:PMID
2708:2020
2682:PMID
2664:ISSN
2604:PMID
2495:PMID
2477:ISSN
2395:PMID
2217:2020
2121:PMID
2109:4083
2025:PMID
1986:PMID
1974:4407
1933:2739
1889:2017
1878:PMID
1870:ISSN
1858:4258
1812:1737
1781:2020
1310:and
1250:List
1178:Bufo
1109:and
1085:and
960:and
945:and
933:, 5
919:IUCN
809:and
781:and
767:ants
669:and
516:and
505:and
477:and
462:and
446:and
408:and
399:Mini
390:and
362:and
337:and
253:vent
208:The
4131:EoL
4118:CoL
4082:ASW
4069:ADW
3994:doi
3953:PMC
3943:doi
3902:doi
3820:doi
3779:PMC
3769:doi
3660:doi
3656:261
3633:526
3564:PMC
3548:doi
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3454:doi
3442:198
3335:doi
3258:PMC
3250:doi
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3180:doi
3119:PMC
3109:doi
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3010:PMC
3000:doi
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2375:doi
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2175:doi
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